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L161— O-1096

FLORA OF PERU

BY

J. FRANCIS MACBRIDE

CURATOR, PERUVIAN BOTANY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART III, NUMBER 2

MARCH 18, 1949

PUBLICATION 622

THE LIBRARY or THE

PRINTED IN THE UNITED STATES OF AMERICA

BY FIELD MUSEUM PRESS

FLORA OF PERU

J. FRANCIS MACBRIDE

GERANIACEAE. Geranium Family

Reference: Knuth, Pflanzenreich IV. 129. 1912.

Annual or perennial herbs or, less frequently, suffrutescent with

opposite or alternate often bistipulate and more or less dentate lobed

or divided, rarely entire, leaves. Pedicels axillary, 1- or 2-flowered

or flowers subumbellate. Sepals persistent. Flowers hermaphrodite,

usually regular with 5 sepals and petals, these rarely 4, or the latter

lacking in Rhynchotheca, mostly equal, imbricate. Stamens 2 (3)

times as long as the sepals, 10 or by abortion 2, 7, or 9 often 5,

alternate ones not antheriferous; filaments rarely completely free.

Ovary 3-5(2-3)-lobed, the 5 styles united around a produced axis

to form a torus or beak, free at tip with narrow stigmas. Ovules

pendulous, 1, 2 or many in each cell. Fruit usually 5 one-seeded

carpels that when ripe spring away from the elongate central axis

and bear the twisted or coiled styles as "tails" or less frequently

these are short, straight and remain attached at tips or the fruit

may be even capsular, 5-valvate, the valves not at all caudate.

Herbs; carpels revolutely caudate, in fruit "tailed."

Stamens with anthers 10; carpel-tails glabrate; leaves often

palmately lobed or parted 1. Geranium.

Stamens with anthers 5; carpel-tails bearded; leaves usually

pinnate 2. Erodium.

Shrubs; carpels ecaudate or the beak erevolute.

Petals none (or obscure) ; mature fruit 5 caudate carpels.

3. Rhynchotheca.

Petals 5; mature fruit capsular with parting but ecaudate valves.

4. Balbisia.

1. GERANIUM [Tourn.] L.

Herbs, annual or perennial, in Peru often cespitose and tufted

or matted, sometimes diffuse. Leaves alternate or opposite, stipu-

late, usually orbicular-reniform, variously palmately lobed or divided.

Flowers 5-merous, actinomorphic, on axillary peduncles, the sepals

and petals imbricate, the former persisting, the latter deciduous and

511

512 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

alternate with 5 glands. Stamens 10 (5), hypogynous, 5 longer

alternating with 5 shorter, the filaments basally united if at all.

Ovary deeply 5-lobed, 5-carpellate. Styles glabrous within, elasti-

cally recoiling when mature and recurving on the 2-ovuled, 1-seeded

carpels.

The following synopsis was basically compiled before the dis-

criminating account of the Mexican species by H. Emery Moore, Jr.,

Contr. Gray Herb. 146. 1943, came to my attention, but I have

followed him in his identification of G. mexicanum and in noting the

petal-venation in the few Peruvian species in which this useful

character, as he pointed out, has been discernible. Several of the

Peruvian species are based upon incomplete material and until the

type localities are revisited their interpretation would seem to be

largely guesswork; I have therefore preferably retained names that

I could identify, restricting the others to their originals. As usual

in this work the characters used in the key have been dictated at

least in part by expediency. The descriptions are mostly after Knuth.

Species of Pelargonium L'HeY. of Africa, similar to Geranium

except that the showy flowers lack the five glands and are somewhat

irregular, are often cultivated as "Geraniums," and, as in warmer

North America, may persist as adventives; for Peru, Pennell 14696

from along the Rio Chillon, Lima, is an example. The "Rose

Geranium" of gardens is a Pelargonium and is a source of some

geranium oil important in perfumery.

Densely cespitose alpines, acaulescent or essentially and forming

mats or polsters, or diminutive and imbedded in puna formation.

Leaves never argenteously sericeous both sides, rarely strigosely

subsericeous, often merely ciliate marginally and on veins or

somewhat pubescent beneath.

Leaf-divisions all with at least 3 linear segments that are clearly

longer than broad G. Humboldtii.

Leaf-divisions at least in part entire, broadly lobed or dentate.

Leaves completely glabrous, the lobes entire and obtuse or

the medial rarely 1-dentate; petals white, not clawed.

G. Jaekelae.

Leaves more or less ciliate, the lobes usually lobulate or

dentate; petals more or less clawed.

Leaf-dentations narrow, acute, rarely lacking the lobes then

acutish; petals usually tinted G. sibbaldioides.

Leaf-dentations broad, obtuse; petals white. G. sessiliflorum.

FLORA OF PERU 513

Leaves argenteous to sordid sericeous but the indument always

distributed evenly.

Indument a tomentulose puberulence or close pilosity that does

not extend ciliately beyond the leaf-margins.

Petals 7-12 mm. long, usually glabrous.

Leaves deeply lobed, the lobes in part 3-dentate.

G. sessiliflorum.

Leaves shallowly lobed, the lobes entire or obscurely

crenate G. Pavonianum.

Petals 14-20 mm. long when fully grown.

Leaf-divisions broad and mostly entire; petals ciliate

basally G. digitatum, G. nivale.

Leaf-divisions narrow, mostly parted; petals (known)

glabrous.

Peduncles to 1.5 cm. long, shorter than the leaves.

G. Weddellii.

Peduncles 2-4 cm. long but the petioles about as long.

G. Staffordianum.

Indument a long-sericeous pilosity, many trichomes extending

loosely beyond the leaf-margins.

Leaf-divisions entire or unequally dentate.

Leaves medially to basally parted, the divisions usually 7.

G. crassipes.

Leaves often less deeply 5-parted G. Lechleri.

Leaf-divisions regularly 3-dentate, at least medial.

Lateral as middle leaf-divisions mostly or all 3-dentate,

subargenteous G. Dielsianum.

Lateral leaf-divisions often entire or 1-dentate, very argente-

ous G. Ruizii.

Loosely cespitose or at least caulescent but the stems sometimes

obscurely developed in juvenile or retarded plants.

Peduncles 1-flowered, solitary, binate or pseudo-umbellate; plants

obviously perennial except for the first species and possibly

G. chinchense.

Petals, fully grown, 10-16 mm. long (cf. G. scissum).

Pubescence more or less spreading and glandular, slightly if

at all retrorse, not appressed, the glands sometimes

deciduous or sparse G. Stuebelii.

514 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Pubescence more or less retrorse-appressed on stems and

peduncles, eglandular, often sparse or minute.

Leaves deeply divided, even the cauline at least medially,

the segments coarsely dentate or subpinnate, canescent

only in G. Raimondii.

Peduncles ebracteate, rarely with 1 bract, or bracted at

base.

Petals at most 12 mm. long; apical peduncles if short

solitary or binate.

Plants green (G. Stuebelii), G. ayacuchense.

Plants canescent G. Raimondii.

Petals about 16 mm. long; apical peduncles short,

subcymose G. Weberbauerianum.

Peduncles bibracteate about medially G. laxicaule.

Leaves shallowly lobed, the segments of the upper leaves

shortly tridentate or merely crenate.

Leaf -lobes crenulate or subentire G. renifolium.

Leaf -lobes in part shortly tridentate G. peruvianum.

Petals 6-7 (8) mm. long, little exceeding the sepals.

Medial leaf-division cuneate-obovate, strongly, even petiolu-

lately narrowed to base, this often nearly or quite to

petiole G. Mathewsii, G. piurense, G. ayavacense.

Medial leaf-division rhomboid-obovate, slightly if at all

narrowed to oblongish base, this rarely extending nearly

to petiole-apex.

Pube'scence eglandular.

Peduncles and leaves subequal.

G. Smithianum, G. Sodiroanum, G. diffusum.

Peduncles 3-5 times longer than leaves G. Harmsii.

Pubescence in part glandular.

Uppermost leaves deeply 3-parted, the segments sub-

entire or tridentate G. huantense.

Uppermost as basal leaves about equally and broadly

5-parted, the segments lobulate or unequally dentate.

G. chinchense.

Peduncles at least in part 2-flowered ; plants often annual, biennial

or short-lived perennials (cf. G. laxicaule}.

Petals nearly 12 mm. long, usually longer.

FLORA OF PERU 515

Leaves, especially the upper sericeous, canescent beneath;

petals about 2 cm. long.

G. scissum, G. matucanense, G. superbum.

Leaves nearly concolor, paler and more pubescent but not

canescent beneath; petals 11-14 mm. long.

G. mollendinense.

Petals rarely if ever 10 mm. long, mostly shorter.

Plants perennial (sometimes short-lived); petals usually ex-

ceeding sepals or the pubescence mostly retrorse-ap-

pressed or usually spreading and glandular above.

Pubescence especially above obviously glandular (there

may be rarely eglandular states); apical peduncles

soon exserted from leaves.

G. Killipianum, G. Seemanni, G.fallax, G. patagonicum.

Pubescence obscurely if at all glandular or the peduncles

short, more or less concealed in the subtending leaves.

Pubescence mostly spreading; petals rather obvious;

plants clearly perennial.

Bracts ovate-lanceolate G. filipes.

Bracts linear-subulate G. limae.

Pubescence mostly retrorse-appressed; petals small;

plants apparently short-lived .G. Herrerae.

Plants annual and introduced; pedicels glandular only in the

obviously annual G. dissectum,G. carolinianum with petals

and calyx subequal.

Sepals obviously mucronate or acuminate-mucronate; seeds

reticulate; leaf -lobes subpinnate.

Peduncles several together or binate; carpels ascending-

villous G. carolinianum.

Peduncles solitary; carpels spreading-hirsute.

G. dissectum.

Sepals not at all or obscurely mucronulate; seeds smooth;

leaf-lobes triparted.

Stamens 10; carpels rugose G. molle.

Stamens 5; carpels smooth G. pusillum.

Geranium ayacuchense Knuth, Repert. Sp. Nov. 18: 291.

1922.

Perennial, with many ascending more or less squarrosely branched

stems about 5 dm. high, the indument short, retrorse, the lower

516 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

internodes to 8 cm. long; petioles retrorse-pilose, to 2 cm. long;

leaves semiorbicular, retuse at base, often 2.5 cm. wide, pubescent

above with minute hyaline trichomes, pilose beneath especially on

nerves, palmately 5-parted three-fourths to four-fifths the long-

cuneate lobes coarsely 3-dentate marginally, the obtuse semioblong

teeth minutely mucronulate; stipules pilose, narrowly lanceolate

from base, long-subulate or capillary acuminate; pedicels axillary,

ebracteate, 1-flowered, slender, 1.5-6 cm. long, often two from fork

of branches; sepals oblong, subhirsute-pilose, 7-8 mm. long with

mucro nearly 2 mm. long; petals somewhat cuneate, the few veins

sparsely branched toward tip, 12 mm. long, pale rose or whitish (or

white with red veins); stamens half as long as calyx, pilose; fruit

18 mm. long, the beak nearly 2 mm. thick, the valves setaceous.—

Section Rupicola but with no close ally (Knuth). But, except for

the larger flowers, it seems to me to be much like G. diffusum at

least in Peru, otherwise like G. laxicaule.

Ayacucho: In grass steppes and small shrubs above Osno, Prov.

Huanta, Weberbauer 5563, type. Pampalca, Huanta to Rio Apuri-

mac, Killip & Smith 22214 (det. Knuth, G. Sodiroanum) .• — Huanca-

velica: Shrub woods, 3,300 meters, Salcabamba, Stork & Horton

10290 (det. Standley, G. peruvianum') . — Apurimac: Rocky hillsides,

3,600 meters, Stork & Horton 10777 (det. Standley, G. Herrerae).

Huancarama, 3,900 meters, West 3750,—Cuzco: Damp shady places,

Quenco Grande, Prov. Acomayo, Vargas 9747? (det. Standley,

G. Herrerae, peduncles in part 1-2-bracted; cf. G. laxicaule). Haci-

enda Paucartica, Calca, Vargas 155 (det. Standley, G. diffusum?).

Hacienda Churu, 3,500 meters, Herrera 1017; 1027a (both distr.

asG. peruvianum). .

Geranium ayavacense Willd. ex HBK. Nov. Gen. & Sp. 5:

231. 1822; 213. G. partitum Willd. I.e.

Perennial from a thick much-branched root that is crowned by

the basal leaves and many diffuse geniculate branching ascending

or procumbent stems that are often several dm. long, their internodes

as much as 1.5 dm. long; basal petioles 6-10 rarely 15 cm. long,

appressed retrorse pilose-puberulent, the leaves somewhat ashy

with a very fine indument, orbicular, palmately 7-parted usually

to base, the lobes irregularly bi- or tri-lacinulate even medially, the

lateral segments often unequal in length, all linear, rather abruptly

acuminate, submucronulate ; stem leaves similar, the middle and

upper short-petioled but scarcely sessile; stipules triangular-lanceo-

FLORA OF PERU 517

late, acutely acuminate, pulverulent; peduncles axillary, 1-flowered,

ebracteate, 3-10 cm. long, sparsely retrorsely appressed-pubescent;

sepals ovate-oblong, slightly acuminate with mucro 1 mm. long,

puberulent especially on nerves and long-ciliate toward base and

membranous margins; petals obovate, whitish, ex neg. about 10 mm.

long or one and one-half times longer than calyx, this exceeding

stamens; valves puberulent as the 18 mm. long beak. — Written by

error "agavacense" and obviously to be corrected to conform to

the source of the name. F.M. Neg. 35705.

Piura: Ayavaca, Bonpland, type. — Huanuco: Wood clearing,

Mito, 1740 (det. Knuth, G. piurense). Cani, 3476 (det. Knuth,

G. piurense). Open waste land, Pillao, Woytkowski 27, Ecuador;

Venezuela.

Geranium carolinianum L. Sp. PI. 682. 1753; 54.

Annual, the 2 to several dm. high more or less branched sulcate

stems puberulent with somewhat recurved trichomes, the lower leaves

alternate, upper opposite, reniform-orbicular, subappressed puberu-

lent, 5-7-parted nearly to the base, the lobes subpinnatifid with

linear obtusish segments; lower petioles 5-10 cm. long; bracts linear-

subulate; peduncles short, biflowered or the flowers nearly umbellate;

sepals ovate or broader, mucro 1-1.5 mm. long or longer, mostly

5-7 mm. long, accrescent in fruit, the petals as long or slightly longer;

valves smooth, nearly setose; seeds favose-punctate. — A weedy

species of North America said to have been found even to Paraguay

and so to be expected.

Peru: cf. note above. North America.

Geranium chinchense Knuth, Repert. Sp. Nov. 28: 4. 1930.

A several dm. tall perennial* (or biennial?), the sparsely branched

stems hirsute with rather long spreading trichomes; lower internodes

1-2 dm. long; basal petioles to 2.5 dm. long, hirsute, those of the

upper leaves densely so, the trichomes in part glandular; stipules

soon marcescent, hirsute margined, 8 mm. long, lanceolate; basal

leaves depressed, 5-angled-rotund, 5-parted four-fifths or deeper,

appressed pilose on nerves both sides, the main lobe cuneate-

rhomboid, the upper lateral two coarsely incised pinnatifid, the

lobules minutely mucronulate, those of the basal leaves semiovate,

those of the upper lanceolate; peduncles 1-flowered, ebracteate,

glandular hirsute, 13 mm. long, not rarely forming in the absence

of developed leaves a small cyme; sepals ovate, 5-6 mm. long,

518 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

mucronulate, densely glandular hirsute, the white petals one and a

half times longer; fruit about 18 mm. long, the beak 1.5 mm. thick,

glandular hirsutulous, the valves sparsely pilose. — Section Rupicola

(Knuth) but cf. G. huantense. My 1404 sepals 4 mm. long, petals

7 mm. long, the veins branched above only.

Huanuco: Chinche, Yanahuanca, 1263, type. Mito, 1404 (det.

Knuth).

Geranium crassipes Hook, ex Gray in U. S. Expl. Exped. Bot.

1: 309. 1854; 83. G. muscoideum Knuth, Bot. Jahrb. 37: 567.

1906; 87. G. sessiliflorum Cav. var. lanatum Knuth, I.e. 565, at

least as to Peru.

Stemless cespitose ashy pubescent perennial, the densely matted

plants as much as a dm. in diameter, the sericeous indument extend-

ing even to the sepals, only the free part of the stipules pilose, these

about two- thirds adnate, 7 mm. long, acutely linear-acuminate;

petioles 0.5-1.5 cm. long; leaves scarcely coriaceous, nearly orbicular,

5-6 mm. across, palmately 7-parted almost to base, the acutish

oblanceolate lobes similar; peduncles solitary, 1-flowered, ebracteate,

at most 5 mm. long; sepals oblong-lanceolate, acutish, coriaceous,

3-nerved, 5-6 mm. long, 2.5-3 mm. wide, the inner often longer;

petals white, 13-15 mm. long, obovate, ciliate at the somewhat

narrowed base, or nearly linear, glabrous as here interpreted;

stamens and pistils shorter than calyx, the puberulent beak of the

fruit finally one and a half times longer. — This is Knuth's description

for his species; he referred Hooker's to G. sessiliflorum. Illustrated,

Pflanzenreich, I.e. fig. 14- F.M. Neg. 26373 (G. muscoideum).

Junin: Near La Oroya among polster and rosette plants, 4,300

meters, Weberbauer 2619 (type, G. muscoideum). Obrajillo, Wilkes

Exped., type. Rocky slope, Huaron, 1148B (det. Knuth, G. sessili-

florum var. lanatum). Tarma to Jauja, 4,500 meters, Killip &

Smith 21968 (det. Knuth, G. Lechleri). Cobracancha Valley near

Cerro de Pasco, 4,200 meters, Grant 7557 (distr. as G. Ruizii).

Cerro de Pasco, (Mathews).

Geranium Dielsianum Knuth, Bot. Jahrb. 37: 563. 1906; 81.

Habit of G. sessiliflorum but densely lanuginose with silvery

sericeous indument; petioles 2-3.5 cm. long; leaves reniform-rotund,

the largest 10 mm. long by 14 broad, palmately 7-parted to the

middle or somewhat farther, the lobes broadly cuneate-obovate,

the three middle ones tridentate, the outermost lateral with 1 tooth

FLORA OF PERU 519

at the outer edge; stipules about 13 mm. long, more than half connate,

pilose marginally, the 5 mm. long free parts acutely acuminate;

peduncles solitary, basal, 1-flowered, ebracteate about 8 mm. long,

densely lanate; flowers medium, not topping the leaves; sepals ob-

long-lanceolate, acutely acuminate, to 7 mm. long, 2.5 mm. wide,

lanate; petals white, 8-10 mm. long, obovate, entire, glabrous even

on the claw. — Cf. G. sessiliflorum var. lanatum Knuth. G. sericeum

Willd., 80, might be sought here but the leaves divided as inG. acaule;

Ecuadorean, it is to be expected. Illustrated, Pflanzenreich, I.e. fig.

14. F.M. Neg. 26368.

Cajamarca: At Pass Coymolache above Hualgayoc on rocks at

4,000 meters, Weberbauer 3959, type; 272.

Geranium diffusum HBK. Nov. Gen. & Sp. 5: 230. 1822; 214.

A perennial, with many ascending or suberect slender sparsely

branched retrorsely subhispid stems 1.5-2.5 dm. tall; middle petioles

2 cm. long, softly spreading pubescent, the uppermost nearly lacking;

leaves appressed hispidulous, round-reniform, palmately 5-7-parted,

the obcuneate lobes apically trilobulate, the middle lobe often

trilaciniate, the lobules all ovate and obtusely mucronulate; stipules

lance-subulate, membranous, ciliate, 3-4 mm. long; peduncles axil-

lary, solitary, ebracteate, 1-flowered, densely spreading pilose, 2-3

cm. long; sepals elliptic-oblong, about 5 mm. long with mucro 0.5

mm. long, pilose; corolla 10-12 mm. broad, the oblanceolate clawed

petals 7 mm. long. — The Peruvian material is scarcely typical;

Knuth accepted and proposed many segregates perhaps with reason.

F.M. Negs. 4776 (Berlin); 35706 (Paris).

Without data, Bonpland, type. — Cuzco: Colinas del Saxaihuaman,

Hen era 1044 (distr. asG. filipes). Near Cuzco, Herrera 2341 (distr.

as G. filipes). Calca (Vargas 155; 3222). — Apurimac: Sunny space

in forest, 3,200 meters, Goodspeed Exped. 10628 (det. Standley,

G. tablasense Knuth?). Ecuador.

Geranium digitatum Knuth, Repert. Sp. Nov. 28: 1. 1930.

Cespitose, forming dense mats as much as 14 cm. across and

rarely with a densely leafy stem 1-2 cm. long; petioles about 1 dm.

long silvery sericeous as the sepals and the numerous leaves, these

2-2.5 cm. wide, 5-lobed nearly to base, the elongate-oblong minutely

mucronate lobes 12 mm. long, 4 mm. wide; stipules 15 mm. long,

free only 4 mm. where lanceolate, acuminate; peduncles 1-flowered,

ebracteate; sepals lanceolate, 10 mm. long, the margins pale, the

520 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

stamens as long; the petals one and a half times longer, violet;

fruit 18 mm. long, the beak and valves minutely sericeous. — Section

Andina; character after Knuth; apparently referable to G. nivale,

as an ecological race, robust, caudices long, leaflets much larger, etc.

Junin: Limestone outcrops, La Oroya, 3,400 meters, 940, type.

Geranium dissectum L. Cent. PI. 1: 21. 1755; 51.

Annual, often with several or many spreading or ascending

usually retrorse hirsute stems 1 to 5 (or more) dm. long; petioles

3-4 times longer than the basal broadly lobed or parted leaves, the

reniform-orbicular middle and upper leaves 5-7-parted nearly to

the base, the segments subpinnatifid with linear divisions; peduncles

solitary, 2-flowered, the upper ones often shorter than the glandular

pedicels; sepals ovate, acuminate, long-mucronate, open and accres-

cent to 9 mm. long; petals obcordate, as long as calyx, bearded

above claw, typically purple; valves smooth, spreading-hirsute, the

seeds regularly and deeply favose-punctate. — Widely spread, includ-

ing much of South America as Ecuador and Chile.

Peru: Will be found in waste places. Generally introduced from

Europe and Eurasia.

Geranium fallax Steud. Flora 39: 439. 1856; 147. G. tucumanum

Knuth, Pflanzenreich IV. 129: 147. 1912?

Stems branched, puberulent, rather weak; petioles pilose; leaves

3-5-parted, the lobes 3-6-lobulately incised, setulose-pilose; peduncles

mostly 1- rarely 2-flowered, hirsute pilose as the calyx, this with

ovate shortly acuminate sepals that are a little shorter than the red

petals. — Not recognizable from description according to Knuth and

scarcely the same as the Tucuman plant, which however was labeled

in Herb. Berlin G. fallax. Ex neg. the trichomes seem to be gland-

tipped; seems to be G. patagonicum or allied. F.M. Neg. 4780.

Puno: Near Tabina, Lechler 1907, type.

Geranium filipes Killip, Journ. Wash. Acad. Sci. 16: 569. 1926.

Rhizome vertical, 4-5 mm. thick, apically with 2 or 3 slender

little-branched erect or ascending densely subretrorse-strigillose

stems 1-1.5 dm. high, exceeding the basal leaves, these as the cauline

orbicular-reniform, 1-1.5 cm. long, 1.5-2 cm. wide, 5-lobed about

to middle (lobes trilobulate, rarely entire, the rounded segments

mucronulate), membranous, appressed-strigillose above, pilosulous

on nerves beneath; petioles 3.5-7 cm. long; stipules linear-lanceolate,

FLORA OF PERU 521

5-7 mm. long, acute, merely ciliate; peduncles solitary, very slender,

2-5 cm. long, retrorse-strigillose, 2-flowered; bracts acuminate,

glabrous; pedicels pilose, 4-6 mm. long; sepals lanceolate, 4-5 mm.

long, conspicuously mucronate, appressed hirsutulous, the subtri-

nerved slightly glandular, pilosulous; petals 5-8 mm. long, deep

pink, few, unbranched, nerves pale; stamens shorter than calyx,

filaments ciliolate, anthers 1 mm. long. — After Killip, who noted:

evidently of section Rupicola, the two other Peruvian species with

more numerous ebracteate white flowers, emucronate leaf -lobes; cf.

also G. magellanicum Hook, f., 70, and G. limae, 74.

Contains a bitter principle used for throat infection, "thrush"

(Herrera).

Cuzco: Hacienda Churu, Prov. Paucartambo, 3,500 meters,

Herrera 1044, type. "Chile-chile," "ujutillo."

Geranium Harmsii Knuth, Bot. Jahrb. 37: 559. 1906; 213.

Tufted from an elongate horizontal branched rhizome, the

branches 1 dm. long or longer, subligneous, 4-5 mm. thick, tortuous

and more or less clothed with stipules and old petioles; stems usually

many, to 3 dm. long, laxly ascending, pseudodichotomously squar-

rose-branched, many-leaved, puberulent; basal stipules lanceolate-

acuminate, finally subulate, the medial 4-5 mm. long; basal petioles

4-5 cm. long, the upper gradually shorter, never obsolete; leaves

puberulent, all reniform-rotund, the basal 12-15 mm. broad, usually

three-fourths to four-fifths regularly palmately 5 (-7) -parted, the

cuneate lobes to one-third trilobed, the lateral lobules oblong-ovate,

the middle one broader, all obtusish or scarcely mucronulate;

peduncles axillary, 1-flowered, ebracteate, at anthesis 3 cm. long,

finally 5 cm. long; sepals 6 mm. long, gradually acuminate with

mucro 1 mm. long, glabrescent within, pilose without, especially

margins and toward base, the oblong-ovate retuse white petals

scarcely 8 mm. long; pistils and stamens shorter than calyx; beak

shortly puberulent.— F.M. Neg. 4782.

Junin: Among scattered shrubs and small grasses east of Palca,

2,800 meters, (Weberbauer 2451, type); 248.

Geranium Herrerae Knuth, Repert. Sp. Nov. 28: 1. 1930.

Many-stemmed annual or apparently short-lived perennial, 2 dm.

or so tall, more or less sparsely appressed pilose, most densely the

pedicels and calyces; basal leaves with petioles about 2 dm. long,

the round blades 3-4 cm. wide drying papyraceous, 7-parted to well

. OF ILL LIB.

522 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

below the middle, the lobes one-third to two-fifths 3-lobulate-dentate,

the anterior lobules rounded and minutely mucronulate; uppermost

leaves nearly subsessile; stipules about 8 mm. long, linear-subulate,

very acute; peduncles biflowered, 8-20 mm. long; bracts 4, subulate,

2 mm. long; pedicels 1 cm. long; sepals broadly ovate, abruptly

acuminate, acute, 3 mm. long, 2 mm. wide, the petals twice as long,

beautiful carmine, the pilose filaments as long; pistils hirsutulous,

the beak minutely puberulent, the valves somewhat appressed-

spreading hirsute. — Section Chilensia Knuth with the habit of G.

pyrenaica and the short small-leaved upper branchlets (Knuth) ; but

it suggests strongly G. carolinianum and G. core-core Steud., 75; type

not seen.

Cuzco: Ollantaytambo, Prov. Urubamba, 3,000 meters, (Herrera

234, type). Near Cuzco, Est. Exp. Kaisa, Vargas 667 (det. Standley,

G. peruvianum?) . — Arequipa: Water courses, Chachani Mountains,

Hinkley 66 (distr. as G. core-core?). — Moquehua: Near Torata,

3,100 meters, Weberbauer 7479 (det. Knuth). "Aguja-aguma,"

"relojito" (both Vargas), "comirachi" (Hinkley).

Geranium huantense Knuth, Repert. Sp. Nov. 28: 8. 1930.

G. choimacotense Knuth, I.e. 9.

A perennial, with many procumbent or scandent furcately

branched stems that attain a meter or more, glabrous below, some-

what sulcate and retrorse or above more or less spreading subhirsute;

petioles to 8 cm. long, densely lanate-hirsute with long more or less

spreading trichomes; stipules nearly subulately contracted from an

ovate base, 4-7 mm. long; leaves to 5 cm. broad, for four-fifths

3-5-parted, appressed hirsutulous especially beneath on nerves, the

lobes rather rhomboid, the two upper lateral deeply pinnatifid with

lobules 3-6 mm. long, 2-6 mm. wide, mucronulate; uppermost leaves

3-parted, the acute lobes entire or 1-toothed; peduncles ebracteate,

1-flowered, 18 mm. long, not rarely in small cymes, densely stipitate,

glandular as sepals, these ovate-lanceolate, about 5 mm. long, 2-2.5

mm. wide, membranous-margined with mucro less than 1 mm. long;

petals reddish-white, cuneate, with rounded or obtuse apex, about

6 mm. long, veins little-branched; stamens and pistils glandular;

fruit 15 mm. long, the valves glabrate, the beak densely glandular.—

Habit of G. laxicaule Knuth but indument diverse (Knuth). I have

included in the description G. choimacotense which as to type seems

to me to be at most only a variant or race with smaller leaves, the

lobes more evenly dentate, and the pubescence rather less glandular;

FLORA OF PERU 523

I choose the name G. huantense as more appropriate. Nearly G.

Knuthianum Macbr. Candollea 6: 7. 1934 (G. elongatum Knuth, 209)

of Ecuador, that with peduncles in part 2-flowered, fruit beak 2.5

cm. long.

Ayacucho: Choimacota Valley, Prov. Huanta, Weberbauer 7579

(type, G. choimacotense) ; same locality, Weberbauer 7619, type.—

Junin: Near Huancayo, Soukup 2760. — Huanuco: Trailing at wood

edge, Tambo de Vaca, -4-407 (det. Knuth, G. choimacotense).

Geranium Humboldtii Spreng. Syst. 3: 70. 1826; 87. G. acaule

Willd. ex HBK. Nov. Gen. & Sp. 5: 231. 1822, not L. or Thunb.

G. potentilloides Willd. ex Spreng. I.e. 71, not L'HeV. ex DC. Prodr. 1:

639. 1824. G. multipartitum Benth. PI. Hartw. 166. 1845, at least

as to Peru. G. hypoleucum Benth. I.e., fide Knuth. G. multipartitum

Benth. var. velutinum Knuth, Bot. Jahrb. 37: 566. 1906? and var.

glabrescens Hieron. ex Knuth, I.e.

Forming low dense mats from the shortly branching ligneous

thick caudex that above may be more or less clothed with the remains

of the often filiformly acuminate stipules, these broad and adnate

below, 8-10 mm. long; petioles about 1 (-2) cm. long, glabrate;

leaves 7-10 mm. wide, glabrous and lustrous above in type and

silvery sericeous beneath, but as interpreted, variable in pubescence,

orbiculate, 5-7-parted, the divisions 2-5-laciniate nearly to base

with thickish linear obtusish segments; sepals ovate-lanceolate and

acute or in the Peruvian forms long-acuminate or sometimes

mucronulate, about 5 mm. long, puberulent or nearly glabrous;

peduncles ebracteate, often only 5-10 mm. long; petals about 7 mm.

long; fruit beak in type to 10 mm. long, tomentose, in vars. shorter

and glabrous or nearly. — In view of the variation accepted by the

monographer himself for the species of Bentham it has seemed im-

possible to consider the plant of Peru as more than a part of a single

species. The Wilkes Expedition specimen, very meager, has leaves

appressed-strigose, merely subsericeous both sides, but nearly gla-

brous in my collections; fide Knuth the leaves in type are not glabrous

as described originally but argenteous-sericeous beneath. F.M.

Neg. 4766.

Cajamarca: Above Hualgayoc, dense shrub-cactus area, 4,000

meters, (Weberbauer 3990, type, G. multipartitum var. velutinum).—

Lima: Obrajillo, Wilkes Exped. — Huanuco: Wet grassy slopes, petals

white or pinkish, 3296 (det. Knuth, G. multipartitum var. glabrescens).

Mossy rocky uplands, flowers bright pink or white, Tambo de Vaca,

4401 (det. Knuth, G. multipartitum var. glabrescens). Ecuador.

524 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Geranium Jaekelae Macbr. Candollea 6: 7. 1934; 87. G.

minimum Knuth, Bot. Jahrb. 37: 567. 1906; 87, not Cav., 1787.

A small densely cespitose nearly completely glabrous perennial,

the stemless individual plants only a few cm. across but with shortly

branched tuberculate caudex 6-20 mm. thick that is closely clothed

above with stipular and petiolar remains just below the many small

leaves and flowers; stipules 4-5 mm. long, two-thirds adnate, the

free point linear-subulate, very acute; petioles 5-15 mm. long,

minutely appressed setulose; leaves semiorbicular or r"eniform-

rotund, 5-7 mm. wide, deeply palmately 7-parted, the broadly obo-

vate often nearly rotund obtusish lobes entire or rarely the medial

1-3-toothed; peduncles solitary, 1-flowered, ebracteate, 5-7.5 mm.

long, glabrescent; sepals 5-6 mm. long, 2.5 mm. wide, lanceolate,

scarcely acuminate, callose-mucronate, not prominently nerved;

petals white, one and one-half times longer than calyx, obovate,

entire, not clawed; filaments about half as long as calyx; fruit

undescribed. — Commemorates in taxonomic botany with exceptional

worthiness the work of Annemarie Jaekel (Mrs. A. Hirsch), principal

photographer for the Field Museum series of type photos. F.M.

Neg. 4789.

Junin: Near La Oroya in polster and rosette plant mats, 4,300

meters, Weberbauer 2623, type; 201, 223.

Geranium Killipianum Knuth, Repert. Sp. Nov. 34: 144. 1933.

Perennial, with many decumbent-ascending pseudodichoto-

mously branched stems, these rather densely stipitate glandular-

hirsute or -hirtellous as also petioles (lower 4-5 cm. long), peduncles

(2-4 cm. long), pedicels (1-1.5 cm. long) and sepals; leaves opposite,

appressed pubescent above setose pilose beneath, especially on

nerves, deeply but usually not to base 5 (-3) -parted, the lobes

coarsely incised, pinnatifid, the middle one 10 mm. long, 3 mm. wide,

the lateral much smaller; stem leaves short-petioled, those of the

uppermost about 1 cm. long; peduncles 2-flowered, most numerous

above; sepals ovate, acutely acuminate, 7-8 mm. long, nearly 3 mm.

wide at base, 3-nerved, little longer than stamens, the roseate

obovate petals about 8 mm. long, the veins nearly free; fruit 23 mm.

long, the beak 1.75 mm. thick, glandular-hirtellous as the valves

dorsally. — Section Mexicana. Apparently with no distinctive

character from G. Seemanni, etc.

Lima: Rio Blanco, about 3,300 meters, Killip & Smith

type.

FLORA OF PERU 525

Geranium laxicaule Knuth, Pflanzenreich IV. 129: 209. 1912.

Many-stemmed from a nearly naked horizontal rhizome 3-7 mm.

thick; stems slender, decumbent or dependent, 5-8 dm. long, little

branched, sparsely retrorse-pubescent, the internodes often 1.5 dm.

long; stipules lanceolate, acute, membranous, pilose; leaves all alike,

the younger as the petioles silvery sericeous, the former in age sparsely

appressed pubescent with hyaline trichomes, the latter spreading

pilose, the basal very lax and 1.5 cm. long or longer; 'leaf -blades

angulate-reniform, irregularly three-fourths or four-fifths palmately

5-parted, the broadly obovate lobes 3-dentate-lobulate, the lobules

minutely mucronulate, the middle one often 5 mm. wide and long;

peduncles filiform, 8-12 cm. long, retrorse-spreading, pilose, bibrac-

teate, uniflowered (so far as known), the lanceolate-linear acute

pilose membranous bracts 3 mm. long; pedicels pubescent and

eglandular like peduncles, 2-5 cm. long; sepals ovate, 4-6 mm. long,

2 mm. wide, 3-nerved, scarious-margined, densely covered with long

hyaline trichomes, mucronate with callus 0.5 mm. long; petals

obovate, twice as long as calyx, reddish white; stamens as long as

calyx, the styles a little shorter. — Allied by Knuth to G. elongatum

Knuth, 209, of Ecuador, with densely glandular-hirsute pedicels and

sepals. G. holosericeum Willd., 104, probably Colombian, seems

from the negatives to be in general similar. F.M. Neg. 4786.

Lima: Southwest of Matucana behind the valley Huillacachi at

a small waterfall, 3,000 meters, (Weberbauer 187, type).

Geranium Lechleri Knuth, Pflanzenreich IV. 129: 80. 1912.

G. sericeum Willd. var. microphyllum Wedd. Chlor. And. 2: 285.

1857. G. microphyllum (Wedd.) Briq. Ann. Cons, et Jard. Bot.

Geneve 11 & 12: 183. 1908, not Hook, f., 1844.

Dense cespitose stemless perennial herb forming mats about

5 cm. across that are densely clothed with a rather long silvery

lanuginose indument which extends, appressed, even to the sepals;

caudex much branched; stipules 5-7 mm. long, to two-thirds adnate,

pale, glabrous except ciliate margins, the free portion triqetrous-

ovate, acute; petioles about 1 cm. long or one and a third times

longer than the leaf blades, these coriaceous, reniform-rotund, the

sinus open or suborbicular, deeply but rarely to the base palmately

5-parted, 5 mm. long, 8 mm. wide, the 3 middle lobes rather obovate,

the lateral oblong, all obtusish, the middle often toothed or with a

lobe, the lateral rarely; peduncles solitary, basal, uniflowered,

ebracteate, barely to 1 cm. long, stout; flowers white, medium size

526 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

scarcely exceeding the leaves; sepals 5.5-6.5 mm. long, ovate-oblong,

barely acute, hardly twice exceeded by the beak. — The type is from

great altitudes in the cordillera of Carabaya. F.M. Neg. 4787.

Puno: Near Agapata, Lechler 1985. Bolivia.

Geranium limae Knuth, Pflanzenreich IV. 129: 74. 1912.

Perennial, with deeply set brown often tuber-like root; entire

plant to sepals more or less canescent with spreading setulose-pilose

indument little if at all glandular, appressed and sparse on the

leaves above, patent on the veins beneath; stems 1-several, sulcate,

prostrate or ascending-erect, about 3 dm. high, branched at middle;

stipules and bracts narrow, acuminate, the former to 5, the latter

to 10 mm. long; basal petioles 12-20 cm. long, about five times

longer than width of leaves, these angulate-reniform, usually three-

fourths 7-palmately parted, 5-6 cm. broad, 3 cm. long, the broadly

cuneate lobes to one-third palmately, 2-5-lobulate (usually medially

3-lobulate), the ovate rounded lobules obscurely mucronulate;

peduncles usually biflowered, slender, 3.5-6.5 cm. long, long white-

pilose; pedicels 7-25 mm. long, most densely pilose beneath calyx;

petals roseate or pink, about 7 mm. long, the few veins little branched,

the sepals broadly ovate, 6 mm. long, 3-nerved, minutely mucronate;

fruit at least 17 mm. long, the beak puberulent, the valves lightly

pilose.— Typically at least a local "species." F.M. Neg. 26369.

Said to be used for toothache.

Lima: San Lorenzo, Gaudichaud, type; Wilkes Exped. San

Gallen Island, (Murphy 3473; 3474). Barranco, rock crevices,

Worth 9117. Lomas Chancay, Raimondi; Ruiz & Pavdn. Lomas de

Dona Maria, Goodspeed Exped. 9255. Atacongo lomas, Goodspeed

Exped. 9288. Viso, Goodspeed Exped. 11526. — Ancash: Lomas de

La Chay, among boulders, Goodspeed Exped. 9207. — Arequipa: Near

Atequipa, Worth & Morrison 15665. "Antaccara," "chili-chili."

Geranium Mathewsii Briq. Ann. Cons. Jard. Bot. Geneve

11 & 12: 188. 1908; 213.

A perennial to about 4 cm. high from a short caudex and with

slender striate flexuous procumbent stems that, like petioles (lower

11-40 mm. long) and pedicels (5-15 mm. long), are finely retrorsely

puberulent or glabrate; stipules short, lanceolate, scarious; basal

leaves to 6 mm. broad, 3-5-parted, the obcuneate segments about

8 mm. wide, often 3-toothed to middle, the teeth narrowly oblong,

subacute, ashy green both sides with appressed puberulence; cauline

FLORA OF PERU 527

leaves few, reduced, shortly petioled; bracts 10 mm. long, very acute;

flowers small, solitary or binate in the axils, the sepals ovate, densely

pilose below, more appressedly above, 4 mm. long including mucro

1 mm. long; petals 4 mm. long, obovate, glabrous, the stamens

shorter; valves shortly appressed puberulent, 8 mm. long, crowned

with the short spreading styles. — Type, Herb. Zurich. In habit

simulates G. sibbaldioides according to Knuth but the type, merely

a scrap, is obviously stemmed, and doubtfully more than a young

or poorly developed G. ayavacense or G. piurense, if these are separable.

Amazonas: Chachapoyas, Mathews, type.

Geranium matucanense Knuth, Repert. Sp. Nov. 28: 5. 1930.

Perennial, with glabrous or sparsely pilose stout sulcate little

branched stems attaining 6 dm. or so, the root a more or less vertical

rhizome as much as 2 cm. thick at top; basal petioles about 15 cm.

long, all especially the upper as the peduncles (1-2-flowered, 4-12 cm.

long) and pedicels (4 cm. long) retrorse appressed pilose; stipules

soon marcescent, 7-8 mm. long; leaves 3-5 cm. broad, nearly to

base 5(-7)-parted, angulate-rotund, sparsely pubescent above with

minute hyaline trichomes, densely silvery or lutescent sericeous

beneath, the lobes coarsely incised pinnatifid, the lobules about

9 mm. long, 2.5 mm. wide, barely mucronulate; bracts linear-subu-

late, 4-5 mm. long; sepals oblong, 1 cm. long, 3 mm. wide, with

linear mucro 2 mm. long, closely sericeous; petals deep blue or blue-

purple, marginally pilose; stamens and sepals equal length, pilose;

gynaecium sericeous. — Section Rupicola. Seems to go toG. superbum

unless for the pilose petals and stamens, these not described from

the latter.

Lima: Matucana, 1+62, type.

Geranium molle L. Sp. PI. 682. 1753; 57.

An annual, typically rather densely pilose with flat trichomes

of unequal length; stems soon procumbent-ascending, often many

and becoming several dm. long; petioles longer than the reniform

leaves, these medially 7-9-lobed, the lobes deeply 3-parted with

broad obtuse segments; peduncles 2-flowered; sepals oblong-ovate,

acuminate, shortly acuminate and obscurely mucronulate, the ob-

cordate petals sometimes twice as long, finely ciliate toward base;

valves glabrous but transversely rugose, the seeds smooth. — Eurasia

and widely spread even to Chile. G. rotundifolium is similar but

petals shorter, valves puberulent, seeds reticulate.

Peru: To be expected. Eurasia.

528 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Geranium mollendinense Knuth, Pflanzenreich IV. 129: 580.

1912. G. multiflorum Knuth, Bot. Jahrb. 37: 561. 1906; 146 not

Gray, 1854.

Character in general that of G. Weberbauerianum; leaves from

base to apex 2 cm. long, 5-7-parted nearly to base, the apically

rounded divisions 3-5-lobulate, the lobules coarsely 3-5-lobulate,

the middle one often 5 mm. long, 7 mm. broad, mucronulate, the

lobes of the upper leaves much narrower, acute; lower stipules acute;

peduncles mostly biflowered, 1.25-2 cm. long, with short spreading

pubescence, recurving in fruit; sepals ovate or oblong-ovate, hirsute

on the 3 nerves, 3 mm. wide, 6-7 mm. long not counting mucro,

this about 1 mm. long; petals 11-14 mm. long, obcordate, emargi-

nate, white; mature fruit 2 cm. long, the valves long-hirsute, the

beak densely spreading puberulent. F.M. Neg. 4790.

Arequipa: Among rocks in the lomas at 400-600 meters near

Mollendo, Weberbauer 1579, type.

Geranium nivale Knuth, Bot. Jahrb. 37: 563. 1906; 79.

Similar in habit to G. muscoideum; leaves 7 mm. long, 10 mm.

broad, subcoriaceous, ashy sericeous with a dense indument, pal-

mately 5-parted nearly to base, the obovate- or lanceolate-spathulate

entire lobes obtusish or rounded and the middle often broader than

the lateral and tridentate, the lower lateral sometimes with 1 lobule

or tooth; stipules often four-fifths adnate; peduncles 5-20 mm. long;

sepals 8 mm. long, 3 mm. wide; petals 14-20 mm. long, long-clawed;

fruit 18 mm. long, the valves pilose. — Illustrated, Pflanzenreich I.e.

fig. 14, page 82. F.M. Neg. 4791.

Junin: Between Tarma and La Oroya on limestone rocks at 40

meters, Weberbauer 2533, type. La Oroya, Kalenborn 88.

Geranium patagonicum Hook. f. Fl. Ant. 252. 1844-47; 74.

G. titicacaense Knuth, Repert. Sp. Nov. 45: 60. 1938. G. tablasense

Knuth, Meded. Rijks Herb. Leiden 27: 68. 1915, at least as to Peru.

Lax with stems 4-6 dm. long, 3 mm. thick, grayish below with

rigid retrorse trichomes, toward the tips these hyaline and more or

less spreading; lower petioles 6-10 cm. long, glandular; stem leaves

similar to basal but shortly petioled, about 5 cm. wide, appressed

pilose-puberulent both sides with hyaline trichomes especially on

the nerves, 5-parted nearly to base, the divisions narrowly elongate-

rhomboid and subpinnate with scarcely mucronulate acutish teeth;

stipules and bracts linear, very acute, the former 8-9 mm. long, the

FLORA OF PERU 529

latter 4 mm. long; peduncles 2-4 cm. long, 2-flowered, densely glandu-

lar as pedicels (5-7 mm. long), sepals and beak of fruit 4-6.5 mm.

long; sepals oblong-ovate with mucro 1 mm. long, the petals 6-8 mm.

long (probably still longer), purple or roseate; beak 2 cm. long, valves

pilose, seeds punctulate-rugose. — This is Knuth's description of

G. titicacaense of which he wrote : distinct from G. patagonicum Hook,

f. in the dense glandular indument of the upper parts (Knuth).

The type was originally referred to Hooker's plant by Knuth him-

self, who however describes that as having the broad leaf-segments

of G. mexicanum; maybe there is one somewhat variable species.

Moquehua: Carumas, 3,000 meters, Weberbauer 7307 (type,

G. titicacaense). — Arequipa: Ubinas, Raimondi (det. Knuth). — Cuzco:

Hacienda Pfuyucalla, 3,400 meters, (Herrera 2981, det. Knuth).

Cerro de Cusilluyoc, in thicket, 3,000 meters, Pennell 14133 (det.

Knuth, G. tablasense). Yucay, Prov. Urubamba, 2,840 meters,

Vargas 712 (det. Standley, G. tablasense). Bolivia. "Rebojillo."

Geranium Pavonianum Briq. Ann. Cons. Jard. Bot. Geneve,

11&12: 183. 1908; 83.

In general like the pubescent form of G. sessiliflorum but densely

tomentulose; petioles 1-1.5 cm. long; leaves 5 by 9 mm., 5-lobed

scarcely to middle, the lobes entire or slightly tricrenate; peduncles

about 2 cm. long; sepals 3.5 mm. long with mucro 0.5 mm. long;

petals 6-7 mm. long, 3 mm. broad.— F.M. Neg. 26375.

Peru(?) : Without data by Pavon in Herb. Delessert, type.

Geranium peruvianum Hieron. Bot. Jahrb. 21: 316. 1895; 160.

Stems sparsely branched, ascending, hirsute-pilose with spreading

or subreflexed trichomes; lower petioles to 4 cm. long, hirsute;

leaves hirsute-pilose both sides, the lower cordate-orbicular, 7-lobed

with cuneate apically tridentate lobes, the terminal teeth broadly

ovate, shortly acuminate, the narrower more elliptic lateral acutish;

basal leaf (one known) 1 cm. long, 18 mm. wide, ultimate depth of

upper sinus about 5 mm.; stem leaves smaller, 5-lobed, similar, or

the lowest lateral lobes entire or with a tooth or lobule; peduncles

apparently all axillary, 1-flowered, bibracteate at base, the bracts

like the narrow scarious stipules, or ebracteate, 1.5-6.5 cm. long;

sepals 3-nerved, oblong, 8-10 mm. long, 3 mm. wide, hirsute, ob-

tusish, mucronate; petals obovate-cuneate, clawed, truncate, about

5-nerved, the veins flabellately branched, said to be about 16 mm.

long including the 4-5 mm. long claw, 7 mm. wide below the apex;

530 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

filaments cilate at base; fruit about 7 mm. long, the hirsute valves

4 mm. long, the glabrous beak about 3 mm. long. — The negative of

the scrappy type shows much smaller flowers than described, actually

only about 12 mm. long but maybe not fully grown; in this case the

species may be compared with the probably distinct G. Weberbauer-

ianum, or if smaller-petaled with the glandular G. Stuebelii or

eglandular similar G. ayacuchense or G. Raimondii. Therefore until

again collected it may best be restricted to type. F.M. Neg. 4792.

San Martin: Between Pacasmayo and Moyobamba, Stuebel 51 d,

type.

Geranium piurense Knuth, Repert. Sp. Nov. 18: 290. 1922.

Much like G. ayavacense; stems about 2 dm. long; internodes

2-3(8) cm. long; lower petioles to 5 cm. long, all nearly filiform;

leaves 2-3 cm. wide, 5-7-parted to base; stipules long-acuminate

from base to filiform tip; peduncles 1.5-3(6) cm. long; sepals 5.5-7

mm. long; petals spathulate, two times longer than calyx, early red-

dish white, finally white with purple red veins; fruit 15 mm. long, the

beak ashy puberulent, the valves glabrate with a few dorsal setae. —

Peraffine G. ayavacense (Knuth), and doubtfully ex char, distinct,

especially as from same region.

Piura: Above Ayavaca, grass steppes and small shrubs, Weber-

bauer 6374, type.

Geranium pusillum Burm. f. Spec. Geran. 27. 1759; 48.

Puberulent annual, the trichomes all about the same length;

stems many, ascending or procumbent, sometimes elongate; leaves

reniform, more than medially 7-9-lobed, the lobes scarcely a third

3-parted, with ovate obtuse teeth; basal petioles elongate; peduncles

biflowered; sepals oblong-ovate, acuminate but little mucronulate;

petals narrowly obcordate, barely or not longer than calyx, purple,

finely ciliate at base; valves and seeds smooth. — Another annual

that is widely spreading over the world and so probably to Peru.

Peru: See above. Eurasia.

Geranium Raimondii Knuth, Repert. Sp. Nov. 28: 9. 1930.

G. canescens Wedd. Chlor. And. 2: 286. 1857; 215, not L'He"r., 1787.

Cespitose, nearly all parts ashy tomentose, the shortly branched

caudices clothed with the remains of the thick petioles; flowering

stems unequal, erect, sparsely branched; leaves at least 15 mm. wide,

mostly basal, long-petioled, densely tomentose both sides, rotund,

FLORA OF PERU 531

deeply 7-lobed, the imbricate obovate-cuneate lobes 3-lobulate, the

lobules broadly ovate, obtuse; free part of stipules linear-subulate,

puberulent or glabrate; peduncles long exceeding petioles, 1-flowered,

ebracteate; flowers rather large (ex neg. 10-12 mm. long), white,

the much shorter sepals 8 mm. long, lanceolate, slenderly mucronate;

beak about 8 mm. long, pubescent. — To judge by the photograph

the pubescence is appressed strigose rather than tomentose; the

type is a scrap. F.M. Neg. 4773.

San Martin (?): Between Cajamarca and Chachapoyas, 3,600

meters, Raimondi, type.

Geranium renifolium Hieron. Bot. Jahrb. 21: 315. 1895; 158.

With the habit of the related G. Stuebelii but much less pubescent

and not at all glandular; basal petioles 2.5 cm. long, appressed

sericeous-pilose, the 7-lobed reniform leaves above sparsely beneath

lightly hirsute or glabrate, the truncate-rounded lobes usually entire;

stem leaves similar, 3-7-lobed, smaller; peduncles 2.5 cm. long,

reflexed sericeous; sepals 6.5 mm. long; stamens unequal, the longer

6 mm. long. — Illustrated, Pflanzenreich I.e. fig. 21.

San Martin: Above Tambo Mayo between Pacasmayo and

Moyobamba, Stuebel 43, type.

Geranium Ruizii Hieron. Bot. Jahrb. 20: Beibl. 49: 31. 1895;

81.

Habit of G. Lechleri; pubescence shorter, appressed sericeous;

petioles to 15 mm. long; leaves 7-parted to about the middle, to

5 mm. long, 6 or 7 mm. wide, the lobes entire or often with a tooth;

sepals about 6 mm. long, 2 mm. broad, mucronulate, glabrous

above; petals cuneate, long-clawed, obtuse, 10-11 mm. long, 4 mm.

wide, 5-nerved; filaments and pistils about 5 mm. long; petal claw

ciliate at base in my collection; fruit about 12 mm. long, hirsutu-

lous. — Knuth suggests perhaps not specifically distinct. Used as an

astringent for the gums.

Junin: Morococha, (Raimondi, det. Knuth). — Huancavelica:

(Raimondi, fide Knuth). — Lima: Grassy upland slope, Rio Blanco,

7184 (det. Knuth).

Geranium scissum Knuth, Repert. Sp. Nov. 28: 4. 1930.

Perennial, with many ascending-suberect stems branched only

above 5-10 cm. where repeatedly forked attaining about 3 dm.;

stems, petioles (the lower often 4 cm. long, the upper leaves nearly

532 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

subsessile) and peduncles more or less hirsute; leaves angularly

rotund, about 3 cm. wide, appressed hirsute above, the nerves there

prominent, three-fourths or in the upper leaves four-fifths 5-parted,

the lobes cuneate-narrowed below the middle, remotely pinnate-

incised with oblong minutely mucronulate lobules; stipules linear

above, very acute, 8 mm. long; peduncles 1-flowered, ebracteate,

2 cm. long, very hirsute beneath the calyx, this with ovate-oblong

glaucous green sepals 1 cm. long, 4 mm. wide, the petals 2 cm. long,

broadly obovate from a narrowed base, apparently pale; filaments

somewhat hirsute below; the gynaecium densely sericeo-hirsute

basally. — Probably ex char, should be included in G. superbum;

peduncles always 1-flowered?

Cuzco: Hacienda Churu, Paucartambo, (Herrera 1916, type).

Geranium Seemanni Peyr. Linnaea 30: 66. 1859; 197. G. mexi-

canum HBK. var. minoriflorum (Briq.) Knuth & var. macranthum

(Briq.) Knuth, Pflanzenreich IV. 129: 197. 1912 fide Moore, Contr.

Gray Herb. 146: 84. 1943.

Perennial, the root sometimes woody, the stems at first erect

elongating to as much as 10 dm., branching-ascending in age, gla-

brate to densely long-pilose-hirsute with flat trichomes, the long

basal petioles, these to 2 dm. long, and the ample leaves, especially

on nerves beneath, similarly pubescent or the indument above ap-

pressed; leaf-divisions 5 nearly equal and extending one-half to two-

thirds to base, cuneate-obovate and twice cleft or dentate or shallowly

so above, the middle division with short oblong obtuse segments;

stem leaves similar but puberulent and long-pubescent, 2-6 cm.

wide, 3-lobed, the segments dentate or cleft above the middle, the

middle one broadly rhombic the latter with a basal lobule; peduncles

1.5-3 or 4 cm. long, pubescent like the leaves and usually somewhat

glandular, 2-flowered, solitary from the upper axils or cymose at

the branchlet tips, the pedicels 3-10 mm., pilose or glandular, erect

in fruit; sepals 4-5 mm. long, awned; petals 6-8 or sometimes nearly

10 mm. long, 2-3 mm. wide, usually tinted, rarely nearly white,

pilose or glabrate at base the veins nearly free except near the tip;

fruit 17-22 mm. long, style beak 1.5-2 mm. long, column hispidulous

and usually glandular, the carpels long-hirsute, the seeds reticulate,

usually shallowly. — Weberbauer noted stems as long as 3 meters.

After Moore, who recognizes the closely allied G. subulato-stipulatum

Knuth, 199, G. guatemalense Knuth, 200, and other segregates.

However I can only cite the following collections here since deter-

FLORA OF PERU 533

mined by Knuth as G. mexicanum var. minor "iflorum. It is amazing

to me that monograph students so often, as here, treat a widely

distributed species-complex as constituting an areal problem.

Cuzco: Ollantaytambo, Cook & Gilbert 717 (det. Knuth). — Puno:

Sandia, (Weberbauer 640, det. Knuth). — Huanuco: Waste land,

Pillao, Woytkowski 115, Mexico. "Chepu-chepu" (Cook & Gilbert).

Geranium sessiliflorum Cav. Diss. 4: 198. pi. 77. 1787; 83.

G. caespitosum Walp. Nov. Act. Acad. Leop. -Carol. 19: Suppl. 1:

315. 1843, fide Knuth. G. razuhillcaense Knuth, Repert. Sp. Nov.

28: 2. 1930?

Cespitose and branched caudices crowned by the many leaves,

crowded stipules (these a cm. or two long), solitary and rather showy

white flowers on retrorsely pilose often silvery sericeous, sometimes

lanate peduncles a half to 2 cm. long; petioles 1-8 cm. long, typically

somewhat pilose; leaves nearly glabrous except for the nerves and

cilia or densely appressed puberulent or sericeous both sides (rarely

very shortly), often irregularly reniform, deeply 5-7-parted the

medial of the usually tridentate lobes rounded or a little elongate

and subpinnatifid; sepals oblong, acute, hirsute, 4-5 mm. long, the

oblong petals usually glabrous, one and a half times longer; fruit

(8) 15 (-17) mm. long, valves pilose, beak puberulent. — A charac-

teristic polster plant of the high punas and grass steppes between

3,400 and 3,600 meters. G. sericeum Willd. of Ecuador, similar,

has 5-7-parted leaves, the lobes medially or basally 2-5-laciniate,

pubescence rather loose; G. ecuadoriense Hieron., 86, leaves of G.

sericeum but indument appressed. The indument of Knuth' s species

of 1930, I.e., according to him "probably a variety but with entirely

different habit and indument" is short, nearly velvety; it can be

scarcely more than a variant, in which case it may be given a simpler

name, for instance var. albatum Macbr., var. nov. Illustrated,

Pflanzenreich I.e. fig. 15, page 84 after Weberbauer, page 202.

Cajamarca: Above Ocros, Weberbauer 224. — Lima: Banos, Al-

pamarca, (Maclean, Pickering). Chicla, (Weberbauer 255, in part).

Hacienda Arapa near Yauli, (Weberbauer 363}; 221. Bunch grass

slopes, Rio Blanco, 808; 29? '4; 3006 (all det. Knuth). Viso, 608

(det. Knuth). — Ancash: Pass Chonta, (Weberbauer 2775}. — Junin:

Tarma to Oroya, (Weberbauer 2549). Huaron, 1148 'A (glabrate, det.

Knuth). Above Tarma, 4,500 meters, Kittip & Smith 21971 (det.

Knuth). La Oroya, Kalenborn 80. East of Huancayo, 3,400 meters,

Stork & Norton 10217 (det. Standley). In puna grass, 4,500 meters,

534 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Prov. Huancayo, Stork 10939 (det. Standley, G. cucullatum); also

Soukup 1863 (distr. as G. Ruizii). — Huancavelica: East of Surcu-

bamba, 3,100 meters, Stork & Horton 10383 (det. Standley).—

Ayacucho: Mount Razuhuillca, small flat cushions in puna, 4,400

meters, Weberbauer 7495 (type, G. razuhilkaense) . — Cuzco: Prov.

Cercado, (Gay}. Saxaihuaman, Herrera 2383. Pucara, Puno to

Cuzco, Prov. Paruro, Vargas 846. Prov. Quispicanchis, Vargas 926

(det. Standley, G. filipes); (Weberbauer 410); 185, 203.— Puno:

Suchez to Poto, 4,600 meters, (Weberbauer 1003a); 219. Araranca,

4,100 meters, Pennell 13439 (det. Johnston). — Moquehua: Volcano

Tacsani, Weberbauer 7326. — Tacna: (Meyen, det. Knuth). To Pata-

gonia; also in Tasmania and New Zealand.

Geranium sibbaldioides Benth. PI. Hartw. 166. 1845; 88. G.

cucullatum HBK. Nov. Gen. & Sp. 5: 231. 1822, not L. G. ciliatum

Willd. ex Spreng.-Syst. 3: 71. 1826, not Cav. and others.

Closely to rather laxly cespitose from a long-produced rhizome,

the caudices to 7 mm. thick supporting mats 5-7 cm. in diameter;

petioles puberulent, 2-3 cm. long, with reflexed trichomes; leaves

many, basal, typically glabrous except the definitely ciliate margins,

round-reniform, 9-15 mm. broad, palmately 5-parted to base, the

lobes 3-parted about one- third, these segments oblong or the medial

broader, all acutely mucronate, the outer two lobes often entire,

acute, oblong-lanceolate; stipules to 8 mm. long, the free parts

glabrous, membranous, acuminate-aristate; peduncles solitary, 1-

flowered, often ebracteate, nearly filiform, little longer (not at all,

my spec.) than the leaves, more or less densely reflexed hirsutulous;

sepals oblong-lanceolate, mucronate, glabrous, about 5 mm. long;

petals obovate, unguiculate, violet-red (apparently sometimes white),

with 3 or 5 more deeply colored nerves (in my 2191, 7 mm. long);

beak of fruit finally 1 cm. long or, fide Knuth, barely one and a half

times longer than calyx, pulverulent. — After Knuth; I have not

seen Bentham's plant. Var. elongatum (Wedd.) Macbr. Candollea

6: 7. 1934, G. cucullatum HBK., var. elongatum Wedd. Chlor. And.

2: 285. 1857, has strongly elongate prostrate branches the leaves

pubescent both sides, smaller and known as yet from Colombia;

Weberbauer 6083 with subentire leaf lobes is a variant of it or dis-

tinct. But it agrees better otherwise with negative of HBK. type

than any other Peruvian specimen seen. Illustrated, Pflanzenreich

I.e. fig. 14e. F.M. Neg. 35704 (G. cucullatum).

Piura: Huancabamba, 3,000 meters, Weberbauer 6083 (var.).—

Amazonas: Chachapoyas to Moyobamba, (Raimondi, det. Knuth). —

FLORA OF PERU 535

Huanuco: Northeast of Huanuco, grassy hillside, 3,500 meters, 2191,

det. Knuth. — Junin: West of Huacapistana in rather swampy places,

3,500 meters, (Weberbauer 2251). — Cuzco: Province del Cercado,

(Gay, fide Weddell).

Geranium Smithianum Knuth, Repert. Sp. Nov. 34: 146.

1933.

Diffuse, the ascending appressed pubescent stems 3-6.5 dm. long,

the middle internodes 5-10 cm. long; lower petioles 7 cm. long the

upper gradually shorter, all appressed retrorse pubescent; leaves

above sparsely and very shortly puberulent, more densely so beneath

and marginally, reniform-orbicular, 2.5 cm. wide, usually 5-parted

to base, the lobes broadly cuneate and variously incised, the medial

coarsely pinnate with about 5 oblong segments 5 mm. long, 2 mm.

wide, the outer with 3-4 segments; stipules narrowly lanceolate,

acute, 4-6 mm. long; peduncles ebracteate, 1-flowered, 1.5 cm. long,

ashy subsericeous with retrorse appressed trichomes; sepals oblong-

ovate, acutish, (4.5) 5.5 mm. long, scarcely nerved, marginally and

basally long white hirsute, slightly exceeding stamens and pistils;

petals obovate, 7-8 mm. long, with few nearly unbranched veins,

rose or whitish; beak about 13 mm. long, 1.5 mm. thick, puberulent,

the valves pilose. — Section Diffusa; affine G. Sodiroani (Knuth) ; and

seems also to resemble G. ayacuchense, but flowers smaller.

Lima: Rio Blanco, 3,300 meters, Killip & Smith 21769, type.—

Ayacucho: Pampola, 3,000 meters, (Killip & Smith 22213; 23248,

both det. Knuth).

Geranium Sodiroanum Knuth, Bot. Jahrb. 37: 557. 1906;

210. G. diffusum HBK. var. subsericeum Hieron. Bot. Jahrb. 20:

Beibl. 49: 32. 1895.

Character in general that of G. diffusum and allies but the petioles

mostly about the same length, 2-5 (8) cm. long and the peduncles

distinctly longer; leaves more or less pubescent only beneath, 15-32

mm. wide; stipules nearly hirsute, 4-6 mm. long; sepals 4.5-5.5 mm.

long, almost as long as the white petals; stamens half as long as

calyx, the fruit two and a half times longer, the valves pilose, the

beak puberulent. — Seems probably only a form or variety along with

other segregates. Illustrated, Knuth, I.e. page 211.

Lima: Chicla, stony places among short plants, Weberbauer 255;

169. Ecuador.

536 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Geranium Staff ordianum Knuth, Repert. Sp. Nov. 40: 216.

1936.

Cespitose from a long simple vertical root that supports a mat

as much as 8 cm. wide, silvery sericeous with closely appressed

trichomes even to the sepals except for their membranous margins;

petioles 3.5-5 cm. long; leaves orbicular, 1.5-1.75 cm. wide, to two-

thirds 5-7-parted, the lobes mostly deeply 3-lobulate with ovate or

oblong obtusish lobules 3 mm. long, 2 mm. wide; peduncles solitary,

ebracteate, 2-4 cm. long; sepals long-oblong, 8 mm. long, equaled

by the stamens, nearly 3 mm. wide with mucro 1 mm. long; petals

to 18 mm. long, somewhat retuse; fruit twice as long as sepals, the

beak 2 mm. thick. — Distinctive in section Andina in the long

peduncles and the large flowers (Knuth) ; the description reads much

like that of G. Weddellii in a more luxuriant state.

Cuzco: At 4,000 meters, (Stafford, type, in herb. Kew).

Geranium Stuebelii Hieron. Bot. Jahrb. 21: 316. 1895; 159.

Diffuse, procumbent, the flowering branches ascending, the

hirsute pubescence dense and the trichomes in part gland tipped

often deciduous or sparse except on the sepals; petioles 3-10 mm.

long; leaves reniform, about 13 mm. by 2 cm., 5-lobed to the middle

or deeper, the terminal lobe a trifle larger, tridentate or rarely entire,

the lateral lobes entire or the upper of these tridentate, the lowest

with one tooth on the lower edge ; stipules scarious, ciliate, hirsutulous

both sides, narrowly ovate, acuminate; peduncles 3.5-4.5 cm. long,

after an thesis recurved, the flowers nodding even in bud; sepals

oblong, obtusish, mucronate (as leaf-lobes), 9 mm. long, about a

third as wide, 3-nerved, green except the narrow margins, the outer

densely the inner sparsely hirsute; petals obovate-cuneate, clawed,

obtusely emarginate, 5-nerved, about 12 mm. long, half as wide;

stamens subequal, dilated at the ciliate base, 5 mm. long; pistils

6 mm. long; valves hirsute. — Flowers white, turning rose. G. chil-

loense Willd., 146, of Ecuador would be sought here; it has larger

leaves with the divisions in part subpinnate. Illustrated, Pflan-

zenreich I.e. fig. 22.

San Martin: Above Tambo Mayo between Pacasmayo and

Moyobamba, Stuebel 43a; type. — Amazonas: Above Balsas, (Rai-

mondi 1793, det. Knuth). — Cajamarca: Near Chota, 2,500 meters,

prostrate, Stork & Horton 10040 (det. Standley, G. bolivianum) .

Near. Conchan, Prov. Chota, 2,500 meters, Stork & Horton 10062

(obscurely glandular, det. Standley, G. chinchense). — Lima: Above

FLORA OF PERU 537

Obrajillo, 3,100 meters, rocky canyon, Pennell 14417 (det. Knuth,

G. choimacotense'). Huaros, rock slide, diffuse, Pennell 14718 (det.

Johnston, G. bolivianum).

Geranium superbum Knuth, Bot. Jahrb. 37: 561. 1906; 140.

Stems 1-4 from a subligneous stout rhizome, simple below but

3-10 cm. above the base pseudo-dichotomously branched, ascending,

2-5 dm. high, retrorsely appressed pubescent to tip; basal leaves

early many, the retrorsely pilose petioles 5-10 cm. long; leaves

appressed pubescent above, sericeous beneath, reniform-rotund, the

lobes coarsely palmate-pinnatifid, the segments linear-oblong, acut-

ish, reddish-brown callused ; stem leaves similar but gradually shorter

petioled; stipules subulate, 6-7 mm. long; peduncles biflowered

(rarely 1-flowered), bracteate, 6-10 cm. long, sparsely retrorse-

pilose; bracts linear-subulate, very acute, 5 mm. long; pedicels 1-3

mm. long; sepals oblong or ovate-oblong, rather densely ashy pilose,

10 mm. long with reddish mucro 1.5 mm. long; corolla 3 cm. broad

the broadly obovate petals 16-21 mm. long, scarcely emarginate;

stamens shorter than calyx; valves long appressed pilose, the beak

spreading puberulent. — G. rupicolum Wedd., 148, of Bolivia has

leaves more sparsely pubescent above, peduncles 1-flowered, ebracte-

ate. F.M. Neg. 4798.

Lima: Between Puente de Arichi and Chicla, grassy brook shore,

3,500 meters, Weberbauer 259, type; 169. Rio Blanco, Killip &

Smith 21758.

Geranium Weberbauerianum Knuth, Bot. Jahrb. 37: 556.

1906; 145.

Stems many, 2 dm. high or higher, a little taller than the basal

leaves, pseudo-dichotomously squarrose-branched, softly puberulent

as petioles (the basal 7-17 cm. long), former emarcidate ones densely

crowning the thick vertical rhizome; leaves appressed puberulent

and beneath on the nerves densely long ashy-pilose, 4 cm. broad and

long, 7-angulate-orbicular, deeply palmately 7-parted, the more or

less broadly rhomboid lobes unequal, the larger middle one 25 cm.

long, 17 cm. wide, all coarsely palmately pinnatifid with obtuse or

rounded segments, the middle often 3-4 mm. long and broad or larger;

stem leaves similar but shorter petioles, the uppermost nearly sessile;

stipules linear-acuminate, puberulent; peduncles 1-flowered, ebracte-

ate, either solitary from the axils of branches or pseudo-umbellately

crowded toward the branch tips, 2-5 cm. long, ashy pilose; sepals

8-9 mm. long, lanceolate or nearly linear-lanceolate, acutely acumi-

538 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

nate, the membranous margins and midnerve long setose-pilose;

corolla 15 mm. broad, whitish, with deep colored nerves, the lanceo-

late-obcuneate petals barely emarginate, 16-18 mm. long; fruit 14

mm. long, the beak minutely puberulent; seeds nearly glabrous dor-

sally. — Easily recognized by the globose habit, the stems below

strictly pseudo-dichotomously branched, the flowers above pseudo-

umbellate while the lower are solitary, the leaf-segments rounded

apically (Knuth). Cf. G. ayacuchense. F.M. Neg. 4801.

Cuzco: At Pucana, rocky canyon in limestone, 3,700 meters,

Weberbauer 450, type; 186. Yucay, 3,000 meters, Soukup (distr. as

G. peruvianum) .

Geranium Weddellii Briquet, Ann. Cons. Jard. Bot. Geneve

11 & 12: 183. 1908; 80.

Compactly cespitose, the stout somewhat branching caudex a

cm. in diameter or larger, the vegetative portion of the plants rarely

more than 2 or 3 cm. above the surface with the large flowers barely

equaling or shorter than the closely sericeous 5-parted leaves, these

deeply and narrowly trilobulate with obtuse segments; petioles

slender, 1-1.5 cm. long; stipules 6-9 mm. long, long-adnate, finally

glabrous; peduncles many, solitary, ebracteate, 1-1.5 cm. long;

sepals oblongish, minutely mucronulate, often reddish, lightly

puberulent, 7 mm. long, the stamens about as long; petals finally

about 15 mm. long, glabrous in Soukup spec.; fruit beak nearly 1 cm.

long, appressed sericeous-pilose. — F.M. Neg. 26382.

Puno: Macusani, 4,336 meters, Soukup 537 (distr. as G. Lechleri?).

Bolivia.

2. ERODIUM L'H<§r.

In general like Geranium but the leaves usually pinnately lobed

or dissected and more or less dentate, rarely merely lobed or even

subentire. Fertile stamens 5, the alternate sterile and scale-like.

Styles bearded inside. Pedicels usually recurved sharply in fruit or

after anthesis.

All the species in Peru are immigrants from the Mediterranean

region.

Leaves cordate-ovate, more or less lobed.

Leaves lightly if at all lobed, eglandular E. malacoides.

Leaves more or less trilobed, very glandular E. geoides.

Leaves pinnate or bipinnate.

FLORA OF PERU 539

Leaflets serrate and sparsely incised; stipules obtuse.

E. moschatum.

Leaflets incisely pinnatifid; stipules acute E. cicutarium.

Erodium cicutarium (L.) L'Her. ex Ait. Hort. Kew. 2: 414.

1789; 274. Geranium cicutarium L. Sp. PI. 680. 1753.

Annual or biennial, the more or less hirsute often reddish leafy

stems procumbent-ascending and finally several dm. long; leaves at

first in a close rosette, oblong-lanceolate, finely bipinnatisect, hirsute-

pilose, the alternate or subopposite oblique-ovate pinnae sessile or

subsessile, the uppermost confluent; peduncles to 12 cm. long, 5-10-

flowered, the medially connate involucral bracts to 2 mm. long;

pedicels 8-15 mm. long, early glandular; sepals ovate, nerved, tipped

with 1 or 2 setae, finally 6 or 7 mm. long, the often unequal rose-

purple bimaculate ciliate clawed petals little to twice as long or

longer; filaments little dilated at base, not toothed; beak of fruit

2-4 cm. long. — Variable vegetatively, the many named varieties and

forms not clearly of taxonomic interest. Adapted to light or barren

soils and often important as a forage plant; known in English as

Filaree, Alfilaria, Pin Clover or Storksbill, this last term applied to

other species also. There are many illustrations. Most of the native

names are after Herrera.

Lima: Obrajillo & Bafios, (Wilkes Exped.}. Lima, Weberbauer

120. — Amazonas: Lamud, Raimondi. — Junin: Rio Blanco, G. S.

Meyers (det. Ferris). La Oroya, Kalenborn 9. — Arequipa: Atiquipa,

Raimondi. Nevado de Chachani, 3,500 meters, Pennell 13269 (det.

Johnston). — Apurimac: Ampuy, Goodspeed Exped. 10630. — Cuzco:

Calca, Vargas 678. Prov. Cercado, 3,400 meters, Herrera ll.—Puno:

Salcedo, Soukup. Sta. Lucia, Sharpe 64- Chuquibambilla, 3,900

meters, Pennell 13401 (det. Johnston). Nearly cosmopolitan.

"Alfilerillo," "montillapafio," "tupu-tupu," "auja-auja," "yauri-

yauri," "trinchi-trinchi," "rchjillo," "agujo" (last two fide Vargas).

Erodium geoides St. Hil. Fl. Bras. 1: 99. pi. 19. 1825; 248.

Much like E. malacoides but much more glandular and densely

pilose-hirsute even from the base of the many branching stems, these

procumbent, 1-3 dm. long, with numerous more or less trisected and

incised-dentate leaves; bracts brown-membranous, glandular- villous;

sepals 5-6 mm. long, exceeded by the small lilac petals; beak about

3 cm. long. — It seems doubtful if the following collection could have

been more than a glandular state of the ruderal E. malacoides since

540 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

the species of St. Hilaire is native to a region rarely represented in

Peru.

Arequipa: Cachendo and Pasco, (Guenther & Buchtien 263; 263a,

both det. Brims). Southern South America.

Erodium malacoides (L.) Willd. Sp. PI. 3: 639. 1800; 245.

Geranium malacoides L. Sp. PI. 680. 1753.

Softly pilose annual or biennial sometimes 3 or 4 dm. high, the

leafy more or less branched stems ascending; basal leaves many,

long-petioled, cordate-ovate, often 4 cm. long, 2.5 cm. wide, un-

divided or sublobed, crenate-dentate; upper leaves gradually reduced,

the uppermost sessile; stipules acute, often 6 mm. long; peduncles to

8 cm. long, 2-8-flowered, more or less glandular as the filiform

pedicels, these 7-17 mm. long; bracts many, pale, glabrate or ciliate,

ovate, obtuse, 2-2.5 mm. long, lightly connate at base; sepals oblong-

ovate, glandular, 5 mm. long, 5-nerved with mucro 1 mm. long, the

concolor (not spotted) petals about as long; fruit-beak 2-3 cm. long,

glabrate. — Willdenow used the name in Phyt. 10. 1794. E. laciniatum

(Cav.) Willd. 241, allied and to be expected, may be known by the

retrorse-hispid stems, the more or less 3-lobed lower leaves, the

uppermost more or less pinnatifid, the bracts only 2, ovate-suborbicu-

lar, the fruit-beak often 5 cm. long or even longer. Often illustrated.

Lima: At Lima, Ruiz & Pavdn. Obrajillo, (Wilkes Exped.}.

Amancaes Hills, Weberbauer 1627. Rio Chillon, Pennell 14483.

Puente de Verrugas, Seler 226. Matucana, 94; 124; Weberbauer

63. — Loreto: Raimondi. — Cuzco: Near Cuzco, Herrera 282. South-

ern Europe.

Erodium moschatum (L.) L'HeY. ex Ait. Hort. Kew. 2: 414.

1789; 281. Geranium moschatum L. Sp. PI. 680. 1753.

Rather similar to the related E. cicutarium but the stems when

developed (the plant is early a close rosette of leaves) glandular and

typically pale, the stipules conspicuous, scarious, the leaflets broadly

ovate, serrate or lobed, petiolulate, the bracts free, to 4 mm. long,

sepals without bristles or merely 1-2-short-setose, filaments winged

at base and with 2 upward pointing teeth, petals glabrous, equal,

not spotted, at least twice as long as sepals; stipules large, obtuse,

the stems becoming stout or fleshy. — As suggested by the name the

original plant presumably was musk-scented but apparently the

weedy races have little if any odor. It has been observed that

Mimulus moschatus is a similar case. There are many illustrations

FLORA OF PERU 541

as in Pammel, Man. Poison. PI. fig. 320. E. Botrys (Cav.) Bertol.

has oblong-ovate leaves the basal ordinarily pinnately cleft into broad

crenate lobes, is scarcely white-pubescent, beak of fruit often 7 cm.

long or longer.

Lima: Along railroad to La Oroya, Weberbauer 97. Common at

Matucana, 204- — Junin: Near Huancayo, Luis A. Chavez 181.—

Arequipa: Atiquipa, Raimondi. Widely dispersed from the Mediter-

ranean region.

3. RHYNCHOTHECA R. & P.

Rhynchothelia Pers. Syn. PI. 2: 92. 1806; Aulacostigma Turcz.

Bull. Soc. Nat. Mosc. 20. 1: 149. 1847.

Shrubs with axillary pedicelled flowers at the tips of the many

branches, opposite entire or lobed leaves. Sepals 5. Petals and

glands none (or early caducous?). Stamens 10, free, all with large

oblong anthers. Ovary 5-lobed, maturing as 5 caudate one-celled

carpels that remain joined at base and apex.

Rhynchotheca spinosa R. & P. Syst. Veg. 142. 1798; 549.

R. integrifolia HBK. Nov. Gen. & Sp. 5: 232. pi. 464. 1822; R. diversi-

folia HBK. I.e. 233. pi. 465. Aulacostigma inerme Turcz. I.e.

About 1 (-2.5) meter high, the slender tortuous flowering branches

4-5 cm. long, early quadrate and pubescent, later glabrate, leafless

and spinescent; leaves many, oblong-ovate, entire or palmately

3-5-lobed, more or less sericeous beneath becoming glabrate, 10-15

mm. long, 3.5-6 mm. wide; sepals obovate, obtuse, mucronulate,

7-9 mm. long, twice as long as the caducous petals, or, apparently,

these usually wanting; fruit 14-17 mm. long, silvery sericeous.—

Knuth recorded three variants: var. integrifolia (HBK.) Knuth,

var. diversifolia (HBK.) Knuth, var. lobata Knuth, I.e. 550, the last

with the most deeply 3-5-lobed leaves, the largest (17 mm.) carpel

beaks; the other two forms are Ecuadorian. Flowers yellow and

lavender (Seibert), in my specimen sepals reddish, anthers greenish-

yellow. Illustrated, Knuth, I.e. pages 550, 551.

Lima: Callao, Ruiz, type. Dense canyon shrub, Chasqui, 3302

(det. Johnston). — Piura: Prov. Huancabamba, 3,200 meters, Weber-

bauer 6321. — Junin: Fundacion, Cerro de Pasco to Huanuco, 3,100

meters, Seibert 2304. — Cuzco: To Sta. Ana, 3,200 meters, among other

shrubs, Weberbauer 4969 (var. lobata); 244. — Puno: Sandia to Cuyo-

cuyo, 2,300 meters, Weberbauer 873a (var. lobata) ; 237. Ecuador.

542 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

4. BALBISIACav.

Suffrutescent, more or less branched and pubescent, the alternate

or opposite leaves mostly 3-parted, rarely entire. Flowers solitary,

terminal, yellow on rather long peduncles, regular, with 5 imbricate

sepals, 5 hypogynous contorted petals. Stamens 10, hypogynous,

free, anthers large. Ovary 5-angled, 5-celled, erostrate; stigmas 5,

ligulate, sessile, ovules many in 2 series in each cell. Capsule apically

dehiscent loculicidally by 5 valves that persist, the many seeds with

thin fleshy endosperm.

Filaments and ovary subequal; sepals about 7-9 mm. long; central

Peru B. verticillata.

Filaments longer than ovary; sepals about 10 mm. long.

Leaflets 1-1.5 mm. wide; involucral leaves setaceous.

Leaflets 5-10 mm. long, usually greenish B. Meyeniana.

Leaflets 3-6 mm. long, silvery B. Weberbaueri.

Leaflets 2-3 mm. wide; involucral leaves linear-lanceolate.

B. peduncularis.

Balbisia Meyeniana Klotzsch, Linnaea 10: 432. 1836; 555.

Ledocarpon Meyenianum (Klotzsch) Steud. Nomencl. ed. 2. 2: 20.

1841.

Somewhat to much branched shrub to about one meter, the ash-

colored branches glabrous, the flowering branchlets densely tomen-

tose, 1 mm. thick; leaves numerous, opposite, 1-1.5 cm. distant,

sessile, mostly 2-3-parted to base, the divisions ashy subhirsute or

sericeous, linear, entire, obtusish, 5-8 mm. long, 1 mm. wide, gradu-

ally narrowed to base; flowers terminal on slender puberulent pedicels

1.75-3 cm. long; involucral leaves 8-10 mm. long, the subulate

lobules more or less spreading hirsute; sepals to 14 mm. long, 3 mm.

wide, lanceolate, gradually and acutely acuminate, ashy puberulent;

corolla orange, 4-4.5 cm. broad, the petals twice as long as the

sepals, 2-2.5 cm. long, 11-13 mm. wide, rounded apically, entire;

filaments twice as long as the white hirsute ovary, the densely villous

capsule half as long as calyx. — After Knuth. It may be recorded

that my European notes read : apparently the same as B. verticillata.

Illustrated, Knuth, I.e. fig. 71.

Chewed under name of "capo" to prevent coughs ( Weberbauer) .

Ayacucho: Coracora, 2,900 meters, Weberbauer 5796 (distr. B.

Weberbaueri). Near Ayacucho, Weberbauer 5509 (det. Knuth). —

Apurimac: Hacienda Catahuacho, 2,900 meters, Weberbauer 5856

FLORA OF PERU 543

(distr. B. verticillata). — Arequipa: Rocky hill, 2,600 meters, Pennell

13188 (det. Johnston, B. Weberbaueri). — Moquehua: Open mixed

formation, 2,700 meters, Weberbauer 7271 (det. Knuth). In rocks

near Moquehua, 1,900 meters, Weberbauer 7430 (det. Knuth). —

Tacna: Meyen in 1831, type. Near Tarata, Metcalf 30361 (det.

Leonard).— Puno: Lake Titicaca, Besser 54- Bolivia. "Picipinto,"

"capo."

Balbisia peduncularis (Lindl.) D. Don, Edinb. New Phil.

Journ. 11: 277. 1831; 555. Ledocarpum pedunculare Lindl. Bot. Reg.

17: pi 1392. 1831. Cruckshanksia cistiflora Hook. Bot. Misc. 2: 211.

pi 90. 1831.

Branches erect, rather strict, the flowering branchlets 1.5-2 mm.

thick, densely tomentulose, the older brown and glabrous; leaves

numerous, opposite or nearly, 1-2 cm. distant, sessile, 2-3-parted

even to base, the canescent to sericeous-villous lanceolate obtusish

entire lobes 7-17 mm. long, 2-3 mm. wide; flowers orange, becoming

as much as 7 cm. in diameter, the pedicels 3-5 times longer than the

leaves, 2-5 cm. long, 1.25-2 mm. thick, puberulent; involucral leaves

7-12 mm. long, the linear-lanceolate lobules ashy sericeous; petals

cuneate-obovate, entire, 3-4 cm. long, 17-20 mm. wide below the

rounded apex; filaments one and a half to two times longer than

niveous ovary, the anthers oblong; capsule appressed sericeous, half

as long as the calyx. — Next to B. verticillata, with which Reiche

merged it, the oldest name in this group of six narrowly defined

closely allied species, races or variants, only one of the three that are

neatly demarcated being in Peru. Illustrated, Knuth, I.e. 556.

Arequipa: Near Mejia, (Guenther & Buchtien 1704, det. Bruns).

Chile.

Balbisia verticillata Cav. Anal. Cienc. Nat. 7: 62. pi 46.

1804; 557. Ledocarpon chiloense Desf. Me"m. Mus. Paris 4: 551.

pi 13. 1818? L. cistiflorum Meyen, Reise 1: 470. 1834?

Bushy-branched shrub becoming about a meter tall, the short

slender very leafy flowering branchlets densely but minutely puberu-

lent and terminating in a 1-2.5 cm. long peduncle to the solitary

showy clear or lemon-yellow flower; leaves 3-parted to base, the

linear leaflets obtusish or acute, more or less attenuate basally, often

10-12 mm. long, about 1.5 mm. wide, but sometimes smaller or a

few somewhat longer, always with rather prominent midnerve

beneath and more or less canescent but rather greenish than white

by the dense puberulence on both sides; involucral leaves setaceous,

544 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

shorter than or about as long as the narrowly ovate subsericeous calyx

lobes, these acutely acuminate, 7-9 mm. long; petals broadly obovate,

apiculate at the subtruncate-rounded apex, 1.5-2 cm. long; filaments

about as long as the densely silvery hirsute ovary, slightly ovate-

oblong anthers about 3 mm. long; pubescent capsules finally little

shorter than the persisting sepals. — This is the shrub common in the

foothills and it seems doubtful that in typical form it occurs in

Arequipa and Tacna as indicated by Knuth; otherwise there is

probably only one species concerned and not several, as Knuth

maintains.

Lima: Between Obrajillo & Canta, Nee, type; Ruiz & Pavdn.

Viso, 2,800 meters, Goodspeed Exped. 11541 (det. Johnston). Valley

Rio Rimac, 90 km. east of Lima, 3,000 meters, Goodspeed & Weber-

bauer 33057 (det. Leonard, B. Weberbaueri). Matucana, in rocks,

176 (det. Knuth, B. Meyeniana); at 2,389 meters, Soukup 2063 (det.

Standley); Rose 18664; Ward; Weberbauer 87; 114; 163, 164.

"Capo."

Balbisia Weberbaueri Knuth, Pflanzenreich IV. 129: 557. 1912.

Much like B. Meyeniana but uniformly with white or silvery

tomentose leaflets that are only 3-6 mm. long; petals truncate

apically, not apiculate, at least 2.5 cm. long when grown; filaments

longer than ovary; anthers oblong. — If distinct it is a local race of

B. verticillata, sensu lat. Used for fuel, the stem at base 12 mm.

thick and called "capo Colorado" (Hinckley).

Arequipa: Arampae to Pampa de Arrieros, 3,200 meters, Weber-

bauer 1408, type; also Rose 18963 (distr. asB. verticillata). Pampa,

3,050 meters, Hinckley 28 (distr. asB. verticillata). Base of Volcano

Misti, Weberbauer 4831; 128, 129. Near Yura, Schmidt (det.

Standley); R. S. Williams 2561 (distr. as B. verticillata). Sandy

loam in Cacti, Mount Chirvata, 2,200 meters, Eyerdam & Beetle

22107 (det. Johnston). "Capo Colorado."

OXALIDACEAE. Oxalis Family

Reference: Knuth, Pflanzenreich IV. 130: 1930.

Herbs with bulbous or fleshy rhizomes, acaulescent or caulescent,

or the Peruvian not infrequently more or less suffrutescent. Leaves

alternate, with or without stipules, digitately or pinnately 1-several-

foliate, the leaflets entire or cordate-incised; leaves rarely phylloide-

ous. Peduncles axillary, basal, cauline or apical, 1-flowered, ebrac-

FLORA OF PERU 545

teate or often cymose- or pseudo-umbellate and bracteate. Flowers

yellow, rose or white, hermaphrodite, regular, 5-merous, usually

heterostylous, the 5 free sepals and petals imbricate or the latter

valvate, sometimes contorted; glands none; stamens 10(15), more

or less biseriate, annulately joined at base, usually all anther bearing.

Ovary 5-lobed, 5-celled with 5 distinct styles except in Hypseocharis

1, the stigmas capitate; ovules 1-several. Capsules loculicidally

dehiscent, sometimes fleshy, 5-lobed, rarely baccate.

Averrhoa carambola L. of the South Pacific, cultivated in Brazil

as "Caramboleiro" and widely in all tropics, is a tree with impari-

pinnate leaves crowded at branchlet-tips, 5-7 pairs of oblique-ovate

leaflets, many small flowers in cyme-panicles, especially, baccate

fruit. The similar and also cultivated A. bilimbi L. has 10-20 pairs

of lanceolate leaflets.

Leaves digitately 1-several foliolate or pinnately 3-foliolate rarely

phylloideous; stamens 10, biseriate 1. Oxalis.

Leaves pinnate at least in part.

Stamens 10, biseriate; styles 5; caulescent herbs. .2. Biophytum.

Stamens 15, more or less triseriate; style 1; subacaulescent herbs.

3. Hypseocharis.

1. OXALIS L.

Character of the family. Petals contorted. Outer stamens

smaller, opposite sepals, inner larger, opposite petals. All Peruvian

species are digitately trifoliolate (except 0. fritillariiformis where

they are phylloidiform) and are sessile or equally subsessile except

as noted.

The important species in Peru, economically at least, is 0.

tuber osa, the edible tubers commonly known as Oca; other common

names and a reference may be found under the species name. The

Shamrock, according to the exhaustive paper on the application of

that common name by Harold N. and Alma L. Moldenke, Journ.

N. Y. Bot. Gard. 47: 49-59. 1946, may as well be Oxalis as Trifolium

or any 3-leaved small plant.

In preparing the following compilation relatively few photo-

graphs have been available or types seen; this as usual has been

indicated by placing the specimen citation in parentheses. It may

be historically useful to remark that the monographer never found

it convenient to permit photography and thus belonged to a small

minority who followed the opinion of some English curators. It

546 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

will be apparent to any user that Knuth's work has not been clear

to me and that I have not concerned myself with extra-Peruvian

names or revision, both, after all, beyond the scope of this work.

The following names are omitted : 0. leptopodes, 0. lespedezoides and

0. Pavonii, all by G. Don, Gen. Syst. 1: 754, 755. 1831. As their

brief characterizations are non-diagnostic they are nomina nuda

and so far as known no collections have been assigned to them, those

upon which they were based in the Lambert Herbarium by Ruiz &

Pavdn not having been identified by the monographer. There would

be no scientific purpose served if they were identified.

Caulescent species, the stems sometimes short, the leaves and

peduncles then forming a fleshy-ligneous rhizome or caudex or

the former even bulbiform. 0. fritillariiformis, not elsewhere

in key, has leaves in part simple and phylloidiform; all other

Peruvian species, leaves with 3 leaflets. (Pseudoacaulescent

annuals are included here.) Acaulescent species, page 552.

Stipules lacking or obscure; middle leaflet petiolulate sometimes

shortly but the lateral sessile or subsessile.

Leaflets acute to acuminate; shrubs.

Plants glabrous or nearly.

Cymes irregular; pedicels to 2 mm. long. ... 0. Tessmannii.

Cymes regularly parted; pedicels 3-3.5 mm. long.

0. Williamsii, 0. Macbridei.

Plants obviously more or less pubescent.

Flowers small, the sepals about 3 mm. long. .0. Spruceana.

Flowers large, the sepals 6-8 mm. long 0. Mathewsii.

Leaflets rounded or retuse at tip, rarely subacute, then plants

more or less herbaceous.

Spreading shrub; leaflets only 3-4 mm. wide and long.

0. peruviana.

Erect or subherbaceous; leaflets usually larger.

Pedicels subumbellate, early congested, finally spreading

or pendent; leaflets never subacute or flowers purple.

Stockily branched shrubs; sepals at least acutish, 4.5-6

mm. long.

Leaflets glabrous above, puberulent beneath.

0. hypopilina.

Leaflets puberulent above, tomentose beneath.

0. velutina.

FLORA OF PERU 547

Strict, herbaceous or partly ligneous with virgate

branches; sepals obtuse, 3-4 mm. long in flower.

0. Poeppigii.

Pedicels soon pendent or reflexing in racemes or bifurcate

cymes, or leaflets subacute, or flowers purple.

Flowers yellow or yellowish, rarely roseate in age.

Leaflets ovate-oblong, subacute (type); half-shrub.

0. juruensis.

Leaflets rounded or emarginate at tip.

Leaflets broadly obovate or obcordate.

Plants not at all arachnoid.

Stems glabrate or hispidulous 0. Cornelli.

Stems viscid-puberulent; leaflets small.

0. Pickeringii.

Plants arachnoid toward apex. . .0. Hochreutineri.

Leaflets narrowly cuneate-oblong, retuse.

0. bartolomensis.

Flowers purple or rose-colored from anthesis.

Sepals about 4 mm. long; leaflets elliptic, rounded both

ends 0. amazonica.

Sepals about 3 mm. long; leaflets subovate, obtuse or

subacute 0. Barrelieri.

Stipules usually obvious, sometimes completely adnate or petioles

merely margined at base, sometimes caducous or obsolete but

leaflets in any case all sessile or equally and minutely petiolu-

late.

Diminutive or fragile sometimes acaulescent annuals, the flowers

3-6 mm. long except in glandular 0. puberula (juvenile

plants 0. bulbigera might be sought here).

Petioles merely dilated at base; plants setulose-villous or

glandular-puberulent 0. micrantha, 0. puberula.

Petioles margined by the apically free scariose stipules.

Flowers solitary; stipules conspicuous, usually fimbriate;

plants dense 0. nubigena.

Flowers 2 or usually several; stipules 1-2-lacerate; plants

stemmed 0. cuzcensis.

Sturdy or succulent, various in duration and habit but, espe-

cially if annual, the flowers larger and plants not glandular-

puberulent.

548 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Petioles articulate often above the base and stipulately

margined, the stipules more or less free at tip; plants

never developing a fleshy caudex or rhizome but roots

sometimes tuber-bearing.

Stems low, more or less wiry below and usually rooting at

the leaf -bearing nodes; leaflets unless in 0. corniculata,

0. adpressa, rarely 5 mm. wide, capsules at least often

becoming erect, by the refracted pedicel, cylindric, as

long as or longer than broad (cf. also 0. Sterribergii}.

Longer filaments glabrous; peduncles often several-

flowered; pods much longer than broad.

0. corniculata.

Longer (or all) filaments hirsute; flowers 1-5; pods much

longer than thick 0. adpressa.

Longer filaments pubescent, flowers rarely binate; pods

(known) about twice as long as thick.

Pedicels often shorter than 1 cm. ; plants forming mats.

0. breviramulosa.

Pedicels usually longer; plants rather open in habit.

0. parvifolia.

Stems usually elongating, sometimes decumbent-ascending

toward the base and rooting there; leaflets usually

wider than 5 mm.; capsules (known) little longer than

broad except 0. Sternbergii and 0. tuberosa.

Sepals about 5 mm. long, the flowers less than twice as

long; stems succulent-herbaceous, not slender or

creeping at base 0. villosula, 0. spiralis.

Sepals (5) 6-8 mm. long, the flowers at least twice as

long; stems often firm, or slender and creeping at

base.

Stems succulent, in herb sulcate, obviously not erect;

leaflets usually more than 2 cm. long; roots

(complete) with one or more tubers; capsules

oblong-cylindric 0. tuberosa, 0. lucumayensis.

Stems firm to ligneous, at least becoming so toward

base, or in any case capsules ovoid (known), as

in the nearly glabrous.

Leaflets mostly about 2.5 cm. long, 1-1.5 cm. wide,

flowers usually many.

FLORA OF PERU 549

Erect strigillose-stemmed plant from fascicle of

tuberiform roots; shortly bifurcate umbel 20-

flowered 0. huantensis.

Scandent or lax plants, at least stems glabrous

below, peduncles soon long-bifurcate.

Leaflets oblong, about three times longer than

wide, villous beneath, stamens and style

glabrous 0. subintegra.

Leaflets cuneate-obovate, glabrous; stamens and

styles puberulent 0. melilotoides.

Leaflets obovate, appressed pubescent beneath;

stamens glabrous 0. medicaginea.

Leaflets mostly about 1.5 cm. long or smaller;

flowers few, borne loosely and irregularly.

Leaflets at least slightly longer than wide, stems

at least below not sericeous villous.

Stems ligneous, exfoliating below; filaments

sparsely ciliate 0. dolichopoda.

Stems slender, firm or merely subligneous;

filaments glabrous or evenly pubescent, at

least typically.

Leaflets obcordate, acute but scarcely cuneate

at base.

Internodes and petioles typically short;

stamens and styles glabrous.

0. mollis(1}, 0. lotoides.

Internodes, petioles short; filaments gla-

brous, styles not 0. medicaginea.

Internodes and petioles longer; filaments

dentate and styles hirsute.

0. phaeotricha.

Leaflets obovate, usually cuneate at base;

longer stamens glabrous.

Leaflets 1-2 cm. long, glabrous at least

above 0. phaeotricha.

Leaflets smaller, somewhat pubescent above.

0. fruticetorum.

Leaflets broadly obcordate, mostly or all wider

than long medially, shortly acute at oblique

550 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

base, plants more or less villous.

0. picchensis, 0. Sternbergii.

Petioles not margined (sometimes dilated at base, this persist-

ing), the stipules free, caducous or somewhat persisting

and more or less densely scarring (sometimes with petiole

base) the often fleshy sometimes ligneous stems or

caudex.

Stipules caducous or at least not scarring a ligneous stem

or a fleshy caudex or rhizome; stems usually procum-

bent toward base where often rooting, not rarely elon-

gating and clambering.

Leaflets narrower at emarginate apex than at middle.

Leaflets obliquely oblong-elliptic, several cm. long.

0. distincta.

Leaflets oblong-obovate, cuneate at base, 1.5-2.5 cm.

long.

Stipules 1 mm. long, caducous, or longer, persisting,

completely free, not at all membranous.

Stems 4-5 mm. thick below; styles puberulent.

Sepals to 5 mm. long; stamens glabrous.

0. villosula.

Sepals to 7 mm. long; stamens as styles hirsutu-

lous 0. San-Miguelii.

Stems about 2-3 mm. thick; stamens and styles

glabrous 0. tabaconasensis.

Stipules membranous, about 2 mm. long, deciduous.

0. mollis.

Leaflets usually much wider (at least as wide) at lunate

to truncate apex than at middle.

Sepals about 6 mm. long; filaments papillose to

puberulent 0. Ortgiesii.

Sepals about 4 mm. long; filaments glabrous.

0. rigidicaulis.

Stipules or (and) petiole bases persisting or scarring a

fleshy or infrequently ligneous stem or rhizome (or

both), the latter rarely bulbiform; peduncles and leaves

apically crowded (the latter deciduous) on the short or

long very erect succulent stems (usually) or these

FLORA OF PERU 551

lacking, leaves thus pseudobasal from more or less

enlarged fleshy to ligneous stems or roots.

Sepals ovate-lanceolate to ligulate or narrower, usually

acute; flowers early more or less umbellate but the

inflorescence often soon more or less bifurcate,

especially in fruit.

Leaflets densely pubescent both sides; stems densely

scaly 0. cinerea.

Leaflets glabrous or only lightly pubescent, at least

above (0. polyantha, unique in Peru by its red

umbellate flowers).

Stems clearly ligneous nearly to apex, densely

scarred; petioles typically broadly dilated.

(0. Weberbaueri), 0. Herrerae.

Stems at least typically succulent unless at or toward

the base, but often firm, less densely scarred;

petioles more or less dilated especially sometimes

(0. peduncularis, 0. Weberbaueri, 0. marcapa-

tensis).

Filaments glabrous; petioles broadly dilated.

Sepals 3.5-4 mm. long, obtuse; cyme umbelli-

form 0. coralleoides.

Sepals to 6 mm. long, narrowed to obtusish

tip; cyme shortly forked. . .0. Weberbaueri.

Filaments more or less pubescent; petioles often

slender, filaments not described in 0. Staf-

fordiana, 0. polyrhiza.

Sepals described (by Knuth) as 4-5 mm. long,

petals 10-13 mm. long.

Stems elongating; cymes usually more or less

bifurcate. 0. ptychoclada, 0. peduncularis.

Stems a few cm. high from thick rhizome;

flowers few, umbellate or nearly.

0. Staffordiana, 0. modestior.

Sepals described as 6-8 mm. long, petals (12)

15-20 mm. long (ex Diels, 0. ptyochoclada

would go here).

Petals 12 mm. long; sepals obtuse or rounded.

0. polyrhiza.

552 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Petals 15-20 mm. long; sepals acute.

0. marcapatensis, 0. paucartambensis.

Sepals, at least the outer, broad and more or less dilated

at base, especially in fruit.

Rhizome or caudex becoming stout, apically brown-

stipulate and in age often producing a scarred

stem or stems, simple or shortly branched.

Bracts small, rarely more than 3.5 mm. long, in any

case slender; inflorescence various.

Flowers usually many, typically 15-20 mm. long.

0. solarensis, 0. megalorrhiza.

Flowers few, sometimes 1-2, rather small.

Petioles elongate. . .0. pachyrrhiza, 0. polyrhiza.

Petioles and peduncles shorter than 3 cm.

0. juninensis.

Bracts 5-7 mm. long, membranous-foliate; flowers

early subumbellate and large, later ones some-

times small 0. sepalosa.

Rhizome small, bulbiform; diminutive plants with

small flowers in umbelliform cymes.

Outer sepals deltoid, wider than long. . . .0. lomana.

Outer sepals triangular-rotund, at least as long as

wide 0. bulbigera.

Acaulescent species usually bulbous, at least clearly so if the flowers

are umbellate, the flowers solitary otherwise. (The introduced

0. pes-caprae and 0. chasquiensis only found once, are for

convenience omitted from key, both acaulescent.)

Peduncles regularly 1-flowered from a bulb or rarely from a

bulbous caudex clothed with petiolar remains; alpine species

often near limit of flowering plants.

Stipules and leaf-bases forming a caudex that is apparently,

not clearly, a bulb 0. eriolepis.

Stipules and leaf -bases not obviously persisting, the plant clearly

from a bulb.

Sepals ecallosed at the often purplish tip; small plants with

peduncles shorter or little longer than the leaves.

0. Philippii, 0. minima.

Sepals with 2 or more calli, sometimes faint, at tip; peduncles

about as long or much longer than leaves.

FLORA OF PERU 553

Stamens apparently glabrous; peduncles and petioles sub-

equal 0. oreocharis.

Stamens pubescent; peduncles usually elongate (1-flowered

plants of 0. moqueguensis, but filaments glabrous).

0. pseudolobata, 0. punensis.

Peduncles terminating in an umbel that is rarely reduced to

1 or 2 flowers, but obviously from a bulb; plants often from

high altitudes but subalpine, rarely lowland.

Leaflets more or less emarginate or parted, little if at all wider

than long or flowers purple.

Bulb simple, the brown outer scales usually more or less

lanate.

Leaflets parted about medially and the lobes spreading;

sepals about 6 mm. long.

Lobes oblongish, usually longer than wide, widely

divaricate 0. bipartite,.

Lobes roundish, little longer than wide, somewhat

divaricate 0. arenaria.

Leaflets approximately obcordate or wider than long and

emarginate or the lobes suberect; sepals sometimes

only 3.5-5 mm. long.

Filaments and sometimes styles puberulent; flowers

several to many.

Sepals typically 3.5-5 mm. long; styles glabrous.

0. latifolia.

Sepals not described; styles as filaments puberulent.

0. carminea.

Sepals in type 6-8 mm. long; styles and filaments

puberulent 0. elegans.

Filaments and apparently also styles glabrous; sepals

6 mm. long.

Flowers white, usually several (3-8) . . .0. acromelaena.

Flowers tinted, few (1-4) 0. moqueguensis.

Bulb compound, that is, composed of many small bulbs.

0. Martiana.

Leaflets broadly deltoid, twice wider than long at subtruncate

apex; flowers white; bulb with thick tubercles.

0. Regnellii.

554 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Oxalis acromelaena Diels, Bot. Jahrb. 37: 423. 1906; 260.

Acaulescent, 7-14 cm. high, from a rotund bulb about 12 mm.

in diameter with gray lanceolate scales; petioles few (3-8), to 8 cm.

long, rather lax, glabrous or at insertion of the leaflets subglabrous;

leaflets glabrous, glaucous-green, broadly triangular, 5-8 mm. long,

often about twice as broad or broader, very shallowly incised;

peduncles only 1-3, 3-8-flowered, glabrous as pedicels and calyces,

5-11 cm. long, with broadly ovate membranous somewhat erose

bracts 2 mm. long; pedicels filiform, 1-1.5 cm. long; flowers 12-14

mm. long, narrowly campanulate from base, the lanceolate sepals

deep purple ("nigro-maculata") at the acute tip, (4-4.5) 6 mm. long

(Diels), the clawed white petals rounded at apex; glabrous. — Near

0. elegans HBK. and 0. articulata Sav. but flowers white, sepals less

acuminate, different leaflets (Diels). 0. pazensis (Rusby) Knuth,

259, has somewhat ciliate leaflets 6-12 mm. long, 10-15 mm. wide,

2-4 mm. incised, flowers only 2-3, about 2 cm. long on pedicels

1.5-2.5 cm. long, sepals about 7 mm. long, obtuse, 5-green-nerved;

0. Buchtienii (Rusby) Knuth, 260, also from as near as La Paz, has

calyx 6 mm. long, the apical tubercle large, red in herb., the leaflets

deeply incised, 8-12 mm. long, 9-15 mm. wide, the flowers rather

many on pedicels to 1 cm. long. But all three plants seem basically

to belong to 0. latifolia sens. lat.

Junin: Stony places, 3,100 meters, Tarma, (Weberbauer 2390,

type); 176.

Oxalis adpressa Knuth, Pflanzenreich IV. 130: 160. 1930. 0.

parvifolia DC. var. pluriflora Knuth, Notizbl. Bot. Gart. Berlin 7:

301. 1919.

Densely leafy annual with many tortuous ascending or pro-

cumbent stems 1-1.5 mm. thick, branched at base, more or less

spreading pilose, often with short branchlets in the leaf -axils;

leaflets sessile, obcordate-reniform, to 10 mm. wide, 5-6 mm. long,

usually one-third to two-fifths incised, with rounded lobes, broadly

cuneate at base, sparsely pubescent both sides, the margins rather

rigidly ciliate; petioles to 2 cm. long, sparsely hirsute; stipules

adnate, 2 mm. long; peduncles also from lower axils, to 2 cm. long,

hirsute or glabrate, with acute linear bracts to 4 mm. long, the umbel

1-5-flowered, its pedicels to 6 mm. long, hirsutulous as the calyx,

this with oblong-lanceolate obtusish sepals, often purplish and 3.5

mm. long; petals pale yellow, about twice as long; stamens hirsutu-

lous, the longer exserted from calyx; ovary hirsute, the capsule 9 mm.

FLORA OF PERU 555

long, 2 mm. thick, with brown rugose depressed seeds 0.75 mm. in

diameter. — Species very variable when small, the peduncles and

pedicels short, 1-flowered, when elongate these well-developed

(Knuth). Unknown to me; may be a mixture or otherwise a variety

of 0. parvifolia as first designated. Bolivian species said to be

related are 0. soldanelliflora Knuth, 160, and 0. Bangii Rusby, 161,

the former with subfiliform creeping stems, deep yellow purple-

veined petals 15 mm. long, the letter with also large flowers but

reddish-pilose. Type from northern Bolivia.

Peru: (Undoubtedly). Ecuador; Bolivia.

Oxalis amazonica Prog, in Mart. Fl. Bras. 12, pt. 2: 503. 1877;

63. Acetosella amazonica (Prog.) Kuntze, Rev. Gen. 1: 92. 1891.

Erect or suberect annual, the firm slender stems strict and

minutely appressed puberulent above where moderately leafy, the

nearly filiform subopposite or fasciculate petioles about 3.5 cm. long;

leaflets glabrous, broadly oblong-ovate or elliptic, rounded both ends,

2-3 cm. long, 1.5 cm. wide, the medial with petiolule 5-8 mm. long;

peduncles often 3-4 in an apical fascicle, erect, 3-5 cm. long, bifur-

cate, the branches in fruit about 1 cm. long, usually 4-6-flowered;

bracts 1 mm. long, subulate; pedicels 2-3 mm. long, recurved in

fruit; sepals lanceolate, acute, minutely puberulent, 4 mm. long, the

narrow-based obovate violet petals about twice as long; filaments

glabrous, the longer with one lateral medial tooth; styles hirsutulous;

capsules elongate-ovate, acute, 7 mm. long, with 3 rugose seeds in

each cell.— F.M. Neg. 32441.

San Martin: Villa Nova and Tocache, Poeppig, type.

Oxalis arenaria Bert. Merc. Chil. 16: 739, 740, ex Colla, Mem.

Ace. Torino 37: 48. 1834; 265. Acetosella arenaria (Bert.) Kuntze,

Rev. Gen. 1: 91. 1891. Sassia tinctoria Molina, Sagg. Nat. Chile

146. 1782, not Poepp. ex Prog, in Mart. Fl. Bras. 12, pt. 2: 479.

1877. 0. pseudoarenaria Knuth, Repert. Sp. Nov. 38: 196. 1935?

Acaulescent from a rotund-oblong bulb to 2 cm. long, the ovate-

lanceolate acutely acuminate scales little or not at all lanate within,

1.5 cm. long; leaves only 3-6, on glabrous or sparsely pilose petioles

6-11 cm. long, the broadly obcordate leaflets usually pilose at least

beneath, in Peru often glabrate, to 2 cm. long, 17 mm. wide, incised

two-fifths or nearly to the middle with acute sinus, the lobes rounded;

peduncles usually 1-several dm. long, subglabrous with simple 2-7-

flowered umbel; bracts deltoid-lanceolate or narrower, acutely

556 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

acuminate, to 3 mm. long; pedicels often lax, subfiliform, to 4 cm.

long; sepals linear-lanceolate, obtuse, or acute, thick-red-callose at

tip, finally 6-7 mm. long; corolla purple, four times longer than calyx;

styles densely puberulent. — 0. pseudoarenaria if recognizable may

include all Peruvian plants, but ex char, there appears to be no basic

distinction; less pilose, bracts nearly linear, acute, 4.5-5 mm. long,

corolla 3 times longer are all variations that may be states of develop-

ment; for example, it is noticeable the calyx is longer as the flower

matures. 0. longiflora L., cited here and also under 0. Acetosella L.

by Knuth, is apparently from type locality, "Virginia," more likely

referable to the latter. F.M. Neg. 19189.

Junin: Huancayo, Soukup 1893 (det. Leonard). — Cuzco: Sailla,

Quebrada de Oropeza, (Herrera 2688}. Hacienda Churu, Paucar-

tambo Valley, 3,500 meters, Herrera 1398 pt. Yucay, 3,000 meters,

Valle del Urubamba, (Herrera 2275)', 1369 (det. Knuth). — Are-

quipa: Raimondi (det. Knuth). Prov. de Anta, Herrera 3632.

Huascao, Herrera 3109a. Near Cuzco, 3,700 meters, (Stafford,

type, 0. pseudoarenaria). Bolivia; Chile; Argentina; Paraguay.

Oxalis Barrelled L. Sp. PI. ed. 2. 624. 1762; 65. Lotoxalis

Barrelieri Small, N. Am. Fl. 25, pt. 1: 49. 1907.

Becoming a dm. to several dm. tall, glabrous or glabrescent or

the leaflets sparsely pubescent beneath, these ovate-oblong-lanceo-

late, usually 1.5-4.5 cm. long, obtuse or acute, the lateral evidently

smaller; peduncles mostly longer than the petioles, the finally

glabrous pedicels 3-4 mm. long; sepals 3-3.5 mm. long, the outer

oblong-lanceolate, glabrous or barbate at tip; petals 6-9 mm. long;

longer filaments appendaged above the middle and there puberulent;

styles pubescent; capsules broadly oblong, 5-7 mm. long. — After

Small.

Peru: (Possibly). West Indies and northern South America.

Oxalis bartolomensis Knuth, Notizbl. Bot. Gart. Berlin 7: 302.

1919; 124.

Slender glabrous erect -simple stemmed annual to a dm. high,

leafy above the middle; petioles often nearly 3.5 cm. long, the

middle petiolule to 2.5 mm. long, the lateral 1 mm. long; leaflets

elongate-obcordate, often 11 mm. long, 7 mm. wide near the ob-

tusely incised tip, the lobes rounded; lateral leaflets mostly oblique,

the outer lobule reduced; peduncle 5-7.5 cm. long, without the

flowering branches 3-4 cm. long; bracts linear, membranous, obtuse,

FLORA OF PERU 557

1 mm. long; pedicels to 2.5 mm. long, often persisting; sepals to 4

mm. long, lanceolate acute; petals yellow or pale, 8.5-10 mm. long,

narrowly lanceolate, rounded; filaments described as glabrous, as

the styles shorter than sepals. — Filaments pubescent in Pennell

plant; probably there is an earlier name for this in the vicinity of

0. frutescens L., 73, or 0. euphorbioides St. Hil., 50.

Lima: In cactus-shrub formation, 1,500 meters, San Bartolome",

Weberbauer 5296, type. Quive, Pennell 11+301 (det. Knuth). Be-

tween Ambar and Huacho, 1,200 meters, Stork 11469 (det. Johnston,

0. lotoides}. — Moquehua: Mount Estuquina, 1,600 meters, Weber-

bauer 7446 (det. Knuth).

Oxalis bipartita St. Hil. Fl. Bras. 1: 125. pi. 25. 1825; 263.

Acetosella bipartita (St. Hil.) Kuntze, Rev. Gen. 1: 92. 1891.

Glabrous except petiole tips or the inflorescence obscurely

puberulent, acaulescent, 1-2 dm. high from a rotund bulb often

8 mm. in diameter, simple and nearly without scales; leaves 5-20,

lax, the petioles 5 (-15) cm. long with a fascicle of brown trichomes

at the juncture with the leaflets, these green, 12-20 mm. long, bilobed

for more than half their length, the lobes divaricate, often 1 cm. long,

2 mm. wide, oblong-linear; peduncles few (1-4), often 5-7 cm. long,

rarely 26 cm. long, the umbels often irregular and 2-12-flowered with

many narrow bracts 3 mm. long, red-callose apically as the oblong-

lanceolate or linear acutish sepals, these 5-6 mm. long; pedicels

2- nearly 3 cm. long; corolla conic-campanulate, at least four times

longer than the calyx, violet, the petals more or less retuse; larger

stamens nearly two times longer than sepals.

Cuzco: Hacienda Churu, Paucartambo Valley, 3,500 meters,

Herrera (det. Knuth). Cuzco, 3,000-3,600 meters, Herrera (det.

Knuth). Calca, Vargas 164. Prov. de Anta, Herrera 3634.—

Apurimac: Upper Marino Valley, Goodspeed Exped. 10637. Prov.

Abancay, 2,800 meters, Vargas 9612. Prov. Grau, Vargas 9782.

Uruguay; southern Brazil.

Oxalis breviramulosa Rusby, Mem. Torrey Club 3, pt. 3: 13.

1893; 165. 0. guaquiensis Knuth, Meded. Rijks Herb. Leiden 27:

63. 1915, fide Knuth.

Suffruticose, about 5 cm. high, the procumbent or ascending

stems to 10 cm. long and 1.5 mm. thick, densely foliose, subligneous

and subglabrous below, densely glandular-subviscous above; leaves

congested on the very short branchlets, the sessile reniform glaucous

558 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

rather fleshy leaflets incised below the middle, 3 mm. wide, half as

long, glabrous above, somewhat lanate beneath; petioles 5 mm. long,

nearly filiform, appressed pubescent, the stipules nearly completely

adnate; flowers solitary on pedicels only 1-2 mm. long; sepals ovate,

obtuse, glabrous, often purple-margined, 2-3 mm. long, the yellow

petals to 9 mm. long, broadly cuneate, more or less retuse; larger

stamens densely ciliate, much shorter than sepals. — Type of 0. gua-

quiensis from stony places on the Bolivian side of Lake Titicaca.

0. bisfracta Turcz., 142, of Bolivia, is apparently related.

Puno(?): Probably about Lake Titicaca. Bolivia.

Oxalis bulbigera Knuth, Notizbl. Bot. Gart. Berlin 7: 306.

1919; 200. 0. microbolba Knuth, Repert. Sp. Nov. 23: 280. 1927.

Acaulescent, 4-5 cm. high from a bulbous root 7 mm. long, 4-5

mm. thick; leaves few to many (about 20), rather fleshy, the equal

leaflets triangular-obcordate, 5-10 mm. long and broad, with very

wide 1 mm. deep sinus, glabrous above, arachnoid beneath; peduncles

sparsely pilose, little if at all exceeding the leaves, bearing a 4-5

(-10) -flowered congested umbelliform cyme with linear obtuse pilose

bracts 2-3 mm. long, the arachnoid pedicels only 1-3 mm. long;

inner sepals lingulate as long as the outer broadly ovate ones, these

2.5 mm. wide and long or 3.5-4.5 mm. in fruit and equaling the

capsule, this with 3-4 seeds in each cell; corolla about 5 mm. long,

yellow.

Lima: Mount Morro, near Chorillos, Weberbauer 5688, type.

Mill by Barranco, lower edge of lomas, Weberbauer 5699 (root not

enlarged, first year, ace. to coll.). Lee of large rocks, north of

Barranca, Worth & Morrison 9112 (det. Johnston). Common over

plains south of Huacho, Goodspeed Exped. 9235 (det. Johnston).

Loma Ancon, 800 meters, Soukup 2126. Near Lima, Rose 19472

(type, 0. microbolba) ; also 18574. Stony slopes seaside hills, Chorillos,

5854 (det. Knuth, 0. microbolba').

Oxalis carminea Knuth, Notizbl. Bot. Gart. Berlin 7: 312.

1919; 261.

Acaulescent, glabrous, from a rotund bulb about 1 cm. in diame-

ter, its lanceolate very acute scales rarely lanate within, to 1 cm.

long; leaves 3-8, the petioles about 4 cm. long, the leaflets broadly

obcordate, often 1.5 cm. wide, 1 cm. long, sometimes larger, one-

fourth to one- third incised with rather acute sinus, the lobes rounded;

peduncles 7-13 cm. long, about 5-flowered, the more or less irregu-

FLORA OF PERU 559

larly ovate bracts with wide membranous margins; pedicels 5-12

mm. long, often refracted; sepals lanceolate, acutish, with two red

thick calli; corolla pale red, three times longer than calyx; stamens

and styles minutely puberulent. — Much like 0. araucana Reiche but

lateral nerves of the leaves obsolete (Knuth) ; apparently belongs to

0. latifolia, sens, lat., but type not seen.

Lima: Below Matucana, 220 meters on stony outcrops, (Weber-

bauer 5205, type).

Oxalis chasquiensis Knuth, Pflanzenreich IV. 130: 437. 1930.

Acaulescent with about 8 leaves and 2 or 3 peduncles from the

neck of a horizontal rhizome 3-5 mm. thick, the roots long, few;

petioles 4-10 cm. long, sparsely pilose, the stipules lanceolate, 4 mm.

long, acute, hirsute, in part adnate; leaflets sessile, equal, more or

less appressed pilose both sides of the margins, densely ciliate and

narrowly marginate, obcordate, obtusely 2 mm. incised, to 12 mm.

long and 10 mm. broad; peduncles 6-8 cm. long (without the 2

flowers), hirsute as pedicels, these filiform, 7-17 mm. long, and

calyces; bracts subulate, very acute, 4 mm. long; sepals narrowly

lanceolate, acute, 8 mm. long; petals yellow, 14 mm. long, long-

clawed; stamens and styles puberulent. — Allied to the Argentinian

0. lasiopetala Zucc. with violet flowers; and thus also to 0. Regnellii

Miq. which compare.

Huanuco: Chasqui, 3288, type.

Oxalis cinerea Zucc. Denkschr. Akad. Muench. 9: 156. 1823-

24; 194. 0. Haenkeana Zucc. Abh. Math. Nat. Cl. Akad. Muench.

1: 233. 1829-30, not Spreng. 1827.

Stems 2.5-5 cm. high, thick, simple or with few basal branches,

densely imbricate-scaly with pubescent rudiments of petioles; leaves

many at stem tips the sessile leaflets acutely incised with rounded

lobes, equal, appressed sericeous villous both sides; petioles erect-

spreading, 4-5 cm. long, villous, the base dilated-amplexicaul ;

peduncles strict, 3-4-flowered with filiform pedicels articulate a

little above the base, villous as the minute subulate bracts; sepals

lanceolate, acute, hirsute; corolla large, sordid yellow with purple

veins, three times longer than the calyx; filaments and styles hirsute.

— The locality "Guanocco" is probably a misreading for Huanuco.

The var. peruviana Walp. Nov. Act. Acad. Leop.-Carol. 19: Suppl.

1: 319. 1843 of 0. Haenkeana has more obtuse leaflets, pedicels two

times shorter than calyx and, as the species was as unknown to

560 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Knuth as to me, it is quite possibly a pubescent state of 0. paucar-

tambensis or 0. peduncularis, sens. lat.

Huanuco: (Haenke in herb. Sternberg, type). — Cuzco: Sicuani,

3,550 meters, Prov. de Canchis, (Hicken).

Oxalis coralleoides Knuth, Pflanzenreich IV. 130: 433. 1930.

Stems many from neck of a branched rhizome, rather fleshy,

glabrous, 5-7 cm. high, leafy above, below clothed with petiolar

bases 2-4 mm. long; petioles to 3 cm. long, broadly dilated, glabrous,

the lanceolate acute stipules ciliate, 4 mm. long; leaflets sessile,

triquetrous-obcordate, 5 mm. wide, 6 mm. long, fleshy, glabrous

above, ciliate beneath and on the often reddish margin, 1-2 mm.

incised with obtuse sinus; peduncles 6-8 cm. long, inflated below

the middle, glabrous, the nearly regular umbel 3-6-flowered; bracts

subulate, sparsely ciliate, 3-4 mm. long; pedicels 6-10 mm. long,

filiform, ciliolate; sepals rather ovate, obtusish, often purplish,

scabriusculous with minute hyaline trichomes; petals to 12 mm. long,

red-veined, yellow, obovate; stamens glabrous; seeds 2-3 in a cell. —

Allied to 0. Herrerae Knuth, fide author. I doubt if the Cuzco

specimen belongs here. The Indians add it to the drink Chicha to

induce sleep (Herrera).

Junin: Tarma, 4,000 meters, 1060, type. — Cuzco: Machu-picchu

to Ollantaytambo, 2,100-2,800 meters, (Herrera 2085). "Macha-

macha" (Herrera).

Oxalis Cornell! Anderss. Vet. Akad. Handl. Stockh. 1853: 246.

1855; 65. 0. sepium St. Hil. var. glandulifera Norlind, Arkiv. Bot.

20A, pt. 4: 9. 1926. 0. ramulosa Knuth, Notizbl. Bot. Gart. Berlin

7: 290. 1919. 0. Haughtii Knuth, Repert. Sp. Nov. 40: 291. 1936.

0. peruviana [Standley] Knuth, Pflanzenreich IV. 130: 67. 1930.

Often flowering as a slender annual, becoming perennial and

subligneous below, usually erect or strict, to several dm. tall or in

some habitats prostrate with slender vine-like stems, glabrescent

but more or less pubescent especially the younger parts with acicular

sometimes viscid trichomes; petioles 1-4 cm. long; medial petiolule

3-8 mm. long; leaflets broadly obovate or nearly rotund, rounded

or slightly retuse apically, usually about 1.5 cm. long, 1 cm. broad,

often smaller and in vigorous plants to 3.5 cm. long, 3 cm. wide,

more or less glaucous, often firm; peduncles upper-axillary and

terminal, ascending or erect, 2.5-6 cm. long, the bifurcate cyme

sometimes 7 cm. long; bracts linear, acute, 1.5-2 mm. long; pedicels

FLORA OF PERU 561

1-2.5 mm. long; sepals lanceolate, 5-nerved, glabrous, 4-5 mm. long,

the orange-yellow obovate rounded petals about twice as long;

larger stamens subequal, the filaments ciliate, the much shorter

smaller stamens glabrous, half as long as the ciliate styles; capsules

pendulous, about 7 mm. long, 5 mm. thick, with several rugose

seeds in each cell. — This follows the interpretation of Svenson, Amer.

Journ. Bot. 33: 455. 1946, who, besides the synonyms above (except

0. Haughtii) includes 0. Barrellieri sensu Knuth, 65, in part which,

according to him, is a "quite different purple-flowered species."

0. Haughtii seems to me to be the vernal state. Apparently 0.

amazonica Prog., 63, sensu Knuth, likewise is referable here. —

Illustrated, Svenson, I.e. 456.

Piura: Toward Hacienda Nomala in half shrub formation, 200

meters, Weberbauer 5956 (type, 0. ramulosa). Paita, Pennell 14816

(det. Knuth, 0. peruviana and 0. sepium, var.). Parinas Valley,

Haught 281 (type, 0. Haughtii). Near La Brea, (Haught & Svenson

11590, det. Svenson). — Huancavelica: Somate, Townsend 827 (type,

0. peruviana). Talara, Haught 46; 282. — Libertad: Entirely pros-

trate with slender stems, Chicama Valley, Smyth 37 (det. Leonard,

0. peruviana [Standl.] Knuth). Ecuador; Galapagos; Brazil?

Oxalis corniculata L. Sp. PL 435. 1753; 146. Xanthoxalis

corniculata (L.) Small, Fl. S. E. U. S. 667. 1903.

Loosely pilose to nearly glabrous, rhizomatous, diffuse to pros-

trate, the usually numerous very foliose stems often congested at

base and 1-3 dm. long, commonly more or less rooting; leaflets

broadly cuneate-obcordate, to one-third incised, pilose, especially

beneath and marginally, small, to about a cm. wide and long, or

larger, the petioles much longer, the 1-6-flowered peduncles 1-7 cm.

long; bracts acute to acuminate, 1-5 mm. long; pedicels short or

often about as long as the long peduncles, hirsutulous-puberulent

as the calyces or glabrate, the sepals ecallose, about 5 mm. long;

corolla 8-10 mm. long, pale yellow; larger filaments glabrous;

capsules erect, slender, 12-15 mm. long, appressed pubescent.—

Widely distributed around the world in both temperate and sub-

tropical regions, including Ecuador and Bolivia. Similar and prob-

ably occurring is 0. stricta L., 143, stems erect or decumbent, corolla

5-10 mm. long, pubescence appressed, leaflets nearly glabrous,

bracts 1-1.5 mm. long, following Knuth's interpretation of these

variable species. But see also Wiegand, Rhodora 27: 113, 133. 1925.

There is a purplish colored form (or distinct species?), 0. corniculata

L. var. atropurpurea Planchon, Fl. Serres 12: 47. 1857, which a

562 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

contemporary without explanation has changed to a variety of

0. europaea Jord. under Xanthoxalis, the former, fide Knuth, 0.

stricta L. var. europaea (Jord.) Knuth. But cf. Wilmott, Journ.

Bot. 53: 172-174. 1915. There seem to be a number of variants,

maybe distinct but not surely in Peru, for example, 0. filijormis

HBK., 169, Ecuador, and many others, even in different sections

by the monographer, maybe correctly, certainly not clearly.

Lima: Irrigation ditch bank, 507 (det. Knuth). — Huanuco: At

Huanuco, Soukup 2230 (distr. 0. stricta?). — Cuzco: Hacienda Churu,

3,500 meters, Prov. de Paucartambo, (Herrera 1382). Nearly

cosmopolitan. "Vinagrillo."

Oxalis cuzcensis Knuth, Notizbl. Bot. Gart. Berlin 7: 303.

1919; 124. 0. ollantaytambensis Knuth, Repert. Sp. Nov. 23: 142.

1926; 124.

Small rather succulent annual normally becoming a dm. or so

high, early abundantly leafy and floriferous nearly from the base,

glabrous or very sparsely puberulent unless in variant; petioles to

2 cm. long with membranous acute stipules to 3 mm. long, long-

adnate; leaflets obcordate, exactly cuneate at base, to 2 mm. incised

apically, pale green, to 11 mm. long, often smaller, nearly as wide;

upper peduncles to 4 cm. long, cymosely parted and often bifurcate,

usually (in type "cymose, globosely stacked at summit"), 8-14-

flowered; bracts 1-2 mm. long, very acute; pedicels filiform, often

nodding, 3-10 mm. long; sepals narrowly lanceolate, obtuse, nearly

membranous, often violet tinged and with a fascicle of trichomes at

tip, the yellow lanceolate-cuneate petals 6 mm. long; styles as long

as sepals, longer than stamens, the often slightly exserted capsule

with few seeds. — In spite of the name of the apparent synonym a

collection from Bolivia is its type. More material may prove me

wrong in regarding 0. cuzcensis as based on undeveloped plants,

probably so small because starved; however some peduncles are

1-flowered. Except as indicated determinations by Knuth, as 0.

ollantaytambensis. The Argentinian 0. membranifolia Knuth, Repert.

Sp. Nov. 40: 291. 1936, allied by the author to the Chilean 0. rosea

Jacq., 126, and to this, has yellow petals two and a half times longer

than the 5-7 mm. long sepals, minor differences that may suggest

variability within 0. rosea and wider distribution, sens. lat. Here

would be sought 0. yungasensis Rusby, 159, hirsute, and apparently

approached by Herrera 2997a.

Lima: Shrubby canyon, 3,000 meters, 3096. — Junin: In rocks,

Yanahuanca, 1216. Huancayo, Soukup 1896 (det. Leonard, 0. ol-

FLORA OF PERU 563

lantaytambensis). — Huancavelica: West of Huanta in gravelly sand,

Stork & Horton 10810 (det. Standley, 0. ollantaytambensis). —

Cuzco: In rock- wall, San Sebastian, Pennell 13633. Valle del

Apurimac, Hen era. Hacienda Pfuyucalla, 3,400 meters, Paucar-

tambo Valley, Hen era 2997 a (det. Knuth, 0. pygmaea). Below

Tres Cruces, shrubby grass steppe, 3,500 meters, Weberbauer 6921,

type. Valle del Urubamba, Ollantaytambo, 3,000 meters, Cook &

Gilbert 350; 354- — Puno: Aziruni, Soukup 1038. Bolivia; Colombia

(?). "Oca-oca."

Oxalis distincta Knuth, Repert. Sp. Nov. 40: 292. 1936.

A succulent herb with subhirsute stems, petioles and peduncles,

the first 3-4 dm. high, erect, leafy from near the 5 mm. thick base,

the second to 8 cm. long, 1.5 mm. thick, the last at least as long to

even 10.5 cm. and bibranching apically into an umbelliform cyme

2 cm. across; stipules linear, 1-2 mm. long, often persisting; leaflets

sessile, subglabrous or pilose especially marginally, rhombic-oblong,

5.5 cm. long, 2.5 cm. wide, subequal, retuse with mucro 1 mm. long;

lower pedicels to 1 cm. long; bracts subulate, very acute, 4 mm. long,

puberulent, persisting; sepals narrowly lanceolate, acute, to 8 mm.

long, shortly pilose, the yellowish cuneate-dilated petals to nearly

twice as long; stamens pubescent and with styles about as long as

sepals. — Section Clematodes and in form of leaves like 0. subintegra

Knuth, but obviously diverse by habit and size of all parts (Knuth).

Junin: Yapas, about 1,500 meters, in thick wood along Pichis

Trail, Killip & Smith 25586, type.

Oxalis dolichopoda Diels, Bot. Jahrb. 37: 423. 1906; 179.

Stems ligneous, simple or rarely forked, 4-10 dm. high, 3-4 mm.

thick, densely foliate above the middle, dark brown and glabrous

below, reddish tomentose-villous above; leaflets pilose but green

above, fuscous sericeous tomentose beneath, nicely obovate, mostly

8-14 mm. long, 5-12 mm. wide, rather acutely to one-fourth incised,

without punctae or calli; petioles 1.5-2 cm. long, reddish tomentulose

and with 4-6 mm. long stipules; peduncles many above, 3 cm. long,

reddish puberulent as pedicels (1 cm. long) and sepals, these 6-7 mm.

long, linear-lanceolate, obtuse or subrotund at tip; bracts acute,

3 mm. long; flowers about 14 mm. long, campanulate, yellow, the

petals retuse-rounded; larger stamens very sparsely pilose or glabrate,

little longer than sepals; styles glabrous.— This may be the older

ligneous state of another species but, as the author observed, ap-

parently distinct in habit, form of leaves and in indument.

564 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Huanuco: Southwest of Monzon, 2,200 meters, (Weberbauer

3707, type)', among low bushes in sphagnum, Playapampa, 3,000

meters, 4484 (det. Knuth); 94; 255. — Cuzco: Cerro de Cusilluyoc,

"Pillahuata," 3,000 meters, Pennell 14107 (det. Knuth). In woods,

"Pillahuata" to Acanacu, 2,600 meters, West 7066 (det. Johnston).

Bolivia.

Oxalis elegans HBK. Nov. Gen. & Sp. 5: 182. pi. 466. 1821;

252.

Acaulescent from a bulb, its scales very long-lanate; glabrous

(or nearly) even to the lanceolate (6-8 mm. long) sepals but these

marked at the acuminate tip by 2 lineate glands; peduncles 5-15 cm.

long, 2-6-flowered, the filiform pedicels 16-28 mm. long; leaflets

subsessile, broadly subrotund-obovate, subtruncate, rather acutely

emarginate, subequal, membranous, violet (glandular, Progel)

beneath, 12 mm. wide, 8 mm. long, the lateral a little oblique;

petioles filiform, nearly glabrous, 7.5 cm. long; flowers subumbellate,

about 2 cm. long, violet with black spots in the throat; bracts

lanceolate, 3 mm. long; petals rounded apically; longer filaments

puberulent above; styles pubescent. — After HBK., the type from

Loxa. Perhaps 0. atroglandulosa, here included in 0. latifolia,

belongs here or is distinct, fide Knuth, by its dark callose instead of

reddish sepal tips and many-flowered umbels, scarcely impressive

specific characters, but type not seen in comparison. All the speci-

mens cited under 0. latifolia for expediency, in spite of the larger

leaflets, probably ex char, belong here, and indeed, the Wilkes Exped.

collection was so named by Gray.

Peru: Probably. Ecuador.

Oxalis eriolepis Wedd. Chlor. And. 2: 290. 1857; 246.

Bulbous acaulescent, the oblong reddish bulb scales with lanate

margins, the stipules and petiole bases persisting; petioles elongate;

leaflets obcordate-bilobed, 5 mm. long, at least twice as wide, sessile,

glabrous but cellulose-lacunose beneath; stipules connate to tips,

lanate on margins; peduncles 1-flowered, glabrous, bibracteolate

below the summit; sepals long or lanceolate, often acute, 5 mm. long,

violet margined, more or less red callose at tip, glabrous or puberu-

lent, less than half as long as the purple or white petals. — The collec-

tion mentioned by Weberbauer, 221, is cited by Knuth as 0. Philippii

but the reference is given under this name. Weddell cites first

Jameson from near Quito, not seen; no bulb on the Mandon photo.

Possibly material is mixed. Apparently 0. pachyrrhizus and 0.

FLORA OF PERU 565

juninensis are allied, and maybe 0. chasquiensis, not seen. It may

well be 0. minima as interpreted by me at least in part. F.M.

Neg. 26311 (Mandon 842).

Cuzco: Sicuani, 3,550 meters, Prov. de Canchis, (Hickeri). Prov.

del Cercado, (Hen era}. — Puno: Between Limbani and Aqualani,

2,700 meters, Vargas 1286? Bolivia; Ecuador. "Occa-occa."

Oxalis fritillariiformis Knuth, Notizbl. Bot. Gart. Berlin 7:

297. 1919; 105.

Erect, 4 dm. high, the simple stem to 3 mm. thick, puberulent

below, more or less densely tomentose-subhirsute above, where

densely clothed with leaves and peduncles but the former not entirely

lacking toward the base; leaves mostly reduced to phylloidea, 10-13

cm. long, 2.5 cm. wide, gradually narrowed to base into a winged

pseudo-petiole 1.5 cm. long, more or less abruptly contracted at tip

and often acuminate, with acumen 3 mm. broad at base, the very

narrow margin minutely but densely ciliolate, often apically bearing

3 leaflets 8 mm. long, half as wide, ovate, obtuse, on petiolules to

1 mm. long; peduncles filiform, 3-4 cm. long, erect, racemosely

bifurcate at tip, the branches 1-1.5 cm. long, densely marked by

pedicel scars, 8-15-flowered; bracts subulate, 1 mm. long; pedicels

capillary, 3 (4) mm. long; sepals lanceolate, acutely acuminate,

puberulent, to 3 mm. long, the yellow petals to three times longer,

clawed, oblong-lanceolate, obtuse; larger stamens longer than calyx,

not very ciliate; capsule 3 mm. thick, lightly depressed, glabrous. —

Nearly 0. daphniformis Mikan, 105, of eastern Brazil but differs

in stems hirsute above, more or less leafy below, larger flowers,

etc. (Knuth). F.M. Neg. 28343.

Rio Acre: San Francisco, Ule 9470, type.

Oxalis fruticetorum Diels, Bot. Jahrb. 37: 424. 1906; 135.

O.Lechleri Knuth, Pflanzenreich IV. 130: 134. 1930.

Scandent in shrubs, otherwise procumbent, the reddish-brown

subligneous stems to 1 meter long, glabrous below, to 2.5 mm.

thick, sparsely spreading, pubescent to subtomentose above, with

rather few divaricate slender subligneous branches that are mostly

naked except toward the flowering tips, the somewhat hirsute leaves

on petioles only about 1 cm. long; leaflets all sessile and subequal,

about 7 mm. long and 3-5 or 6 mm. wide, slightly narrowed at base,

obovate, apically moderately incised with obtuse or acute sinus,

nearly glabrous above; stipules 5 mm. long, about one-fourth free,

566 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

lanceolate; peduncles (1) 3-4.5 cm. long, sparsely spreading pilose

as pedicels, these 7-10 mm. long, refracted in fruit, and sepals, these

oblong-lanceolate, obtuse, or subacute, hyaline margined to 6 mm.

long; umbels irregular, (1) 4-6-flowered; petals at least 12 mm. long,

broadly cuneate to retuse-rounded tip, yellow with reddish nerves;

larger stamens glabrous, scarcely denticulate and little longer than

calyx. — With the Pennell collections before one it appears that

0. Lechleri was based on a diminutive upper portion of the plant

earlier named by Diels which the latter rightly compares with the

Colombian 0. scandens HBK., the Peruvian plant more ligneous

with smaller leaflets pilose beneath. F.M. Negs. 26314; 26329

(0. Lechleri).

Cajamarca: In shrubs, Hacienda La Tahona, Hualgayoc, 3,100

meters, Weberbauer 4028, type; 261. Sandy soil, Socota to Tambillo,

Prov. Cutervo, 3,200 meters, Stork & Horton 10169. Sprawling in

humus or moss, 3,100 meters, Mito, 1845 (det. Knuth). On mossy

trees, 4,000 meters, Tambo de Vaca, 4379 (det. Knuth). — Cuzco:

Mossy bank, 3,800 meters, Cerro de Cusilluyoc, Pennell 13819 (det.

Knuth, 0. Lechleri); also 14100 at 2,800 meters in thickets. — Puno:

Prov. Sachapata, Lechler 2708 (type, 0. Lechleri). Ecuador.

Oxalis Herrerae Knuth, Pflanzenreich IV. 130: 115. 1930.

Becoming suffrutescent, a dm. or so high, with divaricate branches

to 5 mm. thick and dark brown bark; older branchlets glabrous, the

younger nearly aculeate with many lanceolate stipules at base of

petioles, the former 2 mm. long, the latter often 2 cm. long and

variously foliose-dilated, glaucous, glabrous and fleshy like the sessile

rotund-obcordate leaflets that are abruptly contracted toward the

base, often minutely but not conspicuously excised, 5 mm. long, 4

mm. wide; peduncles often 1 dip. long, terminating in a 5-7-flowered

2-branched cyme, the branches to 2 cm. long; bracts subulate, 2 mm.

long, minutely glandular as the unequal pedicels (to -5 mm. long)

and as long, narrowly lanceolate acute sepals; petals reddish, to 12

mm. long, spathulate, more or less retuse; shorter stamens as long

as the sepals, the stamens and styles puberulent. — Species most

distinct in the dilated petioles (Knuth). According to the collector

it may be narcotic. It develops into a gnarled or stockily branched

shrub or the stems sometimes virgate; an extreme form, or distinct

species is villous with subentire leaflets, Herrera 3507, Machu-

picchu, but det. Knuth, 0. coralleoides.

Cuzco: Ollantaytambo, Valle del Urubamba, Herrera 3356;

Pennell 13658 (both det. Knuth); West 6474 (det. Johnston).

FLORA OF PERU 567

Hacienda Silque, Valle del Urubamba, 2,950 meters, (Herrera 1757,

type). — Puno: Rocky shaded stream bank, Oconeque, Prov. Sandia,

Metcalf 30608 (det. Leonard, 0. Herrerae). "Macha-macha."

Oxalis Hochreutineri Macbr. Candollea 6: 9. 1934. 0. patula

Knuth, Notizbl. Bot. Gart. Berlin 7: 302. 1919; 123, not Eckl. & Zeyh.

Enum. 1: 87. 1836.

Annual, distinctive by the long arachnoid indument on the many

stems (these erect or ascending-divaricate, 10-18 cm. long), pedicels

and calyces; leaves sometimes fascicled on the more or less spreading

branches, most abundant above; leaflets glabrous, broadly obcordate,

more or less cuneate at base, rounded or barely incised apically, to

1 cm. long, slightly wider, the petioles often 2-3 cm. long; peduncles

2-5 cm. long, with simple or bifurcate umbel of 3-8 yellow flowers,

the ovate lanceolate somewhat arachnoid bracts 3-4 mm. long, the

pedicels to 5 mm. long or nearly obsolete; petals about 11 mm. long,

nearly clawed, cuneate dilated, retuse; smaller stamens 2, larger

about 3.5 mm. long, the styles 5 mm. long, both ciliolate. Root

vertical, rather stout. — The species name is for my friend who was

director of the botanical institutions at Geneva during my sojourn

there.

Lima: North of Chosica on stony outcrops with Cacti and other

scattered plants, 1,000 meters, Weberbauer 5342, type.

Oxalis huantensis Knuth, Pflanzenreich IV. 130: 432. 1930.

Strigillose firm simple stems about 2 dm. high, 5-6 mm. thick

below; petioles 4-10 cm. long, grayish tomentose puberulent, the

stipules to 4 mm. long, the narrowly triangular free part only 1 mm.

long; leaflets sessile, glabrate and opaque above, appressed gray-

pubescent beneath and on the margins, oblong or broadly ovate-

cuneate, to 2.5 cm. long, 1.5 cm. wide, with obtuse or rectangular

sinus 2-2.5 mm. deep; peduncles 14-20 cm. long, puberulent, with

a more or less bibranched 13-20-flowered umbel, the branches to

7 mm. long; bracts irregular or linear, acute, 4 mm. long; pedicels

nearly filiform, glabrous, 7-10 mm. long; sepals narrowly lanceolate,

acute, pale green, glabrous, 6-7 mm. long; petals brown-nerved,

15 mm. long, long-clawed, obovate; styles of long-styled flowers one

and a half times longer than sepals, the larger stamens as long as

sepals; stamens and styles puberulent. — Allied to 0. spiralis, fide

Knuth.

Ayacucho: Huanta, 3,200 meters, Weberbauer 7519, type.

568 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Oxalis hypopilina Diels, Bot. Jahrb. 37: 424. 1906; 88.

Shrubby, the ultimate branches ligneous, 2 mm. thick, tomentose

and densely leafy toward the tips, the leaves crowded but not

rosulate; leaflets oblong or oblong-ovate, obtuse or more or less

rounded each end, about 3 cm. long and half as broad, glabrous

above with some small white spots, sericeous puberulent beneath;

medial petiolule 5-8 mm. long; petioles 3 cm. long; peduncles nearly

apical, 4-5.5 cm. long, tomentose puberulent as the pedicels (3 mm.

long), bracts (1 mm. long, deciduous) and calyces; sepals lanceolate,

acutish, 5-6 mm. long, the broadly spathulate yellow petals twice as

long; short stamens glabrous, the longer as the styles pubescent, a

little longer than the sepals, the black rounded or slightly depressed

capsule scarcely as long. — Distinct in habit and indument from 0.

Poeppigii (Diels) ; about 1 meter high, the branches thick and densely

scarred above by the numerous fallen petioles. F.M. Neg. 26323.

Amazonas: Tupen, Prov. Luya, stony outcrop, 1,100 meters,

(Weberbauer 4796, type; 155).

Oxalis juninensis Knuth, Notizbl. Bot. Gart. Berlin 7: 305.

1919; 223.

Subterranean caudex stout, sometimes 1 cm. thick, subligneous,

more or less forked at apex with branches extending about 1 cm.

above the ground and densely clothed with petiolar remains; petioles

to 2.25 cm. long, succulent, nearly glabrous or sparsely pilose;

leaflets glaucous, paler beneath and also lacunose-cellulose, glabrous

above, triangular, retuse or very obtusely incised to one-eighth,

3 mm. long and broad; peduncles 1 cm. long, filiform, 1-2-flowered;

pedicels 2.5-3 mm. long, sparsely pilose; sepals foliaceous, the outer

ovate-orbicular, 2 mm. long and broad, rounded at tip; petals to

8 mm. long, yellow. — Spelled by author, 223, "iuninensis" ; cf.

0. megalorrhiza and relatives. 0. androsacea Knuth, 228, of Ecuador

has leaflets pubescent both sides, tomentose acutish sepals 3-4 mm.

long.

Junin: Tarma, open stony places, 300 meters, (Weberbauer 2359,

type).

Oxalis juruensis Diels, Verh. Bot. Ver. Brandenb. 48: 173.

1907; 90.

More or less ligneous, to 1 meter high, with rather spreading

puberulent densely leafy branches, the alternate estipulate petioles

to 6 cm. long, medial petiolule 7-8 mm. long; leaflets ovate, paler

FLORA OF PERU 569

and minutely subsericeous beneath, sparsely pilose above, to 4.5 cm.

long, 2.5 cm. wide; peduncles many, 7-9 cm. long, about one mm.

thick, somewhat pubescent, with a bifurcate umbel of 2-5 flowers;

bracts lanceolate, acute, 1-1.5 mm. long; pedicels nearly filiform,

5 mm. long, somewhat lanate; sepals ovate, obtuse, becoming lanceo-

late and acute, then to 6 mm. long, subsericeous; petals pale yellow,

twice as long as sepals; stamens as styles hirsutulous; capsules gla-

brous or minutely glandular, ovoid, 10 mm. long, 5-6 mm. thick,

with 3 seeds in a cell. — Var. emarginata Huber of Brazil has the

leaflets rounded and emarginate. The Tarapoto collection is placed

here from the similarity of the negatives; this may be an error but

the specimen is certainly not 0. amazonica ex char, or negative.

However other species which perhaps are all phases of one include

0. microcarpa Benth., 63, and 0. sepium St. Hil., 64; the former from

Colombia is said to extend to southern Ecuador and seems to be

the same except for slightly smaller flowers and capsules; the latter,

typically of Brazil, has violet flowers. F.M. Negs. 26327; 26290

( Ule 6714).

San Martin: Road to Tarapoto, Ule 6714 (det. Knuth, 0. ama-

zonica). Brazil.

Oxalis latifolia HBK. Nov. Gen. & Sp. 5:. 237. pi. 467. 1822;

273. 0. araucana Reiche, Bot. Jahrb. 18: 302. 1894, at least as to

Peru. 0. atroglandulosa Knuth, Notizbl. Bot. Gart. Berlin 7: 311.

1919; 253. lonoxalis latifolia (HBK.) Rose, Contr. U. S. Nat. Herb.

10: 113. 1906.

Bulbs attaining a dm. or so in diameter, the brown scales merely

ciliate, many-nerved; plants in flower usually a dm. or 2 high, gla-

brous or the leaflets, petioles and peduncles sparsely ciliate-pilose;

leaflets broadly deltoid, to about 4.5 cm. wide, 2 cm. long or little

longer, often smaller, the broad sinus shallow, the divergent lobes

more or less ovate; petioles slightly succulent, often 8-12 cm. long

or even longer, the basal peduncles sometimes 2 dm. long terminating

in a simple umbel of several to 17 pink or violet flowers, these 8-15

mm. long; bracts more or less abruptly and acutely acuminate, 1-3

mm. long; pedicels 1-2 cm. long; sepals oblong or nearly, 3.5-5 mm.

long, acute, reddish or dark bicallose at tip; typically longer filaments

ciliate and with a lateral tooth; styles glabrous. — With 0. violacea L.,

271, the basic species of a large group of forms centering in Mexico,

and it is possible that all the Peruvian material is distinct or even

divisible but the monographer has not made this clear. F.M. Negs.

570 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

26292 (Gaudichaud, det. Knuth, 0. araucana); 29556 (0. atroglandu-

losa).

Lima: WilkesExped., (Seler 244; 261, all fide Knuth). In rocks,

San Geronimo, 5889 (det. Knuth). Talus slope, north of Barranca,

Worth & Morrison (det. Johnston, 0. atroglandulosa). Lomas de

Dona Maria, Goodspeed Exped. 9252. Amancaes, Soukup 1799 (det.

Leonard). Limestone slopes, Atocongo, Pennell 14778 (det. Knuth,

0. araucana). Mongomarca Mountain, Weberbauer 5682 (type,

0. atroglandulosa). Chancay, Ruiz & Pavdn (det. Knuth, 0. atro-

glandulosa). San Lorenzo, Gaudichaud (det. Knuth, 0. araucana).—

Huanuco: Near Acomayo, 2,100 meters, Woytkowski 5. — Apurimac:

In cultivated fields, 2,500 meters, Vargas 9778? — Cuzco: Yucay,

3,000 meters, Valle del Urubamba, (Herrera 1369). — Arequipa:

Rock slides, south of Chala, Worth & Morrison 15676 (det. Johnston,

0. atroglandulosa). To Mexico (Texas) and the West Indies.

Oxalis lomana Diels, Bot. Jahrb. 37: 426. 1906; 200.

Acaulescent, 4-5 cm. high, from a bulbous caudex 8 mm. thick,

its many basal roots filiform; leaves rosulate, 10 or fewer, with

sparsely pilose petioles 3-4 cm. long, the obcordate, obtusely incised

leaflets to 9 mm. long and broad, opaque and a little pilose or gla-

brate above, paler and somewhat hirsute-arachnoid beneath;

peduncles 2-4-flowered, densely pilose as the bracts (subulate, 4 mm.

long), pedicels (4-8 mm. long) and sepals, these unequal, the larger

ovate-deltoid, to 6.5 mm. long, 4 mm. wide, in fruit equaling the

oblong 9 mm. long capsule; flowers yellow, 10 (-12) mm. long;

stamens glabrous, the styles nearly so. — The petioles and peduncles

are articulate about 1 cm. above the base which persists. The var.

hirsuta Knuth, 200, is the typical form, more or less lanate including

the sepals; in the var. glabrescens Knuth, I.e. these are glabrate.

Among species with unequal sepals easily recognizable by the tuberi-

form rhizome, indument of leaves and inflorescence (Diels).

Arequipa: Lower edge of lomas, Mollendo, Weberbauer 1480,

type; 144; Johnston 3550; 6292 (both det. Knuth). Posco, Cachendo

and Mejia, (Gunther &Bruns 256; 256a; 257, all fide Bruns). — Tacna:

Tacna, 800 meters, Werdermann 726 (det. Knuth); Woitschach 13;

15; 17 (these three var. glabrescens).

Oxalis lotoides HBK. Nov. Gen. & Sp. 5: 241. 1822; 138.

Acetosella lotoides (HBK.) Kuntze, Rev. Gen. 1: 92. 1891.

Supported in shrubs or procumbent, sometimes 7 dm. long or

longer, rather densely divaricately branched, foliate and floriferous

FLORA OF PERU 571

nearly from the base, herbaceous-subligneous, the greenish brown

stems glabrous below, often 2.5 mm. thick, increasingly soft-pubes-

cent from the middle to the tips; petioles 2-2.5 cm. long, the 3-9

mm. long stipules two-thirds adnate, acute; leaves often crowded,

the leaflets all sessile, obcordate, mostly 15 mm. long, 12 mm. wide,

one-eighth incised with obtuse sinus, appressed hirsute both sides

and marginally; peduncles 3-6 cm. long with 2-9-flowered irregular

cyme, the subulate acute bracts 4 mm. long, the pedicels (8-12 mm.

in type) 0.5 to 4 cm. long; petals pale, violet-striate, obovate, 17-22

mm. long, the oblong-lanceolate acutish sepals 6 or 7 mm. long;

styles pubescent (type). — Type from Colombia. 0. pichinchensis

Benth., 138, doubtfully distinct according to Knuth, and the same

according to Weddell, has leaflets glabrate above as sepals, these

6.5-8 mm. long and obtuse or retuse. But my 4281, which I refer

with some doubt to 0. fruticetorum and Knuth here, suggests that

either the species of HBK. is too narrowly defined or is not in Peru;

the Raimondi collection is not now at hand. Knuth describes

stamens and styles glabrous, the former not noted by HBK. Illus-

trated, Hook. Icon., n. ser. 3 pi. 661. F.M. Negs. 36805; 26340

(0. pichinchensis).

Cajamarca: Montana de Nancho, Hualgayoc, 2,500 meters,

(Raimondi, det. Knuth). Colombia to Bolivia; Venezuela.

Oxalis lucumayensis Knuth, Repert. Sp. Nov. 23: 144. 1926;

135.

Procumbent-ascending, the nearly fleshy branches to 4 dm. long

and (in herb.) 5 mm. thick, brownish-yellow tomentose as the leaves,

especially beneath, and the entire plant in greater or less degree

even to the irregular 8-flowered umbels, except for the yellowish

petals that apparently are about twice as long as the 8 mm. long

lanceolate somewhat hirsute sepals; leaflets all sessile, oblong, the

lateral a little oblique at the rounded base, the medial somewhat

cuneate, all bilobed with acute 5 mm. sinus, often 6 cm. long, 3.5

cm. wide; petioles to 9 cm. long, as often also the peduncles; bracts

linear, as densely tomentose as the pedicels. — Imperfectly known.

Cuzco: Lucumayo Valley, 1,800-3,600 meters, Cook & Gilbert

1351, type.

Oxalis Macbridei Knuth, Pflanzenreich IV. 130: 431. 1930.

Suffruticose, glabrous, 6-10 dm. high, the ligneous stems to 5 mm.

thick, subsimple, leafy especially above the middle; petioles to 8.5

572 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

cm. long, firm, purplish toward base; leaflets 3, the middle one

lanceolate, cuneate at base, gradually acutely acuminate, 8-9 cm.

long, 3-3.5 cm. wide, on firm petiolule 2.5 cm. long, the lateral sub-

sessile, often 5.5 cm. long, 2.5-3 cm. wide, abruptly and very acutely

acuminate; peduncles rather numerous, 6-8 cm. long, strict; bracts

linear-lanceolate, acute, 3 mm. long; inflorescence with two branches,

these with two forks, the former to 1 cm. long, the latter 2 cm. long

with 1 or 2 flowers at tip and marked with pedicel-scars; sepals

ovate, obtusish, finally dilated, 4 mm. long, the yellow petals 3

times longer; styles and stamens glabrous. — Nearest, fide the author,

to 0. Tessmannii. Bush to 1 meter high with canary yellow flowers

said to cause irritating skin eruption (Mexia).

Huanuco: Pampayacu, 1,100 meters, 5089, type. Along Rio

Huallaga to Riachuela Chontalagua, Mexia 8295a (det. Standley).

Oxalis marcapatensis Knuth, Repert. Sp. Nov. 29: 216. 1931.

Glabrous; rhizome deeply set, perpendicular, 4 mm. thick, erect,

closely scaly from petiole or stipule rudiments, topped apically by

rosette 7 cm. across of about 8 peduncles and many leaves, the former

mostly 2-3-flowered, as long as petioles, the latter with sessile obovate

leaflets 10 mm. long, 6 mm. wide, about 1 mm. notched at apex;

petioles 5 cm. long, fleshy, dilated, articulate 2 mm. above base;

free part of stipules acute, lanceolate, 3 mm. long; bracts 2 mm. long,

linear, acute, often reddish; pedicels 1-2 cm. long, filiform; sepals

lanceolate, acute, without callus, 6.5 mm. long the larger stamens as

long, sparsely ciliate; corolla tubiform, the petals orange-yellow

veined with red, retuse, about 15 mm. long. — Section Carnosae,

affine 0. paucartambensis (Knuth), and apparently not distinguish-

able.

Cuzco: On rocks, 2,500 meters, Marcapata Valley, at Chilechile,

(Weberbauer 7870, type).

Oxalis Martiana Zucc. Denkschr. Akad. Muench. 9: 144. 1823-

24; 250.

Acaulescent, often 2 dm. high or taller, from a rotund bulb

2-2.5 cm. in diameter composed of many brown bulblets about

4 mm. thick; leaves often many, rosulate, on lax petioles to 2.5 dm.

long that are sparsely hirsute with a few long spreading trichomes

and a small fascicle of trichomes at the insertion of the rotund-

obcordate leaflets, these glabrous, minutely callosed marginally,

2.5-3.5 cm. long and broad, lightly but often acutely incised;

FLORA OF PERU 573

peduncles few (1-5), often 1.5 dm. long, 5-12-flowered, glabrous as

the pedicels and calyces or somewhat hirsute; bracts several, 2 mm.

long, ecallose; pedicels irregularly umbellulate, 1-4 mm. long;

sepals ovate-lanceolate, reddish-callose at acutish tip, 5-7 mm. long;

corolla conic-campanulate, 3-4 times longer than calyx, blue-violet,

paler toward base, the cuneate-obovate petals rounded-retuse, entire;

larger stamens longer than sepals. — The similar 0. bulbifera Knuth,

252, and 0. limosa Prog., 251, of Brazil, the former from Bolivia, are

unique (according to the monographer) in having many stolons from

the elongate bulb, these bearing apically bulblets; the last has a

very irregular umbel, that of the first being regular. Illustrated,

Bot. Mag., pis. 2781; 3896. F.M. Neg. 19199.

San Martin: Near Moyobamba, Klug 3593 (det. Standley).

Lima: Soukup 1916 (det. Leonard). San Geronimo, ditch bank,

5888 (det. Knuth). Brazil; Bolivia; Argentina; and widely estab-

lished in warmer regions.

Oxalis Mathewsii Knuth, Pflanzenreich IV. 130: 87. 1930.

0. tumbezensis Knuth, I.e. 431.

Shrub densely fuscous tomentose even to the umbels, the short

branches 3-4 mm. thick; petioles to 9 cm. long, the lateral petiolules

2-3 mm. long, the medial to 13 mm. long or longer; leaflets all ovate-

lanceolate, cuneate to base and acute apex, the medial often 5 (-7.5)

cm. long, half as wide, the lateral 4-5 cm. long or smaller; peduncles

to 6 cm. long, many as the leaves toward the branchlet tips, the

pseudo-umbellate flowers scarcely or shortly dichotomous; pedicels

1.5-4 (10) mm. long; sepals oblong, obtuse, 6-8 mm. long, the yellow

cuneate-obovate or -oblong petals 10-12 mm. long; larger stamens

as long as sepals, subglabrous, the smaller two times shorter; styles

hirsutulous or rarely glabrous, as long as or longer than sepals. —

The second species of Knuth seems to be only a robust plant, slightly

larger in all parts; the species is apparently very near to 0. psoraleoides

HBK., 87, of Colombia and 0. piauhyensis Knuth, 94, both earlier

names. F.M. Neg. 26333.

Tumbez: Hacienda Chicama, 800 meters, Weberbauer 7639 (type,

0. tumbezensis). — Cajamarca: Near Jae"n, Weberbauer 6196 (det.

Knuth, 0. sublignosa) ; also Raimondi 2229; 4740 (det. Knuth, 0.

piauhyensis). — Amazonas: Chachapoyas, Mathews, type, Herb.

Delessert. — Cuzco: Echarate, Prov. Convencion, Goodspeed Exped.

10515. Hacienda Santa Rosa, Prov. Convencion, Soukup 799.

Hacienda Potrero, 1,300 meters, Prov. Convencion, Vargas 2487.

574 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Oxalis medicaginea HBK. Nov. Gen. & Sp. 5: 242. 1822; 136.

Acetosella medicaginea (HBK.) Kuntze, Rev. Gen. 1: 92. 1891.

Glabrous and procumbent below but the younger ascending

branchlets pubescent probably several dm. long; petioles nearly

glabrous, 2-2.5 cm. long; stipules membranous, adnate below the

tips free, pilose; leaflets subsessile, obovate, emarginate, reticulate,

ciliate, glabrous above, appressed pubescent and subglaucous be-

neath, the lateral 14-15 mm. long, the medial 16 mm. long, 12 mm.

wide; peduncles filiform glabrous, bifid apically, 4-12-flowered, a

dm. long or longer, much exceeding the leaves; flowers becoming

racemose, about 2 cm. long, on filiform pubescent pedicels 6-10 mm.

long, articulate above the base; bracts linear, acuminate, puberulent;

sepals glabrous, oblong-lanceolate, obtuse, a half as long as the

corolla; petals rounded or emarginate; filaments glabrous; styles

pubescent. — After HBK. Progel describes the longer filaments as

with a lateral tooth; Knuth found the filaments and styles hirsute,

the sepals pubescent, acute, to 7.5 mm. long. From the negative

the stems appear to be subligneous. It is noteworthy that the origin

of the type was not surely known. If Knuth's characterization is

correct the name seems to be the earliest for a number of forms.

F.M. Neg. 36808.

Peru: Possibly; according to Knuth Colombia and Venezuela to

Argentina in the Andes but not in Peru.

Oxalis megalorrhiza Jacq. Oxal. 33. 1794; 184. 0. carnosa

Molina, Sagg. Nat. Chile ed. 2. 288. 1810. Acetosella megalorhiza

(Jacq.) Kuntze, Rev. Gen. 1: 92. 1891.

Plant 4-10 cm. high, glabrous (except a var.), from a subligneous-

fleshy horizontal rhizome even to 2 cm. thick, the similar and erect

simple or little forked stem scaly above with scars of fallen petioles,

leafy and floriferous only at the tip; petioles to 8 cm. long; leaflets

glaucous-green, slightly lustrous above and papillose beneath, ob-

cordate, 1-1.5 cm. long, mostly 12 mm. long and broad, lightly

incised; peduncles to 10 cm. long, the umbel 2-5-flowered with

narrowly linear bracts; outer sepals broadly ovate, nearly hastate

at base, acutish, 7 mm. long, 4 mm. wide, the inner narrower, sub-

oblong, and almost hidden by the outer; corolla yellow or orange,

18 mm. broad, the marginate petals broadly connate at base, three

to four times longer than the calyx; seeds many in each cell. — The

var. hirta (Knuth) Macbr. comb, nov., 0. carnosa Mol. var. hirta

Knuth, Pflanzenreich IV. 130: 186. 1930, has the leaves hirsute

FLORA OF PERU 575

beneath. Biology discussed by Hildebrand, Lebensverh. Oxal. 9.

1884. Knuth cites the type of Jacquin as definitely the same as

the plant of Molina but the latter's name may be in the first edition

of his work, 1782, and thus earlier. The type number of 0. arequi-

pensis at Chicago was determined by Knuth as 0. carnosa! I have

placed it with 0. solarensis, doubtless merely a variety with smaller

obtuse or rounded sepals. — Illustrated, Bot. Mag. pi. 2866; Bot.

Jahrb. 18: pi. 9; Knuth, I.e. fig. 14, page 185.

Determinations by Knuth except as noted.

Ancash: In sand among loma boulders, Casma to Huarmei,

Goodspeed Exped. 9162 (det. Johnston). — Lima: near Lima, Rose

18575. San Geronimo, 5986. San Gallen and Viejas Islands,

(Murphy 34.69; 3218, det. Johnston). Chorillos, 5853; 5852. Aman-

caes, Soukup, 2139; 1798 (det. Leonard) ; Mexia 04012 (det. Johnston,

typica). Rocky slopes near Barranca, Worth & Morrison 9118;

(9108) (both det. Johnston). Prov. Huarochiri, Goodspeed & Metcalf

30245 (det. Johnston). Atocongo, Pennell 14769. Chosica, 2871.

Rock crevice, Huariaca, 3106. Rock outcrops and loose soils,

Matucana, 167; 2918; 2956. Rio Blanco, rock outcrops, 3,000

meters, 2973; Killip & Smith 21538. Tacna, (Woitschacli). Gala-

pagos; Chile; Bolivia.

Oxalis melilotoides Zucc. Denkschr. Akad. Muench. 9: 165.

1823-24; 109. 0. Haenkeana Spreng. Syst. 4, pt. 2: 186. 1827.

Acetosella melilotoides (Zucc.) Kuntze, Rev. Gen. 1: 92. 1891.

Glabrous (or nearly except cilia) greenish-red little branched

herb 6-10 dm. high, probably trailing; leaves remote, the leaflets

sessile or nearly, cuneate-obovate, emarginate or lightly obcordate,

pale green above, glaucescent beneath, the midnerve often callus-

tipped, about 2.5 cm. long, 13-17 mm. wide; petioles to 5 cm. long;

stipules adnate, the younger finely ciliate; peduncles nearly twice

longer than the leaves, bifid at apex, to 15-flowered, the laxly race-

mose flowers secund on filiform erect pedicels 2.5 cm. long, finally

spreading-refracting; sepals lanceolate, acute, as the bracts but

these a little ciliate; corolla ample, infundibuliform, the yellow red-

striped petals obovate; larger filaments and styles finely barbate;

ovary 5-angled, glabrous, the capsules ovoid. Flowers 15 mm. long

or longer. — The type locality may be a misreading of "Huanuco."

The Stork & Horton plant was prostrate or pendent, about 1 meter

long, calyx stems and leaves beneath purple as also Woytkowski's,

this with red-orange flowers! F.M. Neg. 19200.

576 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Huanuco: Fog belt forest, Carpish, Stork & Horton 9926. With-

out locality, probably dept. Huanuco, Haenke, type. Sariapampa,

open glade, 3,200 meters, Woytkowski 34291. — Junin: Carpapata

above Huacapistana, 2,700 meters, Killip & Smith 24468 (det.

Knuth, 0. tuberosa).

Oxalis micrantha Bert, ex Colla, Mem. Accad. Torino 37: 50.

1834; 121. Acetosella micrantha (Bert.) Kuntze, Rev. Gen. 1: 92.

1891. 0. chosicensis Knuth, Notizbl. Bot. Gart. Berlin 7: 303. 1919;

125.

Small usually more or less tufted annual ordinarily only a few

cm. tall, in varying degree hirsute and somewhat glandular, especially

the nearly filiform divaricate pedicels (these to about a cm. long) of

the laxly branched several to many-flowered cymes; petioles to 8 cm.

long, about half as long as the peduncles, the broadly obcordate

rather deeply emarginate leaflets sometimes 14 mm. long, 15 mm.

wide, often smaller; bracts caducous, 1-2 mm. long; sepals often

rubescent below and marginally, nearly linear, acute, 3-4 mm. long,

sometimes equaled by the spathulate yellow petals; seeds 1-many

in each cell. — The form designated var. setifera Knuth is more densely

and more harshly pubescent. F.M. Neg. 26335.

Lima: Loma Ancon, 700 meters, Soukup 2127. On stony out-

crops north of Chosica, Weberbauer 5347 (type, 0. chosicensis).

Saldacuna, Soukup 1939 (det. Leonard). — Arequipa: North of Ati-

quipa, Worth & Morrison 15655. Mejia and Posco, (Gunther &

Buchtien 253, det. Bruns). Moquehua: Open mixed formation,

2,200 meters, Torata, Weberbauer 7393 (det. Knuth). Chile;

Argentina.

Oxalis minima R. & P. ex G. Don, Gen. Syst. 1: 760. 1831.

0. Ruizii Knuth, Pflanzenreich IV. 130: 248. 1930. 0. Philippii

Knuth, Meded. Rijks Herb. Leiden 27: 67. 1915, probably, at least

as to Peru.

Bulb about a cm. in diameter, the brown scales fibrous lanate

in age ; peduncles and petioles subequal or the former in type shorter

(maybe this is a diminutive form), a cm. or so long, sometimes 5 cm.

high, the 2 bracts early below the calyx as to type, finally about

medial apparently as the peduncle lengthens; petioles glabrous,

2.5-4 cm. long, more or less margined especially toward the base;

leaflets obcordate, 4-6 mm. long, 5-8 mm. wide, somewhat shorter,

often a little larger; sepals 3-5 mm. long, rather more obtuse than

acute, oblongish, ecallose but sometimes slightly ciliate toward tip

FLORA OF PERU 577

and reddish on the thinner margins; petals pink or white but more

or less tinted in part, usually 12-15 mm. long; stamens puberulent,

so far as observed. — Leaflets described as pubescent both sides

under a lens by Exell, Journ. Bot. 63: 174. 1925, on whose description

of type, Herb. Lambert, I have drawn, but this may refer to the

lacunose condition, especially beneath. The name of Ruiz & Pavon

is to be retained as its first valid use in the genus. The species, at

least sens, lat., is apparently under various names in all the high

Andes. F.M. Neg. 29560 (det. Knuth, 0. Philippii).

Lima: Rio Blanco, 3018 (det. Knuth, 0. Asplundii). Hacienda

Arapa near Yauli, (Weberbauer 307; 31*6, both det. Knuth, 0.

Philippii); 221. — Junin: Cerro de Pasco, (Raimondi, det. Knuth,

0. Philippii). La Oroya, Kalenborn 25. — Libertad: Grassy slopes

of glacial valley, Cerro Huaylillas, Prov. Huamachucho, 3,800

meters, West 8118 (det. Johnston, 0. oreocharis). Huancavelica:

Puna grass slopes, 3,900 meters, Stork & Horton 10840 (det. Standley,

0. Asplundii). Without data, Herb. Madrid, Ruiz & Pav6n, type.

Oxalis modestior Knuth, Repert. Sp. Nov. 40: 293. 1936.

Trunk fleshy-subligneous, 5 cm. high or higher, 5 mm. thick,

simple in the only one seen, scaly by the remains of leaves and

bearing many leaves and peduncles at the tip, both of these (as to

type) glabrous, the former with filiform petioles 2-3 cm. long, the

latter also filiform but one and a half times longer with an apical

umbel of 3-5 flowers on pedicels 5-8 mm. long; stipules lanceolate,

acute, hispidulous, to 5 mm. long; leaflets rotund-obcordate, broadly

cuneate or nearly rounded at base, rounded or lightly rounded-

excised at tip, 3-4 mm. long and broad, in type glabrous and glaucous;

bracts linear, acute, hirsutulous, 2-3 mm. long; sepals narrowly

lanceolate, 4-5 mm. long, 1.5 mm. wide, retuse at the often reddish

tip, glabrous; petals 10-12 mm. long, unguiculate-cuneate from base,

rounded at apex, apparently whitish, the veins purple; stamens in

type puberulent longer than the larger sepals. — Section Carnosae,

but could be a diminutive state of 0. peduncularis; neither the West

nor the Stork and Horton specimens exactly agree with type, the

former with a megalorrhiza type caudex-root, the latter with leaflets

pilose beneath.

Cuzco: Saxaihuaman, 3,200 meters, Herrera 3121, type. Ruins

of Tambomachay, Saxaihuaman, 3,600 meters, West 3856 (det.

Johnston with query 0. megalorrhiza). — Libertad: rock crevices,

Huamachuco, 3,300 meters, West 8093. — Huancavelica: Gravelly

578 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

clay north of Pampas, 3,250 meters, Stork & Norton 10239 (det.

Standley, 0. pygmaea Gray). "Chchahulco" (West).

Oxalis mollis HBK. Nov. Gen. & Sp. 5: 241. 1822; 137. Aceto-

sella mollis (HBK.) Kuntze, Rev. Gen. 1: 92. 1891.

More or less densely villous unless above where puberulent-

tomentose, the stems somewhat ligneous and apparently several dm.

long, sparsely branched and rather densely leafy toward the tips;

leaflets sessile, obovate-obcordate, broadly cuneate at base, to 2 cm.

long, 17 mm. wide (16-18 by 12-14, type), to one-sixth incised with

obtuse sinus, the broad rounded lobes straight, appressed pilose

above, rather appressed lanate beneath; petioles 1.5-4 cm. long

with stipules membranous villous adnate or tips about 2 mm. long;

peduncles about 7 cm. long, lax, slender with a 7-10-flowered cyme,

its branches sometimes 1.5 cm. long, the pedicels 1-14 mm. long;

bracts subulate, acute, to 3.5 mm. long; sepals lanceolate, acutely

acuminate, 4 mm. long, finally 7 mm. long (Knuth), more or less

sericeous-villous, the whitish petals apparently about twice as long;

longer stamens and styles pubescent. — Knuth suggests very probably

a form of 0. lotoides HBK., 138. The Colombian type at Paris is a

young plant, the sepals thus short, petals not developed. 0. mollis-

sima (Rusby) Knuth, 137, of Bolivia, has densely tomentose villous

leaflets 3.5-4.5 cm. long, 2.5-4 cm. wide. The var. glandulosa

Knuth, 433, the Peruvian plant, may not belong here; it has broadly

cymose inflorescence, densely and minutely glandular. F.M. Neg.

36810.

Huanuco: Muna, 1^11. Huacachi near Muna, 4091. Colombia;

Bolivia (Knuth).

Oxalis moqueguensis Knuth, Pflanzenreich IV. 130: 438. 1930.

Bulb scales narrow, ribbed, somewhat lanate, acuminate; leaves

1-5, the leaflets glabrous but more or less callus dotted marginally,

broadly triangular, to 2 cm. wide, 1.25 cm. long, emarginate some-

what rectangulately, petioles slender, 7-14 cm. long, the few

peduncles slightly to about twice as long, terminating in an umbel

of 2-5 (rarely only 1) flowers, their pedicels often 3 cm. long or longer,

pendent in fruit; bracts variable in form, about 2 mm. long; sepals

lanceolate, acute or obtusish, 7-9 mm. long, with 2 or 3 dark linear

calli at tip; flowers purple or rose-colored, sometimes changing white,

17-20 mm. long; stamens and styles glabrous, the longer stamens

equaling or finally longer than the sepals. — Pods linear-cylindric,

FLORA OF PERU 579

nearly 2 cm. long, at maturity pendent. Seems to be near 0. pazensis

(Rusby) Knuth, 259, not seen and may be a local variant of 0. elegans.

Moquehua: Rainy green formation, 3,100 meters, Carumas,

Weberbauer 7255, type; also 7309.

Oxalis nubigena Walp. Nov. Act.' Acad. Leop.-Carol. 19:

Suppl. 1: 320. 1843; 218. 0. pygmaea Gray in U. S. Expl. Exped.

Bot. 1: 322. 1854. 0. aetheria Macbr. Candollea 6: 8. 1934 (0.

pygmaea Gray, not Meyer). 0. yauliensis Knuth, Repert. Sp. Nov.

24: 51. 1927.

A few cm. high (with stem) and more or less compact, or some-

times a. globular little annual, the stem obsolete or very short and

concealed completely in the many crowded leaves and peduncles,

the former 1-3 cm. long with conspicuous scarious stipules a third

to a half as long and soon lacerate at tip, the latter shorter or longer,

1-flowered, with a pair of caducous subulate linear or filiform sub-

medial bracts varying from 1-2 to several mm. long; leaflets broadly

obcordate, glabrous or early with a few marginal cilia, almost minute

to 7 mm. wide, 5 mm. long, usually somewhat smaller, obviously

but moderately retuse; flowers white or probably always with yellow

centers, small, about 3 mm. long or in type said to be 3.5-4 mm.

long, the sepals described as ovate-lanceolate but for plants seen

linear-lanceolate, acutish, membranous, 2-2.5 mm. long, slightly

longer in age; capsules short, subovoid, with about 8 seeds or usually

2 in each cell. — Without the type this interpretation may be incorrect

but the careful Weddell, Chlor. And. 2: 291. 1857, wrote of 0.

pygmaea: "flowers yellow; otherwise character of 0. nubigena."

However Pennell 13766 (det. Knuth, 0. pygmaea) was proximally

yellow and my 918 had a yellow tube. This seems to be a starved

plant; half covered with snow when found, it was (naturally?)

reduced and rubescent.

Lima: On limestone rocks, Pucara, 3,700 meters, Weberbauer 437

(det. Knuth). Bafios and below Culnai (Cullnay), Wilkes Exped.

(type, 0. pygmaea). Rio Blanco, stony barren hillsides, 2971 (det.

Knuth, 0. pygmaea). Alpamina, Weberbauer 5124 (det. Knuth). —

Junin: Yauli, 4,200 meters, rock ledges, 91 8 (type, 0. yauliensis).

On porphyric rocks, 4,400 meters, (Weberbauer 303, det. Knuth).—

Cuzco: Bushy banks, Colquipata, 3,500 meters, Pennell 13766 (det.

Knuth, 0. pygmaea). Hacienda Churu, Valle de Paucartambo,

3,500 meters, Herrera 1043a; 1907b; 1912 (all det. Knuth, 0.

pygmaea). Asuncion Bridge, among mosses, 3,750 meters, Vargas

580 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

11027 (det. Standley, 0. ollantaytambensis). — Puno: Chiquibambilla,

puna streamlet bank, 3,900 meters, Pennell 13394 (det. Knuth,

0. pygmaea). Compuerto, (Weddell). Lake Titicaca, (Meyen,

type). Bolivia; Argentina(?) ; Chile(?).

Oxalis oreocharis Diels, Bot. Jahrb. 37: 423. 1906; 246. 0.

Asplundii Knuth, Repert. Sp. Nov. 24: 53. 1927, at least as to Peru.

Acaulescent, 3-7 cm. high from a rotund bulb 12 mm. in diameter

or smaller, densely squamose with lanceolate scales about 6 mm.

long; stipules membranous, long-pilose, 6-8 mm. long; petioles 2-6

cm. long, lax, glabrous; leaflets glabrous or sparsely pilose, softly

long-ciliate, ovately obcordate, about 8 mm. long and 9-12 mm.

broad, glaucous-green, cellulose-lacunose both sides, the lower sur-

face often completely dark punctate-striate; peduncles only 1-6,

1-4 cm. long in type, about as long as petioles, 1-flowered, glabrous

as the sepals, bibracteolate 5-10 mm. below the tip, the abruptly

acuminate nearly mucronate bractlets (2) 3-6 mm. long, connate

basally; flowers broadly conical, rose-colored, (15) 13-18 mm. long,

the sepals 5-7 (4-4.5) mm. long, oblong, truncate or retuse, bi-

callose at tip, the petals broadly cuneate, retuse, rounded, entire;

longer stamens very sparsely ciliate or glabrous longer than sepals,

the styles pilose below. — Knuth distinguishes his plant, type from

Bolivia, Tuapata, Watkins (det. Knuth, 0. Asplundii), by wide

petioles appearing alate in herb, leaflets sparsely ciliate, sepals

lanceolate, membranous, acute, the calli scarlet slender lines, dif-

ferences if specific not apparent in our material. For that matter

the significance of the characters I have been forced to use in key

without general reduction are unknown to me, and here too, type or

material cited by Knuth not seen.

Lima: Bunch grass slope, Rio Blanco, 3,000 meters, (det. Knuth,

0. Asplundii). — Junin: East of Palca on grass steppes with scattered

shrubs, Huarochiri, 3,000 meters, Hrdlicka (det. 0. Asplundii, by

Knuth); 3,500 meters, (Weberbauer 2493, type); 94; 223; 249. La

Oroya, among polster and rosette plants, 4,300 meters, (Weberbauer

2625, fide Diels).

Oxalis Ortgiesii Regel, Gartenflora 24: 1. 1875; Act. Hort.

Petrop. 3: 117. 1875; 110. 0. excisa Prog, in Mart. Fl. Bras. 12, pt.

2: 496. 1877. Acetosella Ortgiesii (Regel) Kuntze, Rev. Gen. 1: 92.

1891. A. excisa (Prog.) Kuntze, I.e. 0. Raimondii Knuth, Pflan-

zenreich IV. 130: 111. 1930.

FLORA OF PERU 581

Becoming a dm. to several dm. high, rooting along the decumbent

base in age and the fibrous roots with an occasional small perhaps

abortive tuber, the stems more or less densely strigillose especially

toward the leafy tips, in age even 4-5 mm. thick; petioles slender,

2-8 cm. long; leaflets obscurely to rather densely appressed pubescent

especially beneath, thinly membranous in herb., subequal but only

the lateral strongly oblique, the somewhat larger medial broadly

obtriquetrous in younger plants, about 2.5 cm. wide at apex, 1.5 cm.

long, in older or vigorous plants about three times larger, variously

subtruncate to lunate and subacute to rounded at the upper angles;

peduncles somewhat longer than the leaves, the short cyme subum-

bellate, 2-10-flowered, the branches becoming 1 cm. long and often

nodosely scarred; bracts caducous; sepals linear-lanceolate, glabrous

or nearly, acute, 5-6 mm. long; petals yellow, about 10 mm. long;

filaments and styles puberulent, papillose or apparently sometimes

glabrous (0. Raimondii, which, in Knuth's key, I.e. 107, is written

Raymondii). — In view of the use of the presence or absence of

pubescence on the filaments as a taxonomic character I may err in

drawing this diagnosis to include Knuth's segregate (seemingly

younger plants) but there appears to be no concomitance between

size of leaves, shape of leaflets and filament-character. General

revision in this subgroup as elsewhere is needed. Illustrated,

Bailey, Stand. Cycl. Hort. 4: fig. 2687. F.M. Neg. 32453.

San Martin: Tarapoto, Spruce 3999. Zepelacio, Klug 3555;

3383. Pongo de Cainarachi, Klug 2683. Yera, Woytkowski 35306 —

Huanuco: By Tocache in dark woods, Poeppig (type, 0. excisa).

Rio Chinchao Valley, wooded slopes, 5024; Stork & Norton 9860.

Tingo Maria, Stork & Horton 9467. Cushi, 4834 (det. Knuth, 0.

Raimondii). Muiia, 4046 (det. Knuth, 0. Raimondii). — Junin:

Oxapampa, Soukup 2400. Rio Paucartambo Valley, Killip & Smith

25356, San Jose" to Cueva de la Achira, Raimondi; also, shores

Huallaga, Raimondi (type, 0. Raimondii). — Ayacucho: Huanta to

Rio Apurimac, Killip & Smith 22352. — Cuzco: Valle Lares, Diehl

2503?

Oxalis pachyrrhiza Wedd. Chlor. And. 2: 290. 1857; 231.

Low, acaulescent, subcespitose from a thick fleshy more or less

elongate caudex that is ebulbate but imbricate-scaly with lanceolate

ciliate petiolar remains; petioles elongate, dilated at base and

connate with ciliate stipules; leaflets obcordate, 5-12 mm. long and

broad, shortly petiolulate, sparsely hirtellous only at base; peduncles

582 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

tribracteate above the middle, with ciliate bracts; sepals ovate,

obtuse, glabrous, 4-5 mm. long; corolla apparently yellow.

Puno: Near La Compuerta, on the way from Puno to Arequipa,

400 meters, (Weddell, type). Argentina.

Oxalis parvifolia DC. Prodr. 1: 693. 1824; 166. 0. microphylla

HBK. Nov. Gen. & Sp. 5: 245. 1822(?), not Poir, 1816. 0. hirtella

Willd. ex Zucc. Denkschr. Akad. Muench. 9: 160. 1823-24, not

Jacq. 1794. Acetosella parvifolia (DC.) Kuntze, Rev. Gen. 1: 93.

1891. 0. rufescens Turcz. Bull. Soc. Nat. Mosc. 31. 1: 429. 1858,

at least as to Peru.

Herb with vertical root, the many procumbent stems 1-2.5 dm.

long, 1-1.5 mm. thick, sometimes subligneous, bearing verticels of

2-7 leaves and a few peduncles at rather closely spaced nodes;

leaflets obcordate, about 5 mm. long and broad or broader, or often

smaller, rather deeply incised, more or less hirsute beneath or at

least marginally, glabrate above; petioles 1-2 cm. long or longer;

stipules adnate; peduncles 1-flowered, said typically to exceed 1 cm.

(with pedicel), medially with two linear or setaceous hirsute bracts

2-3 mm. long; sepals oblong or rather ovate, obtuse, pilose, 3-4 mm.

long, the somewhat longer yellow corolla infundibuliform, the petals

slightly retuse; stamens pilose, the trichomes of the styles said to be

spreading-erect; seeds 3-6(?). — 0. filiformis HBK., 169, seems to

be the same. F.M. Neg. 36809.

Lima: Dooryard weed, 3039; upland slope, 806 (both det. Knuth).

Yauli, Stork & Horton 10874- — Junin: Huancayo, Soukup 1894. La

Oroya, gravelly places, Kalenborn 87. — Cuzco: Watkins (det. Knuth,

0. rufescens). — Puno: Streamlet bank on puna, 3,800 meters, Chuqui-

bambilla, Pennell 13399. Colombia to Chile and Uruguay.

Oxalis paucartanibensis Knuth, Pflanzenreich IV. 130: 187.

1930. 0. Ledigii Knuth, Repert. Sp. Nov. 40: 292. 1936.

Glabrous except for a few long soft trichomes on the obovate-

cordate leaflets, a slight pubescence on the 2 mm. long subulate

very acute bracts and often a line of cilia on the lanceolate acute

3-6 mm. long stipules; stems succulent, in type 3-4 mm. thick, 7 cm.

high, leafy toward the tip and with 1-5 peduncles, these 4-15 cm.

long or longer terminating in a few-flowered shortly or obscurely

branched cyme; pedicels 5-10 mm. long, slender; leaflets to 13 mm.

long, 9-13 mm. wide, acute with somewhat rectangular acute sinus

1.5-2 mm. deep, its lobes rounded; sepals 7.5-8 mm. long, 2 mm.

FLORA OF PERU 583

wide, pale green, often purple margined, narrowly lanceolate, acute;

petals 15 mm. long, reddish without, yellowish within, cuneate from

clawed base, lightly erose at retuse apex; stamens and styles puberu-

lent. — Stems sometimes 3 dm. tall. The Rio Blanco plant is nearly

0. peduncularis.

Lima: Rock slides, Huaros, Pennell 14712. Rio Blanco, Killip

& Smith 21565 (det. Knuth); my 2972 (det. Knuth, 0. peduncularis}.

— Junin: Acopolco Canon, Huancayo, 3,700 meters, Ledig 53 (type,

0. Ledigii}.— Apurimac: Prov. Canas, Vargas 11045. — Cuzco: Haci-

enda Churu, Paucartambo Valley, 3,500 meters, Herrera 1390,

type. Acomayo, 3,700 meters, Vargas 206 (slightly more pubescent).

"Occa-occa."

Oxalis peduncularis HBK. Nov. Gen. & Sp. 5: 239. 1822; 188.

Acetosella peduncularis (HBK.) Kuntze, Rev. Gen. 1: 93. 1891.

Somewhat pubescent perennial to 5 dm. tall in flower, the fleshy,

rarely ligneous, glabrous stems 5-7 mm. thick, erect or ascending,

simple, to 2.5 dm. high, more densely foliate apically than below;

leaflets about equal, sessile, obovate, lightly incised, to 12 mm. long,

9 mm. wide, glabrous above, little paler but appressed pubescent

beneath and marginally (or glabrous), rather fleshy, green; petioles

4.5-6 cm. long with free lanceolate pilose stipules 2 mm. long;

peduncles to 3 dm. long, erect, usually pubescent above with a

contracted or biforked cyme of 9-16 flowers, the cyme branches

sometimes 3 cm. long; bracts linear-subulate, acute, pubescent, 1.5-

2.5 mm. long; pedicels 2-10 mm. long, filiform, pubescent; sepals

lanceolate-ovate, acuminate, 4-5 mm. long, glabrous or nearly, often

purple-margined, the narrowly cuneate red-striate yellow petals to

13 mm. long; stamens puberulent; styles glabrous. — Determinations

by Knuth except as noted; type, Quito. Apparently it becomes

ligneous, especially in Huancavelica, and then simulates 0. Herrerae.

F.M. Neg. 35682.

At Mito it was distilled in water and vinegar for coughs.

Lima: Mossy rocky slope, 431. — Junin: Open hillside, Tarma,

Killip & Smith 21813. La Oroya, Kalenborn 90 (det. Knuth, 0.

pubescens). Carpapata, 2,400 meters, Killip & Smith 24343. —

Huanuco: Loose sunny soils, Mito, 1581. Near Pozuzo, rocky clay

banks, 4799. Sunny rocky banks, Chaglla, 3649. Red clay slopes,

Maria del Valle", 3553 (det. Knuth, 0. Herrerae). — Huancavelica:

Clay bank, Salcabamba, 3,250 meters, Stork & Horton 10267. Near

Surucubamba, Stork & Horton 10344. — Apurimac: Clay-gravel banks,

584 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Prov. Abancay, 2,200 meters, GoodspeedExped. 10567 (det. Standley,

0. melilotoides) . Rock crevices, Chincheros to Andahuaylas, West

3721 (det. Johnston, 0. crenata). — Cuzco: Apurimac Valley, Herrera

3077. Urubamba Valley, Herrera 3452. Calca, Vargas 3138 (det.

Standley, 0. tuber osa). Paucartambo Valley, Herrera 1467. Yucay,

Soukup 763. Ecuador. "Chchulcu," "chullco-chullco," "chulco."

Oxalis peruviana Norlind, Arkiv. Bot. 20A, pt. 4: 5. 1926; 48.

O.fruticosa Knuth, Notizbl. Bot. Gart. Berlin 7: 297. 1919, not Raddi.

Procumbent or ascending shrub with very irregular branching

stems several dm. long, glabrous below, puberulent- viscous above,

the subsimple branches densely leafy apically, often naked below,

the petioles more or less persisting, these nearly filiform, 11-13 mm.

long; leaflets petiolate, glaucous, minutely glandular- viscous, the

medial reniform-rotund, 3-4 mm. wide, 2-3 mm. long; peduncles

about erect, 1-3-flowered, to 3.5 cm. long, densely linear-bracted

above; pedicels 1-2 mm. long; flowers to 7 mm. long, the sepals

lanceolate, acute, somewhat viscous puberulent, about 3.5 mm.

long; petals spathulate from clawed base, rounded, yellow; stamens

and styles glabrous, the larger stamens shorter than the sepals.—

Perhaps most nearly related to the Brazilian 0. saxatilis St. Hil., 70,

with slightly larger glabrous leaflets.

Ayacucho: Rio de Lomas near Hacienda Pampa in sparse vegeta-

tion among scattered rocks, 1,000 meters, Weberbauer 5746, type.

Oxalis pes-caprae L. Sp. PI. 434. 1753; 297. 0. cernua Thunb.

Diss. Oxal. 14. 1781.

Acaulescent 1.5-3 dm. high from a subsimple root 3 mm. thick;

petioles to 13 cm. long, glabrous or glabrate, often 10 or 12 together,

the cuneate-obcordate leaflets little paler beneath but long-ciliate

and pilose, to 2 cm. long, 3 cm. wide or wider; peduncles 1-3 dm.

long, more or less pilose, 3-8-flowered, erect or ascending; bracts

many, narrowly linear, acute, pilose, 1-4 mm. long; pedicels 1-2 cm.

long, as the calyces somewhat pubescent; sepals oblong-lanceolate,

acutish, 5-7 mm. long; corolla about 22 mm. long, finally campanu-

late, yellow, the narrowly cuneate petals retuse; smaller stamens as

long as sepals. — Knuth himself uses the name "pes-caprae," I.e.

page 36; name without hyphen originally. Illustrated, Jacq. Oxal.

pi. 6; Lodd. Bot. Cab. 12: pi. 1154.

Junin: La Oroya, Kalenborn 180. — Tacna: Woitschach 106.

Africa; widely escaped or established elsewhere. "Foot-of-Goat"

or "goat-foot."

FLORA OF PERU 585

Oxalis phaeotricha Diels, Bot. Jahrb. 37: 424. 1906; 110.

Many stemmed, early glandular but soon merely rusty hirsute

or glabrate, including the petioles and peduncles and leaflets beneath;

stems to 3.5 dm. tall, erect, or, fide Knuth, ascending or suberect,

sparsely foliate below, shortly branched and densely leafy above;

petioles about 2 cm. long, the adnate basal membranous stipules

2-4 mm. long; leaflets glabrous above, elongate- or spathulate-

obcordate, broadly or narrowly incised, 10-15 mm. long, 6-7 mm. wide,

glaucous, narrowly hyaline-margined; peduncles solitary, 2-4-

flowered, 4-9 cm. long, erect; bracts subulate, 2 mm. long; pedicels

about 1 cm. long, recurving; flowers 14-18 mm. long; sepals linear-

lanceolate, narrowed to tip but obtuse, subhirsute, 8 mm. long, 2 mm.

wide, the conical corolla with cuneate-obovate yellow petals with

brownish-red veins; longer stamens shortly dentate, pubescent as

styles, longer than sepals, or, fide Knuth, glabrous. — The var. glabra

Knuth, with type, designates more slender entirely glabrous plants.

Allied by author to 0. mollis and 0. pichinchensis (0. lotoides) but

ex char, apparently nearly 0. medicaginea.

Junin: East of Palca in grass steppes among scattered shrubs,

3,500 meters, (Weberbauer 2486, type) ; 249.

Oxalis Philippii Knuth, Meded. Rijks Herb. Leiden 27: 67.

1915; 247.

Bulb about 1 cm. in diameter, with lanceolate carinate scales

that are villous-lanate within; petioles 3-4 cm. long, glabrous or

nearly; leaflets cuneate-obcordate, lightly incised, glabrous above,

cellulose-lacunose beneath, more or less long-ciliate marginally, to

9 mm. long, 10 mm. wide, often smaller; peduncles 2-5, about as long

as the petioles, bibracteolate a little below the flowers with obtusish,

or erose lanceolate-ovate bracts 2-3 mm. long; sepals mostly deep

purple tinged apically and long-ciliate toward acutish or obtusish

tip, otherwise glabrous, 5-6 mm. long, equaled by the styles, shorter

than the stamens, both stamens and styles minutely puberulent;

petals broadly cuneate, 15 mm. long, said to be violet-blue.—

Author describes sepals in key as "obtuse." Species with others

apparently very near indeed to 0. eriolepis, to which Weberbauer

plants were referred, once. It seems probable however at this

time that the Peruvian plants at least are more properly referable

to 0. minima.

Peru: Fide Knuth; cf. 0. minima. Chile; Bolivia.

586 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Oxalis picchensis Knuth, Repert. Sp. Nov. 29: 214. 1931.

Subhirsute including the sepals; root slender; stem 2-4.5 dm. long,

about 1.5 mm. thick below, procumbent to the middle and sparsely

leafy then erect, subsimple, becoming densely leafy and 2.5 mm.

thick, the upper part nearly villous the lower pilose and not rarely

bearing bulblets 5 mm. thick as in 0. tuberosa; leaflets pale green,

densely appressed-hirsute beneath, sparsely so above, equal or some-

times the lateral somewhat oblique, sessile, broadly obcordate, to

nearly 2 cm. long, a little broader (or smaller), the sinus broad, to

one-eighth incised, stipules 8-9 mm. long, mostly adnate; peduncles

axillary, often 6 cm. long, with apical umbel of 1-3 bright yellow

flowers, the very acute bracts 4-6 mm. long, the almost villous pedicels

10-15 mm. long; sepals linear or nearly, 7 mm. long, acute, shorter

than the styles, subequaling the longer filaments; petals 13 mm.

long. — Section Ortgieseae (Knuth). Apparently distinct from 0.

Sternbergii, greener, more slender; type not seen; other material

except Balls with smaller leaflets; long filaments pilose in specimens

seen.

Apurimac: Strictly erect but with long prostrate runners, open

grassy area, 3,200-3,650 meters, Ampuy, Goodspeed Exped. 10627;

Vargas 760. — Cuzco: Cerro Piccho, (Herrera 2806, type). Huaila-

bamba, Prov. Paucartambo, Balls B6737. Talus slopes, Asuncion

Bridge, Rio Apurimac, 3,730 meters, Vargas 11029. "Occa-occa,"

"Occa-chchullca."

Oxalis Pickeringii Gray in U. S. Expl. Exped. Bot. 1: 323. 1854;

123. 0. inflate Knuth, Notizbl. Bot. Gart. Berlin 7: 302. 1919;

123.

Erect much branched annual about 1 dm. high, glandular-sub-

viscous except for the petals or the faces of the leaflets; root 2-4 mm.

thick, very tortuous; leaflets obcordate or reniform, to 4 mm. wide

and long, glaucous, cordate or retuse at base, broadly incised apically;

middle petiolule 1-2 mm. long; petioles often 2 cm. long; peduncles

1.5-2 cm. long, cyme branches to 2 cm. long; pedicels 1-4 mm.

long; sepals narrowly lanceolate, acute, 4 mm. long; petals yellow,

10 mm. long to three-fourths connate in an inflated tube; stamens

and styles pubescent, the larger stamens longer than the styles,

shorter than the petals; capsule ovate, 3-3.5 mm. long, with 3-4

seeds in each cell. — This is Knuth's description of his species, which

he noted as probably a synonym of 0. Pickeringii but not certainly;

however the characteristic corolla of 0. inflata seems to be evident

in the Gray plant.

FLORA OF PERU 587

Lima: Near Bafios, Wilkes Exped., type. — Huancavelica: Above

Pisco between Huauyanga and Pampano, stony places with Cacti,

etc., 1,000 meters, Weberbauer 5385 (type, 0. inflata).

Oxalis Poeppigii Prog, in Mart. Fl. Bras. 12, pt. 2: 511. 1877;

90. Acetosella Poeppigii (Prog.) Kuntze, Rev. Gen. 1: 93. 1891.

0. ovalis Ruiz ex Knuth, Notizbl. Bot. Gart. Berlin 7: 294. 1919;

91.

Strict herb to 7 dm. tall, or shrubby with strict terete herbaceous

branches the younger hirtellous with subappressed or early more or

less spreading trichomes; petioles little longer to twice as long as

the leaflets, these oval or oblong-oval or slightly obovate, rounded

both ends or very shortly acute at base, ashy tomentose and nervose

beneath, 2.5-4 cm. long, 1.5-2 cm. wide, peduncles 3-7 cm. long,

tomentulose; inflorescence subumbellate; pedicels 2-5 mm. long;

sepals ovate, rather hirsute, 3 mm. long, 5-7 mm. long in fruit;

corolla flavescent, 5-7 mm. long; filaments glandular at base, the

larger more or less barbate with somewhat spreading trichomes, not

denticulate; styles hispidulous or, in short-styled flowers, glabrous;

capsules ovoid-rotund, glabrous in the Peruvian form, about equaling

the calyx, the cells 2-3-seeded; this is var. segetalis [Poeppig] Prog.,

the leaves pubescent above with long-appressed trichomes and some

obscure and minute stellate ones; var. canescens Prog., Brazilian

plant, has the leaves ashy stellate pubescent above. — 0. hedysaroides

HBK., 57, reported as near as Ecuador, in the typical Colombian

form has ovate leaflets minutely retuse at tip. F.M. Negs. 19204;

32455 (var. segetalis); 29559 (0. ovalis).

San Martin: Tocache, Poeppig (type, var. segetalis). Juanjui,

King 3886 (det. Standley). Zepelacio, King 3463 (det. Standley).—

Junin: Slender shrub with elongate branches, near Perene" Bridge,

Killip & Smith 25274- Rio Seco near La Merced, Soukup 2504-—

Loreto: Balsapuerto, King 2859. Without locality, Ruiz & Pavdn

(type, 0. ovalis). Brazil.

Oxalis polyantha Walp. Nov. Act. Acad. Leop.-Carol. 19:

Suppl. 1: 319. 1843; 130. Acetosella polyantha (Walp.) Kuntze, Rev.

Gen. 1: 92. 1891.

Sometimes a dm. high or higher and the 1-5 more or less palmately

branched stems somewhat ligneous, the trifoliate leaves many at the

branchlet tips and well exceeded by the umbels of 4-10 rose-colored

flowers; petioles glabrous or very sparsely pilose, to 6 cm. long;

588 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

leaflets obcordate, acutely incised one-third to two-fifths, glabrous

or minutely pilose beneath, often 7 mm. long, 6 mm. wide; bracts

more or less pilose, linear-subulate, to 2 mm. long; pedicels filiform,

to 5 mm. long, sepals ovate, acutely acuminate, 3-4 mm. long ex-

ceeded by the larger stamens; petals about 10 mm. long, narrowly

cuneate, retuse-rounded; styles densely puberulent; seeds solitary

in each cell. — The var. peruviana Walp. has smaller leaflets pilose

beneath. 0. trichocalyx Steud. Nomencl. ed. 2. 2: 242. 1841, and

Flora 443. 1856, may, fide Knuth, be the same. F.M. Neg. 26342.

Puno: Toledo, 5,000 meters, Meyen (type, var.). Chile.

Oxalis polyrhiza Knuth, Repert. Sp. Nov. 29: 217. 1931.

Caudex often 5 cm. across, fleshy ligneous, articulately branched,

bearing many little stems 3-10 cm. long mostly 2-3 mm. thick above,

or apically with rather many leaves and peduncles equal in length,

the latter 2-6-flowered the former glabrous, and with subsessile

broadly cordate somewhat oblique leaflets often 9 mm. wide and

long, with obtuse sinus 1-2 mm. deep; petioles often 1 dm. long,

densely white hirsute below; bracts linear-subulate, hirtellous, to

3 mm. long; sepals oblong-lanceolate, obtuse or rounded, 6 mm.

long, glabrous, not rarely reddish, the purple petals twice as long,

broadly obovate, veined, the larger stamens a little shorter.—

Section Carnosae according to the author but ex char, apparently

another race in the 0. peduncularis complex; he did not describe the

filaments.

Cuzco: Hills of Saxaihuaman, 3,600 meters, (Herrera 2726,

type).

Oxalis pseudolobata Knuth, Pflanzenreich IV. 130: 244. 1930.

Acaulescent, 8-17 cm. high, from a subrotund bulb about 7 mm.

thick that is densely covered with sparsely lanate lanceolate scales

about 6 mm. long; peduncles only one from each bulb (as to type,

this without leaves), almost thread-like, to 13 cm. high, 1-flowered,

glabrous as the sepals and pedicel, the opposite linear-subulate acute

bractlets 1-2 mm. long, minutely red-callose at tip; corolla crateri-

form, 13 mm. long, yellow, with retuse rounded, broadly cuneate

entire petals; sepals lanceolate, acute, 4-5 mm. long, tipped like

bracts; stamens and styles hirsutulous. — Name originally written

with a hyphen. The type is mixed with 0. arenaria and is so broken

that the flowers are doubtfully precocious; the leaves may have been

detached, and lost; otherwise except for slight differences, especially

FLORA OF PERU 589

in measurements, this seems to be essentially 0. punensis to which

I would refer it, that being a preferable name, same date. 0. lobata

Sims of Chile, 243, is quite different, its bulbs densely fibrous, its

sepals obtuse.

Cuzco: Paucartambo Valley, Hacienda Churu, 3,500 meters,

Herrera 1398 (part), type.

Oxalis ptychoclada Diels, Bot. Jahrb. 37: 425. 1906; 187.

Glabrous, except the narrow ciliate-pilose short stipules and leaf-

lets beneath, sometimes 1.5 dm. long from a much forked root to

5 mm. thick with stout woody tortuous rootlets, the stems forked or

branched from the base, to 7 cm. high, 4 mm. thick, procumbent-

erect, ligneous, densely scaly below with the petiole bases and

stipules, the leaves rosulately congested with the peduncles at the

tips; leaflets obcordate, thin-margined, (6) 7-10 mm. long, 5-8 mm.

wide, lightly incised with broad sinus, glaucous and glabrous above,

pilose especially near midnerve beneath; petioles 1-5 cm. long;

peduncles few, 5-7 cm. long, the 4-10 flowers on lax filiform pedicels

about 1 cm. long; bracts very acute, 2-3 mm. long; sepals oblong-

lanceolate, acute, (4) 6-7 mm. long, the entire apically rounded

petals 12-14 mm. long, reddish-brown striate; longer filaments

subglabrous, dentate, 6-7 mm. long, the styles puberulent. — Knuth

designates both Weberbauer specimens as "type," and distinguishes

the species from his 0. paucartambensis by the shortly forked instead

of umbelliferous peduncle, and from 0. peduncularis by fewer flowers,

shorter forked peduncle!

Junin: Tarma, in rocky outcrops, 3,000 meters, (Weberbauer

2357, type); 176; 180. La Oroya, on rocks, 3,700 meters, (Weber-

bauer 2568, fide Diels).

Oxalis puberula Phil. Anal. Univ. Chile 2: 332. 1865; 122.

A densely leafy little annual much like 0. micrantha but obviously

glandular-puberulent even to the narrowly lanceolate sepals, and

the flowers to about 10 mm. long; leaflets usually 6 mm. long and

broad; petioles to 5 cm. long, nearly as long as the furcate several-

flowered peduncles; bracts setaceous, 1 mm. long; pedicels 5-10 mm.

long; stamens pilose; seeds 2 in each cell. — If the following determina-

tion was correct it extends the species north from Atacama, Chile.

Areq'uipa: Near Mejia, (Gunther & Buchtien 258, det. Bruns).

Chile.

590 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Oxalis punensis Knuth, Pflanzenreich IV. 130: 246. 1930.

0. quispicanchensis Knuth, Repert. Sp. Nov. 29: 218. 1931.

Acaulescent, glabrous, a few cm. to 2 dm. tall or taller, from a

rounded bulb about 1 cm. in diameter, the many pale brown scales

lightly lanate; peduncles and leaves few, the former 3-20 cm. long,

1-flowered, 10-17 mm. below the apex with 2 basally connate acutish

or obtusish ovate bractlets about 2 mm. long, the latter with lax

petioles 3-18 cm. long; leaflets broadly obcordate, obtusely 2 mm.

excised, to 1 cm. long, 1.5 cm. wide, cellulose-lacunose below,

marginally dark callose-striate; sepals lanceolate, acutish, about

6 mm. long with 2 slender calli at tip; petals broadly cuneate, retuse-

rounded, entire, pale purple, forming a conical corolla 18 mm. long;

styles of long styled flowers longer than sepals. — Seems too much

like 0. oreocharis; in the key the author describes the petals as

yellow! Cf. also 0. pseudolobata which in all probability should be

included but more material from type localities desired. Since

leafy the Apurimac specimen is cited here, but it has the smaller

flowers of 0. pseudolobata and resembles closely the Bolivian 0.

platylepis Wedd., 262, which ex char, has glabrous bulb scales, shorter

sepals, maybe doubtful characters. Ex char. 0. quispicanchensis

seems very doubtfully to be specifically distinct unless these differ-

ences are taxonomically sound: bulb hirsute-lanate, leaflets 6 mm.

long, 7-8 mm. wide, obovate with wide sinus, bracts 1 or 2, filiform,

corolla white, 16 mm. long, sepals 5-5.5 mm. long, stamens sparsely

ciliate, all in my experience, admittedly slight, reasonable variations

for any specific unit.

Cuzco: Marcapata, 3,100 meters, grassy places in woods, Weber-

bauer 7808 (type, 0. quispicanchensis). — Apurimac: Sandy clay

canyon, Andahaylas, 3,050 meters, Stork & Horton 10724- — Puno:

Occa Pampa, Prov. Huancane, Shepard 69, type, Gray Herb.

Oxalis Regnellii Miq. Linnaea 22: 545. 1849; 205. Acetosella

Regnellii (Miq.) Kuntze, Rev. Gen. 1: 93. 1891.

Acaulescent, 1-2.5 dm. high in flower; rhizome about 5 cm.

long, densely imbricate-squamose and with tubercles 7 mm. thick;

leaves rosulate, only 4-9 or few, the glabrous petioles 10-15 cm. long,

not dilated at base, the leaflets deltoid, even very broadly so, minutely

incised or subentire with divaricate obtuse lobes, to 2.5 cm. long,

twice as broad, often smaller, pale green, glabrous above, sparsely

or scarcely pilose beneath; peduncles 10-15 cm. long, only 1-4,

glabrous as pedicels (2-4 cm. long) and calyces, the umbel simple,

FLORA OF PERU 591

3-7-flowered; sepals linear-lanceolate, minutely reddish-bicallose at

acute tip, 5-5.5 mm. long, the larger stamens longer and as the styles

hirsute; corolla white, at least 4 times longer, with oblanceolate

rounded apically entire petals; capsule columnar, nearly glabrous,

with many seeds in each cell. — Illustrated, Knuth, I.e. fig. 16, page

213, and Norlind, Arkiv Bot. 14, pt. 6: pi. 1. F.M. Neg. 35688.

Loreto: Middle Ucayali, (Tessmann 5469, det. Knuth). Brazil

to Argentina and Paraguay.

Oxalis rigidicaulis Knuth, Repert. Sp. Nov. 40: 290. 1936.

Stems densely strigillose-pilose even to the trailing base where

leafless and rooting, 2 or 3 dm. long to 3 mm. thick at base, simple,

more or less erect above, the alternate leaves finally remaining only

on the upper third and often 1.5-2 cm. remote, their slender glabrous

petioles 2-3 cm. long; leaflets subequal but the lateral mostly strongly

oblique, all sessile, green, to 14 mm. long, 16 mm. wide, very obtusely

incised to 3 mm., the semiovate lobes obtuse; stipules none or minute;

peduncles often 4 cm. long not rarely with two branches of 3-6

yellow flowers; bracts acute, 1 mm. long; pedicels 3-7 mm. long;

sepals 4 mm. long, linear-lanceolate, acutish, the spathulate-obovate

petals nearly to quite twice as long; long-styled flowers with styles

exceeding sepals; filaments glabrous. — Section Ortgieseae, according

to the author, who describes the plant as completely glabrous; the

lower part of the ascending stems are more or less coated with soil;

the type appears abnormal, probably a state of 0. Ortgiesii Regel.

Junin: Dos de Mayo, Pichis Trail, 1,800 meters, Killip & Smith

25873, type. — Huanuco: Moist open slopes, 3,100 meters, Saria-

pampa, Woytkowski 297.

Oxalis San-Miguelii Knuth, Pflanzenreich IV. 130: 432. 1930.

0. machupicchuensis Knuth, Repert. Sp. Nov. 40: 289. 1936.

Perennial, the tomentose stems about 3 dm. high, simple or a

little branched at base where about 5 mm. thick, medially and

below clothed only with the very acute linear-subulate 4 mm. long

free stipules; petioles 1.5-3.5 cm. long, densely lanate- tomentose;

leaflets sessile, equal, densely sericeous, lanate beneath, sparsely

pilose at maturity above, broadly ovate or obovate, with more or

less cuneate base, 17-20 mm. long, about 11 mm. wide, the shallow

sinus 1-2 mm. deep; peduncles including the bibranched 2-6-

flowered umbel to 7 cm. long, the umbel branches often 4-10 mm.

long; bracts subulate, 1-4 mm. long, hirtellous; pedicels filiform,

592 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

2-6 mm. long, glabrous as the oblong-lanceolate obtuse pale green

sepals, these 7 mm. long; petals narrowly obovate with long claw,

beautiful yellow, 12-15 mm. long; stamens and styles spreading-

hirsutulous. — Allied by author to 0. melilotoides Zucc. Knuth

distinguishes his second species from those of section Ortgieseae by

the "obovate leaflets, lanuginose indument, manifestly bibranched

cymes and obviously erect simple stems"; a similar plant from La

Paz, Bolivia, has been determined by Johnston 0. crenata Jacq.

but since the monographer includes that species in 0. tuberosa I

do not adopt here the Jacquin name, unknown to me.

Cuzco: Valley of the San Miguel, Machu-picchu, 2,400 meters,

Herrera 2044, type. Valle de Urubamba, Machu-picchu, 2,200

meters, Herrera 3315 (type, 0. machupicchuensis). Bolivia?

Oxalis sepalosa Diels, Bot. Jahrb. 37: 426. 1906; 186. 0.

Buchtienii Bruns, Mitt. Inst. Allgem. Bot. Hamb. 8: 51. fig. 7. 1929.

Habit of the related 0. megalorrhiza but in flower 2.5-3.5 dm.

high and the 7-15 cm. tall stems especially beneath the apical

foliage densely scaly with the brown foliaceous stipules; leaflets

sometimes arachnoid beneath, often 2.5 cm. long, 3.5 cm. wide and

one-fourth incised with broad sinus; petioles 12-15 cm. long;

peduncles 12-20 cm. long with pale foliaceous ovate acute bracts

to 7 mm. long and glabrous, 6-20-flowered umbels (in age 2-3-

forked), the pedicels 10-17 mm. long; sepals unequal, the larger

broadly ovate or subdeltoid, obtuse or rounded, pale margined,

10-12 mm. long, 7.5-9 mm. wide, two narrowly oblong, 2 mm. wide;

corolla wide-open, the yellow rounded entire petals 2 cm. long (or

late flowers much smaller); larger stamens glabrous, longer than

sepals, styles minutely puberulent. — Bruns compared his species

with 0. dolichopoda; it is probably restricted to the lomas and (if

not an earlier described Chilean species) related as Diels noted to

0. megalorrhiza but seems to be well distinct by the large stipules

and almost foliaceous bracts. The Bruns plant, nearly past flowering,

has open forked inflorescence, flowers only 6 mm. long, differences,

it seems to me, from Diels plant as described due to age. F.M.

Negs. 29561; 20923 (0. Buchtienii).

Arequipa: On rocks in the lomas, 400 meters, Weberbauer 1519,

type; 94; 146. Loma north of Atiquipa under bushes, Worth &

Morrison 15669 (det. Johnston, 0. lomana); also west of Atiquipa,

15656. East of Chala, Worth & Morrison 15608 (det. Johnston,

FLORA OF PERU 593

0. lomana). Ravine near Mejia, (Gunther & Buchtien 254, type,

0. Buchtienii). Cachendo, (Gunther & Buchtien 255, det. Bruns).

Oxalis solarensis Knuth, Notizbl. Bot. Gart. Berlin 7: 306.

1919; 201. 0. arequipensis Knuth, Pflanzenreich IV. 130: 188.

1930. 0. ornata Phil. Fl. Atac. 13. 1860, at least as to Peru.

Perennial, with a thick shortly branched vertical rhizome 10-15

mm. across above, ligneous, bearing many leaves and peduncles at

the neck, the latter sparsely pilose as the petioles and with an irregular

dichotomous cyme of 4-14 flowers; leaves to 9 cm. long, fleshy, the

equal leaflets cuneate- or deltoid-obcordate, often a little oblique,

subretuse but obscurely incised, to 2 cm. long, 2.5 cm. wide, mostly

smaller, arachnoid and hyaline papillose beneath, glabrate above;

bracts 2-3 mm. long, acute; pedicels 3-10 mm. long, glabrous;

outer sepals evidently triangular-dilated at base, rounded at tip,

4-5 mm. long or a little longer in age, 2.5 mm. wide, the inner

narrower; corolla three to four times longer, yellow, the petals

spathulate-lingulate; styles 6 mm. long, the larger stamens shorter. —

Apparently with reason Johnston noted in herb, "perhaps not distinct

from 0. megalorrhiza." The Huanuco specimen, entirely out of

range, has smaller flowers, smaller leaflets.

Lima: Mount Morro, Chorillos, at lower edge of lomas, Weber-

bauer 5690, type. Above Chosica, Horton 10991 (det. Johnston).

Below Matucana, stony outcrops, Weberbauer 5201. Eulalia Valley,

Goodspeed & Stork 11498 (det. Johnston). — Huanuco: Common on

adobe walls, also on open grassy slope, 2,400 meters, Woytkowski

261 (doubtful). — Arequipa: Yura, stony places with small shrubs,

Weberbauer 6841 (det. Knuth). Tingo, Pennell 13117 (det. Knuth,

0. carnosa}. Rock ledges above Arequipa, Pennell 13192, type,

0. arequipensis. Rocky ravines, Chachani Mountains north of

Arequipa, Hinkley 27 (det. Knuth, 0. ornata}. North of Atiquipa,

Worth & Morrison 15631 (det. Johnston., 0. megalorrhiza). "Chunco"

(Hinkley).

Oxalis spiralis R. & P. ex G. Don, Gen. Syst. 1: 755. 1831;

107. 0. pubescens HBK. Nov. Gen. & Sp. 5: 240. 1822, fide Knuth,

not Stokes, 1812. 0. bifida Willd. ex Zucc. Denkschr. Akad. Muench.

9: 162. 1823-24 (name only), not Thunb. 1781.

Succulent-herbaceous, 3-5 dm. high, early erect or ascending,

or trailing in age, little branched, sparsely lanate pubescent including

the leaves below and peduncles, the latter not crowded apically,

594 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

6-12 cm. long and bearing a bifurcate 6-12-flowered cyme; leaflets

equal, sessile, obcordate, the tips openly emarginate, often 2 or about

2.5 cm. long, 1.5-2 cm. wide (in 0. pubescens, 1 cm. by 1 cm., pubes-

cent both sides), glabrate above; petioles to 7 cm. long; cyme

branches 2-5 cm. long; bracts linear-lanceolate, acute, 2-4 mm. long,

densely pubescent as the 3-8 mm. long pedicels; sepals 5-6 mm.

long, obtusely acuminate, often purplish and glabrate above; petals

about 6-9 mm. long, spathulate, pale yellow; longer filaments as

styles puberulent, a little longer than the sepals; capsule glabrous,

ovoid-cylindric with many (3-4 in 0. pubescens) seeds in each cell.

Type locality of 0. pubescens "Peru." There is a var. glandulosa

(Knuth) Macbr. Candollea 6: 9. 1934 (cyme glandular) in Colombia

and Venezuela. F.M. Neg. 35684 (0. pubescens).

Lima: Raimondi. — Piura: Chorro Blanco, succulent stems ramb-

ling for 1 meter, Stork 11408 (det. Johnston, 0. crenata). — Cajamarca:

Raimondi. — Huanuco: San Rafael, Soukup 2242, Puente Durand,

Stork & Horton 9585 (det. Standley, 0. melilotoides)', also 9379,

river bed south of Huanuco. Above Pillao, Woytkowski 34030.—

Ayacucho: Ccarrapa, 3,000 meters, Killip & Smith 22277 (det.

Killip). Pampalca, Killip & Smith 22255 (det. Killip).— Cuzco:

Urubamba Valley, Cook & Gilbert 952. Ollantaytambo, (Cook &

Gilbert 600). Machu-Picchu, (Cook & Gilbert 851). "Pillahauta,"

Pennell 13986 (det. Knuth). Paucartambo Valley, Herrera 1814.—

Puno: Sicuani, (Cook & Gilbert 152). Limbani, Prov. Sandia,

Metcalf 30427 (det. Leonard). Without locality, Ruiz & Pawn,

type ; Bonpland (type, 0. pubescens) . Chile and Argentina to Panama.

"Chuco" (Quechua, Stork and Horton).

Oxalis Spruceana Prog, in Mart. Fl. Bras. 12, pt. 2: 481. 1877;

53. 0. frutescens R. & P. ex G. Don, Gen. Syst. 1: 754. 1831, not

L. 0. sublignosa Knuth, I.e. 95 (name change for 0. frutescens).

Suffruticose, to 3 meters high, the stems squarrosely much

branched above, the densely leafy branches arcuate-spreading-erect,

minutely but densely gray-puberulent with crisped trichomes, this

indument extending to sepals and capsule (or these even sericeous),

sparse only on the upper side of the leaflets which are lanceolate,

acute or obtusish, the medial 5-10 mm. long petiolate, usually 4-6

cm. long, 1-2 cm. wide, the lateral smaller; petioles 1.5-4 cm. long;

peduncles many, 2-4 (6) cm. long, umbellately 3-8-flowered; bracts

subulate, to 2 mm. long; pedicels 1-3.5 mm. long; sepals lanceolate,

acute, about 3 mm. long, the pale yellow (often with 2 orange-red

FLORA OF PERU 595

basal marks) narrowly cuneate petals 5-7 mm. long; larger stamens

puberulent, longer than sepals, the globose capsule scarcely as long

and with 1 seed in each cell.— Knuth describes stamens of 0. sub-

lignosa as glabrous but they are puberulent in, type. 0. rhombifolia

Jacq., known from Mexico to Colombia and Venezuela, is sparsely

branched with erect branches, has somewhat longer sepals. F.M.

Negs. 26348; 26353 (0. sublignosa).

San Martin: Tarapoto, Spruce 3919, type; Ule 6443. Without

locality, Poeppig. — Junin: Rio Perene", Killip & Smith 25200;

25418. Oxapampa, Soukup 2402. La Merced, openly wooded slope,

5334.— Huanuco: More open rocky slopes, Pozuzo, 4607 (det.

Knuth); Ruiz & Pavdn (type, 0. frutescens). North of Tingo Maria,

gravelly bank, Stork & Horton 9450 (det. Standley). — Cuzco:

Echarate, Prov. Convention, edge of forest, Goodspeed Exped. 10502

(det. Standley).

Oxalis Staff ordiana Knuth, Repert. Sp. Nov. 38: 195. 1935.

Similar to 0. megalorrhiza and 0. paucartambensis ; petioles 1-3.5

cm. long, glabrous with broad basal marcescent stipules about 3 mm.

long; leaflets glabrous or at margins and beneath on veins sparsely

ciliate-pilose, 1-2 mm. incised, 6 mm. long, 5 mm. wide, subsessile,

ecallose; peduncles 1-3-flowered, 1.5-3 cm. long, in herb, nearly

filiform as the 1 mm. long bracts; pedicels 6-8 mm. long; sepals obtuse

or nearly retuse, to 5 mm. long; petals orange, beautifully purple-

nerved, long-cuneate, to 13 mm. long; larger stamens a little longer

than the sepals. — Affine 0. paucartambensis according to the author,

who did not describe the filaments!

Cuzco: In rocks, 3,000-4,000 meters, (Stafford, Herb. Kew, type).

Oxalis Sternbergii Zucc. Denkschr. Akad. Muench. 9: 156.

1823-24 and Abh. 1: 220. 1829-30; 171. 0. Commersonii Pers. Syn.

PI. 1: 519. 1805 as to Peru plant. 0. petrophila Knuth, Notizbl.

Bot. Gart. Berlin 7: 300. 1919; 159. 0. longicalyculata Knuth,

Repert. Sp. Nov. 23: 279. 1927; 170. 0. colquipatensis Knuth,

Pflanzenreich IV. 130: 433. 1930.

More or less sericeous villous even the sepals minutely, these

linear-lanceolate or subligulate, obtusely acute, 7-10 mm. long;

stems procumbent-ascending, to 2.5 dm. long or longer, about 2.5

mm. thick below where defoliate and in old plants ligneous, with

the petiole bases more or less persisting, densely foliate above the

filiform or slender petioles 3-7 cm. long little exceeded by the 1-3-

596 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

flowered, rarely irregularly 3-5-flowered peduncles, these with 2 subu-

late bractlets one or two cm. below the calyx, there refracting in fruit;

leaflets broadly obcordate, the lateral often oblique, all slightly wider

than long, rather acutely emarginate (1-2 mm. or so), usually 10-12

mm. wide, sometimes smaller, usually less pubescent above than

below; stipules membranous free only at the scarious tips, these

sometimes 2-3 mm. long; petals yellow or red dotted toward base,

15-17 mm. long; stamens and styles pubescent, sometimes minutely

and obscurely. — If I may judge from an earlier observation and now

from the negative the plant of Zuccarini is not the same as that of

Persoon; but maybe the Munich Haenke specimen (actual type,

herb. Sternberg) was not correctly named. In any case I see no

fundamental differences between the three species of Knuth cited.

0. Commersonii apparently has much more deeply incised usually

smaller leaflets, shorter sepals, whitish flowers. F.M. Neg. 19211.

Used according to Herrera, for treatment of "thrush," an oral

fungous infection.

Cuzco: Hacienda Churu, Valle de Paucartambo, Herrera 101+3

(det. Killip, 0. petrophila). Cuzco, 3,000-3,600 meters, Herrera

(type, 0. longicalyculata) . Cerro de Colquipata, rocky cliff, 4,100

meters, Pennell 13737 (type, 0. colquipatensis) . Between Pisco and

Paucartambo, 4,100 meters, Weberbauer 6918 (type, 0. petrophila).

Acanacu, 3,500 meters, among shrub, West 7053 (peduncles 2-5-

flowered, det. Johnston, 0. pubescens). Without locality, Soukup

1184- Without locality, Haenke, type. Bolivia? "Occa-occa,"

"occa-chchullcu. "

Oxalis subintegra Knuth, Notizbl. Bot. Gart. Berlin 7: 298.

1919; 133.

Scandent, the few stems about 1 meter long, 3-5 mm. thick,

glabrous, the rather remote short branches scarcely ligneous, their

internodes 3-7 cm. long, the nodes slightly enlarged; branchlets

slenderer, leafy, to 8 cm. long; leaves rosulate or alternate, on gla-

brous or somewhat pilose petioles to 3 cm. long; leaflets rhomboid-

or lanceolate-oblong, cuneate at base, lightly incised with rounded

lobes, beneath and marginally villous, subglabrous above, about 3 cm.

long, 10-12 mm. wide, the 1 mm. long petiolules hirsute; stipules to

9 mm. long, villous, nearly completely adnate; peduncles erect, about

5 cm. long, glabrous as pedicels, bracts, and calyces, the first 5-12

mm. long, the second acute to 3 mm. long, the last 8 mm. long, the

sepals narrow and very acute; flowers more or less nodding on two

FLORA OF PERU 597

branches, orange yellow or violet (Mathews spec.), to 17 mm. long,

the cuneate petals retuse; larger stamens little shorter than calyx,

glabrous as the styles. — F.M. Neg. 26352 (Mathews).

Piura: In shade of shrubs and trees, Prov. Huancabamba, 2,900

meters, Weberbauer 6092, type. — Amazonas: Chachapoyas, Mathews

3015. — Huanuco: Huacachi, forming a tangled mass, 4092 (det.

Knuth). Ecuador.

Oxalis tabaconasensis Knuth, Notizbl. Bot. Gart. Berlin 7:

299. 1919; 134.

Procumbent or scandent, sometimes about 6 dm. long, the some-

what sulcate stems about 2 mm. thick, pubescent especially on the

leafy 2-10 cm. long branches; leaflets all 1 mm. long, petiolulate,

papyraceous, green but rather densely and softly appressed pilose,

often 15 mm. long, 9 mm. wide, cuneate oblong-obovate, the rounded

tips 1 mm. emarginate; petioles 2-2.5 mm. long, pilose-hirsute, with

1 mm. long soon deciduous stipules; peduncles about 2 cm. long, pilose

below in type, glabrous above as the pedicels, bracts, and calyces,

the 1-5-flowered cyme irregular; bracts linear-subulate, to 3.5 mm.

long; pedicels 5-12 mm. long; sepals pale green, elongate-oblong,

obtuse, 7 mm. long slightly exceeded by the larger glabrous stamens,

the short-styled flowers with styles glabrous; petals orange-yellow

or yellow, 12-15 mm. long, broadly dilated from a narrow base.—

Unknown to me but apparently the following Huanuco specimens

belong here or possibly to 0. mollis, but the hirsute stipules are free,

not membranous.

Cajamarca: Edge of grass steppe formation above Tabaconas,

Prov. Jae*n, (Weberbauer 6305, type). — Huanuco: Exposed rock

bank, Muria, 4281 (det. Knuth, 0. lotoides). Sariapampa, Woyt-

kowski 34297. Pillao, Woytkowski 34161.

Oxalis Tessmannii Knuth, Repert. Sp. Nov. 23: 141. 1926; 83.

Arborescent, attaining 2 meters, the stem 3 cm. thick at base,

simple to about 1 and a half meters where squarrosely branched, the

younger densely leafy branchlets puberulent; lateral leaflets sub-

sessile, the medial with slender petiolule about 2 cm. long, all lanceo-

late, gradually and acutely acuminate, entire, glabrous or beneath

on nerves minutely puberulent, the lateral 7-8 the medial 11 cm.

long, 3-3.5 cm. wide; peduncles scarcely 1 dm. long, glabrous, with

apical irregular but not bibranched cyme of 6-8 flowers, the pedicels

unequal in length, filiform, often 2 mm. long; sepals 2.5 mm. long,

598 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

ovate, obtuse, the yellow petals one and a half times longer.—

Apparently too near 0. polymorpha Zucc., 78, of Brazil or related

"species" as 0. suffruticosa Salzm., 80, as also 0. Macbridei, 431.

A revision of the genus by an able taxonomist is much needed.

Loreto: Mouth of the Santiago in flooded highwood at edge of

the Rio Maranon, (Tessmann 4484, type).

Oxalis tuberosa Molina, Sagg. Nat. Chile 3: 109. 1782; 109.

Acetosella tuberosa (Molina) Kuntze, Rev. Gen. 1: 92. 1891. 0.

crenata Jacq. Oxal. 27. 1794, fide Knuth.

Rhizome branches often tuberiform the tubers even 4 cm. long,

3 cm. thick (or nearly twice as large in cultivation) and bearing a

number of triquetrous scales; stems densely pubescent, erect, about

3 dm. high, simple or little branched, densely foliate especially at

and above the middle; petioles 7-10 cm. long, like the much longer

peduncles sparsely hirsute or glabrate; leaflets equal, pale green,

more or less densely hirsute both sides, fleshy, sessile, about 2.5 cm.

long, 22 mm. wide, the very broadly rounded tips to one-eighth

incised, the lobes rounded; stipules to 15 mm. long, adnate; peduncles

often binate, to 17 mm. long, bearing regular or nearly regular 5-8-

flowered umbel of pale yellow flowers usually about 15-22 mm. long

on villous pedicels 7-20 mm. long; bracts very long; larger filaments

pubescent; capsule narrowly oblong. — Last flowers in cymes may be

smaller, as in my 2090. Determinations by Knuth except as noted.

W. H. Hodge, Journ. N. Y. Bot. Gard. 47: 214-224. 1946, has

given a good account with excellent photographs of this species (and

Arm, Tropaeolum tuberosum) commonly cultivated (to at most 3,800-

4,000 meters, fide Weberbauer) for its starchy tubers, which, rich

in crystals of calcium oxalate, must be cured several days by leaving

in sunlight when they may be eaten. They are usually boiled as

potatoes. When they are to be stored a chuno, called cjaya, is

prepared in the manner used for that of Solarium tuberosum, described

by Herrera, Sinop. Fl. Cuzco 1: 284. 1941; also by Hodge, I.e. 218.

The former listed "Lluchcho-occa" as well as "Sapallu," "Chacha-

pea," "Pauccar" and "Mestiza" as native names for tubers colored

red, yellow, gray, red, and black. Hill, Bull. Misc. Inf. Kew 169-

173. 1939, found the color differences of tubers in Bolivia associated

with long-styled flowers (yellow), medium (white), short (red).

Cajamarca: Trujillo to Cajamarca, Raimondi. — Lima: Above

Matucana, Weberbauer 5276; also my 108; 2955. — Huanuco: In

underbrush, 2090 (det. Knuth, 0. melilotoides). Among rocks,

FLORA OF PERU 599

Yanahuanca, 1231. Erect from mossy forest floor or pendent from

trees, Tambo de Vaca, 4417. Mossy bunch grass slope, to 1.5

meters high, Chasqui, 3287. — Jtmin: Stream bank, Tarma, Killip

& Smith 21917 (det. Standley). — Huancavelica: Near Cordova,

Metcalf 30271 (det. Leonard). — Apurimac: Part shade in forest,

Ampuy, Goodspeed Exped. 10606 (det. Standley). — Cuzco: Machu-

picchu, Herrera 3318. Sheltered ravine, Acanacu, West 7054 (det.

Johnston, 0. pubescens). "Pillahuata," Vargas 13 (det. Standley).

Huaros, Pennell 14712. Cuzco, Herrera 434,' 454- Hacienda Churu,

Herrera 3138. — Puno: Commonly cultivated at Cuyo-cuyo (Weber-

bauer, 295). Above Sicuani, Cook & Gilbert 152; 152a (det. Knuth,

0. carnosa). Bolivia to Colombia. "Occa-occa," "bueno grille."

Oxalis urubambensis Knuth, Repert. Sp. Nov. 29: 216. 1931.

Stem simple, ligneous, about 4 dm. high, scarred below every

5-7 mm. by the fallen petioles, foliate and densely yellowish tomen-

tose above; stipules linear-subulate, acute, deciduous; petioles 3 cm.

long, hirsutulous, soon glabrate; leaflets sessile, equal, broadly ovate,

more or less oblique and cuneate at base, 2 mm. long, 1.25 cm. wide,

the sinus 2-3 mm. deep, above sparsely beneath densely appressed

pilose or subtomentose; peduncles 2-4, densely puberulent or gla-

brate, 5 cm. long, apically forked with many flowered branches 1.25

cm. long; bracts 2-4 mm. long; sepals lanceolate, pale green, glabrous,

5 mm. long, the long-clawed petals twice as long; smaller stamens

half as long as larger, all glabrous; styles a little shorter than the

small stamens. — Section Myriophyllum Knuth according to the

author but ex char, comparable to 0. San-Miguelii or possibly 0.

Herrerae.

Cuzco: Maquina, Valley of the Urubamba, 2,050 meters, ( Herrera

2699, type).

Oxalis velutina Diels, Bot. Jahrb. 37: 425. 1906; 86.

Squarrosely branched shrub, 1 meter high, the ligneous outer

branches 4-5 mm. thick, densely tomentose above as the petioles

(4 cm. long) and peduncles, these axillary in the many leaves toward

the tips but not apical, 4.5-5.5 cm. long, the lower part of the

branches densely marked by scars of fallen petioles; leaflets ovate,

the lateral little smaller, about 3.5 cm. long, 2 cm. wide, rounded

both ends or even retuse at base, not rarely dotted or spotted with

white, puberulent above, lanuginose- tomentose beneath; medial

petiolule 7-10 mm. long; pedicels, bracts and calyces densely villous-

600 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

i- i

tomentose, the first 3-5 mm. long, the second 2 mm. long, the 5-8

umbellate flowers 10-12 mm. long; sepals lanceolate, acutish, 5.5-7

mm. long, the corolla twice as long, the broadly spathulate petals

yellow; larger stamens scarcely longer than the sepals, minutely

puberulent, the rounded or slightly acuminate black capsule about

as long. — Seems, ex char., not specifically distinct from 0. hypopilina

and 0. psoralioides HBK., 87, of Colombia, which as to type has

emarginate slightly smaller leaflets, shortly bifid cymes. Diels

remarked: new unless 0. hypopilina but leaflets pubescent above,

nerves prominent, the most tomentose of all Peruvian species. 0.

loxensis Knuth, 86, of southern Ecuador is similar but the longer

leaflets are obtuse or less rounded apically, the sepals more or less

acuminate, the petals to 2 cm. long, the stamens tomentulose.

Cajamarca: In open shrub-Bromeliad formation, 1,500 meters,

below Santa Cruz, (Weberbauer 4126, type); 94; 189.

Oxalis villosula Knuth, Repert. Sp. Nov. 38: 195. 1935.

Perennial, with long oblique root and one to few simple erect

rather fleshy stems to 4 mm. thick, 2-4 dm. high, glabrous below

but densely lutescent-villous above the middle; leaflets subequal,

oblong-obovate, cuneate at base, the very broad sinus 2 mm. deep

with rounded or obtuse lobes, to 15 mm. long, 10 mm. wide, sub-

glabrous above, densely tomentose-sericeous beneath; petioles to

3 cm. long, dilated, tomentose; peduncles as inflorescence glabrous,

5-7 cm. long, in herb. 1.25 mm. thick, the cyme 5-7-flowered ; lower

bracts 5-7 mm. long, subulate, acute; pedicels to 15 mm. long, nearly

filiform; sepals lanceolate, drying nearly membranous, 3.5-5 mm.

long, 1.25-1.5 mm. wide, the yellow-orange petals two and a half

times longer, oblong-lanceolate; stamens all glabrous, the larger

equaling the sepals, the smaller the puberulent high-connate styles. —

Section Ortgieseae (Knuth).

Cuzco: Machu-picchu, Valle del Urubamba, 2,100 meters,

(Herrera 2953, type).

Oxalis Weberbaueri Diels, Bot. Jahrb. 37: 425. 1906; 189.

Two to 3.5 dm. tall in flower the more or less densely scaly

stems about 5 mm. thick, 4-14 cm. high, probably firm-succulent

in life but described as ligneous, the leaves and peduncles apically

crowded, the former with petioles to 4 cm. long, the leaflets some-

what appressed pubescent beneath, often about 10-15 mm. long,

7-12 mm. wide, openly emarginate, the latter usually much longer,

FLORA OF PERU 601

glabrous or sparsely pilose as the nearly filiform 5-10 mm. long

pedicels, these as many as 15 in a tardily branched umbel; sepals

6 mm. long, gradually acuminate to obtusish tip; corolla long-narrow-

ed to base, the yellow petals glabrous, retuse, entire, 18-20 mm.

long; filaments subglabrous, styles glabrous. — Seems to be perhaps

too near 0. peduncularis but the one specimen seen has glabrous

filaments. The type was probably prepared by the alcohol method,

which may account for its brittle ligneous quality. The author

compared it to the Chilean 0. ornata with unequal sepals.

Piura: Above Palambla in grass steppes with scattered shrubs,

Weberbauer 6023. — Ancash: Pampa Romas in close grass-shrub

formation, (Weberbauer 3205, type); 170; 173. In rocky places,

Huaraz, 2,200 meters, (Weberbauer 3002, probably, Knuth). "Paza

tullmasch."

Oxalis Williamsii Knuth, Repert. Sp. Nov. 38: 195. 1935.

Suffrutescent, entirely glabrous, the subsimple erect ligneous

brown stem 5 dm. tall, 3-3.5 mm. thick below; leaves scarcely

numerous, the subequal leaflets ovate-lanceolate or lanceolate, more

or less cuneate at base and narrowed to the acute tip the middle one

(petiolule to 2.5 cm. long) to 9 cm. long, 3.5 cm. wide, the 2 mm.

petiolulate lateral little smaller, the common petiole to 1 dm. long;

peduncles axillary in upper leaves, 3.5 cm. long, 0.5 mm. thick,

bibranched apically, the 2 cm. long branchlets not rarely biparted,

scarred by fallen flowers and with only a few open flowers near tips;

pedicels filiform, 3-3.5 mm. long; sepals 2.5-3 mm. long, ovate-

lanceolate, acute, finally ovate; petals yellow, lanceolate-obovate,

scarcely 1 cm. long; larger stamens 5-6 mm. long. — Section Tham-

noxys, affine 0. rhombeo-ovata St. Hil., 78, of Brazil (Knuth).

San Martin: San Roque, 1,400 meters, Williams 7751, type;

also 7204.

2. BIOPHYTUM DC.

Caulescent, usually more or less ligneous plants with the leaves

all at the top of the stem or in lateral fascicles and always multi-

pinnate with a terminal seta. Sepals imbricate, the petals contorted.

Capsule valves spreading from the axis in maturity and finally free.

Otherwise like Oxalis and the species similarly either very much

alike or not understood.

Leaflets 3-6 pairs, at least the terminal wider than 1 cm.

Sepals linear-lanceolate, subulate-acuminate B. somnians.

602 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Sepals oblong, acuminate, with seta 0.5-1 mm. long B. Foxii.

Leaflets (at least most leaves) more than 6 pairs, often smaller.

Peduncles 1-17 cm. long.

Leaflets 8-12 pairs, the middle about 2 cm. long; peduncles to

17 cm. long B. amazonicum.

Leaflets often more numerous or smaller; peduncles 1-8 cm. long.

Leaflets often 3 cm. long; flower heads often 1 cm. across.

B, globuliflorum.

Leaflets about half as long or smaller as also the flower heads.

Peduncles more or less hirsute.

Leaflets rounded or subobtuse apically; flowers white or

pale in dense heads 4 mm. thick B. peruvianum.

Leaflets acutish; flowers violet in rather lax heads.

B. dendroides.

Peduncles very densely setose-hispid B. juninense.

Peduncles obsolete B. Tessmannii.

Biophytum amazonicum Knuth, Pflanzenreich IV. 130: 416.

1930.

Subacaulescent, often 18 cm. high from a deep root to 5 mm.

thick; petioles 1 cm. long; leaves to 15 cm. long with 9-12 pairs

of broadly oblong-rectangular leaflets, the lower cordate, the upper

obliquely broadly ovate, all glabrous except sparsely ciliate margins,

glaucous, the medial often 2 cm. long, 10-12 mm. wide, retuse, tip

retuse-rounded with mucro to 0.5 mm. long; peduncles to 17 cm. long,

subhirsute as leaf-rachis, the subulate 4-5 mm. long bracts forming

apical head 1.75 cm. thick; flowers often 5-15 in the head on sub-

glabrous pedicels 2-3 mm. long; sepals 5-6 mm. long, acutely

acuminate, strongly costate; petals 10 or 11 mm. long, obovate;

larger stamens of medium styled flowers equaling calyx, subhirsute.—

Allied by author to B. Foxii.

Loreto: Pongo de Manseriche, (Tessmann, type).

Biophytum dendroides (HBK.) DC. Prodr. 1: 690. 1824; 399.

Oxalis dendroides HBK. Nov. Gen. & Sp. 5: 250. 1822.

Stems creeping at base the simple erect portion terete, ligneous,

7-30 cm. high, 2-4 mm. thick and yellowish-tomentose toward apex

where are crowded together the 20 or more leaves, 3-11 cm. long;

leaflets 10-20 pairs, 10-18 mm. long, gradually larger toward the

apex of rachis, ciliate, appressed pubescent above, hirsute especially

FLORA OF PERU 603

on midnerve beneath and more or less glaucescent, the lowest ovate,

acutish, the medial oblongish with an angle at inner base, the upper-

most broader and obovate-subtrapezoid; petiole and rachis hirsute;

peduncles 5-many, hirsute, apical, 2-5 cm. long, 1-4-flowered, the

many capitulate but scarcely globulate-congested bracts linear-

setaceous, 4-5 mm. long, acute, carinate, the pedicels little longer;

sepals subequal, lanceolate, acute, nervose-striate, hirsute, half as

long as the infundibuliform violet corolla, the cuneate petals rounded

at tip; larger stamens pubescent, exceeding the glabrous or pubescent

styles; capsule barely exserted, ovate-globose, 5-angled, pubescent

especially toward apex, the cells 1-2-seeded. — "Mexico to/ Peru and

Brazil" (Knuth), but no specimens cited; cf. B. peruvianum but it

seems to me doubtful that there is more than one slightly variable

species concerned. Illustrated, Knuth, I.e. 400. F.M. Neg. 36802.

Peru: Probably, sens. lat. Brazil to Mexico.

Biophytum Foxii Sprague, Kew Bull. 343. 1911; 416.

Herbaceous, erect, the appressed pilose stems about 5.5 cm.

high with an apical cluster of about 8 leaves 3.5-8 cm. long; leaflets

3-6 pairs the terminal obliquely obovate, 2-3 cm. long, rarely 2 cm.

wide, the remaining gradually reduced, truncate at base, the inter-

mediate subtrapezoid, the lower triangular-ovate; peduncles 4 cm.

long, crisply pubescent, many-flowered; sepals oblong, acuminate,

with seta to 1 mm. long, 7-9-nerved, ciliate and papillate-pilose,

about 6 mm. long; corolla white with tube 6 mm. long, the reflexed

lobes 4.5 mm. long, 3.5 mm. wide; smaller stamens glabrous, about

3 mm. long, larger pilose, nearly 5 mm. long, styles 1.75 mm. long,

minutely pilose for about 1 mm. — Cf. B. somnians; type presumably

in Herb. Kew.

Peru: (W. Fox, type).

Biophytum globuliflorum Knuth, Pflanzenreich IV. 130: 403.

1930.

Stout, the stems 5-8 mm. thick, 1.5-5 dm. high; leaves 15-20,

to 2.5 dm. long, with about 15 pairs of narrowly rectangular-lanceo-

late leaflets the medial often 3 cm. long, a third as wide, retuse at

base, acutely and somewhat obliquely acuminate, sparsely pubescent

above, moderately pilose beneath, the lower obcordate, the upper

ones obliquely or falcately ovate-lanceolate; peduncles mostly 1 dm.

long, densely puberulent, the very many 7 mm. long subulate-filiform

bracts forming a head 1.5 cm. in diameter, the flowers 1-4 on pedicels

604 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

2-3 mm. long; sepals 6-7 mm. long, acutely acuminate, strongly

costate; petals 10-12 mm. long, oblanceolate, pale yellow with pink

tips or canary yellow (Mexia). — Similar but smaller isB. bolivianum

Knuth, I.e., the leaflets obtuse or nearly rounded and hirsute. The

Ecuadorian B. Heinrichsae Knuth, Repert. Sp. Nov. 38: 198. 1935,

has leaflets retuse at base, rounded at tip, 2.5 cm. long, 8 mm. wide,

stems often biparted, peduncles setose. F.M. Neg. 26289.

Loreto: Flood-free wood, Pongo de Manseriche, Tessmann 4037,

type; also 3886 below Pongo de Manseriche. Ridge to right of mouth

of Rio Santiago, floor of dense forest, Mexia 629 J^. (det. Standley).

Bolivia. "Ambrosocco" (Mexia).

Biophytum j immense Knuth, Repert. Sp. Nov. 38: 198. 1935.

Perennial, the simple rhizome with many brown roots; stem

strict, 1-1.5 dm. high, 3 mm. thick, very densely tomentose; leaves

apical, 13-20, to 12 cm. long, leaflets 15-25 pairs, narrowly rectangu-

lar, to 12 mm. long, 3.5 mm. wide, glabrous except the medial nerve

long-setulose, obliquely retuse at base, obliquely rounded at tip,

finally all elongate-obovate, lower leaflets gradually reduced toward

base of rachis the lowest obliquely triquetrous; petioles nearly lack-

ing; peduncles 4.5-6.5 mm. long, 0.5 mm. thick, very densely setose-

hispid; bracts subulate, 2-2.5 mm. long; flowers unknown. — Section

Dendroidea, aff. B. dendroides but recognized by the densely setose

rachis (Knuth). This reads like a race of B. peruvianum if that itself

is distinct from B. dendroides.

Junin: Cahuapanas, Rio Pichis, 340 meters, (Killip & Smith

26755, type).

Biophytum peruvianum Knuth, Pflanzenreich IV. 130: 402.

1930.

Similar to B. dendroides; leaflets subequal, usually 8-12 pairs,

rarely 16 mm. long, 4 mm. wide, obtuse or rounded, obscurely or

not at all mucronulate, subglabrous both sides; bracts minute, glo-

bosely or cylindrically congested the resulting head about 3 mm.

thick and 3-5 mm. long; flowers white or cream-colored. — Cf.

Knuth who accepts but defines without clear distinctions one Bolivian

and several Colombian plants as species (described previously) that

like this may be variants of B. dendroides; however at present for

reasons of expediency, since the Peruvian plant is not typical, Knuth 's

name, actually based on a specimen from Bolivia (Bang 1397!}, may

be used.

FLORA OF PERU 605

San Martin: Chazuta, Klug 4043- Zepelacio near Moyobamba,

1,100 meters, Klug 3700. — Junin: Rio Seco near La Merced, Soukup

2505. Puerto Bermudez, 375 meters, Kittip & Smith 26575 (det.

Killip). — Huanuco: Between Monzon and the Huallaga, 700 meters,

open wood, Weberbauer 8583; 285. Pozuzo, 4549. Tingo Maria,

Stork & Horton 9489; Soukup 2217. Hacienda San Carlos, Rio

Ysabel, 1,000 meters, Mexia 8139. — Ayacucho: Kimpitiriki, Rio

Apurimac, Killip & Smith 23029 (det. Knuth). — Loreto: Yurimaguas,

Killip & Smith 28910; Spruce 4588. San Antonio de Cumbaso,

Ule 6533. Bolivia; Brazil.

Biophytum somnians (Mart. & Zucc.) Knuth, Notizb. Bot.

Gart. Berlin 7: 317. 1919; 416. Oxalis somnians Mart. & Zucc. ex

Zucc. Denkschr. Akad. Muench. 9: 181. pi. 4- 1823-1824.

Perennial, with ligneous stem rooting at the prostrate base then

ascending 3-5 dm., simple, terete but somewhat enlarged at nodes,

subtomentose with short brownish appressed trichomes; leaves

subverticillate at nodes and apex, petioled, the short petioles thick

and rugose-sulcate at base; leaflets unequal, to 4 cm. long, 2.5 cm.

wide, the smaller lower obliquely subcordate, acute, the larger upper

obovate rounded at tip, all slightly or obscurely and bluntly extended

at the lower angle of the truncate base, typically appressed pubescent

with short trichomes both sides, subciliate, the rachis densely villous

especially beneath; peduncles solitary, typically 7-12 cm. long, pubes-

cent, many-flowered with numerous linear-lanceolate acuminate

pubescent dry brownish bracts about the base; flowers small, semi-

concealed in the crowded lanceolate subulate-setaceous ciliate-hispid

bracts; sepals linear-lanceolate, glabrous but apically subciliate,

subulate-acuminate, the little longer oblong obtuse petals roseate,

white at base; stamens all much shorter than the pilose styles, the

larger pilose above, the smaller glabrous and scarcely equaling the

pubescent ovary. — After Knuth; the Peruvian plants are less

pubescent with shorter peduncles and apparently slightly larger or

more exserted flowers and probably represent B.Foxii which however,

ex char., seems to be a variant. Illustrated, Knuth, I.e. fig. 25,

page 411. F.M. Neg. 19208.

Loreto: In forest, La Victoria, Williams 2910. Mishuyacu near

Iquitos, Klug 183 (det. Killip). Rio Mazan, Jose Schunke 143 (det.

Standley). Northwestern Brazil. "Ambrosoco."

Biophytum Tessmannii Knuth, Pflanzenreich IV. 130: 404.

1930.

606 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Allied to B. dendroides but subacaulescent, the stems only a cm.

or two high from a tortuous ligneous root to 5 mm. thick; leaflets

12-15 pairs, rectangular, the medial 17 mm. long, 6 mm. wide, retuse

at base, more or less oblique at the mucronulate tip the mucro 0.5

mm. long, margins ciliate, surfaces minutely appressed pilose, the

lower nearly cordate, the upper obliquely ovate; peduncles obsolete

or 5 mm. long, in an apical head 5 mm. thick; bracts 3-4 mm. long,

numerous; flowers often one or two, the pedicels 0.5 mm. long, gla-

brous as sepals, these pale or pale rose, very acute, 6 mm. long;

petals 14 mm. long, oblong, white except above the middle where

roseate.

Loreto: In moss and selaginella on large stones in flood-free

wood, Pongo de Manseriche, (Tessmann 4741, type).

3. HYPSEOCHARIS Remy

Essentially acaulescent herbs with spreading pinnate leaves, or

the few upper ones pinnately incised, the leaflets subentire or 3-lobu-

late or pinnate-incised. Peduncles cymosely 1-many-flowered, the

flowers orange. Sepals and petals 5, the former imbricate, the latter

contorted, glands none. Stamens 15, subtriseriate, coalescent at base.

Ovary 5-lobed, 5-celled, the style one, the stigma capitate, terminal,

the seeds many in each cell. Capsule loculicidally dehiscent. — The

genus is not clearly a member of this family but its position is then

uncertain; cf. Knuth, Bot. Jahrb. 41: 170. 1908.

Leaflets all obtusely bilobed at tip, the peduncles 1-flowered.

H. bilobata.

Leaflets entire or minutely tricrenate-denticulate H. tridentata.

Leaflets dentate or pinnately incised; peduncles 1-4-flowered.

H. pedicularifolia.

Leaflets bipinnately incised; peduncles 5-7-flowered. . . . H. Pilgeri.

Hypseocharis bilobata Killip, Journ. Wash. Acad. Sci. 16: 570.

1926; 439.

Root cylindric, elongate, dark purplish, 1.5-2 cm. thick; petioles

as sometimes the leaf rachis puberulent, otherwise glabrous, the

former 5-10 mm. long, the latter 2-6 cm. long, with alternate or

subopposite sessile or subsessile leaflets the lateral oblong-orbiculate,

cordulate at base, 3-6 mm. long, 2-5 mm. wide, the terminal ovate-

orbicular, obliquely cordate at base, 6-10 mm. long, 5-8 mm. wide,

all shallowly bilobed at apex, the sinus to 1.5 mm. deep, the erect

FLORA OF PERU 607

lobes obtuse; peduncles slender, 1-flowered, 1.5-2 cm. long; sepals

oblong, obtuse, about 4 mm. long, 3 mm. wide; ovary broadly ovoid;

corolla not seen. — After author, who notes that in the similar H.

tridentata the root is not strongly thickened and the leaflets are

3-dentate.

Cuzco: Near Cuzco, 3,000-3,600 meters, Herrera, type.

Hypseocharis pedicularifolia Knuth, Bot. Jahrb. 41: 173.

1908; 429.

Leaves 5-7 cm. long, the glabrous petioles broadly dilated at

the fleshy base, the 13-17 pairs of leaflets alternate or opposite,

sessile, very broadly ovate or obovate, 5-7 mm. long, coarsely or

lobulately incised on each side, with one or often both sides also

with a basal obtuse lobe; peduncles scapiform, not more than 2 cm.

long, 1-2-flowered the flowers less than 3 cm. across; petals cuneately

broadly-obovate, pale red (Fiebrig). — H. Fiebrigii Knuth, I.e. has

elliptic serrate-dentate leaflets.

Arequipa: Above Chivay in heather formation, 3,900 meters,

Weberbauer 6899 (det. Knuth). Bolivia.

Hypseocharis Pilgeri Knuth, Bot. Jahrb. 41: 174. 1908; 429.

Leaves 7-11 cm. long, pinnate but toward the apex only pinnately

incised; petioles glabrous, the broadly dilated bases nearly mem-

branous margined; leaflets 13-19, alternate or subopposite, to 2 cm.

long, sessile, broadly ovate, coarsely pinnate-incised the lobes more

or less deeply cleft with obtuse or obtusish divisions; peduncles

5-7-flowered forming a few-flowered corymb, the flowers themselves

unknown. — In Goodspeed collection "flowers white, falling very

readily," the oblong obtuse sepals about 5 mm. long, nearly 3 mm.

wide.

Lima: Stony places near stream between Norquinia and Matu-

cana, 2,300 meters, Weberbauer 95, type; 166. Below Matucana,

rock-outcrops among xerophytes, 2,300 meters, Weberbauer 5200

(det. Knuth); 2,400 meters, Goodspeed 33072 (det. Leonard).

Hypseocharis tridentata Griseb. Goett. Ges. Wiss. Nachr. 493.

1877 et Abh. 24: 73. 1879; 428.

Leaves 3-10 cm. long, with basally broadly dilated glabrous

petioles, the margins nearly membranous; leaflets 11-17, alternate

or subopposite, sessile, the lateral 5-14 mm. long, elliptic or oblong,

the terminal 7-18 mm. long, ovate, all entire or with three obtusish

608 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

teeth at apex or, more accurately, emarginate with a medial mucro

or tooth; peduncles 6 cm. long, 1-flowered, the flowers unknown.—

H. pimpinellifolia Remy, 428, has 15-21 oblong-lanceolate or elliptic

leaflets 2-3 cm. long, those in the upper part unequally crenate-

dentate, below mostly entire, the lower trilobed to base; the leaves

to 17 cm. long, the corolla to 4 cm. broad. It is possible that this,

the type species, is polymorphic and the other described plants

variants. Determinations mostly by Knuth. Vargas 9722 has

oblong obtuse sepals nearly 6 mm. long and about half as wide,

petals 12 mm. long, nearly 6 mm. wide at the somewhat broader tip.

Huancavelica: Between Cuchicancha and Huaytara, Raimondi.

Huancavelica, Raimondi. — Ayacucho: Near Cangallo, Raimondi. —

Cuzco: At 3,400 meters, Herrera 476. Ocongate, 3,350 meters,

Vargas 9722. Bolivia; Argentina.

TROPAEOLACEAE Juss. Nasturtium Family

Reference: Buchenau, Pflanzenreich IV. 131. 1902.

Scandent herbs (two Chilean species procumbent) with nearly

always long-petioled often peltate leaves that are entire, lobed or

divided to the petiole, and solitary axillary well-peduncled irregular

showy flowers (one Ecuadorean species, T. umbellatum Hook.,

flowers umbellate). Stipules rarely conspicuous, often slender and

caducous, sometimes wanting. Sepals 5, imbricate or valvate, the

lateral often broader. Petals usually 5, entire to variously lobed or

dentate, eciliate to fringed-ciliate; axis of sepals and upper petals

produced into a straight or curved spur. Stamens 8, free; anthers

bicelled, erect, laterally dehiscent. Ovary 3-celled; cells 1-ovuled;

stigmas sessile, 3, papillose within. Fruit 3 spongy or rather fleshy

follicles that are usually ridged dorsally.

1. TROPAEOLUM L.

The only genus, and important in floriculture since 1843 with the

discovery of T. peltophorum Benth.; T. minus L. and T. majus L.

were however popular in gardens from the middle seventeenth

century, the former introduced into Germany in 1570, the latter

more than a hundred years later (Buchenau). Hybrids, particularly

of these species, adorn the gardens of the world as "Nasturtiums."

Only about a dozen species, at one timfe and another, have been in

cultivation.

FLORA OF PERU 609

The buds and immature fruits of T. majus and other species, cured

in vinegar, are used as a substitute for capers. The latter especially

suggest the caper pickle (flower bud of the African Capparis spinosa)

in shape and are pleasant as well as peppery in taste; Ruiz and Pavon

also found "people of refinement adding the flowers to salads which

attain a not unpleasant flavor." The tubers of T. tuberosum, cooked,

are much appreciated in the middle and higher Andes; cf. note under

that species.

Besides the following, Buchenau recorded T. ciliatum R. & P.,

20, from Peru on the basis of a Dombey specimen without exact

locality; since the species is today unknown except in Chile this is

doubtless another of the Dombey specimens mislabeled. In compil-

ing this account, from Buchenau, I.e. and D. K. Hughes, "serrato-

ciliata" group, Kew Bulletin 63-85. 1922, I have become impressed

with the late Stapf's remark in his foreword to the latter's paper:

"The range within which the characters used for discrimination

actually vary, and consequently their reliability and perhaps even

the claim of some of the units recognized to the status of species

remain open questions. Observations in the field and cultivation

both have to be called in aid to answer them." It is probable that

there is hybridization in the field as in the garden and a taxonomic

study with the aid of genetics is very much in order. Kuntze used

the name Trophaeum, the resulting synonyms not cited here as his

action now seems clearly to be only of historical interest. My

friend Killip has added to the cited specimens from the collections

in his charge (nearly all determinations were made or verified by

him), and his assistance in the disposal of critical material is gratefully

acknowledged.

Petals at least in part lobed, serrate- or aristate-ciliate, sometimes

not exserted (cf. T. minus, lower at least ciliate apically).

Leaves more or less deeply 5-7-lobed.

Leaf-lobes extending distally at least half way; stipules subulate

and parted, often caducous; sepals sometimes gibbous at

base.

Petals lobed, at least the upper.

Lobes of upper petals obtuse, eciliate.

Plants glabrous; leaf -lobes obtuse; petals irregularly

lobed T. peregrinum.

Plants sparsely pilose; leaf -lobes acute; petals 3-lobed.

T. Seemanni.

610 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Lobes of petals narrow, 3, dentate-aristate . . T. Haynianum.

Petals not lobed, puniceous ciliate T. Smithii.

Leaf -lobes extending less than half way; stipules rather semilu-

nate; petals serrate-ciliate; sepals partly gland-appendaged

at base T. bicolor.

Leaves entire to angulately lobulate, rarely 3-lobed.

Stipules none, minute or caducous.

Leaves entire to angulate; spur straight or merely curved.

Leaves longer than or about as long as broad; flowers

usually 3.5-4.5 cm. long.

Leaves glabrous, often somewhat papillose beneath.

T. cirrhipes.

Leaves sparsely pubescent above, pruinose-papillose be-

neath T. papillosum.

Leaves distinctly broader than long; flowers to 2 cm. long.

T. olmosense.

Leaves obtusely but obviously 3-lobed; spur coiled at tip.

T. Weberbaueri.

Stipules conspicuous, persisting; leaves papillose beneath.

T. Matthewsii.

Petals entire, emarginate or obscurely to coarsely crenulate or the

3 lower ciliate basally (rarely apically, T. minus), rarely wanting.

Leaves more or less deeply 5-7-lobed; petals eciliate basally.

Leaves pedately parted; petals 2, the 3 lower wanting.

T. dipetalum.

Leaves distally lobed, peltate; petals 5.

Flowers 4-5 cm. long; leaves nearly medially 7-lobed with

rounded open sinuses between the obovate lobes.

T. longiflorum.

Flowers to 4 cm. long; leaves rarely 7-lobed (to 7-parted),

but not as above.

Flowers red and orange, tubular-funnelform, the throat

about 1 cm. wide at an thesis; petals typically entire;

roots tuberous, perennial; leaflets about one- third

5-lobed T. tuberosum.

Flowers broadly funnelform, 1-2 cm. wide; petals in part

1-3-crenate; roots (known) annual; leaflets often 6-7-

lobed.

FLORA OF PERU 611

Petals yellow-red or orange; leaves often 5-lobed.

T. crenatiflorum.

Petals purple or violet; leaves often 6-7-lobed.

Flowers 2-2.5 cm. long T. purpureum.

Flowers 3-3.5 cm. long T. Vargasianum.

Leaves very shallowly lobed, subentire or entire; petals at least

lower 3, ciliate or fringed basally.

Flowers conic-tubulate, the lower petals coarsely crenate, ciliate

at base T. peltophorum.

Flowers open funnelform, the lower petals often ciliate apically,

all fringed at base.

Leaf -nerves and petals not mucronulate T. majus.

Leaf-nerves and petals mucronulate but sometimes minutely

or caducously T. minus.

Tropaeolum bicolor R. & P. Fl. Peruv. 3: 76. pi. 313. 1802; 22.

Rather stout, glabrous, high-climbing with firm semilunate acute

incised-ciliate stipules and peltate nearly truncate-based leaves

parted to two-fifths into 7 broadly (ovate) oblong shortly mucronate

lobes; peduncles two to three times longer than the leaves; lower

petals a little longer than the sulphur yellow calyx, the smaller upper

ones yellow the larger lower puniceous-red, all serrate-ciliate; spur

yellow, straight, cylindric-conic, 43 mm. long above, 6 mm. thick at

base; filaments purplish, anthers violet; lower petals eciliate on

claw; sepals obtusish, the lower gland-appendaged at base, 15-18

mm. long; leaves slightly paler beneath to about 5 or 7 cm. wide,

3.5-5.5 cm. long; lower petals about 15 mm. long, the upper about

13 mm. — Name originally bicolorum. F.M. Neg. 12664.

Huanuco: In the mountains near Mufia, Ruiz & Pav6n, type.

In thickets at Tambo de Vaca, 4422. — Junin: above Tarma, Ruiz

& Pavdn. Rio Masamerich, Weberbauer 6636; also 6724. Concep-

tion to Satipo, 1,800 meters, Seibert 2382.

Tropaeolum cirrhipes Hook. Icon. PI. 5: pi. 411. 1842; 23.

Glabrous except for the hirsutulous nodes; stipules minute or

caducous; stems angled and sulcate; petioles curved or cirrhose, to

11 cm. long; leaves long-ovate, but distinctly peltate, entire but

subrepand at base, the basal angle rounded, obtuse at tip, to 9 cm.

long, 5-6 cm. wide at base, glaucous and minutely or obscurely

pruinose-papillose beneath; pedicels strongly cirrhose, to 2 dm. long,

612 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

slightly enlarged below the flowers, these 4.5 cm. long, the red spur

3.6 cm. long, 4 mm. thick at base, obtuse; sepals green, rounded-

ovate; petals not exserted, 4-5 mm. long, dentate-ciliate above,

yellowish-green. — T. menispermifolium Buch., Bot. Jahrb. 34: Beibl.

78: 11. 1904, has leaves slightly broader than long, densely pruinose-

papillose beneath, pedicels a little pubescent, flowers about 5 cm.

long; the Bolivian T. Kuntzeanum Buch., 23, more resembles T.

cirrhipes but has leaf-angles obtuse instead of rounded, pedicels

7-10 cm. long, flowers 5-5.5 cm. long; T. cuspidatum Buch., 23,

likewise of Bolivia, differs from all of these species, which it in general

resembles, by the obscurely or not at all peltate leaves.

Amazonas: Chachapoyas, (Matthews 3177, type).

Tropaeolum crenatiflorum Hook. f. Bot. Mag. 72: pi. 4%45-

1846; 20.

Annual from fusiform root; stipules subulate, deciduous; leaves

reniform-suborbicular, peltate, straight or a little repand at base,

5-lobed about one-fourth (rarely some leaves 6- or 7-lobed), the

obtuse segments mucronate; peduncles very slender, far exceeding

the leaves; flowers 3-4 cm. long; sepals ovate, acute; spur about

15 mm. long above, nearly cylindric, 4 mm. thick at base, abruptly

acuminate; petals entire or the upper merely repand, pale yellowish

red, red-lined, the longer lower orange-yellow, clawed, eciliate, some-

what bicrenate. — Similar species included the Bolivian T. cocha-

bambae Buch., 20 (this fide Buchenau probably with earlier name,

T. Klotzschii Warscewicz), and the Ecuadorean T. Buchenavianum

Hieronymus, 21 (not, it may be noted, invalidated by T. Buchenavii

Phil., the different ending sufficing according to the International

Rules; T. Hieronymi Buch., 21, is therefore a superfluous name);

both species are distinguishable from T. crenatiflorum by the smaller

flowers (20-24 mm. long) ; the first is very slender with many orange

flowers, petals undulate-sinuate, the upper with a dark velvety spot;

the petals of the second are carmine, entire, or obscurely crenulate.

T. septemlobatum Heilb. (cf. under T. purpureum) of Ecuador is

very near but petals entire, more red, spur about 2 cm. long, 5-6

mm. thick at base, leaves regularly 7-lobed. — Illustrated, Fl. Serres

2: pi. 166.

Huanuco: Near Pillao, Lobb, type. Near Chaglla, Lobb. Cani,

3484. — Huancavelica: Ampurco, 2,600-2,700 meters, Weberbauer

6501. — Cuzco: Below Lares, Prov. Calca, Weberbauer 7896.

FLORA OF PERU 613

Tropaeolum dipetalum R. & P. Fl. Peruv. 3: 77. pi. 313.

1802; 18. Chymocarpus stipulaceus Klatt ex Otto, Hamb. Gart. &

Blum. Zeit. 259. 1859.

Distinctive by the conspicuously large 3-lobed stipules, these

deciduous, and the red flowers that lack the 3 lower petals; leaves

broadly spathulate, 5-7-lobed, the obtuse lobes mucronate; petals

cuneate; spur slender, slightly curved, 28-30 mm. long above,

cylindric; calyx persisting at base of the fleshy fruit. — Leaves firm,

with 7 sometimes irregular or lobulate lobes that extend below the

middle, deeply cordate at base, epeltate, to 7 cm. wide, about 4 cm.

long; petioles 8-10 cm. long; peduncles about 16 cm. long; spur

about 2 cm. long, sepals and petals nearly as long. — Illustrated,

Kerner, Hortus Sempervirens 21: pi. 252. F.M. Neg. 12668.

Huanuco: Wooded places in the mountains of Pillao, Ruiz &

Pavdn, type. "Monte-massua."

Tropaeolum Haynianum Bernhardi, Allg. Thuer. Gartenz. 2:

73. 1843; 25.

Glabrous, without stipules and the petioles little longer than the

nearly three-fourths acute or acuminately 5-lobed leaves, the blade

repand at base, the lobes often incised; flowers 30-33 mm. long, the

spur about 2 cm. long above, curved, subulate-acuminate, 4.5 mm.

thick at base, yellow and red-dotted or -lined as the ovate acute

sepals, these shorter than the deeply ciliate-incised 3-more lobed

petals, the larger upper ones orange yellow or scarlet toward edges

or dotted or striped, the smaller lower ones yellow and less dotted.

— The narrow lobes of the petals merge with the capillary or ciliate

teeth, which much exceed the primary divisions. The flowers are

often only 28 mm. long; the spur is extremely slender toward the

tip.— Illustrated, Buchenau, 26 (petals). F.M. Neg. 18268.

Huanuco: Apparently described from cultivated specimens, origin

near Huanuco. — Cuzco: Echarate, Prov. Convencion, sunny edge

of forest, Goodspeed Exped. 10498 (det. Standley). Lares Valley,

Prov. Calca, Weberbauer 7942.

Tropaeolum longiflorum Killip, Journ. Wash. Acad. Sci. 24:

48. 1934.

Entirely glabrous; stipules linear, coriaceous, subpersisting,

about 3 mm. long; petioles 3-4.5 cm. long; leaves peltate, 7-lobed

nearly to middle, 2-3 cm. long, 2.5-4 cm. wide, lobes obovate, to

1 cm. wide, rounded and minutely callose-mucronulate, papillate

614 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

beneath; peduncles 12-13 cm. long; flowers 4-5 cm. long, spur

cylindric, 3.2-3.5 cm. long, about 3 mm. thick, conspicuously

nerved, tip straight or slightly curved; sepals oblong, obtuse, 8-10

mm. long, 3-5 mm. wide; petals similar and subequal, 13-15 mm.

long, 6-9 mm. wide, subtruncate, subentire, the 2 upper ones scarlet,

the 3 lower orange with scarlet veins, all with dark brown veins at

base. — Leaves general outline of T. septemlobatum Heilb. of Ecuador

and T. purpureum but lobes obovate, flowers much larger and petals

more nearly uniform; leaf-shape and size of flower that of T. bicolor

of the serrato-ciliata group (Killip).

Junin: Near Atax, 3,400 meters, Rio Masameric, Weberbauer

6641, type.

Tropaeolum Matthewsii Hughes, Kew Bull. 69. 1922.

Young parts (as to type) more or less ashy villous-puberulent,

the slender stems somewhat compressed and contorted toward the

tips; stipules persisting, semiorbicular, to 1 cm. wide, entire or

essentially, like the leaves scarcely papillose above, pruinosely so

beneath; petioles subcirrhose, nearly 3 cm. long; leaves broadly

triangular-ovate, 2.5-3 cm. long and wide or a little wider, broadly

5-lobed, the lobes emucronate; pedicels cirrhose, scarcely exceeding

the petioles; flowers sparsely pubescent, to 2.5 cm. long, spur straight,

conic, 1-1.8 cm. long, about 4 mm. thick at base; sepals broadly

ovate-oblong, rounded-obtuse, nearly 8 mm. long, 5-6 mm. wide;

petals not at all exserted, dentate-ciliate above, the upper broadly

spathulate-oblong, 6 mm. long, lower obovate, attenuate to claw,

4 mm. long. — Nearly T. stipulation Buch. & Sod., Bot. Jahrb. 34:

Beibl. 78: 12. 1904, of Ecuador, stems and petioles sparsely spread-

ing hirsute to glabrous, stipules 1 (rarely 2-3) cm. wide, leaves

epapillose, glabrous, nerves beneath red, and more or less red

punctate; flowers glabrous, 2.5-3 cm. long, spur 18-22 mm. long,

entirely red, petals dark red, after Heilborn, Arkiv. Bot. 23A, pt. 9:

7. 1931.— Illustrated, Hughes, I.e. 70.

Amazonas: Chachapoyas, (Matthews, type). — Libertad: Mixiollo,

2,900 meters, Weberbauer 7034 (det. Mansfeld).

Tropaeolum ma jus L. Sp. PI. 345. 1753; 21.

Like T. minus but coarser, the leaves orbicular and the nerves

as the petals not mucronulate; spur barely curved, 26-28 mm. long

above, 3-4 mm. thick at base; petals orange or red to yellow, the

lower obscurely if at all spotted.

FLORA OF PERU 615

This is the popular cultivated "nasturtium" or at least the source

in hybridization usually with T. minus or T. peltophorum. The

Worth and Morrison collection suggests the latter in pubescence

but has the petals of T. majus. Its tallos are used to dye yellow

(Herrera); its buds and young fruits pickled; its flowers used in

salads (Ruiz & Pavon, who found it at Lima, Chancay and Huanuco).

Lima: Wet places near Barranco, Weberbauer, 148. Near Lima,

Wilkes Exped. Pueblo Libre, Soukup 2573, fide Killip. — Arequipa:

Granite slides, Chalca, 300 meters, Worth & Morrison 15680 (det.

Johnst.). At Arequipa, Soukup 991 . Without data, Ruiz & Pavon.—

Cuzco: Near Yucay, 3,000 meters, Herrera 1459. Peru to Colombia.

"Mastuerzo," "mastuercillo" (Ruiz & Pavon).

Tropaeolum minus L. Sp. PI. 345. 1753; 21.

Nearly glabrous vine similar to T. peltophorum; leaves orbicular-

reniform, the nerves terminating (sometimes obscurely) in mucronate

points; spur usually obviously curved; petals mucronulate, the lower

ciliate apically and with a conspicuous darker blotch. — T. pinnatum

Andrews, Bot. Rep. 8: pi. 535. 1808 vel 1809, is said to be a

hybrid of this and T. peregrinum L. ; garden forms are usually hybrids

of this and T. majus L. The native plant has citrus or golden yellow

petals, the upper ones red-veined but the lower with a large brownish-

red spot; like the similar T. majus L. the first two leaves are opposite,

one- third 3-lobed, and have white- tipped linear-subulate stipules,

rose-tinted at base, a tone that streaks the young stem and the

younger leaf-nerves. — Material from Cerro de San Agustin is not

clearly referable to this or T. majus and maybe it could be noted as

a variant since the leaves are obscurely mucronulate, the flowers

scarcely 2.5 cm. long, but probably it is only a local race of one or

the other of the similar species. — Illustrated, Bot. Mag. 3: pi. 98.

Cajamarca: Near Cascas, Raimondi. — Ancash: Huaraz, Rai-

mondi; Weberbauer 3220 (det. Loesner). Pampa Romas, Weberbauer

3199 (det. Loesner). — Lima: Moist places, Feuillee. Lomas of Lima,

covering large areas, flowers yellow, Verne Grant 7439. Near Viscas,

Pennell 14477; Soukup 2572. Cerro de San Agustin, Weberbauer

5716.— Ayacucho: Huanta, 2,800 meters, Killip & Smith 23341.

Tropaeolum olmosense Mansf. Notizbl. Bot. Gart. Berlin 9:

52. 1924.

Stem nodes puberulent as also petioles (about 6 cm. long) and

peduncles or the last two glabrescent; stipules none; leaves broadly

616 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

deltoid, broader than long, entire or subrepand-sinuate, very shortly

apiculate, about 3 cm. long, 5 cm. wide; peduncles to 6 cm. long;

spur conical, crimson, about 16 mm. leng upper side, lower 14 mm.

long, 4 mm. thick at base, puniceous; petals yellow, serrate-ciliate,

the upper with a brown spot, little exceeding the red sepals, the

lower also ciliate above on the claw and about twice as long as the

sepals. — Near T. Moriztianum Klotzsch of Colombia and northwards

(Mansfeld), with suborbicular leaves, orange petals, a species not

placed by Miss Hughes in her revision (under direction of Stapf)

Kew Bull. 63-85. 1922 of species 14-29 of Buchenau's treatment,

a curious omission, indeed, for Buchenau, Bot. Jahrb. 34: Beibl.

78: 11. 1904, indicated that its upper petals are in reality serrate-

ciliate. T. repandum Heilb. Arkiv. Bot. 23A, pt. 9: 5. 1931, of south-

ern Ecuador, has larger flowers, scarcely exserted petals, even the

calyx red.

Lambayeque: Above Olmos, 1,100 meters, in stream brush,

Weberbauer 7098, type. — Piura: Chanchaque, Huancabamba, 1,250

meters, Stork 11410.

Tropaeolum papillosum Hughes, Kew Bull. 71. 1922.

Glabrous, the slender striate stems subcompressed and contorted,

the stout petioles somewhat enlarged below and cirrhose, to 4 cm.

long; stipules not apparent; leaves broadly rotund-ovate, subentire,

6.5 cm. long and broad, green and red-punctate, as well as papillose

and sparsely pubescent above, glaucous, obscurely mottled and

densely pruinose-papillose beneath; pedicels very slender, scarcely

cirrhose, to 7 cm. long; flowers glabrous, nearly 3.5 cm. long, the

straight abruptly attenuate spur 2.5 cm. long, 4 cm. thick at base;

sepals ovate; petals dentate, not at all exserted, 4.5 mm. long, the

upper spathulate-oblong, long-ciliate, the lower clawed, smaller,

shortly ciliate. — The illustration, I.e. 72, shows both lower and upper

petals with cilia equally long and about as long as the blade. T.

pubescens HBK., 25, of Colombia and Ecuador and to be expected,

has pubescent flowers 2-2.5 cm. long, petals violet, leaves, at least

early, pubescent, as well as papillose; other Ecuadorian species that

might be sought here include T. fulvum Buch. & Sod., Bot. Jahrb.

34: Beibl. 78: 11. 1904, and T. adpressum Hughes, I.e. 80, illustrated,

84; both have pubescent leaves, punctate or mottled with red, but

epapillose and hirsute flowers, these (3) 4.5-5 cm. long, leaves of

former long-ovate, subentire, of the latter, broadly ovate, subtrilobed.

Another Ecuadorian species, similar if not related, is T. glaucum

FLORA OF PERU 617

Heilb. Arkiv. Bot. 23A, pt. 9: 4. 1931, flowers 4-5 cm. long, leaves

5-angulate-lobed, truncate at base.

Peru(?): Tambo grande (Huanuco?), Andre. Ecuador.

Tropaeolum peltophorum Benth. PL Hartw. 127. 1843; 21.

T. Lobbianum Veitch, Bot. Mag. 70: pi. 4097. 1844.

More or less pubescent (rarely glabrous) fusiform rooted annual

or perhaps sometimes perennial with peltate entire or merely repand

suborbicular mucronate or nearly aristate-tipped leaves often 3.5 cm.

across and rather large (about 3 cm. long) flowers, their lower petals

coarsely crenate-dentate, ciliate at base, orange or scarlet; sepals

ovate, obtusish, somewhat ciliate, spur cylindric-subulate, sub-

curved, 3-4 mm. thick at base, 2.5-3 cm. long above. — Upper petals

large, nearly entire. Weberbauer 6414 seems to be referable here and

may be named var. calvum Macbr. var. nov., foliis glabris ad 8 cm.

latis, circa 7 cm. longis, petalis haud ciliatis. — Probably scarcely

worthy a name since, as Killip has called to my attention, there are

already four variants named in Bailey's Cyclopedia of Horticulture,

which however are unknown to me. T. huigrense Killip, Journ. Wash.

Acad. Sci. 19: 194. 1929, of Ecuador has yellow flowers, spur 1 cm.

long, 1.5 mm. thick at base. — Illustrated, Fl. Serres 2: pi. 3. F.M.

Neg. 26392.

Piura: Huancabamba, 1,900 meters, Weberbauer 6060. Hacienda

Lagunas, Prov. Ayavaca, Weberbauer 6414 (type, var. calvum). —

Libertad: Raimondi. Ecuador; Colombia.

Tropaeolum peregrinum L. Sp. PI. 345. 1753; 27.

Allied to T. Haynianum; leaves weakly peltate, the base indented

or scalloped, the upper part regularly 3-7-lobed, the middle lobes

often on both sides (the others on one side) 1-lobate, the lobes mostly

obtuse but often mucronate; peduncles shorter than the leaves;

sepals obtuse; upper petals broadly spathulate to nearly round,

stiped, irregularly lobate-incised, the lobes usually obtuse, the lower

petals long-stiped, narrowly cuneate, laterally 4 or 5 aristate-capil-

late, all longer than the calyx, sulphur or lemon yellow, the upper

red-dotted near base, to 17 mm. long, 12-14 mm. wide, calyx and

spur greenish-yellow, the latter 12 mm. long above, 5 mm. thick at

base, pyramidate, strongly curved at end. — Illustrated, Buchenau,

26 (petals) ; Bot. Mag. 33: pi. 1351 ; Bot. Reg. 9: pi. 718. Cultivated

and used in medicine according to Herrera. Found at Lima, Chancay

and Huanuco by Ruiz & Pavon.

618 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Lima: Near Lima, (Feuillee). San Rafael, 3134. — Huanuco:

Ambo, 3172 and Mito, 3275.— Junin: Tarma, 3,054 meters, Stork

10946 (det. Standley). La Oroya, Kallenborn 158. Near Tarma,

1039; Killip & Smith 2177 4-— Apurimac: Chincheros, 2,500 meters,

yellow with purple spots, West 3687; Stork & Horton 10728 (det.

Standley ).- — Cuzco: Saxaihuaman, Pennell 13566. Huilabamba,

4,000 meters, Balls 6792. Urubamba, 2,860 meters, Vargas 11088

(det. Standley). Near Cuzco, Herrera 826; 245. Prov. Cercado,

(Herrera 49). Ollantaytambo, 3,000 meters, Cook & Gilbert 402.

Sicuani, Cook & Gilbert 96. Huasao, Herrera 708. Bolivia. "Pax-

jarito," "malla" (FeuillSe), "huallpa-huallpa" (Herrera), "quita-

ano" (West), "afiu-anu" (Herrera).

Tropaeolum purpureum Killip, Journ. Wash. Acad. Sci. 24:

48. 1934.

Glabrous; stipules linear-subulate, 1-1.5 mm. long, soon decidu-

ous; petioles 3-6 cm. long; leaves peltate, 2.5-4 cm. long, 3-4.5 cm.

wide, 7-lobed, the lobes rounded or truncate, mucronulate, epapillose,

glaucous beneath; flowers 2-2.5 cm. long, spur brown-red, cylindric-

conic, about 12 mm. long, 3.5 mm. wide, tip slightly curved, petals

a little clawed, crenate at tip, purple, 5-6 mm. wide, the upper about

1 cm. long, lower 1.5 cm. long. — Nearest T. crenatiflorum Hook. f. and

T. septemlobatum Heilb. Arkiv. Bot. 23A, pt. 9: 9. 1931 of Ecuador

but flowers purple and, as to the former, with 5-lobed much smaller

leaves; from the latter it differs further in the crenate petals (Killip).

Cuzco: Near Marcapata, 3,200 meters, Weberbauer 7788, type;

Vargas 9667 (det. Standley, T. minus).

Tropaeolum Seemanni Buchenau, Bot. Jahrb. 15: 226. 1892;

28.

Allied to T. peregrinum but sparsely pilose; leaves long-petioled,

peltate, 3-5-lobed the oblong lobes entire or lobulate, rather acute,

mucronate; petals all orange, clawed, flabellate-spathulate, the upper

to about three-fourths trilobed, these lobes shortly and obtusely

3-lobulate, lower petals stiped, about medially trilobed, the lobes

3-more serrate-ciliate; flowers about 3 cm. long, the peduncles longer

than the leaves, spur straight, conic-subulate, 15 mm. long above

and below, 3 mm. thick at base; sepals acute, much shorter than

the orange petals. — Illustrated, Buchenau, 26 (petals).

Lima: In gardens, (Miers). — Arequipa: In maize fields, (Mark-

ham; Cook & Gilbert 61). Near Arequipa along irrigation ditch,

Pennell 13046 (det. Killip). Bolivia.

FLORA OF PERU 619

Tropaeolum Smithii DC. Prodr. 1: 684. 1824; 22.

Resembles T. bicolor but with thin subulate 2-3-parted stipules

and leaves divided about three-fourths into 5 acute, sometimes

incised lobes; sepals acutish, gibbous-appendaged ; calyx and spur

scarlet, this 2 mm. thick at base, subcurved, about 18 mm. long,

the abruptly attenuate tip green; petals orange, puniceous-ciliate,

about one and a half times longer than sepals, with red veins and

red-tipped cilia. — Illustrated, Bot. Mag. 74: pi. 4385; Fl. Serres,

4: pi. 384.

Libertad: Parcoy, 3,200 meters, Weberbauer 7084 (det. Mansfeld).

To Colombia and Venezuela. "Pajarito."

Tropaeolum tuberosum R. & P. Fl. Peru. 3: 77. pi 314.

1802; 28.

High-climbing, glabrous, the often red-tinged stems (as petioles,

peduncles) from whitish-yellow usually red-marked obconic or pyri-

form tubers that are sometimes 8 cm. long; stipules subulate, decidu-

ous; leaves repand at base, nearly orbicular-peltate, about one- third

5-lobed, the rounded or very obtuse lobes mucronate; peduncles

usually definitely exceeding the leaves; spur red, rather thick,

cylindric-subulate, slightly curved near the tip, about 2 cm. long,

3.5 mm. thick at base; sepals broadly ovate, obtuse, red, slightly

shorter than the entire orange or reddish petals, the upper of these

nearly orbicular the lower narrower. — Other tuberous species are

probably Chilean except it may be noted that Buchenau, without

data, suggested that T. azureum Miers, 31, and T. tenuirostre Steud.,

33, may be found within Peru; both have small tubers, the leaves

divided to the base; the first has short spur, blue petals, the second,

conical spur, broad opening, yellow petals. Illustrated, Buchenau,

30 (petals); Hooker, Icones 7: pi. 653; Fl. Serres 5: pi. 452.

Commonly cultivated for the sweetish edible tubers for which

Herrera recorded the following native names which refer to the

variety in color: "occe-anu, yana-anu, puca-anu, chchecche-anu,

yurac-anu, ckello-anu, sapallu-anu, muru-anu." An interesting

account of Anu and Oca (Oxalis tuberosa) with plates is given by

W. H. Hodge in Journ. N. Y. Bot. Gard. 47: 214-224. 1946.

Lima: Huariaca, 3097. Valley Rio Rimac, 3,400 meters, Good-

speed Exped. 33132. Rio Blanco, 2975; Killip & Smith 21575.

Matucana, 123; 402. — Ayacucho: Prov. Lucamas, 2,800 meters,

Metcalf 30299.—- Huanuco: Tambo de Vaca, 4423.— Junin: Tarma,

Killip & Smith 21913. — Apurimac: Rio Incabamba, 3,000 meters,

620 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Stork & Horton 10731 (det. Standley, T. Matthewsii).— Cuzco: Valle

del Vilcanota, 3,500 meters, (Cook). Prov. Cercado, 3,450 meters,

near Urubamba, Vargas 11087 (det. Standley, T. Matthewsii).

Valle de Paucartambo, 3,500 meters, (Herrera 482). Below Lares,

Weberbauer 7886. Apurimac Valley, Herrera 3068. Sicuani, Cook

& Gilbert 194; 230. — Puno: near Limbani, 3,200 meters, Metcalf

30484- Ecuador; Bolivia. "Massua" (Weberbauer), "mashua"

(Sodiro), "arm" and "apina-mama" (Herrera), "mayua."

Tiopaeolum Vargasianum Soukup, Diez Anos Serv. Bot.

Univ. Cuzco 50: pi. 17. 1946.

Ex char, similar to T. purpureum except flowers 3-3.5 cm. long,

spur 13-15 mm. long, 4 mm. wide at throat, sepals oblong, about

6 mm. long, 4-5 mm. wide, petals 1-3-crenulate or subdenticulate

(according to plate), the upper 7 mm. long, 6 mm. wide, the lower

about twice as long and approximately 8 mm. wide, all very broadly

spathulate and slightly clawed. — The author does not state if the

leaves are papillose but apparently they are larger than those in

type of T. purpureum, described 3-3.5 cm. long, 34-48 mm. wide;

they are nearly truncate at base, one-fourth to one-fifth 6-7-lobed

(one leaf in plate with only 5 lobes).

Cuzco: Near Quellomayo, 3,200 meters, Panticala to Lucumayo,

(Vargas 4826, type, Herb. Univ. Nac. Cuzco).

Tropaeolum Weberbaueri Loesner, Bot. Jahrb. 45: 462. 1911.

Branchlets, petioles (to 6 cm.) and peduncles (to 4.7 cm.) under

a lens more or less viscid pilose; leaves peltate, 3-lobed but 5-angled-

obtruncate, basal angle rounded, lobes acuminate-mucronate, acutely

angled; flowers solitary, narrowly infundibuliform, 26-30 mm. long,

spur about 15 mm. long, 3 mm. thick at base; sepals lanceolate-

elliptic, obtusish, the petals scarcely two times longer, lemon-

yellow, the upper flabellate, lobate-incised, the lobules obtuse, lower

linear-cuneate, long-stiped, apically long-ciliate or subaristate-

fimbriate. — Affine T. peregrinum L. with leaves deeply 5-lobed

(Loesner) but remarkable as spur is coiled at tip, upper petals

reflexed. T. flavipilum Killip, Journ. Wash. Acad. Sci. 19: 192.

1929, of Colombia would key here but is like T. adpressum (cf.

under T. papillosum) but leaf-lobes rounded, flowers 5.5 cm. long,

petals deep purple, indument dense, yellowish. F.M. Neg. 12684.

Cuzco: Valle Urubamba in evergreen shrubs near Torontoy,

2,100 meters, Weberbauer 5065, type; also 6547. — Huanuco: On

shrubs at Mufia, 4343.

FLORA OF PERU 621

LINACEAE [DC.] Dumort.

Humiriaceae Juss.

Various in most characters including habit, thus annual herbs to

trees, erect or climbing but the leaves simple, entire to toothed,

usually alternate. Stipules variously developed or lacking. Pubes-

cence usually simple, 1-celled trichomes. Flowers perfect, the calyx

and corolla ordinarily in 5's, the former usually persisting, the latter

caducous, sometimes appendaged. Stamens as many as petals,

often more numerous, all fertile or the epipetalous staminodal, the

filaments more or less united; anthers versatile. Disk lacking or

interstaminal, free or attached to tube, or extrastaminal, forming a

ring attached to tube. Ovary sessile, cells same number as petals,

2 or 3 often with more or less developed septa. Styles free or more

or less united. Ovules mostly 2 and pendulous from apical inner

angle. Fruit a capsule often dividing into as many parts as cells,

a 1-seeded drupe or sometimes fleshy, with 1 seed. — A very ancient

family therefore now with many gaps and the sequence of the under-

groups not discernible (Winkler).

Flax, Linum usitatissimum, one of the most ancient of cultivated

plants, is no longer known "wild" or native.

Stamens as many as petals; herbs, sometimes woody toward base.

1. Linum.

Stamens 2-several times as many as petals; shrubs or trees.

Styles 3-5; stamens 10-15.

Styles 5; stigmas small; petals appendaged 2. Hebepetalum.

Styles 3; stigmas large; petals unappendaged 3. Roucheria.

Style 1; stigmas 10, 20 or many.

Stamens 10-20; sepals nearly free.

Ovules 1 in each cell; anthers globose, 2-4-celled.

4. Sacoglottis.

Ovules 2 in each cell; anthers 2-celled 5. Houmiria.

Stamens 50-180; sepals more or less connate; anthers 4-celled.

6. Vantanea.

1. LINUM [Tourn.] L.

Reference: Planchon, Lond. Journ. Bot. 6: 588-603. 1847; 7:

165-186, 473-501, 507-528. 1848.

Annual or perennial herbs or the latter often ligneous toward

base, sometimes shrubby, the sessile usually alternate often narrow

622 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

leaves with 1-several parallel nerves. Stipules glanduliform or lack-

ing. Flowers various in color, solitary or often disposed dichasially,

or racemose. Sepals and petals 5, the former entire or fimbriate or

glandular, the latter fugacious. Stamens more or less connate, 5,

and with minute or setiform staminodia and with 5 nectar glands.

Ovary 5 (2)-celled, 2 ovules to each cell. Styles 5 (2), sometimes

more or less cohering or connate to the middle, rarely to the tip.

Fruit a capsule with ordinarily five 2-seeded imperfectly septate cells.

— There is no modern study upon which to draw the following compi-

lation of the species reported for Peru; it is possible that earlier names

given to species in Ecuador and Chile if not North America may

prove to refer to the same plants. Fruiting material may simulate

Hypericum with pellucid-puncticulate leaves.

Petals shorter than the sepals L. selaginoides.

Petals longer than the sepals.

Flowers large, blue (except white color form); leaves several cm.

long L. usitatissimum.

Flowers yellow or white; leaves 1-2 cm. long, rarely some of them

a little longer.

Stems minutely scabrous-glandular; annual (biennial?).

L. parvum.

Stems glabrous or obscurely scabrous; more or less ligneous-

based perennials, or even shrubby.

Styles connate to tip; basal leaf glands soon callose.

L. Macraei.

Styles free, connate at base or more or less cohering, especially

below the middle; species poorly understood or variable,

the following key arbitrary, ex char.

Leaves to 1 mm. wide; pedicels to 8, seeds to 1 mm. long;

tufted alpine L. andicolum.

Leaves (1)1.5-3.5 mm. wide; plants soon lax or shrubby;

seeds of L. Weberbaueri 1.2 mm. long.

Pedicels 6-12 mm. long; middle leaves in type 1-1.5 mm.

wide L. Weberbaueri.

Pedicels finally at least in part somewhat longer than

calyx; larger leaves 2-3.5 mm. wide. .L. prostratum.

Linum andicolum Krause, Bot. Jahrb. 40: 278. 1908.

Perennial glabrous herb rarely attaining 10 cm., the many stems

divergent from a common base, often decumbent, simple, or sparsely

FLORA OF PERU 623

branched, striate; leaves approximate, persisting, alternate or the

lower opposite, 1-nerved, linear or linear-lanceolate, shortly acumi-

nate, with a pair of globular purplish glands at the subacute base,

the intermediate 4-6 mm. long, scarcely 1 mm. wide, much longer

than the internodes; flowers terminal and solitary or 2-3 in cincinni;

pedicels slender, erect, 3-8 mm. long; sepals broadly ovate, longish

acuminate, 3-5-nerved but the lateral nerves often obscure, 3-3.5

mm. long, nearly 2 mm. wide; petals obovate, subacute or obtuse,

gradually narrowed to base, many nerved, 7-8.5 mm. long, nearly

5 mm. across above, yellow; staminodia small; staminal tube hardly

1 mm. long, with linear acute teeth little shorter between the fila-

ments, these filiform, dilated below, about 3.5 mm. long, the oblong

anthers obtuse, lightly sagittate at base, 1-1.2 mm. long; styles

slender, erect, connate below, 4-4.5 mm. long, stigmas capitellate;

capsule subglobose, little longer than the calyx, about 3.5 mm. broad,

the compressed ellipsoid seeds somewhat curved, reddish, to 1 mm.

long, scarcely 0.5 mm. broad. — Perhaps is a young tufted early

flowering state of L. prostratum. In habit recalls L. Chamissonis

Schiede, Linnaea 1: 69. 1826 but that species with no conspicuous

leaf glands (Krause) ; but this character in all species in Peru seems

to vary. F.M. Neg. 18010.

Junin: Grass steppes south of Tarma, 3,300-3,700 meters, Weber-

bauer 2407, type; 182.

Linum Macraei Benth. Bot. Reg. 16: sub. pi. 1326. 1830; 489.

Glabrous, shrubby below, the virgate or somewhat branching

upper stem herbaceous, striate, often 3-4 dm. long; lower leaves

crowded, 10 mm. long, 2 mm. wide, intermediate and upper narrowly

linear-lanceolate or subulate, rigid, pungent, opposite or alternate,

10-16 mm. long, 3-4 mm. wide; pedicels about 5 mm. long to twice

as long in fruit; sepals ovate, acutely acuminate, margins glandular;

petals twice as long as the calyx equaled by the style, this com-

pletely connate to the quinfid apex; stigmas globose; capsule mucro-

nate by the style base. — The Peruvian plant, scarcely distinct, may

be designated L. Macraei forma peruviana Macbr., forma nov.

foliis ad basin bicalloso-glandulosis floralibus alternis; sepalis haud

glandulosis obscure fimbriatis. According to Planchon, I.e. the

leaves of L. Macraei are eglandular at base but apparently, if one

may judge from modern materials referred here, the character is

variable. Specimens from Arequipa placed in this work with doubt

in L. prostratum resemble this species more, except for the connate

styles.

624 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Ancash: Ocros, 2,800 meters, Weberbauer 5812 (type, forma).

Chile.

Lin u in parvum Johnst. Contr. Gray Herb. 85: 172. 1929.

Erect herbaceous annual (or apparently sometimes enduring),

4-9 cm. high, the solitary minutely stipitate glandular stem laxly

and sparsely cymose-branched above; leaves few, oblanceolate or

linear, 5-10 mm. long, 1.5-2 mm. wide, costate but enervate, entire,

acute, sparsely glandular and with 2 brown globose glands at the

attenuate base; flowers solitary and apical or in unilateral cymes or

these dichotomous, 3-6-flowered; pedicels 0.3-0.9 mm. long; sepals

3- or rarely 5-nerved, broadly lanceolate, entire, 3 (-5) mm. long,

herbaceous and recurved at tip, sparsely pubescent within; petals

yellow, 4.5-5 mm. long, 1.5 mm. wide, oblong-lanceolate, rounded

or emarginate; stamens to 0.5 mm. long-connate, scarcely longer

than the styles, these 2.5 mm. long, more or less connate below the

middle, the stigmas capitate; capsule globose, 3-3.5 mm. broad,

minutely stipitate glandular, bay-colored above the middle. — Rela-

tionship uncertain but no other western South American annual

species has the abundant stipitate glands that, minute, are con-

spicuous on this annual of the loma formation (Johnston). Glands

extremely minute, obscure in Stafford specimen, which, moreover,

is surely more enduring than annual.

Arequipa: Local, sandy places, lower edge of fertile belt in hills

of Mollendo, (Johnston 3549, type); also, the yellow flowers red-

striate, Stafford 823.

Linutn prostratum Lam. Encycl. 3: 525. 1791; 488. L. poly-

galoides Planch. Lond. Journ. Bot. 7: 487. 1848, ex char.

Glabrous perennial, the much divided caudex with many slender

prostrate-ascending stems 7-30 cm. high, woody below, striate,

often rubescent, the leafy (in age woody and leafless below) flowering

ones dichotomously branched toward the tips; leaves oblong-lanceo-

late, acutish, somewhat or little attenuate at base, 10-15 mm. long,

(2) 3-5 mm. wide, often soft-membranous, pale green, lower sub-

opposite, the upper alternate, gradually reduced; basal glands

inconspicuous or obsolete or sometimes solitary or binate and better

developed; flowers typically opposite reduced leaves on short

branchlets, subsessile or apparently, sometimes well-pedicelled ; fruit-

ing pedicels 2-4 mm. long, perhaps rarely longer; styles shortly

connate below; sepals obviously unequal, or subequal, acute, or

FLORA OF PERU 625

abruptly acuminate, rigid, entire, obscurely nerved except midnerve

at base, subglandular-fimbriate, the longer exceeding the depressed

ovoid brownish capsule. — Except for sepals this after Planchon in

part to include his species, his type in herb. Hook. & Lindley not

seen. The Arequipa material is shrubby with longer pedicels. F.M.

Neg. 18103.

Lima: In dry hills near Chancay, Dombey; also Ruiz & Pavdn,

type. Purruchucha, Cuming 586 (det. Planchon, L. polygaloides).—

Cajamarca: Near Celendin, 2,900 meters, petals vivid yellow,

Woytkowski 15? — Junin: Cerro de Pasco, (Mathews 615, type, L.

polygaloides). — Huancavelica: Near Pampas, gravelly hills, 3,200

meters, flowers yellow, Stork & Norton 10249. Prostrate in clay,

flowers yellow, brown above, near Cordova, Metcalf 80254. — Cuzco:

Huancaro, Vargas 3162. — Ayacucho: Toward Lucamas, 3,200 meters,

Metcalf 30321. — Arequipa: Without data, Isern 2503. Mount

Chiwata, shrub 0.5 meter high, Eyerdam & Beetle 22128. Near

Mejia, (Guenther & BuchtienlSS; 188a). Pasco, (Guenther & Buchtien

Linum selaginoides Lam. Encycl. 3: 525. 1791; 178.

Glabrous, the many stems more or less woody and leafless toward

the base, where simple or often branched, ascending, densely leafy

toward the usually corymbose tip, commonly 1-2 dm. long or longer;

leaves alternate, imbricately crowded, subulate, mucronate-fili-

ferous; flowers subsessile, solitary or few, terminal; petals whitish-

pink, clawed; capsule obovoid-globose, obtuse or depressed, com-

pletely 10-celled. — The var. chilensis Planch., I.e. 179, has a much

divided subterranean caudex, larger globose capsules. — Winkler,

Pflanzenfam. ed. 2. 19b, includes Peru in the species' range without,

however, data, and I rather doubt its occurrence, but a Lechler

specimen cited in Mart. Fl. Bras. 12, pt. 2: 470 might be from

Puno. F.M. Neg. 35184.

Peru: Fide Winkler. Chile; Uruguay; Brazil.

Linum usitatissimum L. Sp. PI. 277. 1753; 165.

Typically annual, the tall stem simple at least at base, often

several dm. high, the sessile linear or linear-lanceolate acute 3-nerved

leaves 1-3 cm. long; pedicels finally 2 cm. long or longer; sepals

acuminate, the elliptic outer 7-9 mm. long in fruit, the smaller

inner more ovate, all 3-nerved at base; petals somewhat crenate;

styles nearly or quite distinct; capsules barely longer than calyx,

626 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

the septa or partial septa glabrous; seeds to about 4 mm. long.—

It is interesting that the widely cultivated flax varies in duration,

size of seeds, and otherwise. One of the most ancient of intentionally

grown plants, the generic name itself derived from one of great

antiquity.

Puno: Near Puno, 3,830 meters, rarely, as here, in cultivation,

Soukup 87. Origin probably oriental; cultivated in all temperate

regions. "Lino."

Linum Weberbaueri Krause, Bot. Jahrb. 40: 277. 1908.

Suffrutescent, glabrous, erect, to 4 dm. high with many sub-

virgate stems and branches sparsely leafy especially below, where

more or less ligneous, herbaceous above, scarcely 1 mm. thick and

8-10 lineately striate; leaves alternate or the lower binate, approxi-

mate and subopposite, tardily deciduous, linear-lanceolate, acumi-

nate, narrowed to base, this with a pair of brownish-purple sub-

globose glands, 1-nerved, the intermediate 6-8 mm. long, 1-1.5 mm.

wide, mostly little longer than the internodes; flowers solitary at

branchlet tips on pedicels 6-12 mm. long; sepals 3-nerved, the mid-

nerve prominent, ovate, acuminate, entire, biglandular at base,

2-2.3 mm. long, hardly 1.5 mm. wide; petals broadly obovate to

rhomboid, obtuse or lightly emarginate, somewhat clawed, 6-7 mm.

long, the upper third to 4.5 mm. wide, yellow; staminodia inconspicu-

ous; staminal tube 0.8 mm. long, with 1 acute linear tooth between

the filaments, these little dilated below, 4.5 mm. long, the ovate-

oblong little emarginate anthers 1 mm. long; styles erect, 3.5-4 mm.

long, one-quarter connate, stigma capitellate; ovary ovoid, the

capsule little longer than calyx, about 3 mm. across, the strongly

compressed brown seeds about 1.2 mm. long, 0.6 mm. wide.—

After Krause. Allied by the author to L. ramosissimum Gay, Fl.

Chile 1: 463. 1845, of northern Chile from which it differs (he

thought) in habit, long pedicels, mostly alternate leaves, more

prominent glands; but these are variable characters. Also, Reiche

regarded the Gay plant as only a variety of L. Chamissonis; cf.

L. andicolum, L. prostratum. All the Peruvian material of Linum

has similar sepals, outer essentially 1-calloused, nerved at base, inner

faintly to obviously fimbriate. Also to be considered here are the

Ecuadorean L. filiforme Urban, Linnaea 41: 643. 1877 and L. oligo-

phyllum Willd. ex. Schult. Syst. 6: 758. 1820, "calyx ovate, acumi-

nate, stems branched, striate, scabrous, leaves linear, acute, squami-

form, flowers solitary," origin unknown. Whether or not this is

L. Weberbaueri it seems sensible to drop the name as essentially

FLORA OF PERU 627

a nomen nudum and a source of confusion in view of the well-

described plants. P.M. Neg. 18015.

Junin: In mountains near Palca, 2,500 meters, Weberbauer 1785,

type; 248. — Huancayo: Near Huancayo, 3,317 meters, Soukup

2946.— Lima: Below Surco, 2,000 meters, Weberbauer 3215 (det.

L. oligophyllum, herb. Dahlem). — Ayacucho: About 3,000 meters,

Weberbauer 5523 (det. Dahlem). — Libertad: Cachicadan, Stork &

Horton 9954 (det. L. Macraei, Standley). Near Angasmarca, West

8169 (det. Johnston, L. oligophyllum).

2. HEBEPETALUM Benth.

Glabrous trees, with alternate coriaceous entire or lightly un-

dulate-crenate leaves and small very caducous broadly triangular

stipules. Flowers small, orange-yellow (or white?) in panicled

cymules, terminal or in the upper axils. Sepals and petals 5, the

former very unequal, the latter villous within and with a tiny lamella

or appendage above the claw. Stamens 10, connate at base and

coalescent with more or less developed glands. Ovary 3-5-celled,

1-2 ovules in each cell; styles 5, connate below or perhaps also free.

Drupes ovoid or globose. — Apparently there is no fundamental

difference between this South American tree and species of Hugonia

L. from Africa and Malay except that they have larger flowers, a

comparative situation that recalls certain genera in the Ericaceae.

It seems open to doubt that the petal appendages, alike for both

Hugonia and Hebepetalum, developed independently; probably only

one genus is concerned.

Hebepetalum humiriifolium (Planch.) Benth. in Benth. &

Hook. f. Gen. PI. 1: 242. 1862. Roucheria humiriifolia Planch, in

Hook. Lond. Journ. Bot. 6: 143. 1847.

Branches terete, leafless below; petioles 2-5 cm. long, margined,

canaliculate above, reddish resiniferous at base; leaves few, spreading,

oblong, obtuse or shortly acuminate, obtusely serrate, 8-13 cm. long,

3.5-5 cm. wide, chartaceous, lustrous, the midnerve impressed above,

acute beneath, the lateral nerves slender, spreading, nearly straight,

the fine veins parallel-striatulate; panicles terminal, shorter than the

leaves, with 4 or 5 angulate-compressed branches scarcely 5 cm. long;

pedicels nodiform; calyx cupulate, 2 mm. high, pellucid-punctate;

fruits tipped with the 5 styles, blue-black. — Flowers orange, tree

8-22 meters (Klug).

628 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Loreto: Florida, King 2339 (det. Sandwith). Mishuyacu, King

683. To the Guianas.

3. ROUCHERIA Planch.

Similar to Hebepetalum but petals not clawed, stamens 10-15,

the high tube apparently lacking glands without, and the styles

3 with stigmas much broader. Ovary 3-celled but 2 cells often

collapsed. Fruit incompletely known. — Named for the romantic

French poet Roucher, politically unfortunate.

Roucheria calophylla Planch, in Hook. Lond. Journ. Bot. 6:

142. 1847.

Branches terete; petioles 6 mm. long, leaves attenuate to base,

5-7.5 cm. wide, 2.5-3 dm. long, paler beneath, rigid-chartaceous,

nerves prominent both sides, under lens obsoletely serrulate; bracts

and calyces minutely pubescent; flowers in fascicles of 7 or 8, the

narrow petals fugacious; drupe ovoid.

Peru: (Probably). Brazil; Guiana.

4. SACOGLOTTIS Mart.

Rather similar to Houmiria but stamens 10 or 20, sometimes

connate only at base, often with setiform staminodia, the 5 longer

sometimes apically 3-dentate and 3-antheriferous. Anthers 2-4-

celled. Intrastaminal disk dentate, parted or squamate. Ovary

5-celled, the 1-ovuled cells episepalous. Flowers sometimes yellow.—

For a key to the Amazonian species of Brazil see Ducke, Archiv.

Inst. Biol. Veg. Rio Jan. 4: 28-29. 1938.

The large fruits of S. amazonica Mart, (and probably other

species) are common drift fruits according to R. 0. Williams, Fl.

Trinidad & Tobago 1: 151. 1929.

Sacoglottis Uchi Huber, Bol. Mus. Paraense 2: 489. 1898.

A large tree, glabrous except the inflorescence, the branchlets

slender; leaves distichous, oblong-lanceolate, more or less acuminate

each end, mostly 9-17 cm. long, dentate or crenate-serrate (early

marginally glandular); peduncles glabrous or nearly, the more or

less regular cyme trichotomous or dichotomous in the leaf-axils of

the new branches or in the newer leaves of the past year's branches,

all together forming veritable bouquets of many small yellowish-

green flowers mostly in 3's, the branchlets hirsutulous; sepals in-

FLORA OF PERU 629

distinctly imbricate, rounded; petals oblong or nearly, pubescent

both sides; stamens all fertile, the larger 4, the 2 smaller with globose-

elliptic anthers, filaments papillose, variable as the long-extended

connectives in shape, sometimes irregular or partly serrate; disk

composed of 10 flat free ovate-lanceolate scales half enclosing the

ovary; style as long, stigma 5-lobed; drupe greenish, 7 cm. long,

oblong-ellipsoid. — Illustrated, Huber, opposite page 490 (flowers).

Perhaps most likely to occur within Peru as, fide Ducke, widely

distributed from Para to the Purus and Solimoes. Said to attain

30 meters. Cultivated in Amazonian area for the fruits. Also

expected to occur within Peru is S. reticulata Ducke, Archiv. Inst.

Biol. Veg. Rio Jan. 1: 206. 1935, with 20 fertile stamens, the 5 larger

trifid with 3 anthers.

Peru: (Probably). Amazonian Brazil.

5. HOUMIRIA Aublet

Somewhat balsamiferous trees or tall shrubs with entire or

crenate leaves that are also glandular-punctate on the margins, and

white or greenish flowers that usually are in axillary or lateral cymes.

Sepals nearly free. Petals 5, narrow. Stamens 12-20, more or less

connate, the free part of filaments densely papillose or warty; anther

cells bearded. Interstaminal disk composed of 20 more or less

united scales. Ovary 5 (4) -celled, the 2-ovuled cells epipetalous;

stigma 5-parted. — Caribbean name Touri, also Houmiri (Aublet);

the author wrote Houmiri which Jussieu, Gen. PI. 435. 1789 latinized

to Houmiria; others, but later, have made other suggestions (St.

Hilaire, Humiria; Richard, Humirium, etc.).

Leaves usually subsessile, eglandular* above at base.

Leaves not auricled H. floribunda.

Leaves auriculately produced at base H. balsamifera.

Leaves petioled, 6-8 glandular above at base H. crassifolia.

Houmiria balsamifera Aublet, PI. Guian. 1: 564. pi. 225.

1775.

Large tree (or shrub) with distichous oblong-lanceolate leaves

that bear a row of approximate glands at the margin beneath be-

tween the obsolete crenulations; branchlets acutely wing-angled by

the decurrent petioles, these 1-2 mm. long, and grooved by the base

of the leaves, this produced a little auriculately and thus rounded-

amplexicaul, the blade 3-4 cm. wide, (6) 9-11 cm. long, eglandular

630 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

above; peduncles little branched, the 30-50-flowered dense cymes

2.5-5 cm. long; pedicels articulate, 1-3 mm. long; calyx glabrous,

0.5-0.8 mm. long, lobes suborbicular, scarcely 1 mm. wide, the

inner much smaller, obscurely or not imbricate; petals free, 4.5 mm.

long, narrowed to base, glabrous or sparsely puberulent; filaments

connate below, densely scabrous-papillate, the globose anther cells

with a few long straggly trichomes, the ligulate connective much

extended above; ovary apically barbate about or at base of the angled

style. — To 20 meters high or taller, the crown much branched.

Illustrated, Mart. Fl. Bras. 12, pt. 2: pi. 92 (flower and leaf).

Loreto: Balsapuerto, shrub in dense woods, Killip & Smith

28681. To the Guianas.

Houmiria crassifolia Mart. Nov. Gen. & Sp. 2: 143. pi. 198.

1826.

Leaves alternate, with petioles 2-2.5 cm. long and broadly wing-

appendaged; peduncles trichotomous, the branches bifurcate; calyx

lobes much imbricate; filaments densely tuberculate. — Type locality,

Serra de Arara Coara, Rio Japura, is near the Peru frontier (Amshoff).

Peru: (Undoubtedly). Brazil; British Guiana.

Houmiria floribunda Mart. Nov. Gen. & Sp. 2: 145. pi. 199.

1826.

Shrub or tree with fragrant balsamiferous bark; petioles alternate,

obsolete or rarely to 2 cm. long; leaves eglandular above at base but

more or less closely so beneath in the entire or repand-crenulate

margin, sometimes also puberulent, chartaceous, lustrous above,

finely veined both sides, variable in form but usually about elliptic,

the larger 2-6 cm. wide, 2.5-10 cm. or even 15 cm. long, shortly

and obtusely (or acutely) acuminate to emarginate; peduncles lateral,

alternately forked, wing-angled, the inflorescence 2-8 (14) cm. long;

pedicels 1-5 mm. long; calyx glabrous or pubescent, 0.7-1.5 mm. long,

the lobes more or less imbricate, ciliolate; petals 4.5-6 mm. long,

white or greenish; filaments connate to middle, densely papillose-

scabrous; globular anther cells with liguliform extended keeled con-

nective; ovary apically barbate about the filiform obscurely angled

style; drupe narrowly to broadly ellipsoid or obovoid, 8-14 mm.

long, glabrous, the flesh thin, seeds 2-4. — There are several named

varieties of doubtful merit. Tree 18 meters high, flowers white

(Williams); pale yellow ( Weberbauer) ; orange (Klug). Common

throughout the hyle"a (Ducke). Illustrated, Mart. Fl. Bras. 12,

pt. 2: pi. 92 (fruit).

FLORA OF PERU 631

San Martin: Campana near Tarapoto, Spruce 4335; Raimondi.

Moyobamba, Weberbauer 4478. Zepelacio, Klug 3706 (det. Stand-

ley). — Loreto: Mishuyacu near Iquitos, Klug 1815; 685 (det. Mol-

denke). Balsapuerto, Klug 2846 (det. Standley). Yeveros, Rai-

mondi. Brazil. "Umiry." "apacha-rama."

6. VANTANEA Aublet

Trees, often tall, or shrubs, with entire leaves and white or red

flowers. Sepals more or less connate and often with a dorsal gland;

stamens many, basally connate, the inner sometimes staminodal,

the anther cells 4. Intrastaminal disk fleshy, subentire or irregularly

dentate. Ovary 4-6-celled the 2-ovuled cells episepalous. Stigma

obsoletely 4-6-lobed. Drupes smooth, rugulose or rarely tubercu-

late. — Ducke, Archiv. Inst. Biol. Veg. 4: 31. 1938, has given a key

to the Amazonian species of Brazil.

Vantanea cupularis Huber, Bol. Mus. Goeldi 6: 83. 1910.

Dicania celativenia Standley, Field Mus. Bot. 17: 254. 1937, fide

Ducke.

Branchlets slender, gray-barked, few leaves toward the tips;

petioles 1-2 cm. long, grooved above; leaves distichous, glabrous,

elliptic, acute at base, shortly and obtusely complicate-rostrate,

6-10 cm. long, 3-5 cm. wide, subcoriaceous, lustrous and prominently

nerved and veined both sides, the reticulation lax; inflorescence

terminal or in the upper axils, 2-3 puberulent commonly 4-dicho-

tomous cymes 5 cm. long and broad, peduncles 2-2.5 mm. long, bracts

and bractlets early deciduous, pedicels to 2 mm. long; buds 6-7 mm.

long, the calyx 1 mm. long, broadly cupulate, subglabrous, margin

scarcely undulate; petals whitish, linear, 8 mm. long, 2 mm. wide,

obtuse or acutish or cucullate, ashy hirtellous without, soon decidu-

ous; stamens about 80, the longer little shorter than petals, anther

connective acute, elongate; disk denticulate, tomentose as shorter

ovary. — Probably in Peru as Ducke records it as nearly throughout

the Amazon in non-flooded forests, Para to Rio Solimoes, a tree

6-30 meters tall or taller. V. tuberculata Ducke, I.e. 31, from the

upper Amazon, has a tubercled fruit. Standley's plant, ex char.,

differs in stout brown-barked branchlets, rigid-coriaceous obovate-

oblong leaves 6-11.5 cm. by 3.5-6 cm., broadly rounded at tip and

lightly emarginate, nerves manifest but not elevated, the veins

obsolete both sides, fruiting inflorescence sparsely pilose, fruits

sessile, the immature 2 cm. long, subglobose, densely puberulent-

632 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

tomentose and sparsely hirsute, or 3 cm. long, oblong-ovoid, acute;

and if it belongs here the description of V. cupularis must be changed

to include the above data.

Peru: (Probably, as noted). Brazil.

Vantanea peruviana Macbr. Candollea 5: 371. 1934.

Glabrous except the inflorescence; branches densely branched,

the stout branchlets more or less tuberculate; petioles 2.5 mm. long;

leaves broadly elliptic or elliptic-obovate, shortly cuneate at base,

rounded-retuse at apex, to 8 cm. long, 4.5 cm. wide or mostly about

5 cm. long, 3.5 cm. wide, rigid-coriaceous, lustrous above, opaque

beneath, the nerves scarcely obvious; inflorescence 3 cm. long, 5 cm.

broad; branchlets and very short pedicels angled, sparsely and

minutely puberulent or glabrate; calyx shortly 5-lobed, the lobes

eciliate; buds nearly linear, 10 mm. long; staminodia none(?). — In-

completely described but thought to be nearest V. minor Benth.

of British Guiana, the flowers notably shorter, the calyx glabrous.

A tree about 12 meters high.

Loreto: Mishuyacu, near Iquitos, Klug 1130, type.

ERYTHROXYLACEAE [HBK.] Lindl.

Reference: 0. E. Schulz, Pflanzenreich IV. 134. 1907.

Glabrous shrubs or trees with entire usually alternate leaves, a

single often conspicuous interpetiolar persistent or caducous stipule

(rarely, not in Peru, stipules binate), and small flowers ordinarily

a number or many borne in the axils of leaves or branchlets. Pedicels

more or less enlarged apically, 5-angled, with 2, exceptionally 4,

basal bracts. Flowers actinomorphic, 5-merous, almost always

hermaphrodite. Calyx persisting, sometimes divided to base, the

division somewhat imbricate. Petals 5, free, usually with a scale

or appendage within at base. Stamens 5, biseriate, the filiform

filaments more or less connate; anthers cordate, 2-celled, longitudi-

nally dehiscent. Pistils 3, free or partially connate, usually only

1 cell of the ovary fertile, the pendulous ovules 1 or 2. Fruit

drupaceous, the seed with or without albumen.

1. ERYTHROXYLUM P. Brown

Essentially the only genus — the other African with a single

species — and thus character of the family. Branchlets somewhat

compressed. Petals oblong or nearly, often obtusish, obviously un-

FLORA OF PERU 633

guiculate. Filaments urceolately connate. Ovary often truncate.

Drupe at maturity red. — The name, being composed of the Greek

words meaning "red" and "wood," has been written "Erythroxylon."

Some species, as E. Coca, are of great economic and social signifi-

cance, as the leaves contain the important alkaloid cocaine. The

use of the leaves in Peru was described by Monardes, a physician

of Seville, in 1580, but botanical specimens were first brought to

Europe by the French botanist Joseph de Jussieu, who, on the ex-

pedition of Condamine, observed its daily use by many of the

inhabitants of the Andes. Schulz gives a partial bibliography; the

early account of J. H. Uanul, 1794, in Lima, became a classic in

the literature on coca.

The following descriptions are essentially or mostly after Schulz

and the key is only suggestive; there may be too many species

names — or too few — for some collections determined by the late

monographer, or at least with his assistance, agree poorly with

characters; after nearly fifty years' accumulation of materials since

Schulz's revision it is not surprising that a new study is in order.

The publication of E. Coca var. Spruceana Burck, Teysmannia 1 :

455. pi. 2. 1890, based on a cultivated shrub supposed to have

originated in Peru, has not been seen; its origin apparently was an

error, for a cited synonym is E. truxillense Rusby, Amer. Drug.

Circ. & Chem. Gaz. 44: 220. 1900 and 45: 49. 1901.

The term "leaves bilineate" in the key refers to lines apparently

in the leaf-nervation that parallel the midnerve; they are constant

enough in some species to furnish a convenient key character (as

by Schulz himself, I.e. 70 and 71) but that they are discernible is

not of taxonomic significance; cf. Schulz, I.e. 4, who quotes Nevinny,

the first to show that the lines are merely marks or pleats which

result from the manner in which the young leaves are developed.

Stipules obviously striate.

Calyx lobes somewhat imbricate especially in bud, in anthesis

this mostly apparent by a pleat in the tube.

Petals longer than the calyx lobes; flowers often many (20-40).

E. floribundum.

Petals and calyx subequal or the latter longer; flowers usually

fewer.

Calyx lobes lanceolate, to 6 mm. long; stipules 2.5-5 cm.

long; leaves long-cuneate obovate, to 4 dm. long or longer.

E. macrocnemium.

634 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Calyx lobes ovate, shorter; stipules shorter; leaves smaller,

mostly more abruptly and shortly narrowed to base.

Calyx lobes rather acuminate; leaves often a dm. and a

half long or longer, nervation rather prominent.

E. macrophyllum.

Calyx lobes acute; leaves mostly shorter, rather obscurely

reticulate E. lucidum.

Calyx lobes distinct.

Leaves more or less acuminate or acute.

Stipules marginally fimbriate and the setae fimbriate-ciliate.

E. fimbriatum.

Stipules and setae not at all or obscurely fimbriate.

Staminal tube^half as long as calyx; petioles slender, 6-10

mm. long E. amplum.

Staminal tube little if at all shorter than calyx; petioles

3-6 mm. long, usually stout.

Stipules conspicuously long and long-bisetose.

E. comosum.

Stipules shortly trisetulose.

Pedicels 1-3 (5) mm. long; leaves often large, shortly

to very shortly acuminate, mostly elliptic-obovate.

E. amazonicum.

Pedicels mostly 4-8 mm. long; leaves mostly long-

acuminate, about 1 dm. long.

E. acuminatum, E. paraense.

Leaves rounded and often minutely retuse at tip.

Leaves broadly elliptic, 3-5 cm. wide E. Raimondii.

Leaves more or less oblanceolate, 2-3 (4) cm. wide.

E. deciduum.

Stipules smooth, estriate unless faintly in E. Ulei.

Leaves rounded at tip but often acutish or (and) mucronate,

mostly 3-6 cm. long.

Leaves rarely 18 mm. wide E. Tessmannii.

Leaves, at least mostly, 2-5 cm. wide.

Stipules faintly striate E. Ulei.

Stipules not at all striate, smooth.

Leaves obscurely if at all mucronate, not bilineate.

E. hondense.

Leaves mucronate, often bilineate E. Coca.

FLORA OF PERU 635

Leaves more or less acuminate, sometimes shortly and obtusely,

mostly 7-10 cm. long or longer.

Leaves obtusely acuminate, chartaceous to coriaceous.

Stipules faintly striate; leaves usually less than 7 cm. long.

E. Ulei.

Stipules not at all striate.

Leaves elineate, usually long-acuminate.

E. luculentum, E. Mamacoca.

Leaves bilineate, coriaceous, very shortly acuminate.

E. Shatona.

Leaves acutely acuminate, obviously membranous.

E. gracilipes.

Erythroxylum acuminatum R. & P. Fl. Peruv. 4. pi. 399.

1802; 45. E. patens Ruiz, ex 0. E. Schulz, Pflanzenreich IV. 134: 45.

1907.

Becoming a tree 6 meters high, the slender branches gray, opaque,

little verruculose, the erect-spreading branchlets subangled, 1.5 mm.

thick; stipules subpersisting, 3 mm. long or on innovations to 9,

between leaves to 11 mm. long, striate, broadly lanceolate, long,

3-setulose at the obtusish apex; petioles 3.5-4 mm. long; leaves

obovate or broadly elliptic, acutish at base, obtusely short-acuminate,

7-12 cm. long, 3-6 cm. wide, papyraceous, gray-green above, paler

or subrusty and opaque beneath, the midnerve there very obtuse,

the subhorizontal laterals elevated, these sulcately impressed above;

flowers 4-12; pedicels 4-8 mm. long; bracts ovate, acute, 1 mm. long;

calyx lobes lanceolate, obtusish, 1.5-2 mm. long; petals 3-3.5 mm.

long, one third clawed, the ligule one-third as long as blade; lateral

auricles three times longer than posterior; stamen cup shorter

than calyx the ovary and cup subequal the latter 10-crenate, the

stamens 4.5 mm. long, the styles 2 mm. long or in long-styled flowers

4.5 mm., the stamens 1.5 and 2.5 mm.; drupes about 12 mm. long,

4.5 mm. across, oblong-ovate, acutish, subtrigonous (after icones).—

The species having been illustrated with details it is to be accepted,

following precedent, in this and other works. The flowers are said

to be very fragrant. Suggests E. paraense but perhaps differs in

more persisting, rather long-setose stipules, proportionately broader,

slightly thinner leaves, longer pedicels. However, I have not been

able to see the monographer's character of "more persisting stipules"

and it is possible that the type sheets ought to be interpreted more

636 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

liberally, which would perhaps permit the inclusion of at least the

Peruvian material of E. paraense.

Huanuco: Puente of Pillao, Ruiz & Pavon in 1788, type; also at

Chachahuasi. Ecuador.

Erythroxylum amazonicum Peyr. in Mart. Fl. Bras. 12, pt. 1:

167. 1878; 34.

Much like E. paraense; branchlets said to be more strongly com-

pressed, 1.5-2.5 mm. wide; stipules caducous, obtusish, very shortly

3-setulose, 6-15 mm. long; petioles 3.5-5 mm. long; leaves somewhat

obovate, cuneate-narrowed to the acutish base, 6-13 cm. long, 3-6

cm. wide, coriaceous, not diaphanous, the lateral nerves both sides

but especially beneath little prominent; flowers 3-9, the pedicels

1-5 mm. long, stout, enlarged toward apex; calyx nearly one-half

divided, the lobes 1-1.2 mm. long; petals 3-4 mm. long; staminal

tube about as long as calyx; mature drupes 12-16 mm. long. — As

remarked by the monographer both these species comparable to

E. citrifolium St. Hil., 36, widely distributed from Mexico to the

Guianas and southern Brazil; it seems possible that a single variable

species is concerned and in this case the name of St. Hilaire has

precedence, or perhaps even E. acuminatum R. & P. However,

according to Schulz E. amazonicum is distinct from both E. paraense

and E. citrifolium by its coriaceous not pellucid leaves, pedicels

strongly enlarged at apex, cup and calyx subequal, drupes large,

while E. citrifolium is said to differ from E. paraense in less obtuse

stipules, leaves long acuminate, lateral nerves manifest beneath, all

dubious distinctions, it seems to me. Cf. also E. amplum and

E. acuminatum. F.M. Neg. 19451.

Loreto: Mishuyacu near Iquitos, Klug 82; 549; 610; 1144;

Williams 3759. — Rio Acre: tree 25 meters high, Rio Macauhan,

Krukoff 5526; 5748 (so distributed). Brazil to Trinidad.

Erythroxylum amplum Benth. in Hook. Lond. Journ. Bot. 2:

372. 1843; 34.

Much like E. floribundum, fide Schulz, but the shorter triangular

calyx teeth 1.5 mm. long and not at all imbricate; petioles 6-10 mm.

long; leaves 13-24 cm. long or longer, 4-8.5 cm. wide or wider;

flowers 12-30, the petals 4-4.5 mm. long, scarcely more than one-

third clawed, the coarsely denticulate lateral auricles twice as long

as the posterior; brachystylous flowers with obovate truncate ovary,

the style 1.2 mm. long, the dolichostylous with episepalous stamens

2.2 mm. long. — Apparently a shrub or small tree. Notwithstanding

FLORA OF PERU 637

his comparison Schulz placed this with the segregates of E. citri-

folium, which is noted here under E. amazonicum and to which species

I have referred with some reluctance the specimens I determined at

Dahlem, with the friendly help of Schulz, as "E. amplum"; but

the petioles of that as to type are longer and, especially, the staminal

cup is only about half as long as the calyx; it is possible of course

that these characters are not significant. F.M. Neg. 12606.

Peru: (Perhaps). Brazil; British Guiana.

Erythroxylum Coca Lam. Encycl. 2: 393. 1788; 83.

A smooth densely leafy shrub ordinarily between 1 and 2 meters

high, the reddish-brown bark closely verruculose, the many suberect

branchlets compressed, about 1.5 mm. thick; stipules persisting,

3-3.5 mm. long, acutish and very shortly bisetulose, many at base

of innovations and between the leaves; petioles 2.5 mm. long; leaves

elliptic or somewhat obovate, acute at base, acutish or rarely rounded

at tip but obviously mucronate, usually 4-7 (10) cm. long, 3 (2)-

4 (4.5) cm. wide, the midnerve impressed above, obtusely prominent

beneath, the approximate slender lateral nerves equally conspicuous

both sides and reticulate with the secondary, membranous and

diaphanous with longitudinal lines more or less apparent, pale

green and slightly lustrous above, glaucescent or slightly brownish

and opaque beneath; flowers 6-12 (-20) in axils of leaves and branch-

lets, the pedicels 3.5-4 mm. long; calyx to three-fourths parted, the

segments ovate, acute, 1 mm. long; petals 4.5 mm. long, broadly

elliptic, shortly clawed, with ligule one-half its length; lateral auricles

denticulate, the much shorter posterior minutely tridenticulate;

staminal cup as long as or longer than calyx, 10-crenulate; stamens

in short-styled flowers 4 mm. long, the styles only half as long, in

long-styled flowers 2 and 3 mm. long, the styles 4 mm. long; drupe

orange when ripe, 7-8 mm. long, 3.5-4 mm. thick, oblong-ovoid,

acutish, trigonous, sulcate. — Illustrated, Schulz, 84; Bot. Mag.

pi. 7334.

Commonly cultivated even in antiquity and either native or

established in warm moist regions at middle elevations; the well

known "coca" and the source of the drug cocaine; cf. Gosse in Mem.

Cour. Acad. Roy. Sci. Belgique 12. 1861, a standard work. The

active principle was isolated as cocaine in 1858 in Berlin by Niemann

who was aided by the collections made by Spruce.

San Martin: Near Tarapoto, Williams 5814, et al. — Junin:

Puerto Bermudez, Killip & Smith. — Huanuco: Chinchao, Dombey.

638 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Pillao, Ruiz & Pavdn. Rio Chinchao, 4972. Huanuco, Haenke.—

Amazonas: Moyobamba, Stuebel 65. — Loreto: Yurimaguas, Williams

4647. Iquitos, Klug 1117,et al. — Ayacucho: Near Aina, in thickets,

Killip & Smith 22746. — Cuzco: Cosnipata, Weberbauer 6 941 a. San

Miguel, Urubamba Valley, Cook & Gilbert 1127; 1089; also at Santa

Ana, 1574 and Hen era 2638.— Puno: San Govan, Lechler 2220.

Without locality, Jussieu, type. Bolivia; Brazil. "Coca," "cuca-

cuca," "el coq."

Erythroxylum comosum 0. E. Schulz, Pflanzenreich IV. 134:

32. 1907.

Much branched shrub, the tips of the young branchlets comose

because of the long (to 2 cm.) stipules that equal or exceed the new

leaves; stipules linear-lanceolate, striate, the bifid apex terminating

in setae 2-4 mm. long; petioles 2-2.5 mm. long; leaves ovate or el-

liptic, acute at base, shortly and nearly obtusely acuminate, mucronu-

late, about 3 cm. long, 13-16 mm. wide, the midnerve above evidently

acute, prominent and subobtuse beneath, the lateral nerves slen-

der and little conspicuous even below, coriaceous, subdiaphanous,

lustrous especially above; flowers 3-5 in leaf-axils on pedicels 2-3

mm. long, the outer striate bractlet 5 mm. long, 1-setose; calyx

three-fourths parted, the lanceolate free segments 1.5 mm. long;

petals 3.5-4 mm. long, rounded at tip from claw one-half the length,

the ligule as long; lateral auricles joined with inflexed lobe and half

as long as posterior; in long-styled flowers stamen cup and ellipsoid

ovary at least as long as calyx, 10-dentate, the stamens 1.5 and 2

mm. long; style 3 mm. long. — The similar E. Kirkianum 0. E.

Schulz, Repert. Sp. Nov. 30: 179. 1932, Amazonian, is a tree with

narrowly obovate leaves 4.5-7 cm. long, 2-3 cm. wide or larger,

pedicels 1-1.5 mm. long, staminal cup shorter than calyx. Illus-

trated, Schulz, 32.

Loreto: Cerro de Escaler, 1,200 meters, Ule 6531, type.

Erythroxylum deciduum St. Hil. Fl. Bras. 2: 95. 1829; 57.

Shrub or small tree, the grayish or reddish-brown bark of the

branches densely verrucose with rather narrow white lenticels, the

approximate suberect slightly compressed branchlets 2.5-3 mm.

wide; stipules narrowly ovate, striate, minutely 3-setulose, 3-5 mm.

long; petioles 2.5-3.5 mm. long; leaves somewhat oblanceolate,

acutish at base, obtuse and often a little emarginate, reddish-

mucronulate, mostly 6-11 cm. long, about 2-4 cm. wide, coriaceous

FLORA OF PERU 639

(Peru specimen submembranous), diaphanous, lustrous above,

opaque beneath, the midnerve slender and impressed above at

base, obtuse and very prominent beneath, the lateral nerves especially

above obvious, the secondary conspicuously and densely reticulate

both sides; pedicels few to many, 4-20 mm. long; calyx three-

fourths parted, the semiovate acutish lobes about 1.5 mm. long,

distinct; petals 4-5.5 mm. long, broadly oblong, the double partly

3-denticulate appendage barely one-third as long; staminal tube in

brachystylous flowers shorter than calyx and ovary, the stamens

4-5 mm. long; drupes typically 10-13 mm. long, 4.5-5 mm. broad,

subattenuate at base, very obtuse. — The Peruvian specimen, follow-

ing Schulz key, apparently may be assigned readily to this variable

species of southern Brazil and Argentina; the monographer gives

several varieties accounting for variations in foliage, size of flowers

and drupes. Illustrated, Mart. Abh. Math. Nat. Cl. Akad. Muench.

3, pt. 2: pi. 10 (asE. nitidum). F.M. Neg. 35195.

Junin: Huacapistana, shrubby tree with white flowers, Sandeman

4531. Southern Brazil to Paraguay.

Erythroxylum fimbriatum Peyr. in Mart. Fl. Bras. 12, pt. 1:

162. 1878; 38.

Branches terete but with elevated acutely angled decurrent

lines, the pale brown bark obscurely verruculose; branchlets com-

pressed, erect-spreading, about 1.5 mm. thick; stipules persisting,

little shorter than the stout petioles (2.5-4 mm. long), ovate, obtusish,

decurrent, striate, terminated by 3 fimbriate setae, the outer 2

recurving and as long or longer than the body of the stipule, the

middle about half as long; leaves elliptic or oblong-obovate, some-

what narrowed to the obtuse or acutish base, shortly and acutely

acuminate, 6.5-12 cm. long, 2.5-4 cm. wide or wider, callose-punctate,

membranous, very diaphanous, grayish green and lustrous above,

brown and opaque beneath, the acute extremely slender midnerve

sulcate at base above, the many lateral nerves there impressed,

densely reticulate, nervose, with the secondary only beneath; flowers

3-6 in leaf and branchlet axils; fruiting pedicels 5 mm. long; bracts

ovate, striate, long-setose; calyx segments not imbricate, about 1 mm.

long; drupe apparently ovate, about 8 mm. long, 4 mm. thick.—

Stipule illustrated, Schulz, 3. F.M. Neg. 19459.

Rio Acre: Ule 9471 (det. Schulz). Amazonian Brazil.

Erythroxylum floribundum Mart. Beitr. Kenntn. Erythrox.

1840; Abh. Math. Nat. Cl. Akad. Muench. 3, pt. 2: 398. 1843; 25.

640 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Becoming a stout densely branched shrub or small tree, the

terete branches to 5 mm. thick, the erect-spreading compact branch-

lets 2 mm. thick; stipules finally deciduous, 9-15 mm. long, lanceo-

late, acutish, 3-setulose, closely striate; petioles stout, 5-9 mm. long;

leaves oblong-elliptic, acute or rounded at base, more or less acumi-

nate but not mucronate, 10-20 cm. long or longer, about 4-8 cm.

wide, coriaceous but not rigid, subpellucid, gray-green and lustrous

above, glaucous-reddish and dull beneath, the midnerve above

acutish and prominent at tip, very obtuse and red-violet at base

beneath, where the lateral nerves are prominent and with secondary

reticulate; flowers 15-40 on pedicels 2-7 mm. long; outer bract

3 mm. long, ovate, acuminate, striate; calyx divided nearly to base,

the segments especially in bud imbricate, 2-3 mm. long, broadly

ovate, acuminate; petals oblong, broadly clawed, 3.5-4 mm. long,

the ligule half as long; lateral auricles nearly 3 times longer than

posterior, these joined by an inflexed lobule; short-styled flowers

with stamen cup half as long as calyx, 10-crenulate, stamens 3 mm.

long; style 1.5 mm. long; long-styled, the stamens unequal, the styles

3 mm. long; drupe (fide Moore) 7 mm. long. — F.M. Neg. 19461.

Peru: (Probably). Brazil; Colombia.

Erythroxylum gracilipes Peyr. in Mart. Fl. Bras. 12, pt. 1:

159. 1878; 82.

Gray older branches densely verruculose; branchlets erect-spread-

ing, compressed, 2 mm. broad; stipules persisting, ovate, obtuse, not

setulose, 1-2 mm. long, only 1 or 2 at branchlet base and between

leaves and these very remote; petioles 5-8 mm. long; leaves oblong-

elliptic, acutish at base, acute or shortly acuminate and mucronulate,

11.5-18.5 cm. long, between 4 and 6.5 cm. wide, diaphanous and

more or less membranous, pale green and somewhat lustrous above,

rather glaucescent and dull beneath, longitudinal lines obviously

areolate, midnerve rounded and prominent at base beneath as the

lateral nerves, the secondary rather prominent both sides, closely

reticulate-nervose; flowers 12-16 in axils, pedicels 5-10 mm. long,

very slender; bract ovate, acute, 1.5 mm. long; calyx nearly three-

fourths parted, the acuminate semiovate segments 1.2 mm. long;

lateral auricles denticulate; short-styled flowers with stamen-cup

about as long as calyx, 10-crenate, stamens 3 mm. long, style 1 mm.

long; drupe ovate, acute, 7 mm. long. — The var. exareolatum Schulz,

confused withfi1. Mamacoca, has petioles 4 mm. long, leaves 10-14 cm.

long, 4-4.5 cm. wide, pale green beneath, without lines and not areo-

FLORA OF PERU 641

late, the pedicels only 1 mm. long (perhaps a species?). Three to

ten meter tree, flowers white or yellow, fruit edible (Klug). F.M.

Neg. 12621.

Loreto: Near Iquitos, Klug 1440. Amazonian Brazil; Colombia.

"Sacha mangua."

Erythroxylum hondense HBK. Nov. Gen. & Sp. 5: 176. 1822;

95.

Shrub or tree to 6 meters high with grayish white verruculose

slender branches, the subhorizontal compressed branchlets about

1 mm. thick; petioles 1.5-3 mm. long, little longer than the stipules,

these triangular, not nerved, the obtuse tips 3-setulose, dense at

base of innovations; leaves elliptic or obovate, acutish at base,

rounded or somewhat emarginate at apex, 3-11 cm. long, about

2-5 cm. wide, membranous, pale green and scarcely lustrous above,

dull and glaucescent or reddish beneath, the midnerve there very

prominent, the lateral nerves so above, the reticulate venation not

conspicuous; flowers 1-3 in axils of branchlets; pedicels 1.5-6 mm.

long, the ovate acute bracts about 1 mm. long; petals 3-4.5 mm.

long, about one-third clawed, the ligule to one-half the length of the

blade; lateral auricles three times longer than posterior; stamen-

cup in short-styled flowers equal to calyx, 10-crenulate, stamens 3.5-4

mm. long, style 1.5 mm. long, stigma scarcely broader; in long-styled

flowers stamens 1-1.4 and 1.5-2 mm. long, style 2.5-3 mm. long;

drupes ovate, 8 mm. long, not sulcate. — In the long-styled flowers

the styles are sometimes long-connate as in the case of species

grouped by the monographer in a separate section designated

Microphyllum. Leaves in type elineate, obovate-elliptic, rounded-

retuse and obsoletely mucronulate, 24-30 mm. long, 14-17 mm.

wide; I have not seen recently the Poeppig specimen nor the type

and the species to me is obscure. F.M. Negs. 12622; 35194.

Huanuco: On Cerro de San Cristobal, (Poeppig 1336). To

Colombia. "Mama-coca."

Erythroxylum lucidum HBK. Nov. Gen. & Sp. 5: 179.

1822; 24.

Shrub or small tree sometimes 6 meters high, the erect trunk

5-7 cm. in diameter, the grayish-brown more or less verruculose

branches 4.5 mm. thick, the erect-spreading somewhat compressed

branchlets 2.5 mm. thick; stipules narrowly lanceolate, obtusish

and shortly bisetulose, densely striate, 8-14 mm. long, finally

642 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

deciduous; petioles 5-8 mm. long; leaves oblong-elliptic or obovate,

acute or subrotund at base, acute or shortly acuminate, 7-14 cm.

long, 3-6.5 cm. wide, diaphanous, the older coriaceous, greenish-

brown and rather lustrous above, dull, paler and somewhat brown

or glaucous beneath, densely reticulate-venose, the midnerve little

prominent above, the lateral nerves obscure both sides; flowers

7-11; bracts broadly lanceolate, acute, striate, 2.5-3.5 mm. long;

calyx segments imbricate, broadly ovate, acute, 2.5-4.5 mm.

long, the whitish petals 2.5-4.5 mm. long, the oblong blade about

one-half clawed, the ligule half as long; lateral auricles twice as long

as the posterior; short-styled flowers with stamen cup shorter than

calyx, 10-denticulate, stamens 4-5 mm. long, style 1.5 mm. long,

the depressed stigma a little broader; stamens of long-styled flowers

1.2 and 2.2 mm. long, styles 3.2 mm. long; drupes subglobose or

ovate, 5-7.5 mm. long, 4 mm. thick. — Illustrated (flower), Schulz,

26. F.M. Neg. 12626.

San Martin: Tarapoto, Ule 6389; Williams 5635; 5865; 6668.

Juanjui, King 3767 (det. Standley). San Roque, Williams 7275 —

Loreto: Yurimaguas, Williams 4685; 7827; 3796; Killip & Smith

27540. Middle Maranon, Tessmann 3799. Bolivia to Mexico.

"Murcu tarilla Colorado," "urcu-ynguina" (Williams).

Erythroxylum luculentum Macbr. Candollea 5: 372. 1934.

Shrub, all parts lustrous, the long slender sparsely branched

branches more or less conspicuously lenticellate; stipules persisting,

not nerved, thick-carinate, 2.5 mm. long, apparently shortly setulose;

petioles slender, 5-6 mm. long; leaves oblong-elliptic, mostly about

12 cm. long, 4 cm. wide, acute at base, obtusely acuminate, lustrous

especially above and there olivaceous except for the yellowish mid-

nerve and veins, yellowish green and paler beneath, laxly reticulate

both sides, long lines not evident, chartaceous-membranous; pedicels

5 mm. long; drupes 10-12 mm. long, somewhat curved at tip. —

Probably allied to E. Ruizii Peyr. of Ecuador, which however has

dull leaves; its long and slender petioles possibly distinguish it from

similar Peruvian species as E. Mamacoca but type without flowers

and doubtless should not have been described.

Loreto: La Victoria, Williams 2788, type.

Erythroxylum macrocnemium Mart. Beitr. Kenntn. Ery-

throx. 1840; Abh. Math. Nat. Cl. Akad. Muench. 3, pt. 2: 402.

pi. 1. 1843; 23.

FLORA OF PERU 643

Branches, branchlets and petioles stout, the latter only a few

mm. long, wing-margined above by the decurrent cuneately based

obovate leaves, these 2.5 to 4 dm. long or longer, 8-14 cm. wide,

conspicuously reticulate-nervose beneath; stipules caducous, 3-5 cm.

long; bracts 3-7 mm. long, long-acuminate; calyx segments lanceo-

late, 6-6.5 mm. long; petals 5.5 mm. long the lateral auricles three

times longer than posterior; stamen cup crenate; long styles 4.5 mm.

long, stigma broader than styles. — One to ten meter tree with white

or greenish flowers, rarely yellow (Klug); leaves at summit, flowers

on stem below them (Killip & Smith).

San Martin: Pongo de Cainarachi, Klug 2738 (det. Standley,

E. amplum).—3umn: Puerto Yessup, Killip & Smith 26375 (det.

Standley, E. amplum). — Huanuco: In woods, Cuchero, Poeppig 46.—

Loreto: Yurimaguas, at edge of woods, Poeppig 1760; 2166; Williams

4039; '4149; 4356; Killip & Smith 29081 (det. Killip). On the

Maranon, Tessmann 5053. Santa Ana on the upper Rio Nanay,

Williams 1230. Rio Putumayo, Peru-Colombia boundary, Klug

1616 (det. Standley, E. amplum}. Iquitos, Tessmann 5351; 5053.

Mouth of Rio Santiago, Tessmann 4244- "Maggisapa" (Klug).

Erythroxylum macrophyllum Cav. Diss. 8: 401. pi. 227.

1789; 25.

With the stout branches and branchlets of E. macrocnemium but

the broad calyx segments of the closely related E. lucidum (apparently

only a variety) from which it may differ otherwise as follows: petioles

7-12 mm. long; stipules 7-21 mm. long, minutely 3-setulose; leaves

typically obtusish or shortly acute, 12-23 cm. long, 5-12 cm. wide,

lateral nerves more obvious beneath; flowers 12-18 (25), calyx

segments (2) 4-4.5 mm. long, acuminate, petals 4.5 mm. long, lateral

auricles thrice longer than posterior, stamen cup and ovary equal

in short-styled flowers.— Illustrated, Schulz, 26. F.M. Neg. 29348.

San Martin: Tarapoto, Williams 6591; 6585. — Huanuco: Tingo

Maria, 700 meters, Stork & Horton 9476 (det. Standley).— Loreto:

Pampa del Sacramento, Castelnau. Pamayacu, Klug 3185 (det.

Standley).— Rio Acre: Seringal Auristella, Ule 9473 (det. Dahlem).—

Cuzco: Cosnipata, Weberbauer 6962 (apparently toward E. lucidum).

To the Guianas. "Yutabanco."

Erythroxylum Mamacoca Mart. Beitr. Kenntn. Erythrox.

1840; Abh. Math. Nat. Cl. Akad. Muench. 3, pt. 2: 365. 1843; 79.

Shrub with slender branches that are obscurely verruculose and

erect-spreading branchlets that are little compressed, 1.5 mm. wide;

644 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

stipules persistent, triangular, 1.5-2 mm. long, broadly bicarinate,

estriate, shortly trisetulose at the rounded tip, crowded at base of

innovations and between leaves; petioles (3) 3.5-6 mm. long; leaves

oblong-elliptic, sometimes broadly so, acutish or subrotund at base,

rather obtusely acuminate, 6-10 cm. long, 3-4.5 cm. wide, charta-

ceous, diaphanous, grayish-green above, brown beneath, somewhat

lustrous both sides, the midnerve thick and obtuse beneath, the

lateral nerves not conspicuous above but prominent and reticulate

beneath; flowers 1-3 on pedicels enlarged and 5-6 mm. long in fruit;

calyx lobes 1.2 mm. long; stamen cup little longer than calyx, 10-

crenate; drupe 14 mm. long, 5 mm. broad, 4 mm. thick, ellipsoid,

obtusish. — Leaves darken in drying. The name said to be for the

wife of the king Mayta-Capac. F.M. Neg. 12628.

Huanuco: In mountains near Pillao, Ruiz. Hacienda San An-

tonio, Raimondi. Cuchero, in stony woods, Poeppig 1760. — Loreto:

Yurimaguas, Williams 4639; 4664- Near Iquitos, Williams 8049.

"Mama-cuca," "motelo-caspi" (Williams).

Erythroxylum paraense Peyr. in Mart. Fl. Bras. 12, pt. 1:

164. pi. 30. 1878; 35.

Shrub or small tree with slender terete branches, the bark gray

lenticellate, obscurely verruculose; branchlets erect-spreading, com-

pressed, to 2 mm. broad; stipules finally deciduous, lanceolate, 2.5-5

mm. long, obtuse, 3-setulose, striate, membranous, loose at base of

innovations; petioles 5-6 mm. long; leaves oblong-elliptic, narrowed

or subrounded at base, plicate and mucronulate at the acuminate

apex, 6 (5)-12 cm. long, about 2-4 cm. wide, long lines subareolate,

chartaceous, diaphanous (ex char.), lustrous and gray green above,

rather opaque and rusty beneath, the midnerve and subhorizontal

lateral nerves little prominent both sides, obscurely reticulate;

flowers 3-14; pedicels 2-4.5 mm. long, not enlarged; calyx lobes

1-1.2 mm. long, acute; petals 3 mm. long, ligule half as long, claw

one-third of blade; lateral auricles little longer than posterior;

stamen cup in short-styled flowers about as long as calyx, entire or

10-crenulate, stamens 3 mm. long, ovary ovate, style 1 mm. long,

the stigma a little broader. — E. amazonicum Peyr., 34, has leaves

6-12 cm. long, 3-6 cm. wide, obovate, coriaceous, not pellucid,

pedicels strongly enlarged, drupes 12-16 mm. long; it may be as

near E. amplum. All of the Peruvian specimens (at least) perhaps

should be included in a somewhat more broadly defined E. acumi-

natum.

FLORA OF PERU 645

San Martin: Tarapoto, Ule 6437; Williams 6319; 6325; 6614; 6672;

6673. Near Moyobamba, 900 meters, Weberbauer 4524; Klug 3258

(det. Standley). Amazonian Brazil; Bolivia. "Puca-llaja."

Erythroxylum Raimondii 0. E. Schulz, Notizbl. Bot. Gart.

Berlin 10: 727. 1929.

Branchlets compressed toward tips, 2.5 mm. in diameter, densely

leafy, the blackish bark verruculose-lenticellate; stipules persisting,

3 mm. long, striate, subfimbriate, shortly bisetulose; petioles stout,

2-3 mm. long; leaves elliptic, rounded at base, lightly emarginate

at the mucronulate tip, at maturity 6-9 cm. long, 3-4.5 (5) cm.

wide, subcoriaceous, scarcely revolute, brownish above, subglauces-

cent beneath, the approximate nerves reddish and conspicuously

reticulate; pedicels slender, to 3 mm. long; calyx segments distinct,

narrowly ovate, acutish, 1 mm. long; petals 2.5 mm. long, the

ligule half as long; staminal cup shorter than calyx; fruit unknown.—

Allied to the more northern E. rufum Cav. but the petioles shorter,'

the leaves rounded at base. E. opacum Rusby and E. venosum

Rusby, both Mem. N. Y. Bot. Gard. 7: 270. 1927 of Bolivia, could

not be this plant; the first, ex char., has leaves acute both ends

(stipules estriate) and in the latter they are smaller (Schulz) but

smaller leaves may be no character; flowers from Cook & Gilbert

not dissected. It seems to be nearer E. passerinum Mart, than

E. rufum and, like certain other species, simulates some with estriate

stipules. F.M. Neg. 12641.

Cuzco: Santa Ana, Prov. Convention, Raimondi, type; also

Cook & Gilbert 1463. "Coca-coca," "coco-coco."

Erythroxylum Shatona Macbr. Candollea 5: 371. 1934.

Glabrous, the stout branchlets densely verruculose and lenticel-

late; stipules persisting, not at all striate, obscurely if at all setulose,

4 mm. long; petioles 3-4 mm. long; leaves ovate-elliptic, shortly

acute, 7-10 cm. long, 3-5 cm. wide, coriaceous, slightly lustrous

both sides, glaucous green above, the veins obscure, slender midnerve

impressed, beautifully areolate, long lines conspicuous, brown

beneath in herb.; flowers few; pedicels about 3 mm. long; calyx

segments narrowly ovate, 2 mm. long; styles apparently connate;

drupes oblong-ellipsoid, subangulate, acutish, 12 mm. long, 3.5 mm.

thick. — Perhaps related to E. carthagenense Jacq., 87, of Colombia

but distinguishable, apparently, by its shortly pointed leaves.

Among Peruvian species it most resembles E. gracilipes with thin

646 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

and acuminate leaves. The somewhat related E. Ruizii Peyr., 90,

of Ecuador, has smaller submembranous leaves very lustrous beneath,

shorter stipules, longer pedicels.

San Martin: Tarapoto, Rio Maya, Williams 6212, type. "Shatona

Colorado."

Erythroxylum Tessmannii O. E. Schulz, Notizbl. Bot. Gart.

Berlin 9: 263. 1925.

Much-branched shrub the brownish branches long-lenticellate,

the erect-spreading branchlets 1-2 mm. across; stipules 1.2 mm.

long, estriate, persisting, shortly 3-setulose; petioles 2-3 mm. long;

leaves oblong-elliptic-lanceolate, narrowed both ends, less so toward

the obtuse tip, 4-7 cm. long, 10-18 mm. wide, chartaceous, di-

aphanous^), gray-green and lustrous above, opaque and glaucous

beneath, the slender midnerve conspicuous both sides, the lateral

nerves loosely reticulate; flowers 1-3, in axils of leaves and branch-

lets; pedicels 2-4 mm. long; bracts ovate, acute, 1-1.5 mm. long;

calyx segments ovate, acute, 0.75 mm. long; petals 3 mm. long,

ligule half as long, white with green tips; stamen cup as long as

sepals in long-styled flowers, the stamens 1 and 1.8 mm. long, styles

3 mm. long. — Near E. exaltatum Bong. var. ellipticum (Peyr.) 0. E.

Schulz, 104, of southern Brazil, but leaves narrower, tip not at all

acuminate, opaque beneath, stipules, pedicels and flowers smaller

(Schulz) ; the narrow elongate leaves obtusely acuminate distinguish

it from E. Coca. It is possibly a variant of Bongard's species but the

firmer leaves are obviously opaque beneath and, while narrowed

apically, are obtuse at the tip; doubtfully distinct too fromE". lenti-

cellosum Huber, Bol. Mus. Goeldi 5: 420. 1909, this with rather

similar foliage but with longer subciliate stipules, pedicels (in type)

5-6 mm. long. F.M. Neg. 12551.

Loreto: Rocky shores, middle Aguaytia on the Ucayali, Tessmann

3163, type.

Erythroxylum Ulei 0. E. Schulz, Pflanzenreich IV. 134: 62.

1907.

A 1-5 meter shrub or tree, the branches with brown, evidently

verruculose bark and about 3 mm. thick, the erect-spreading little

compressed branchlets about 1.5 mm. broad; stipules persisting,

triangular, obtuse and very shortly 3-setulose, obscurely and sparsely

striate, 1.5-2 mm. long, lax at base of innovations and between the

FLORA OF PERU 647

leaves, these with petioles 2-3 mm. long, the blades narrowly ovate

(apparently usually broadly so) or elliptic, somewhat acute, the

obtusish tip obscurely mucronulate, the base slightly acute, about

3.5-8 cm. long, 1.5-4 cm. wide, chartaceous, diaphanous, gray-green

and somewhat lustrous above, more or less rusty and opaque beneath,

the lateral nerves obvious both sides, densely reticulate-nervose;

flowers 1-3, the little enlarged pedicels 1-4 mm. long; bracts setulose,

ovate, 1.2 mm. long; calyx segments 1.5 mm. long, broadly lanceo-

late; petals 3.5 mm. long, the ligule half as long, the lateral auricles

twice longer than the posterior; stamen cup (short-styled flowers)

shorter than calyx, entire, stamens 3.2 mm. long, style 1 mm. long,

the stigma as wide; drupes 7.5 mm. long, 3.5 mm. broad, ovate,

acute, subcurved. — The recent collections cited are somewhat

aberrant, the leaves soon subcoriaceous, mostly broadly elliptic,

rounded or scarcely at all acute. F.M. Neg. 12653.

San Martin: Tarapoto, Ule 6346; Williams 644.4. Tocache, in

dry mountain woods, Poeppig. Without data, (Mathews 680;

2022}. — Junin: La Merced, 5408. Colonia Perene", Killip & Smith

25073. Between the Huallaga and Pizana, Raimondi. — Cuzco:

Santa Ana, Cook & Gilbert 1698. Bolivia; Colombia. "Monte cuca."

ZYGOPHYLLACEAE R. Br.

. Shrubs, half-shrubs, rarely trees or herbs, with nearly the floral

and fruit-characters of the closely related Rutaceae but the opposite

or alternate leaves with persisting often spinescent or noduliform

stipules and without pellucid punctae. Stamens usually 2, some-

times 3 times as many as the petals. Ovary commonly 4-5-celled,

rarely 2-12. Fruit various, ordinarily capsular and composed of

few to many parts, conveniently at least termed follicles, carpels or

nutlets. — Descole, O'Donell and Lourteig, Lilloa 5: 258. 1940,

have distinguished the tribes of the family by the nature of the

pollen grains; I am indebted to them for this character, and I have

compiled generally from their careful synopsis of the species of

Argentina. Their illustrations, as cited, are repeated in Descole,

Gen. & Sp. Argent. 1. 1943.

"Lignum vitae," popularly known as one of the hardest woods,

is an American species of Guaiacum; it is not found in Peru but it

has a counterpart it is said in the Wood of Porlieria which I found

greatly prized about Huanuco for the manufacture of small articles,

such as combs and spoons.

648 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Carpels spiny or beaked; endosperm absent; pollen superficially

reticulate with a pore in each cavity; herbs or little ligneous.

Fruits radiate, 5-parted, spiny or winged; ovary developing trans-

verse septa 1. Tribulus.

Fruit convex, (5) 8-12-parted, espinose; ovary without septa.

2. Kallstroemia.

Carpels not beaked or spinose; endosperm present; pollen smooth

or granulate with 3 pleats and 3 pores; often ligneous or partly.

Leaves digitately 3-5-foliate; stamens unappendaged; half shrubs.

3. Fagonia.

Leaves pinnate (leaflets rarely reduced to 2); stamens appendaged;

shrubs.

Leaves pinnate; carpels alate or glabrous.

Petals yellow; carpels compressed, alate 4. Bulnesia.

Petals blue; carpels 3, globose, glabrous 5. Porlieria.

Leaves bifoliate or nearly; fruits 5 villous nutlets. . .6. Larrea.

1. TRIBULUS [Tourn.] L.

Mostly prostrate herbs with pinnate, usually opposite, leaves,

membranous stipules and solitary flowers on axillary peduncles.

Sepals and petals 5; stamens twice as many, the filaments unappend-

aged, the 5 episepalous sometimes sterile. Ovary 5-celled (cells

3-5-ovuled), terminating in a columnar stigmatic body, the flat

fruit composed of 5 carpels or nutlets each of them usually spinescent

or winged leaving no central axis when falling. — The name is thought

to have been suggested by the fancied resemblance of the carpels

to an ancient 3-pointed tool.

Tribulus terrestris L. Sp. PI. 387. 1753.

More or less sericeous pubescent annual, the leaves with 6

(5-7) pairs of oblong mucronate leaflets 7-13 mm. long, 4-7 mm.

wide, and typically longer than the peduncles; sepals caducous;

petals yellow or lemon-yellow, 4-5 (7) mm. long; carpels ordinarily

with 2 strong spines on each one, sometimes with 3 or 4, some

smaller ones, also tubercled and somewhat pubescent. — According

to Svenson the Peruvian plant is var. sericeus Andersson and varies

from densely hirsute to sparsely pilose. To be expected sooner or

later as another introduction is T. cistoides L. perhaps a variant

but said to be more erect, perennial, peduncles half as long to longer

FLORA OF PERU 649

than the leaves, these somewhat larger, and petals much larger,

at least 1.5 cm. long; Weberbauer 5943 was so determined by Krause,

apparently intermediate in character. The fruiting spines are so

hard that they may puncture bicycle tires. Only a few recent

authors have sacrificed tradition for erudition (their own, at any

rate) by "correcting" Linnaeus' name to "terrester" contrary to

euphony. Introduced from southern Europe. Illustrated, Lilloa

5: 261; Amer. Jour. Bot. 33: 456.

Piura: La Brea, (Haught & Svenson 11568}. Near Piura, Weber-

bauer 5943. — Cajamarca: Cascas, Raimondi. — Lima: Street in

Chosica, Mexia 04086. — Huanuco: Near Huanuco, 2059; 2337.—

Apurimac: North of Abancay, Goodspeed Exped. 10540. — Moquehua:

At 2,000 meters, Weberbauer 7437. Widely spread from southern

Europe.

2. KALLSTROEMIA Scop.

Similar to Tribulus but the fruit consisting of 8-12, rarely 5,

merely tuberculate usually 1-seeded follicles that separate at

maturity from the more or less persisting styliferous central axis.

Ovary without transverse septa. Stigma capitate. — The origin of

the name is personal, Kallstroem.

Kallstroemia adscendens Andersson, K. Sw. Freg. Eugen.

Resa. Bot. 245. 1854. K. tucumanensis Descole, O'Donell & Lourteig,

Lilloa 4: 218. pi. p. 219. 1939, fide Svenson, Amer. Jour. Bot. 33:

457. 1946.

A prostrate or diffuse annual the branching stems more or less

finely pubescent and somewhat hirsute; leaves with 3 (2) pairs

(rarely 4?) of obliquely elliptic obtusely rounded slightly mucronate

leaflets, appressed pubescent beneath, mostly 1-2 cm. long, 5-9 (13)

mm. wide; peduncles 1-2 cm. long; sepals lanceolate, hirsute; petals

white (or pale yellow?), obovate, 5 mm. long; fruit appressed

strigose, 7-8 mm. long, the conic broad beak about as long as the

body, the follicles tubercled dorsally, usually reticulate laterally.—

This disposition of the Haught specimens follows Svenson, I.e.; in

Field Mus. Bot. 8: 119. 1930 I suggested that more collections are

needed to establish the significance of the characters now regarded

in the genus as specific and thought that the Peruvian plant might

be a form of K. caribaea Rydb. N. Amer. Fl. 25: 111. 1910 of the

West Indies and Colombia, similar but with narrower sepals, petals

6-7 mm. long; revision of the genus is needed. Illustrated, Lilloa

5: pi. 2, opposite page 266.

650 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

Piura: Near Piura, Haught 161; 51; (147). Galapagos; Ecuador;

Argentina(?).

Kallstroemia maxima (L.) T. & G. Fl. N. Amer. 1: 213. 1838.

Tribulus maximus L. Sp. PI. 386. 1753. K. tribuloides (Mart.)

Wight & Arnott, Prodr. Fl. Ind. Or. 145. 1834, by inference. Ehren-

bergia tribuloides Mart. Nov. Gen. & Sp. 2: 73. pi. 163. 1826?

Similar in general as defined here (sens, lat.) to K. adscendens

but leaves with mostly 4 (3-6) pairs of leaflets, sepals narrow, petals

yellow, 7-8 mm. long, carpels glabrous but tubercled, or, including

the above, sepals narrowly ovate, petals orange, 9-11 mm. long,

carpels smooth and glabrous; character in general, stems more or

less sericeous and pubescent, leaflets mostly 1.5-2 cm. long, 5-13

mm. wide, sepals acuminate, hirsute-pilose, petals about as wide

at tip as long. — The Peruvian material differs from both the above

variants in having strigillose carpels, thus approaching K. boliviano,

Standl. Field Mus. Bot. 11: 161. 1936 but that as to type with petals

12-18 mm. long, fruit 8-10 mm. long, and also K. brachystylis Vail,

of southwestern United States; at least some specimens from Argen-

tina referred here have strigillose carpels; cf. too K. adscendens.

Until there is a generic revision, probably of necessity based upon

or made with the aid of genetic studies, it seems just as well to treat

the Peruvian plants under one perhaps aggregate name, neverthe-

less understandable, with this explanation. It may be noted that

Wight & Arnott, I.e. wrote: "Kallstroemia Scop, (containing Trib.

maximus Lin. and Ehrenbergia tribuloides Mart.) differs," etc.

Weberbauer found the plant used for stock at Mantaro as

"jepo."

Cajamarca: Near Jae"n, Raimondi. — Lima: East of Sayan, Prov.

Chancay, Goodspeed 33036 (det. Leonard). On the railroad at 1,300

meters, Weberbauer 5265. Matucana, 396. — Huanuco: Near Hua-

nuco, 2355. Ambo, flowers orange, Stork 11444 (det. Johnst.).—

Apurimac: Carpeting sandy places, flowers yellow, Goodspeed Exped.

10527. — Huancavelica: Mantaro, 1,300 meters, Weberbauer 6516

(det. Mansf., K. tribuloides}. — Cuzco: Sandy rocky slopes, Chuyani,

Vargas 9725 (det. Standl.). Valle de Apurimac, Hacienda Huantaro,

Bues 758 (det. Herrera, K. tribuloides). Widely distributed. "Jepo."

3. FAGONIA [Tourn.] L.

Reference: Johnston, Proc. Calif. Acad. Sci. ser. 4. 12: 1049-

1053. 1924.

FLORA OF PERU 651

Much branched, more or less woody at base, sometimes shrubby,

with opposite branches and opposite simple or digitately 3-foliate

leaves, often spinescent stipules and solitary usually roseate or

purplish flowers. Sepals and petals 5, caducous. Stamens 10,

filaments unappendaged. Style subulate, stigma simple. Ovary

5-celled, with 2 ovules in each cell near the base. Fruit composed

of 5 1-seeded carpels that finally separate along the inner edge. —

Named for G. C. Fagon, French botanist of the seventeenth century.

Fagonia chilensis Hook. & Arn. in Hook. Bot. Misc. 3: 165.

1833; 1049. F. aspera Gay, Hist. Nat. Chile 1: 470. 1845.

Typically glabrous except for some scabrosity on the fruit;

branches diffuse, repeatedly dichotomous, angled; stipules spinescent,

stout, 2-4 mm. long, 2-3 times shorter than the petioles; leaflets 3,

obovate to linear-lanceolate, acute or acuminate, spine-tipped, 2-8

mm. long, 2-3 mm. wide, the lateral often oblique; pedicels 3-4 mm.

long, deflexed, shorter than the fruits; sepals oblong-lanceolate, 3-4

mm. long, acute; petals pink, 6-8 mm. long; fruit about 5 mm. high,

a little scabrous, beak 2-3 mm. long. (After Standley.) — The variant

aspera (cf. Johnston, I.e. 1051) is typically more or less scabrous;

the Peruvian plant is lightly pilose-pubescent, the loose trichomes

rather dense, especially on the fruits; leaflets 2-4 mm. wide and may

be worth designating var. pubecarpa Macbr., var. nov., carpellis

dense pilosciusculis, circa 4 mm. latis; pedicellis ad 7 mm. longis;

Weberbauer 7431, type. The Chilean species was treated at one

time by Engler as a variety of the Mediterranean F. cretica L.

which has fruits about 7 mm. wide and has recently been referred

to by Johnston as "a species doubtfully separable," but this author

earlier, I.e., followed Standley, Proc. Biol. Soc. Wash. 24: 244. 1911,

who pointed out that all the American plants have noticeably smaller

fruits. The American plants furthermore are extremely variable

vegetatively but nevertheless relatively constant in size of the fruit

and probably constitute one species, at least within Peru.

Lima: Near Obrajillo, Wilkes Exped. Near San Bartolome*,

Weberbauer 5268. Hillside above Santa Eulalia, 1,500 meters,

Goodspeed 33093. — Moquehua: Near Moquehua, Weberbauer 7431

(type, var. pubecarpa}. Chile; California?

4. BULNESIAGay

Trees or shrubby, the numerous upper branchlets sometimes (in

Peru) leafless or nearly except at time of flowering and from this

652 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

character as well as from habit of growth often broomlike in appear-

ance. Sepals and petals 5, the former deciduous, the latter longer,

clawed. Stamens 10, with long dentate or lacerate appendages.

Capsule 3-5-celled, the fruit generally consisting of 5 broad winged

1-seeded carpels each of which dehisces by a ventral suture.

The flowers are suggestive of Cassia. Genus named for Bulnes,

a Chilean president (1841), soldier and statesman.

Bulnesia Retama (Gillies) Griseb. PI. Lorentz. 58. 1874. Zygo-

phyllum Retama Gillies, ex Hook. Bot. Misc. 3: 166. 1833.

Typically a tree or tree-like shrub with early pubescent terete

brittle branchlets from which the small leaves, borne at the flower

buds, are strongly caducous; internodes 3-10 mm. long; leaflets 2-4

pairs, oblong, mucronulate, appressed pubescent, rare or lacking

after floration, to 5 mm. long, 3 mm. wide; stipules ovate, acute,

2 mm. long; flowers about 2, dichasial at each node, peduncles 6-8

mm. long, pubescent; sepals glabrate, 5-7 mm. long, 3-6 mm. wide;

petals obovate, 7-8 mm. long, 4-5 mm. wide; stamen appendages

deeply ciliate-lacerate; ovary glabrous; fruit 5-winged, 3 cm. long,

2.5 cm. wide. — The Peruvian plant is var. Weberbaueri Engler, a

shrub to 3 meters high with long internodes, 2-3 pairs of thick oval

leaflets 2 mm. wide, 3 mm. long, found at the edge of the Prosopis

juliflora formation. — Illustrated, Lilloa 4: 287 (drawing; also pis. 8,

9, photos of type spec, and of a tree).

The wood is hard and useful; a tea prepared from the flowers

of the variety is used locally in the treatment of gonorrhoea (Weber-

bauer).

lea: Toward Hacienda Ocucaje, 360 meters, Weberbauer 7193;

7196a. Between Otoca and Ingenio, 1,050 meters, Weberbauer.

Also, Raimondi. Argentina. "Calato" (Weberbauer), "retamo."

5. PORLIERIA R. & P.

Rigid shrubs with stout branches on which the opposite pinnate

leaves of many leaflets are borne so as to spread open toward the

light, closing in cloudy weather. Stipules small, spinescent. Flowers

1-3, short-peduncled in the leaf-axils. Sepals and petals 4 (5),

imbricate in bud. Stamens 8-10, the filaments enclosed in a short

or long divided or serrate sheath or appendage. Ovary 2-5-celled

with 2-5-sulcate style, simple stigma, the ovules 4 in superimposed

pairs in each cell. Fruit coriaceous, 1-seeded. — Genus commemo-

rates Don Antonio Porlier, a government nobleman (marquis) who

FLORA OF PERU 653

aided the authors in their work. I retain the original spelling,

R. & P. Prodr. 55. pi. 9. 1794.

The wood is said to be like true lignum vitae (Guaiacum sp.).

Porlieria hygrometra R. & P. Syst. Veg. 94. 1798.

Stocky intricately branched shrub with many short branchlets;

leaves with 6-8 pairs of oblong leaflets 5-8 mm. long, 2 (1.8)-3 mm.

wide, more or less loosely pubescent, especially toward the margins,

or ciliate, glabrate; peduncles about 2 mm. long (apparently some-

times to 10 mm. long in fruit), like the flower-buds densely fulvous

pubescent; sepals 3 mm. long, the petals a little longer; filaments

margined toward base; drupes, according to authors, (3) 4, oblong,

gibbous, connate. — In Chilean material they may be 8 mm. long.

Type material from Huanuco and Chile; it is possible that there is

only one variable species, but see P. chilensis Johnst. Journ. Arnold

Arb. 19: 253. 1938, leaflets 5-7 pairs, ovary sparsely villous. The

broad leaflets and large angled fruits of Chilean material are con-

trasts to the mature globose dull-black small (4-5 mm. thick) fruits

of specimens from central Peru and the similar in size but brown and

lustrous fruits of P. microphylla, see below. It is noteworthy that

undeveloped fruits of the last are distinctly angled, and that possibly,

as regards this feature, there is considerable change when the fruit

matures. Other names proposed are: P. arida Rusby, Mem. Torrey

Club 6: 15. 1896 and P. Steinbachii Standl. Field Mus. Bot. 11: 162.

1936, this with ciliate rather broad leaflets (15 pairs), glabrous ovary.

Illustrated, R. & P. Fl. Peruv. 4. pi. 343.

Piura: Huancabamba, Weberbauer 6286. — Cajamarca: Cascas,

Raimondi. — Ancash: Raimondi. — Ayacucho: Prov. Huanta, Rai-

mondi.— Huanuco: Pampayacu, Sawada P53. Hacienda San Roque,

1,800 meters, Soukup 3107. Near Huanuco, 2382; 3221; Dombey;

Ruiz & Pavdn, type. Chulki, Sawada Pi 27. Argentina? Bolivia?

Chile? "Murucho."

Porlieria microphylla (Baillon) Desc., O'Don. & Lourt. Lilloa

5: 329. 1940. Guaiacum microphylla Baillon, Adansonia 10: 315.

1872. P. Lorentzii Engler, Pflanzenfam. 3, 4: 84. fig. 51. 1897.

Drupes nearly globose, about 4.5 mm. thick, lustrous, light

brown; leaflets 5-20 pairs, 4-7 mm. long, 1-1.3 mm. wide, obtuse,

appressed pubescent beneath; flowers 1 or 2, pubescent peduncles

about 1 mm. long (to 4 mm. in fruit); sepals ovate, 2.5-3 mm. long,

filaments winged nearly to the sagittate anthers. — After Descole,

654 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

O'Donell & Lourteig; Johnston has suggested that the species is

the same as that of Ruiz & Pavon and at least for Peru that is

probably the case as my note made in Dahlem for Weberbauer 374.1

reads "rather P. hygrometraT' F.M. Neg. 12657. Illustrated,

Lilloa 5: 331.

Ancash: High steppes with cacti, shrubs, annual grasses, 2,500

meters, Puccha Valley, Prov. Huari, Weberbauer 3741 (det. Harms,

P. Lorentzii); 174. Argentina; Bolivia. "Cucharero," "chukupi"

(Argentina)..

6. LARREACav.

Resiniferous evergreen shrubs with persisting stipules and op-

posite sometimes pinnate leaves but these in Peru with only 2

slightly connate spreading leaflets. Flowers yellow, solitary on short

terminal (pseudoaxillary) peduncles with 5 deciduous imbricate

sepals and petals and 10 stamens, the filiform filaments surrounded

by variously dentate or cleft basal scales. Ovary shortly stiped,

globose, hirsute-pilose, 5-celled, attenuate into a subulate some-

what 5-parted style with minute stigmas; ovules pendulous. Fruit

villous, consisting of 5 indehiscent 1-seeded nutlets. — The conserved

name honored Juan Antonio Hernandez de Larrea, Dean de Zaragoza.

Larrea divaricata Cav. Anal. Hist. Nat. Madrid 2: 122. pi. 19.

1800. Covillea divaricata (Cav.) Vail, Bull. Torrey Club 22: 229.

1895. Schroeterella divaricata (Cav.) Briquet, Verb'ff. Geobot. Inst.

Riibel. 3: 663. 1925. Neoschroetera divaricata (Cav.) Briquet,

Candollea 2: 514. 1926.

Younger angulate branchlets and leaves more or less appressed

ashy pubescent; stipules noduliform, not at all foliose; leaflets

diverging from the slightly connate base with a short mucro between

them, striately nerved, mucronulate, oblongish, nearly straight, 7-8

(15) mm. long, 2.5-3 (4.5) mm. wide; sepals pubescent, concave,

obtuse, obviously unequal, 4-6 mm. long, 3-4 mm. wide; petals

4-8 mm. long or larger; filament scales variously dentate, 6 or 7 mm.

long; fruits globose, villous-hirsute, easily separating into 5 nutlets. —

This is the typical shrub of Argentina to which the Peruvian material

was referred in Dahlem, probably correctly. A tea is prepared from

the plant for pulmonary disorders (Weberbauer). Illustrated,

Lilloa 5: 309.

Arequipa: Valle de Mages, Aplao, 1,000 meters, Weberbauer

6845; Raimondi. To Argentina. "Yarilla," "chamanilla," "jarilla."

FLORA OF PERU 655

RUTAGEAE Endl. Citrus Family

Reference: Engler in Mart. Fl. Bras. 12, pt. 2: 77-196. 1874.

Glandular punctate usually pungently odorous shrubs trees or

rarely herbs with estipulate often opposite leaves that may be

simple but are ordinarily 1-7-foliate or pinnate, sometimes bi- or

tripinnate, the leaflets commonly but not always entire, the petioles

now and then glandular or modified at base, the flowers mostly

cymose and axillary, frequently perfect but not always regular

especially if the petals are more or less connate. Anthers versatile

or adnate the connective often glandular, sometimes extended or

appendaged, the filaments rarely squamate. Disk variously de-

veloped, infrequently wanting. Carpels connate in greater or less

degree, even free, or solitary, the baccate or rarely drupaceous fruit

4-5-celled, the cells or carpels usually 1-seeded and ordinarily

elastically bivalved the endocarp often adhering to the seed and

falling with it. Albumen none or fleshy. — The berry or orange type

fruits are a special development within the scope of this highly

diversified family.

A complete account of citrus fruits is available in The Citrus

Industry; History, Botany, and Breeding; editors, Webber &

Batchelor (taxonomy, W. T. Swingle) 1: 1943; also Citrus Products,

McNair, Field Mus. Bot. 6: 1926-1927.

Two of the largest Amazonian trees belong to this family but

have not yet been found within Peru: Hortia Vandelli has an egg-

shaped stone fruit which presumably allies it to Amyris but it has

long narrow anthers, simple leaves, 5-merous flowers with pubescent

petals, 5-lobed disk, the wood, according to Ducke similar to that

of "Poa amarello" which compare below under Adiscanthus; native

name "Cachaceiro"; Sohnreyia Krause is an equally distinctive tree

but by reason of its ample leaves with many narrow leaflets and its

winged fruits.

Leaves simple or simply parted, sometimes unifoliate; carpels,

except Citrus, 1-2-seeded.

Stamens many; fruit orange-like 1. Citrus.

Stamens 3-10; fruit usually dry, rarely a berry or drupe.

Anthers often as broad as long, ovate, broadly elliptic, sometimes

cordate; petals and stamens free.

Stamens 6-10; fruit a berry or drupe; leaves (Peru) pinnate;

flowers usually perfect.

656 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Fruit a berry; flowers large 2. Murraya.

Fruit a drupe; flowers small 3. Amyris.

Stamens 3-5 (-8) or wanting in female flowers; fruit dry.

Disk short or obscure; stamens or styles wanting or reduced;

carpels glandular-punctate; leaves usually pinnate.

4. Zanthoxylum.

Disk annulate or cupulate; flowers perfect; carpels rugose

or aculeate, rarely smooth; leaves simple or 1-7-

foliate.

Flowers in panicles, often racemiform; ovary usually

tubercled; fruit a ligneous capsule of 4-5 often

radiate parts.

Petioles opposite or in 3's, dilated and often appendaged

at base 5. Metrodorea.

Petioles usually alternate, unappendaged.

6. Esenbeckia.

Flowers spicate or racemose; ovary smooth; fruit 1-5,

often basally united carpels 7. Pilocarpus.

Anthers often longer than broad, linear-lanceolate or narrow;

petals usually united at least at base; stamens rarely all

entirely free.

Calyx lobes or sepals equal or uniseriate.

Petals and stamens both free; disk lacking. 8. Adiscanthus.

Petals at least basally or medially connate; disk present.

Flowers 4-merous; stamens free 9. Leptothyrsa.

Flowers 5-merous; stamens at least lightly affixed in part.

Flowers regular, with long tube; stamens nearly all

completely free, rarely all fertile 10. Ticorea.

Flowers somewhat irregular; stamens never all com-

pletely free, rarely all fertile.

Calyx colored, large; stamens nearly completely

attached; fruit-parts 2-seeded.

11. Erythrochiton.

Calyx small, green, stamens more or less free; fruit-

parts 1-seeded, tardily separating except Cus-

paria.

Stamens somewhat coalescent and joined to long

corolla- tube, the anthers appendaged.

12. Galipea.

FLORA OF PERU 657

Stamens unappendaged; petals nearly free or

shortly united.

Petals only basally connate; filaments lightly

affixed; carpels tardily free. . . .13. Rauia.

Petals connate into short tube; filaments affixed

medially; carpels free from first, or reduced

to 1 14. Cusparia.

Calyx lobes very unequal or foliate.

Tubular corolla concealed in calyx; herb or half-shrub.

15. Monnieria.

Funnelform corolla exserted; shrubs or trees. . .16. Ravenia.

Leaves bi- or tripinnate; carpels each with 3 or more seeds.

Glaucous herb or half-shrub 17. Ruta.

Ashy pubescent shrub 18. Dictyoloma.

1. CITRUS L.

Reference: Swingle, The Citrus Industry 1: 129-474. 1943.

Small trees, the younger branchlets angled but soon terete with

single spines in the leaf-axils, these often lacking later. Leaves

1-foliate; lateral veins few and without conspicuous veins beneath,

the petioles usually more or less winged and articulated with the

blade, except in C. medica. Flowers solitary or in short axillary

corymbose racemes, perfect or staminate. Calyx cupulate, 4-5-lobed,

petals 5 (4-8), thick, linear, gland-dotted, imbricate in bud. Stamens

usually about four times as many as the petals, sometimes many

more. Disk annulate. Ovary cells 10-14 (8-18) with 4-8 or more

ovules in each cell in 2 collateral rows. Style cylindric, abruptly

expanded into the globose or oblate-sphaeroid stigma (after W. T.

Swingle, The Citrus Industry 1: 386. 1943). His excellent descrip-

tion of the fruit is as follows: fruit a hesperidium with the segments

containing seeds near the inner angle and the rest of the space filled

with stalked fusiform pulp-vesicles, filled with a very watery large-

celled tissue; around the segments is a white endocarp outside of

which is the peel, dotted with numerous oil glands and turning

yellow or orange at full maturity; seeds obovoid or flattened obovoid,

more or less angular, containing 1 or many embryos, either white

or green. In young developing fruits short-stalked club-shaped

slime-secreting organs develop rapidly with the pulp-vesicles; these

soon mature and liberate slimy matter which probably enables the

developing pulp-vesicles to slide freely over one another.

658 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

The remarkable genus Rhabdodendron Gilg & Pilger of the

Amazonian region would be sought here on account of its numerous

stamens with long slender anthers; the calyx is reduced to a rim of

the fleshy conical pedicelled axis, petals caducous, style from base

of ovary, fruit firm, ovoid, sessile in the axis, with 1 seed; the known

species are R. macrophyllum (Spruce) Huber and R. amazonicum

(Benth.) Huber, the leaves of the former with very prominent nerve,

this obsolete in the leaves of the latter.

Williams as myself found sour and sweet oranges persisting after

cultivation had been abandoned; he also collected C. reticulata

Blanco, the Mandarin orange, as an escape at Pebas, where known

as "Tansharina."

Citrus fruits were introduced into Peru by the Spanish. Garcilaso

de la Vega was quoted as saying in 1609: "All these fruits grow

there today in abundance." They are still however mostly if not

entirely absorbed by the local markets. Key and descriptions after

Swingle, I.e.; determinations by Standley.

Petioles wingless; flowers perfect or often male C. medica.

Petioles winged, clearly articulated with the blade.

Stamens usually more than 4 times petals; flowers perfect, often

male; petiole wings very narrow C. Limon.

Stamens usually 4 times petals; flowers usually perfect.

Fruits usually 4-6 cm. in diameter, yellowish-green; petiole

wings narrow C. aurantifolia.

Fruits usually 5-9 cm. in diameter, scarlet-orange; petiole

wings broad .C. Aurantium.

Citrus aurantifolia (Christm.) Swingle, Journ. Wash. Acad.

Sci. 3: 465. 1913; 401. Limonia aurantifolia Christm. in Linn.

Pflanzensyst. 1: 618. 1777.

Branchlets with short stiff very sharp spines; petioles narrowly

winged, spathulate; leaves elliptic-ovate or oblong-ovate, rounded

at base, obtusely acuminate, crenulate, 5-7.5 cm. long; racemes

2-7-flowered; petals 8-12 mm. long, 2.4-4 mm. wide; stamens 20-25;

ovary globose, not merging into the early deciduous style; fruits

small, peel very thin, greenish yellow when ripe. — In places appears

to grow without cultivation (Williams).

The Sweet Lime seems not to be distinct taxonomically; here

treated under C. Limon, for convenience of nomenclature, the name

C. Limonia Osbeck, Reise Ostind. & China 250. 1765, not disposed

FLORA OF PERU 659

of by Swingle; Annetta Carter, Univ. Calif. Herb., kindly discovered

for me that their English edition, 1771, contains no such name;

according to Merrill, Amer. Jour. Bot. 3: 579, this name appears

only in the German edition, 1765.

Loreto: Lower Itaya, Williams 177. Lower Nanay, Williams

285; 471. Upper Itaya, Williams 3268. Pebas, Williams 1800 —

Cuzco: Santa Ana, Cook & Gilbert 1556 (det. Tanaka). East Indian

Archipelago. "Lime," "limon agrio" (Williams), "lima."

Citrus Aurantium L. Sp. PI. 782. 1753; 402.

Branchlets with single often short slender spines or stout ones

3-8 cm. long on vigorous shoots; petioles 2-3 cm. long, rather

broadly winged, often 10-18 mm. wide at top; leaves ovate, rounded

to cuneate at base, obtusely acuminate; flowers large, very fragrant,

5-12 per cent male; peel thick with a rather rough surface becoming

brilliant orange with a reddish tint at maturity; cells 10-12 with

sharply acid pulp and many seeds. — Fruit becomes hollow at center.

Introduced into the Mediterranean region by the Arabs about the

eleventh century and for about five centuries was the only orange

known to Europeans. It was the much appreciated orange of medie-

val Europe, used for flavoring marmalade; the flowers were used for

perfume. The closely allied and commonly cultivated sweet orange

is C. sinensis (L.) Osbeck, probably native to southern China but

no longer known in a wild condition; it first became known about

A.D. 1500. According to Williams the orange is sometimes culti-

vated. The related grapefruit is C. paradisi Macf., origin probably

West Indies.

Junin: Hacienda Schunke, Killip & Smith 24733. — Ayacucho:

In clearing, Killip & Smith 22816. — Loreto: Pebas, Williams 1774.

From southeastern Asia. "Sour or Seville orange," "naranja acida,"

"naranjo."

Citrus Limon (L.) Burm. f. Fl. Ind. 173. 1768. C. medica L.

var. Limon L. Sp. PI. 782. 1753. C. Limonum Risso, Ann. Mus.

Hist. Nat. Paris 20: 201. pi 2. 1813.

Small thorny trees, the young leaves and flower buds reddish;

petioles narrowly winged or margined, plainly articulated with the

leaf -blade, this pale green, long-ovate, acuminate, serrate; flowers

white above, purplish below; stamens 20-40; ovary tapering into

thick deciduous style; fruit oval with 8-10 segments, yellow when

ripe. — Brought to the Mediterranean region by the Arabs about

660 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

A.D. 1000 and like the sour orange soon much used in medicine;

now known as the source of the recently isolated vitamin P; regard-

ing this and its status as a species cf. Swingle, I.e. 399. The origin

of the Lemon is unknown; it may be either acid or "sweet" var.

Limetta (Risso) Engl., or, as distinct, C. Limetta Risso, flowers white,

fruit not mammillate, the Sweet Lime; cf. remark under C. aur anti-

folia.

Loreto: Lower Nanay, Williams 266. La Victoria, Williams;

2933. Origin unknown. "Limon," "limon dulce."

Citrus medica L. Sp. PI. 782. 1753; 396.

Very much like C. Limon but the petioles wingless or merely

margined and not obviously articulated with the blades, stamens

30-40 or even 60; style sometimes persisting. — According to Swingle,

Citron was the first citrus fruit to reach the Mediterranean, brought

by the armies of Alexander the Great apparently about 300 B.C.

Theophrastus called it the Persian or Median Apple; Engler, Pflan-

zenfam. ed. 2. 19a: 338. 1931, wrote: Since the fruits had the same

uses as the wood of the Sandarac Tree, the name of this wood,

"Citrus," was transferred to the fruit as Mala citrea. So the Median

Apple became the Citrus Apple and today we have the common

term citrus for Citron and all its many relatives. The candied peel

is well known as a "dulce."

Ayacucho: Clearing near Aina, Killip & Smith 22815 (det.

Swingle). — San Martin: San Roque, Williams 7211. — Loreto:

Iquitos, Williams 1464; Killip & Smith 27420.—- Cuzco: Santa Ana,

Cook & Gilbert 1666 (det. Tanaka). San Miguel, Cook & Gilbert

1078 (det. Tanaka). From China and British India, southward.

"Limon cidra," "cidra."

2. MURRAYA Koenig

Unarmed trees with imparipinnate leaves of alternate leaflets

and rather showy 5-merous flowers crowded in axillary or terminal

corymbs or panicles. Sepals nearly distinct or united below the

middle. Petals imbricate. Stamens 10, free, disk annulate or

cushion-like. Ovary ovoid, 2-5-celled with 1 or 2 ovules super-

imposed or subcollateral in each cell. Style slender, finally deciduous,

stigma capitate. Fruit a small berry with mucilaginous pulp.

Murraya paniculata (L.) Jack, Malay Misc. 1: 31. 1820.

Chalcas paniculata L. Mant. 68. 1767. M. exotica L. Mant. 2: 563.

1771.

FLORA OF PERU 661

Branchlets slender, finely crisp-puberulent toward the tips;

leaves with 2 or more pairs of obliquely obovate subsessile leaflets

about 2-3.5 cm. long, 1-2 cm. wide; flowers white, 13-18 mm. long;

stamens alternately shorter; berries ovoid, orange-colored, about

13 mm. long, with 1 or 2 villous seeds. — Often cultivated in warm

regions; elsewhere in greenhouses; native to southeastern Asia.

San Martin: Near Tarapoto, Williams 5946. — Loreto: Iquitos,

Williams 3696. Yurimaguas, Williams 4055. "Naranjilla." Asia.

3. AMYRIS [P. Br.] L.

Trees or shrubs, usually glabrous, often aromatic, with alternate

or opposite compound, sometimes unifoliate, pellucid-punctate

leaves and many small whitish bibracteate perfect flowers mostly

in 3's in axillary or terminal panicles. Sepals and petals 3 or 4 (5),

the former persisting, the latter imbricate in bud, spreading in late

anthesis, the stamens twice as many from base of obscure disk;

filaments filiform. Carpels in female flowers on a thick disk; ovary

with 2 ovules pendulous from apex of the single cell. Fruit a fleshy

drupe with 1 seed. — In flower may simulate species of Zanthoxylum,

that genus however with usually unisexual flowers.

Amyris pinnata HBK. Nov. Gen. & Sp. 7: 37. pi 610. 1824.

Young branchlets and panicles minutely crisp-puberulent;

stipules none; petioles opposite, 2.5-3 cm. long; leaves imparipinnate

with 2-3 pairs of opposite ovate (or terminal elliptic) leaflets, the

lateral sometimes oblique, all obtuse or obtusely subacuminate,

rounded at base, obsoletely crenate or undulate, reticulate veined,

membranous, glabrous, lustrous above, paler and dull beneath,

3.5-7 cm. long, 2-4 cm. wide, the lower smaller, petiolules about

4 mm. long; panicles axillary, solitary, much branched, peduncles

to about 7 cm. long, flowers 1-3, the short pedicels medially and

minutely bracteolate; calyx urceolate, typically acutely 4-lobed,

glabrous but puncticulate glandular as the 4 (type) petals, these

white, elliptic, somewhat narrowed below, rounded at tip; stamens

8, 4 longer, anthers all alike; ovary glabrous but punctate; stigma

sessile. — Type, "trop. Amer." A. balsamifera L., West Indies,

Cuba, Colombia and Ecuador, has a pubescent ovary; its hard wood,

known as Torch Wood, is aromatic and popular to burn for the

fragrance.

Amazonas: Region of San Carlos, 1,400 meters, Weberbauer

7146. Colombia.

662 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

4. ZANTHOXYLUM [Catesby] L.

Fagara L. Syst. Nat. ed. 10. 897. 1759.

Shrubs or trees, the branches and leaves often armed with straight

or recurved spines, the leaves often imparipinnate, rarely 1-3-foliate

with usually opposite petiolulate entire or crenate pellucid-punctate

leaflets. Cymes often panicled, crowded, axillary or terminal, the

dioecious or polygamous flowers usually white or greenish, the fruits

of 1-5 carpels usually aromatic and glandular-punctate. Calyx

3-5-parted, rarely deciduous, petals 3-5, rarely none. Disk short

or obscure. Male flowers with 3-5 (-8) stamens alternate to the

petals or opposite the sepals (Fagara), in the female wanting or

staminodal. Carpels oblique, 1-celled, the style sublateral, stigma

capitate; ovules 2. — The mature fruit parts are ordinarily bivalvate

to the middle and expose the single reddish or black lustrous seed

pendulous on slender funicle. The name is spelled variously, the

substitution of X for Z alone being noteworthy since alphabetically

significant.

In Fagara, according to Engler, Pflanzenfam. ed. 2. 19a: 215.

1931, the flower parts are in alternation while in Zanthoxylum the

5-8 stamens alternate with the 5-8 petals. This may be significant

morphologically — and thus historically — but, as Engler also hints,

the division of the otherwise homogeneous group on this basis is

scarcely expedient in view of the fact that the diagnostic character

is not observable in pistillate plants. However, so far as known all

the Peruvian species are actually referable to Fagara or, more

practically, to section Fagara (L.) Macbr., comb. nov.

The species' characters seem to be intangible; Engler, in Mart.

Fl. Bras. 12, pt. 2: 156-158. 1874, had to resort in his key to such

doubtful distinctions as presence or absence of spines, development

of leaf-crenation, etc.; in Pflanzenfam. ed. 2. I.e., he remarks that

the number of carpels may prove to be significant and this is my

impression: the nature and position of the inflorescence may be

further clues to the relationship of the species together with the

usual petal number, which however like the leaflet number is variable

within the same species and in itself of questionable value. The

following key then is merely academic and the validity of the species

has, as usual of necessity in my compilations, not been proved.

Many species, it may be remarked, are incompletely known, lacking

fruit or some other part; surely one cannot intelligently write

"armed" or "unarmed" when one has only a flowering branchlet!

FLORA OF PERU 663

Fagara peruviana Willd. ex Roem. & Schult. Mant. 3: 228. 1827

is Z. affine HBK. of Mexico according to Index Kewensis, and in

spite of the name is therefore omitted here; the supposed origin of

the species is apparently an error, as has happened before.

Ruiz and Pavon in their journal gave a name that was never

published to a species from Chacahuasi and Pozuzo, Huanuco,

which could be several species, perhaps most probably Z. Culantrilo,

known, according to them, as "culandro" or "culantro" because

the odor is similar to that of coriander.

Leaflets sessile or subsessile, often narrow or small, except Z. mantaro,

about 2.5 cm. wide or narrower.

Pubescence lacking or simple; flowers racemose or pseudoracemose.

Leaflets more or less crenulate, mostly 2 cm. wide or wider;

petals 4.

Leaflets 7-11, crenulate, usually to 2.5 cm. long; branches

usually thorny; spikes usually only to 15 mm. long.

Z. Fagara.

Leaflets 5-13, crenate, mostly 3.5 cm. long or longer; spikes

usually 2.5 cm. long or longer; spines none or few.

Z. Culantrilo.

Leaflets subentire, to 4.5 cm. wide; petals 5 Z. mantaro.

Pubescence minutely stellulate; flowers paniculate .Z. Ruizianum.

I Leaflets obviously petiolulate, to 4 cm. wide or wider.

Leaves glabrous.

Inflorescence terminal; petals 3 or 5.

Leaflets crenulate, 3-5 pairs.

Petals 3 Z. acreanum.

Petals 5 Z. juniperinum.

Leaflets entire or nearly, often more than 5 pairs.

Leaflets obtusely short-acuminate; pedicels to 2 mm. long.

Z. Riedelianum.

Leaflets acutely acuminate; pedicels about 0.5 mm. long.

Z. valens.

Inflorescence axillary; petals 4 or 5; leaflets entire or nearly.

Panicles nearly racemiform, congested in fruit. . . .Z. Sprucei.

Panicles open, the spreading branches elongate.

Z. Weberbaueri.

Leaves more or less pilose Z. tumbezanum.

664 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Zanthoxylum acreanum (Krause) Macbr., comb. nov. Fagara

acreana Krause, Notizbl. Bot. Gart. Berlin 6: 143. 1914.

Scandent branches and slender branchlets terete or the latter

lightly compressed apically, early sparsely puberulent, densely lenti-

cellate and with a few short erect conic acute spines; petioles 5-6 cm.

long; leaves alternate, 3 dm. long, with 3-4 pairs of opposite petiolu-

late thin-coriaceous glabrous oblong leaflets, narrowed to base,

obtusely acuminate (acumen 10-12 mm. long), often little unequal,

mostly 8-12 cm. long, about 4-4.5 cm. wide, or the terminal a little

larger, all lightly crenulate, with 12-14 lateral nerves very promi-

nent beneath; flowers subsessile, in ample dense many-flowered

terminal panicles 12-15 cm. long; flowering branchlets puberulent,

with small ovate acute bracts; calyx lobes nearly free, scarcely 0.5

mm. long, sparsely pilose below; petals 3, white, oblong, obtuse,

three or four times longer than calyx and a little shorter than the

3 filaments; anthers ovoid, subcordate; rudimentary ovary with

short style. — Belongs near F. subserrata Engl. and F. Warmingii

Engl. but leaflets long-petiolulate (Krause). But it seems to be

essentially Z. juniperinum (from which Engler distinguished it on

less crenulate leaflets) unless the flowers are constantly 3-merous.

F.M. Neg. 26412.

Rio Acre: San Francisco, Ule 9552, type.

Zanthoxylum Culantrilo HBK. Nov. Gen. & Sp. 6: 2. 1823.

Fagara Culantrillo (HBK.) Krug & Urban, Bot. Jahrb. 21: 574. 1896.

Much branched, unarmed; younger branchlets hirtellous; leaves

ternate or imparipinnate with 2 (or more) pairs of oblong obtuse

glabrous crenate (punctate-glandular between the crenations) leaf-

lets, subcuneate at base, obsoletely reticulate, membranous, paler

beneath, 24-32 mm. long (3.5-8 cm., fide Urban), 10-14 mm. wide,

sessile except the terminal in the ternate leaves, the 2 lower smaller

in the pinnate, rachis obscurely margined, petiole not at all; inflores-

cence axillary or subterminal (lateral and terminal, speciform or

branched, Urban), a little longer than the petioles, scarcely 2.5 cm.

(or longer), with 2-4 spreading branches; hirtellous flowers sub-

sessile, few; calyx lobes and petals 4, the former rounded the latter

4 times longer, white; male flowers with 4 stamens, anthers elliptic,

bifid at base, obtuse, the female with no stamens, ovary sessile,

glabrous, 1-celled, stigma capitate; carpel subglobose, smooth, gla-

brous, the seed globose. — A slender tree, 4-6 meters (Killip &

Smith); my collection, a strictly branched tree. If distinct from

FLORA OF PERU 665

Z. Fagara probably includes most if not all of the material referred

to that species. The name was originally written with one "1".

The var. paniculata Engl. is the typical form.

Cajamarca: Jae"n de Bracamoros, Bonpland, type. Nancho,

Raimondi. — San Martin: Region of Moyobamba, Weberbauer 4498;

289. Tarapoto, Spruce 4293 (det. Urban). — Junin: Perene", Killip

& Smith 25032. San Ramon, Schunke Al 02. — Huanuco : near Pozuzo,

4773. San Carlos, Mexia 8212. Colombia; Venezuela; Jamaica.

"Culantrilo," "culantro," "contrevenosa."

Zanthoxylum Fagara (L.) Sarg. Gard. & For. 3: 186. 1890.

Schinus Fagara L. Sp. PI. 389. 1753. Fagara Pterota L. Syst. Nat.

ed. 10. 897. 1759.

Intricately branched shrub or small tree, the branches commonly

thorny, the branchlets early puberulent; petioles and leaf-rachises

winged, the leaves several to about 10 cm. long with 3-4 pairs of

obovate-elliptic (or 7-11 leaflets, Urban) sometimes somewhat orbicu-

lar leaflets mostly 1.5-2.5 cm. long, narrowed to petiolulate base,

rounded or retuse at apex, lustrous above, crenulate, the glands

mostly below the crenations; flowers in short contracted axillary

spikes (early capituliform-racemose) 7-15 mm. long, 4-merous, the

yellow-green petals 2-3 mm. long, the male flowers with 4 exserted

stamens; carpels 2, globose, 3-4 mm. in diameter, the seed black.—

Doubtfully in Peru unless Z. Culantrilo is not distinct. Illustrated,

Sarg. Silva 1: pi 32.

Cajamarca: Raimondi. — San Martin: Tarapoto, Williams 5510;

5511 ; Vie 6355. Near Moyobamba, Klug 3729. — Junin: La Merced,

Soukup 2489. — Loreto: Yurimaguas, Williams 4751; 4449. Florida

to Venezuela and Colombia; and apparently Brazil; Galapagos?

Peru? "Shapilleja" (Williams); "una de gato."

Zanthoxylum juniperinum Poepp. & Endl. Nov. Gen. & Sp.

3: 77. 1845; 167. Fagara juniperina (Poepp. & Endl.) Engl. Pflan-

zenfam. 3, 4: 117. 1896.

Glabrous except the younger puberulent panicles; branches long-

rugose with stout spines, the terete branchlets purplish; petioles

acutely grooved above, 1.5-3 dm. long, internodes 2-7 cm. long;

leaves coriaceous, green and lustrous above, glandular and sometimes

pellucid punctate at margin, mostly abruptly pinnate with 3-4 pairs

of indistinctly crenate gland-margined oblong-obovate obtusely or

rather acutely acuminate espinose leaflets, the larger 1.5 dm. long,

666 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

7 cm. wide, acumen to 1 cm. long; panicles 1-1.5 dm. long and nearly

as broad, glabrous in fruit; pedicels 1-2 mm. long; white petals and

stamens 1.5 mm. long; calyx coriaceous, minutely puberulent lobes

rounded; carpel solitary, subglobose, brown, 3-4 mm. in diameter,

the glands small. — Type from "a tall tree." Similar but with aculeate

petioles is Z. quinduense Tul., 168, panicles glabrous, and Z. macro-

spermum Tul., 168, panicles velutinous, both species Colombian.

F.M. Negs. 12447; 26417.

Loreto: Maynas, Poeppig 2465, type. Yurimaguas, Williams

5205. Santa Rosa, Williams 4911. "Walaja."

Zanthoxylum mantaro Macbr., comb. nov. Fagara mantaro

Macbr. Candollea 5: 372. 1934.

Similar to Z. Culantrilo but except for the lightly pulverulent

young branchlets nearly glabrous and the leaflets subentire; branches

in type unarmed; leaves dense, 8-15 cm. long, the puberulent

teretish petiole complanate above but not at all winged, the 3-5

pairs of leaflets sessile, elliptic, obliquely rounded at base (or slightly

and obtusely acuminate), typically rounded at tip, to 6 cm. long,

half as wide, especially the lower smaller, all coriaceous, lustrous,

epunctate, the midnerve very prominent beneath, lateral nerves

nearly parallel, scarcely prominent, the margin subentire or remotely

and obscurely dentate; racemes little branched, about 3 cm. long,

minutely puberulent as the pedicels; calyx segments ovate; petals 5,

elliptic-oblong, sparsely and most minutely ciliate, 2.5 mm. long;

stamens little exserted; fruit consisting of 1-3 stoutly stiped or

sessile ovate-rotund somewhat compressed sparsely foveolate carpels

4.5-nearly 6 mm. wide. — In spite of the fact that I have not found

an earlier name for this shrub-tree there probably is one; my collec-

tions at one time referred to Brazilian species having leaflets 10-12 cm.

long. The Colombian Z. amoyense Tul. has smaller oblong leaflets

obliquely acute at base; Z. macrospermum Tul. may be near, but its

leaflets are minutely but distinctly crenulate-serrulate and the rather

similar Z. quinduense Tul. has pyramidal panicles. If this proves

to be a valid species the name may be "corrected" to mantaranum,

which probably was intended, the lack of ending unnoticed in proof.

Junin: Rio Mantaro, Weberbauer 6585, type. Huanuco: Cani

near Mito, 3440. Mito, 1732. Cuzco: Torontoy, Urubamba Valley,

Cook & Gilbert 1772.

Zanthoxylum Riedelianum Engl. in Mart. Fl. Bras. 12, pt.'2:

162. 1874. Fagara Riedelianum Engl. Pflanzenfam. 3, 4: 117. 1896.

FLORA OF PERU 667

Trunk straight, conically aculeate; younger branchlets densely

ashy pilose, older long-rugulose with gray bark; petioles broadly

canaliculate above, 1.5-2 dm. long; leaves abruptly, rarely impari-

pinnate with 4-6 pairs of subentire subequal (except 2 small lowest

ones) leaflets the middle and terminal oblong-elliptic, shortly acumi-

nate, obtusish, acute at base into canaliculate petiolule 5-8 mm. long,

mostly 9-12 cm. long, 2.5-4 cm. wide; medial and lateral nerves as

the reticulate veins prominent beneath; panicles terminal, shortly

and densely pilose, compound, the branches spreading, the lower

to 1 dm. long; flowers glomerulate on glabrate slender pedicels 2 mm.

long; calyx glabrous except for cilia on the 5 ovate acute lobes, the

5 acutish petals 5 times longer; stamens 4 mm. long, twice as long

as the petals; female flowers unknown. — The Peruvian specimen

distributed as this is aberrant, the inflorescence axillary, the leaflets

sessile, variations known to occur in other species; however here the

collection is referred to Z. mantaro. Illustrated, Engl. I.e. pi. 35.

F.M. Neg. 19227.

Peru: Probably. Ecuador; Brazil (Sao Paulo).

Zanthoxylum Ruizianum ([Klotzsch] Engl.) Macbr., comb,

nov. Z. obscurum Engl. var. Ruizianum [Klotzsch] Engl. in Mart.

Fl. Bras. 12, pt. 2: 170. 1874. Z. rhoiifolium Lam. var. sessilifolium

Engl. I.e.

Branches and branchlets probably armed, the flowering tips

sparsely and minutely lenticellate, drying rugulose, finely puberulent,

a few similar trichomes, mostly minute, on leaf-rachises and under

side of subtending leaves, these soon glabrous, 1.5-2 dm. long in-

cluding the short petioles; rachises angulately margined by decur-

rent-based leaflets, these mostly alternate, 8 or more pairs, oblong-

lanceolate, strongly oblique at subsessile base, shortly and broadly

acuminate the tip itself truncate or minutely retuse, subequal (un-

less the lower pair), mostly about 6 cm. long, 15-18 mm. wide,

obviously, even strongly crenate with rather prominent indentation-

glands, drying brown-green, paler beneath, slightly lustrous above,

sparsely pellucid-punctate; panicles about 8 cm. long and broad,

the primary widely divaricate branches 3-6 cm. long, secondary

about 1 cm. long; pedicels scarcely 1 mm. long; calyx lobes 5, rounded,

minutely ciliolate; petals probably less than 1.5 mm. long; ovary

subglabrous; carpels (young) solitary, brown, sparsely glandular-

Descriptions from scraps of type and Williams material in bud and

young fruit; detached leaves probably belonging are sparsely aculeate,

668 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

the spines 8-10 mm. long. Seems to differ from Z. obscurum with

about 6 pairs of leaflets, their petiolules 4-5 mm. long;Z. caribaeum

Lam. of this affinity is apparently glabrous; in Pflanzenfam. ed. 2.

19a: 220. 1931, Engler keys out that species and F. Ruiziana,

which thus becomes questionably published, as completely gla-

brous; the latter however as to type is as described above and perhaps

should be included in Z. rhoiifolium Lam. at least as interpreted by

Engler but, ex char., it has ovate acute calyx lobes and the type

locality is said to have been the East Indies; in view of the un-

certainty of the application of the name I retain, albeit reluctantly,

the Klotzsch plant as distinct, as apparently Engler in his most

recent study considered it; Z. aculeatissimum Engl., 176, of Bolivia,

may be an available name but apparently it is not, as its leaflets

are somewhat broader, more ovate-oblong and longer attenuate to

tip. F.M. Neg. 12460 (Z. obscurum var.).

Huanuco: Macora, Ruiz & Pavon (type, Z. obscurum var. Ruizi-

anum). Without data, Ruiz & Pavdn, Herb. Lambert, fide Engler.—

San Martin: Tarapoto, Williams 5559; 6500; 6669. San Roque,

Williams 7235. — Loreto: Mouth of Rio Santiago, Tessmann 4126.

"Quillu-casha," "shapilloja," "huillca."

Zanthoxylum Sprucei Engl. in Mart. Fl. Bras. 12, pt. 2: 167.

1874. Fagara Sprucei Engl. Pflanzenfam. 3, 4: 117. 1896.

Branches rather stout, with gray smooth bark armed with many

spines; petioles 2-3 dm. long, aculeate, grooved above, internodes

sometimes 5 cm. long, with a few aculei 2-3 mm. long; leaves in

age coriaceous, early sparsely punctate, glabrous, lustrous above,

imparipinnate or abruptly pinnate, with 3-5 pairs of essentially

entire or lightly crenate subequal ovate-oblong or oblong scarcely

acuminate or shortly and falcately obtusish leaflets that are oblique

to the 2-3 mm. long petiolules, the larger 12-14 cm. long, 5-6 cm.

wide; lateral nerves as veins prominent both sides, the midnerve

so beneath where aculeate; flowering branchlets axillary, compound-

paniculate, in fruit 4-6 cm. long, branched from base, lower branches

2 (-4) mm. long, the stout branchlets angled, puberulent; calyx

lobes short, obtuse; petals 4 (always?), broadly ovate, 1.5 mm.

long; carpels 3 or 4, ovate-rounded, strongly compressed, 2.5 mm.

long and broad, not at all stiped, brown foveolate-rugulose.— Tree

20 meters high, 6 dm. in diameter (Tessmann). F. caudata Huber,

Bol. Mus. Goeldi 5: 422. 1909, has petiolules to 1 cm. long, leaflets

6 cm. long, 2.5-3 cm. wide, caudate-acuminate and solitary carpel

about 1 cm. long, 6 mm. wide. F.M. Neg. 12461.

FLORA OF PERU 669

San Martin: Tarapoto, Spruce 4187, type; Williams 5681.

Zepelacio, Klug 3373 (det. Standley). — Loreto: Yurimaguas, Wil-

liams 4006. Puerto Limon, Tessmann 3866. Pumayacu, Klug

3161 (det. Standley). Rio Acre: Ule 9506 (det. Krause). "Valajol,"

"ualaja" (Tessmann).

Zanthoxylum tumbezanum Macbr., comb. nov. Fagara

tumbezana Macbr. Field Mus. Bot. 8: 119. 1930.

Branches glabrous, apparently little or not at all armed, the

younger as the leaves both sides more or less densely short-pilose;

leaves crowded, very unequal, sometimes sparsely yellowish-aculeate

beneath, the 3 or 4 pairs of short-petiolulate leaflets oblong-elliptic

or subovate-elliptic, shortly acutish acuminate, obtuse at the oblique

base, entire or minutely crenulate, 3-5 cm. long, 1.5-2 cm. wide,

mostly 8 cm. long, 3 cm. wide, chartaceous-coriaceous, the nervation,

including the reticulate veins, inconspicuous both sides; panicles

subracemose, axillary, minutely pilose, 4 cm. long; pedicels slender,

3 mm. long; calyx segments ovate, about 0.5 mm. long; petals 5,

narrowly ovate-oblong, glabrous, 2 mm. long, filaments nearly as

long, the cordate anthers conspicuous; ovary rudimentary, attenuate

into short style. — Perhaps related to F. Riedeliana Engler which

has been found as near as Ecuador but that as to type glabrous or

only lightly pubescent. It resembles, too, Andean material referred

to Z. rigidum Humb. & Bonpl. ex Willd. Sp. PI. 4: 756. 1806 of

"middle America" which, at least as to type, has emarginate leaflets,

aculeate and pubescent beneath only on veins.

Tumbez: Mountains east of Hacienda Chicama, Weberbauer 7642,

type.

Zanthoxylum valens Macbr. ex L. Williams, Field Mus. Bot.

15: 228. 1936. Fagara valens Macbr. Candollea 5: 373. 1934.

Said to be a large tree but the branchlets and petioles unknown;

leaves about 3 dm. long and 13-foliolate; leaflets subcoriaceous, very

lustrous above, little so beneath, subequal, obliquely oblong-elliptic,

strongly oblique or also acute at base, abruptly caudate-acuminate

(acumen 7-10 mm. long), mostly 13 cm. long, 5.5 cm. wide, entire,

the very prominent midnerve most minutely pulverulent, reticulate

veined especially beneath, the lateral nerves prominent both sides,

minutely and densely puncticulate; petiolules 7-10 mm. long;

flowering branches pulverulent, 3 dm. long, paniculate, with spread-

ing branchlets; pedicels very short; calyx parts short, ovate, acute;

petals 5, subovate, glabrous, scarcely 1.5 mm. long, stamens filiform,

670 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

nearly twice as long. — Material incomplete but perhaps nearest

Z. cuyabense Engl. and Z. Sprucei Engl. ; it could be the staminate

plant of the latter but the leaflets are more numerous than recorded

for that species. The rachis of Williams 574 is sparsely aculeate.

Loreto: Pebas, Williams 1962, type. In forest south of Nanay,

Williams 574. "Raia-caspi."

Zanthoxylum Weberbaueri (Krause) Macbr., comb. nov.

Fagara Weberbaueri Krause, Repert. Sp. Nov. 2: 26. 1906.

A much branched erect shrub, glabrous except the shortly pilose

inflorescences; branchlets armed with compressed brown spines about

5 mm. long; petioles mostly unarmed, upper half sulcate, petiolules

scarcely so, these about 1 cm. long; leaves imparipinnate, with 3-4

pairs of entire subcoriaceous sparsely pellucid-punctate leaflets

drying dark above, paler beneath the midnerve and laterals promi-

nent, elliptic, 11-13 cm. long, 4-5 cm. wide, acute at base, apically

contracted into acumen about 1 cm. long; flowers white, in compound

racemes (or paniculate); pedicels about 1 mm. long; calyx cupulate,

the very short segments broadly ovate; petals 4, 3-nerved, the lateral

ones less distinct, ovate, 2-3 mm. long, the stamens about twice

as long, filaments a little thickened toward the slightly dilated base, ,

anthers yellow, ellipsoid; stigma sessile, bifid. — To 8 meters high.

Klug specimen is nearly glabrous, no spines. Section Macqueria,

in the vicinity of Z. spinifex (Jacq.) of the West Indies and Vene-

zuela (Krause); so distinguished by Engl. Pflanzenfam. ed. 2. 19a:

218. 1931 fromZ. Culantrilo by the entire leaflets; in these it resembles

Z. mantaro but its leaflets are petiolulate, its petals apparently 5.

F.M. Neg. 26424.

San Martin: East of Moyobamba near the Rio Mayo, 800 meters,

Weberbauer 4769; 287. — Loreto: Cachipuerto, between Balsapuerto

and Moyobamba, Klug 3126.

5. METRODOREA St. Hil.

Similar to Esenbeckia but the petioles broadened at base and with

small appendages or lanceolate leaves at or near it, the leaves them-

selves always opposite. Petals valvate in bud. Ovary entirely im-

mersed. Fruit capsular, protuberances unequal in size. — Com-

memorates Metrodoro Sabio who, according to Pliny, was the first

to accompany plant descriptions with illustrations. Apparently the

genus, considered taxonomically, is a part of Esenbeckia, but many

species have a rather different facies.

FLORA OF PERU 671

Metrodorea flavida Krause, Notizbl. Bot. Gart. Berlin 6: 146.

1914. Esenbeckia coriacea A. C. Sm. Bull. Torrey Club 60: 358.

1933.

Branches and branchlets slender, terete, usually densely minutely

lenticellate as the stout petioles, these about 2 to nearly 3 cm. long,

strongly dilated at base, the apex little enlarged; leaflets usually 3,

sessile, thin-coriaceous, glabrous, a little lustrous, drying brown

beneath, oblong or obovate-oblong, obtusely acuminate, subacute

at base or the lateral slightly oblique, 1-1.5 dm. long, 4.5-6.5 cm.

wide, lateral nerves 9-12, prominent beneath; flowers whitish-

yellow, many in terminal and axillary lax panicles 1-2 dm. long,

to 1 dm. wide; flowering branches mostly opposite, sparsely pilosulous

or glabrate, pedicels puberulent, subequaling the globose buds; calyx

segments 6, broadly ovate, 1-1.2 mm. long, lightly puberulent and

ciliolate, glabrous within; petals oblong, subacute, sparsely pilose,

3-3.5 mm. long, glandular-punctate, about twice as long as the

stamens, these with subulate filaments, ovate-cordate anthers; disk

thick including the attached ovary, tubercled, the style as long.—

The petiole is not only enlarged basally but extended within the

axil as a thick free appendage. Differs from the southern Brazilian

M. nigra St. Hil. in the yellow-white flowers; from M. pubescens St.

Hil. & Tul. by the longer slenderer petioles and smaller leaves

(Krause). Type, 3-18 meters high; 30 meters (Krukoff). Fruit

very rugose, 3 mm. long, 4 mm. wide, cells 1-seeded (Smith); the

measurements of Smith are very slightly at variance with those of

Krause, given above.

Rio Acre: Mone Mo, Seringal San Francisco, Ule 9491, type.

Rio Macauhan, Krukoff 5517 (type, E. coriacea). Brazil.

6. ESENBECKIA HBK.

Polembryum Juss. Me*m. Mus. Hist. Nat. Paris 12: 519. 1825.

Trees or tall shrubs with crowded, alternate, sometimes opposite,

simple to 3 (5) -foliate pellucid-punctate leaves without distinctive

stipules or dilated petioles. Branchlets and pedicels of the sparsely

or densely flowered inflorescences with a pair of opposite bracts.

Sepals, petals and stamens 4 or 5, the first finally deciduous, the

second imbricate or nearly valvate in bud, the third between the

8-10 lobes of the annulate or cupulate disk, or this entire. Ovary

sessile or immersed, 4-5-lobed and -celled with usually 2 collateral

ovules. Fruit sometimes nearly keg-shaped, rarely smooth, breaking

into 4 or 5 bivalved 1-2-seeded often radiately disposed carpels.

672 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Endocarp elastic, finally splitting and recurving to eject the exal-

buminous seeds. — Named for Nees von Esenbeck, better known in

botanical work as Nees.

Leaves simple; petioles not articulate.

Carpels cornute medially; leaves pilose E. cornuta.

Carpels cristate; leaves glabrous or nearly E. grandiflora.

Leaves 1-trifoliate; petioles articulate; carpels cornute apically if at

all.

Petioles and inflorescence densely pilose E. Warszewiczii.

Petioles and inflorescence glabrous.

Leaves trifoliate or mostly rounded apically; petals 5 mm. long.

E. venulosa.

Leaves mostly 1-foliate, obtuse; petals 2.5-3 mm. long.

E. pilocarpoides.

Esenbeckia cornuta Engl. in Mart. Fl. Bras. 12, pt. 2: 146.

1874.

Branchlets terete, scarcely 3 mm. thick, ashy pubescent, the

younger as the new leaves and flowering branchlets (these to 1 dm.

long with cymose-paniculate many-flowered inflorescences) densely

pilose; leaves above sparsely, beneath closely and very shortly ashy

pilose, simple, oblong-elliptic, attenuate to obtuse acumen about

1 cm. long, mostly 10-12 cm. long, 7 cm. wide, the terete petioles

2-2.5 cm. long; midnerve stout, the lateral moderately prominent;

flowering branchlets terminal and axillary, pedicels 1-1.5 mm. long,

as long as the globose buds, densely ashy pilose as the calyces and

petals; sepals acutish, nearly 1 mm. long; petals spreading, 2 mm.

long, half as wide at base, glabrous within, stamens nearly as long;

disk annulate, 10-plicate, longer than the sericeous ovary; carpels

ligneous, subtrigonous, about 17 mm. long, 1 cm. broad, finally

glabrate, dorsally horned medially, the horn 5 mm. long and dehiscing

to it, ventrally nearly to base. — F.M. Neg. 12512.

Cajamarca: Jae"n de Bracamoras, Warszewicz, type.

Esenbeckia grandiflora Mart. Nov. Gen. & Sp. 3: 85. 1829;

146. PolembryumJussieui Schott, Rutac. 11. pi. 6. 1834, fide Engler.

Branches grayish, the many short densely leafy branchlets

brownish-red; petioles 3-6 (-15) mm. long, transversely rugose;

leaves simple, sometimes subopposite, more or less obovate-oblong

or also ovate, cuneate at base, obtuse or rounded at tip, typically

FLORA OF PERU 673

about 1 dm. long, 4 cm. wide, or often much smaller, coriaceous,

glabrous or beneath sparsely pubescent, the middle and lateral nerves

there prominent; flowering branchlets from the axils of the longer

upper leaves, opposite or alternate, 3-5 cm. long, paniculate or

racemose; pedicels 1 cm. long, enlarged below calyx; bracts sericeous,

those in panicles 5-10 mm. long, in racemes smaller, the 2 below the

globose buds ovate-lanceolate, acute, ciliate; sepals imbricate, seri-

ceous-pilose and ciliate, 1.5 mm. long and broad; petals thick-coria-

ceous, rufescent, ovate-lanceolate, sericeous without, puberulent with-

in, 4-5 mm. long, 2.5-3.5 mm. wide; disk urceolate, fleshy, 10-plicate,

ovary strongly depressed, tuberculate; style exserted 2 mm., stigma

capitate; capsules ligneous, 5-celled, the carpels medially dehiscent,

longer than 2 cm., less than 1.5 cm. across, with pyramidal tubercles

2-3 mm. long. — The Peruvian specimen was given (in herb.) an

untenable species name by Standley and may be distinct from

Martius' plant of central Brazil; recent collections from there

however seem to be intermediate, so perhaps it is feasible to treat

the following as var. peruviana Macbr., var. nov. foliis ad 15 cm.

longis, 6.5 cm. latis; capsulis ubique tuberculis ad 5 mm. longis

dense obtectis. — A 5 meter tree with cream-green flowers (Klug).

F.M. Neg. 19225.

San Martin: Zepelacio near Moyobamba, Klug 3722 (type, var.

peruviana}. Brazil.

Esenbeckia pilocarpoides HBK. Nov. Gen. & Sp. 7: 248.

pi 655. 1825; 144.

Typically glabrous except the puberulent-hirtellous inflorescence,

and the leaves simple, but variants are pilose or the leaves trifoliate;

petioles 1.5-3 cm. long, semiterete, complanate and narrowly winged

or margined above; leaflets membranous, pellucid-punctate, lustrous

both sides, scarcely paler but more prominently nerved beneath,

ovate-oblong or elliptic-oblong, rather obtusely acuminate (acumen

5-10 mm. long, rounded or acute), and articulate, with petiole 1-1.5

dm. long, 4-5.5 cm. wide; flowering branches terminal and axillary,

as long as the leaves, the short branches and pedicels puberulent;

calyces and petals red-glandular, the ovate-rotund sepals ciliate,

strongly imbricate, 1.5 mm. long, 2 mm. wide; petals white-green,

elliptic-subrotund, 2.5-3 mm. long and broad; disk deeply 5-plicate;

stamens 1.5 mm. long; style very short, clavate-capitate; ovary

depressed, fleshy-tuberculate.— After Engler as other descriptions.

He included E. maurioides Mart. Nov. Gen. & Sp. 3: 82. pi. 232.

674 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

1829 as var. maurioides Mart, ex Engl., I.e. from central Brazil,

with upper leaves trifoliate, the leaflets smaller; similar and equally

with small flowers is E. alata (Karst. & Triana) Triana & Planch.,

145, leaflets more often trifoliate, thus approaching to E. venulosa;

the var. guianensis Engl. has unifoliate leaves densely white pilose

beneath as also the flowering branchlets, pedicels and calyces. The

following (flowers gray-green and dark violet [Klug]) specimens are

scarcely adequate for certain determination but apparently belong

here; the type was from Venezuela. F.M. Neg. 36820.

San Martin: Zepelacio, 4 meter shrub, Klug 275; 3380. To

Trinidad.

Esenbeckia venulosa Macbr. Candollea 5: 376. 1934.

Glabrous with terete branchlets and mostly trifoliate subequal

numerous leaves on subangulate petioles 3-5 cm. long, somewhat

complanate above; leaflets unequal, oblong-elliptic or sometimes

elliptic-obovate, rounded-obtuse or retuse, the lateral obliquely

cuneate to the scarcely petiolulate base, to 7 cm. long, 3 cm. wide,

the larger intermediate distinctly petioled and cuneate at base, to

1 dm. long, half as wide, all chartaceous-coriaceous, but pellucid-

punctate, opaque, conspicuously reticulate, especially beneath;

panicles subracemose, obscurely and sparsely puberulent, 1.5-3 cm.

long; pedicels very short, calyx segments rounded, minutely ciliate,

nearly 1 mm. long; petals narrowly ovate-oblong, 5 mm. long, ob-

scurely pulverulent within, glabrous but punctate without; stamens

about 1.5 mm. long; anthers large; disk fleshy, 4-plicate, including

the ovary, and crowned by the capitellate style. — Relationship not

determined but it may well prove to be a glabrous state of E. Wars-

zewiczii, the flowers of which were not known.

Junin: Rio Mantaro, 2,500 meters, Weberbauer 6584, type.

Esenbeckia Warszewiczii Engl. in Mart. Fl. Bras. 12, pt. 2:

148. 1874.

Branchlets terete, 4-5 mm. thick, shortly white sericeous-pilose,

densely leafy; leaves subcoriaceous, subglabrous above, sparsely

pilose beneath except the white sericeous midnerve, green, very

densely glandular-punctate, 3-foliate; petioles terete, 4-5 cm. long,

half as long as the intermediate leaflet, this nearly 1 dm. long, 4-4.5

cm. wide, little larger than the oblique lateral (7-8 cm.), oblong-

elliptic, obtusish; panicles racemiform, 3-10 cm. long, rachis and

short branchlets densely pilose, pedicels obsolete; sepals broadly

FLORA OF PERU 675

ovate, subobtuse, pilose below, glabrate above or faintly ciliolate,

about 1 mm. long; petals suboblong, ashy pulverulent within,

sparsely and minutely setulose medially without, 5 mm. long;

stamens about 2 mm. long; disk fleshy, style capitellate; carpels

rugose, truncate above with a dorsal apical horn, dehiscing only

above and ventrally, obtuse at base, 1 cm. long and broad. — Seems

to be allied to E. pumila Pohl of Brazil (Engler) ; but cf . E. cornuta

from the same locality, i.e., northern Peru. E. Dielsiana G. M.

Schulze, Bibl. Bot. 116: 100. 1937, of central Ecuador is less pilose,

panicles merely puberulent, pedicels about 1.5 mm. long, petals

dark red, ovate, acute, 5-6 mm. long, carpels ecornute about 18 mm.

high. A 3 meter shrub, the very hard wood used for canes (Weber-

bauer). Flowers described from Weberbauer 7645. F.M. Neg. 12519.

Cajamarca: San Felipe, 2,000 meters, Weberbauer 7110. Sonda,

Warszewicz, type. — Tumbez: Mountains east of Hacienda Chicama,

Weberbauer 7645. "Angohuara" or "ongahuara" (Weberbauer).

7. PILOCARPUS Vahl

Small trees or shrubs with opposite, alternate or whorled 1-3-

foliate or imparipinnate coriaceous or membranous pellucid-punctate

leaves, crowded at the tips of the branchlets. Flowers small, in

elongate axillary or terminal spikes or racemes, the pedicels, when

present, bracteate as the calyx at base, this with 4-5 teeth or sub-

entire, the 4-5 petals coriaceous, minute. Stamens free, 4 or 5,

below the annulate disk, with glabrous subulate filaments and

versatile ovate anthers. Ovary depressed-globose, deeply 4-5-lobed;

style short, stigma capitate, 4-5-lobed. Carpels 1-5, nearly separate,

lunate, loculicidally bivalved, 1-seeded, more or less ribbed.

Pilocarpus spicatus St. Hil. ex DC. Prodr. 1: 728. 1824: 133.

Glabrous except the spiciform racemes these early pilose-puberu-

lent, becoming glabrate, 3 dm. or so long; branching dichotomous,

the branchlets 3-4 mm. thick; petioles alternate, typically 1-2 cm.

long, lightly canaliculate; leaves simple, membranous, pale green,

reddish glandular pellucid, oblong-elliptic, typically obtuse, acute

at base, revolute, midnerve beneath and lateral nerves as reticulate

veins prominent both sides; pedicels 2-4 mm. long; calyx lobes

obtuse, ciliolate; petals greenish, 2 mm. long; carpels mostly 5,

compressed, dorsally and ventrally carinate, reddish glandular, 7 mm.

long and broad. — Williams 4878 in fruit seems to be the same as

Ule 9500 in flower and if so they probably represent a distinct

676 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

species; at present they may be designated var. peruvianus Macbr.,

var. nov. petiolis circa 5 mm. longis, foliis breviter acuminatis,

carpellis (Williams spec.) circa 10 mm. longis et latis. A 2-8 meter

shrub with yellowish-green flowers (Ule). Illustrated, Nees & Mart.

Nov. Act. Acad. Leop.-Carol. 11. pi. 30.

Loreto: Santa Rosa, Williams 4878 (apparently, in fruit). — Rio

Acre: Seringal San Francisco, Ule 9500 (det. Krause, P. spicatus;

type, var. peruvianus). Brazil. "Sapote yaru" (Williams).

8. ADISGANTHUS Ducke

A shrub or small tree with short-petioled obovate-lanceolate

leaves narrowed cuneately at base and long-peduncled 2-3 times

dichotomous few-flowered inflorescences of brownish-red or white

flowers, the calyx 5-toothed, the petals 5, nearly free, the 5 stamens

completely so with linear-lanceolate anthers about as long as the

filaments. Disk lacking. Style 1, stigma obscure. Carpels 1-5,

united to a little above the middle, keeled both sides, 1-seeded, trans-

versely rugulose; endocarp falling with seed. — Apparently not

generically distinct from Euxylophora Huber, Bol. Mus. Goeldi 10:

84. 1910, with obovate leaves, inflorescence 3-4 times dichotomous,

linear anthers longer than filaments ;E. paraensis Huber, fide Ducke,

Archiv. Jard. Bot. Rio Jan. 3: 183, attains 40 meters and yields

good timber known as "Pao Amarello."

Adiscanthus fusciflorus Ducke, Archiv. Jard. Bot. Rio Jan.

3: 187. 1922.

Completely glabrous except petals within, the unifoliate leaves

crowded toward tip of the rather stout striate branchlets, articulate

with the stem, subsessile or the petiolar base to 4 cm. long; blades

obovate-lanceolate, abruptly acuminate, the larger 3.5-5 dm. long,

8-10 cm. wide, alternate, with smaller 2-3 dm. long, 5-7 cm. wide,

firm-membranous, paler and with prominent midrib beneath the

slender nearly horizontal parallel secondary nerves prominent both

sides, as the fine reticulate veins; floral branches 3-3.5 dm. long,

often branched, the flowers few in short cincinni; pedicels strongly

rugose, to about 7 mm. long, elongate in fruit, to 1.5 cm. long;

calyx 2 mm. long, 4 mm. wide; petals about 1.5 cm. long and densely

bearded within to above the middle with rusty trichomes; carpels

about 1 cm. long, reddish brown, the endocarp yellow. — Flowers

brown-red (Ducke); white (Klug). Illustrated, Ducke, I.e. 4: pi. 8

(flowers and fruit). F.M. Neg. 12347.

Loreto: Mishuyacu near Iquitos, Klug 723. Brazil.

FLORA OF PERU 677

9. LEPTOTHYRSA Hook f.

A weak or straggling simple-trunked little tree with whitish

bark, the large obovate-lanceolate nearly sessile epunctate leaves

alternate and crowded at the tip. Flowers large, white, subracemose

on elongate slender axillary peduncles. Calyx cupulate, obscurely

4-toothed. Petals medially connate, lanceolate, spreading, valvate

above. Stamens 4 at base of the entire disk, the filaments filiform,

glabrous, free, exceeded by the nearly linear anthers. Ovary de-

pressed, 4-celled, with 1 short connate style, 4-lobed stigma. Carpels

1 or 2, bivalved, 1-seeded, coriaceous, transversely rugose.

Leptothyrsa Sprucei Hook. f. in Benth. & Hook. f. Gen. PI.

1: 284. 1862; 131.

Glabrous; leaves subcoriaceous, drying paler and brown punctate

beneath with many conspicuous parallel secondary nerves that are

mostly about 1 cm. apart and join at or very near the margins, 3-7

dm. long, only about 5 mm. wide at the cuneately narrowed base

but broadened to 5-13 cm. below the abruptly acuminate apex, the

acumen itself 1-2 cm. long, 3-4 mm. wide; racemes 3-4 dm. long,

floriferous above the middle with 3-5-flowered 2-5 mm. long branch-

lets; pedicels 3-4 mm. long; calyx 1.5 mm. long; carpels rhomboid-

conchiform, pale brown, 1 cm. in diameter. — One to 2 meters high,

aspect of Erythrochiton brasiliense, the white flowers to 18 mm. long;

illustration in Fl. Bras, evidently from young flowers (Ducke).

Illustrated, Mart. Fl. Bras. 12, pt. 2: pi. 28. F.M. Neg. 12473.

Loreto: Near Iquitos, Killip & Smith 29844; Klug 147; (Ducke).

Upper Mouth of Rio Santiago, Tessmann 3526. Amazonian Brazil.

10. TICOREA Aublet

Shrubs, the younger branches somewhat sericeous with golden

yellow trichomes, the leaves long-petioled with lance-elliptic long-

acuminate leaflets, the flowers large, yellowish green or white and

many borne at the tips of the branchlets in narrow elongate panicles,

the pedicels bracteate. Calyx short with 5 teeth. Corolla with

long tube, the 5 lobes valvate, similar in size. Disk urceolate.

Stamens 5, united nearly completely with the corolla tube, all fertile,

the anther connective with short bilobed appendage at base. Ovary

5-lobed, 5-celled with 2 superposed ovules in each cell; style filiform

with thicker obscurely lobed stigma. Carpels connate at base,

laterally compressed, the endocarp bivalved. — Nearly Galipea unless

the lack of sterile stamens is distinctive.

678 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Ticorea longiflora DC. Me'm. Mus. Hist. Nat. Paris 9: 146. pi

9. 1822; 92.

Slender branchlets and very membranous leaves soon glabrous

except the midnerve and the pilose or ciliate margins, the evanescent

pubescence yellow or golden sericeous; glands numerous, mostly

small; petioles about 1 cm. long, a little pilose, lateral petiolules

3-4 mm. long with leaflets oblong-lanceolate, oblique at base, the

intermediate petiolule 1.5 cm. long with elliptic leaflet 1.5 to nearly

2 cm. long, much attenuate both ends, the nervation prominent

both sides, reticulate; flowering branchlets pubescent, about as long

as leaves, pedicels as long as calyces, these cupulate, pilose, with

acute lobes 1 mm. long; corolla appressed yellow sericeous, tube

narrowly infundibuliform 4-5 cm. long, 2 mm. thick, lobes 7-10 mm.

long, 2 mm. wide; anthers exserted from throat, appendage 2 mm.

long, bilobed; ovary glabrous, disk membranous. — Perhaps is T. foe-

tida Aublet? Shrub or small tree to 3 meters, often in argillaceous

terrain, the foliage with unpleasant odor but the white flowers

perfumed (Ducke).

Loreto: Puerto Mele*ndez, Tessmann 4806. To the Guianas.

"Agapurana," "acapu-rana," "cua-acapoc."

11. ERYTHROCHITON Nees & Mart.

Single stemmed or nearly unbranched shrub or small tree with

the long spathulate simple or foliate leaves and showy red and white

racemose flowers crowded at the summit of the slender trunk.

Flowers hypophyllous or peduncled, the peduncles leafy or usually

elongate and angled. Calyx large, tubular, typically scarlet, angled,

subbilabiate with 3-5 cleft lobes exceeded by the subequal spreading

white or roseate corolla lobes. Disk urceolate or annulate. Stamens

5, all perfect or 1-3 anantherous, the filaments affixed to tube.

Ovary 5-celled with exserted 5-lobed stigma. Capsule with 5 2-valved

scarcely cohering carpels, each with 2 seeds. — Name derived from

Greek for red tunic, the usually colored calyx enclosing the corolla

tube.

Leaves simple.

Leaves 4-5 dm. long or longer E. brasiliense.

Leaves at most 2.5 dm. long E. macropodum.

Leaves trifoliate E. trifoliatum.

Erythrochiton brasiliense Nees & Mart. Nov. Act. Acad.

Leop.-Carol. 11: 170. pi. 25. 1823; 107.

FLORA OF PERU 679

Simple stemmed shrub or tree 0.5-5 meters high, with dark bark

and 1-foliate horizontally spreading membranous pale green lustrous

obovate-lanceolate acutely acuminate leaves that are very long

cuneate-attenuate to the short petiolar base, 4-5 dm. long or longer

(acumen 1.5-2 cm. long), a dm. or so wide at the broadest part,

the lateral nerves prominently reticulate both sides; flower branches

equaling the leaves, trigonous, ebracted below but with lanceolate

acute pubescent bracts at the base of the glabrous pedicels, these

1-1.5 cm. long, the short floriferous branchlets of 3-8 flowers scar-

roughened in age; calyx scarlet, 2.5 cm. long, 1 cm. in diameter,

the ovate acute divisions 4 mm. wide enclosing the 5-celled capsule;

corolla snow white, tube 3 cm. long, 2.5-3 mm. in diameter, sub-

equaling the calyx segments, viscid; ovary scarcely 2 mm. long,

style glabrous; carpels barely coherent, lustrous, crustaceous, 1.5

cm. long. — A handsome plant sometimes cultivated for ornament.

Has a remarkable distribution: tropical middle Brazil, subandean

eastern Peru, and, rarely, north of the lower Amazon (Ducke);

Pilocarpus may be a similar example. Illustrated, Mart. Fl. Bras.

12, pt. 2: pi. 22.

Huanuco: Wet rocky open place, Pozuzo, 4606. — San Martin:

Near Moyobamba, King 3668. Tarapoto, Spruce 3917, Vie 6611;

Williams 3761. — Junin: Colonia Perene", Killip & Smith 25153.

Brazil.

Erythrochiton macropodum Krause, Notizbl. Bot. Gart.

Berlin 6: 146. 1914.

Younger terete branchlets, petioles and leaves minutely pilose,

soon glabrous; petioles slender, a little applanate above, subgenicu-

lately enlarged at apex, 3-6 cm. long; leaves simple, membranous,

nitidulous, narrowly oblong or oblanceolate-oblong, acutely acumi-

nate (acumen 12-18 mm. long), sometimes slightly oblique toward

the cuneately narrowed base, 1.5 to nearly 2.5 dm. long, 6-7.5 cm.

wide, lateral nerves 12-14, prominent beneath; flowers few, shortly

pediceled in a terminal pseudo-raceme; peduncles nearly 1 dm. long,

pilosulous, pedicels slender, pilose, 6-10 mm. long, the basal bractlets

pubescent; calyx divisions white, unequal, acuminate, glabrous,

1.5-2.2 cm. long, 5-7 mm. wide, glandular punctate; corolla white

or drying yellowish, with curved cylindrical tube, about 3.5 cm.

long, densely appressed sericeous-pilose lobes oblong, obtuse; disk

truncate; style slender. — Erect shrub to 1 meter high. Nearest

E. trifoliatum Pilger but leaves entirely simple; distinguished from

680 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

E. brasiliense Nees & Mart, by the smaller longer petioled leaves,

from E. hypophyllanthus Planch. & Lind. by the inflorescence

(Krause). F.M. Neg. 12476.

Rio Acre: In woods at Porto Carlos, Ule 9498, type.

Erythrochiton trifoliatum Pilger, Verh. Bot. Ver. Brandenb.

47: 153. 1905.

Shrub 1-2 meters high; petioles 7-15 cm. long, with 3 elliptic

usually somewhat unequal leaflets, shortly narrowed or subrotund

at base, shortly and obtusely subcaudate, thin-coriaceous, 1-2 dm.

long, 3-7 cm. wide, lateral nerves nearly straight, anastomosing

toward margin and prominently reticulate with the veins both sides;

flowers at tip of the stout ebracteate puberulent 10-14 cm. long

peduncles in dense terminal panicles 5-6 cm. long, the short pedicels

as the white calyx minutely villous, slightly exceeding the flowers

in bud, the calyx segments free, membranous, broadly lanceolate,

about 2 cm. long; petals at first scarcely exserted, finally nearly 3 cm.

long, villous, connate to the middle or so, lobes obtuse; stamens 5,

2 or 3 fertile, filaments connate to tube nearly to apex; disk annulate,

upper margin irregularly sinuate; ovules 2, style 8 mm. long, stigma

rotund; carpels 5, subalate dorsally, the calyx divisions nearly as

long. — Remarkable in the distinct calyx lobes; differs from E.

Lindenii also by the trifoliate leaves (Pilger). The Klug specimen

from a 4 meter tree, the fully developed flowers to 3 cm. long, the

carpels with dorsal wings 5 mm. high, 6 mm. wide at base. F.M.

Neg. 12479.

San Martin: Pongo de Cainarachi, Klug 2729. — Ayacucho: Rio

Apurimac, near Kimpitiriki, Killip & Smith 23408. — Loreto : Woods

at Cumbaso, 700 meters, Ule 6715, type; also 6863.

12. GALIPEA Aublet

Trees or shrubs with bright green thin simple or trifoliate leaves

on terete or winged petioles mostly crowded at the tips of the gla-

brous branchlets. Flowers perfect, rather large, in axillary and

terminal cymose or panicled racemes. Calyx small with 5 erect

acute segments alternate to the united petals, these free only at

the middle or above. Stamens 5-8, adnate to the corolla tube, the

3-6 sterile sometimes gland tipped, the anther connective1 basally

appendaged. Disk cupulate, enclosing the 4-5-lobed ovary; style

long with 3-5-lobed stigma; ovules 2 in each cell, superposed. Carpels

united at base and tip, keeled, 1-seeded. — Named for the Galibis,

FLORA OF PERU 681

a people of French Guiana. Some species of the related genus

Raputia Aublet may yet be found within Peru; the known species

have flowers sericeous without, tomentose within, filaments attached

below the narrow medially connate petals, carpels laterally joined

each with 2 seeds. The red-flowered Naudinia amabilis Planch. &

Lind. of Colombia would also be sought here but is essentially a

Cusparia except for its long corolla tube.

Calyx lobes 6-8 mm. long; flowers about 4 cm. long; petioles elongate.

G. longiflora.

Calyx lobes almost minute; flowers about 2 cm. long; petioles 2 cm.

long G. trifoliata.

Galipea longiflora Krause, Notizbl. Bot. Gart. Berlin 6: 144.

1914.

Branches and branchlets terete, the younger as the petioles

densely pubescent becoming glabrate and striate; petioles slightly

complanate above, enlarged at base, 12-16 cm. long; leaflets 3,

sessile, early pilose beneath on the there prominent 12-14 lateral

nerves, glabrous and lustrous in age, ovate-oblanceolate or -lanceo-

late, narrowed to base, acutely and mostly a little obliquely acumi-

nate (acumen 3-4 cm. long), entire, the middle one 2.5-3 dm. long,

8-11 cm. wide, lateral somewhat smaller; flowers shortly pedicelled

or subsessile in long axillary many-flowered panicles, rachis strongly

complanate, angulate, long-sulcate to tip, glab*rate, 2.4 dm. long;

calyx angled, narrowly lanceolate lobes 6-8 mm. long; corolla white,

slender tube 3.5-4.2 cm. long, subsericeous-pilose, glabrous within,

oblong subacute lobes 3-4 times shorter; upper 2 anthers linear-

oblong, lower 3 linear with apical globose glands; disk annulate;

style barely exserted. — Sometimes arborescent, 5-15 meters high,

the white flowers extra large (Krause). G. bracteata (Nees & Mart.)

Engl., 99, of eastern Brazil, has deeply lobed calyx but lobes only

2.5-3 mm. long; Amazonian species to be expected include G. tri-

foliata Aublet, 96, and G. grandifolia Engl., 98, both with very short

calyx teeth; the simple leaves of the latter attain 2.5 dm., the leaflets

of the former scarcely 1 dm.

Rio Acre: Ule 9497, type.

Galipea trifoliata Aublet, PI. Guian. 2: 662. pi. 269. 1775; 96.

Branches terete, slender, the strict densely leafy branchlets

purpurescent; petioles 2 cm. long; leaflets unequal, the intermediate

one largest, 7-9. (12) cm. long, 2-3 (4) cm. wide, cuneate or arcuately

682 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

attenuate to base, the lateral more or less oblique, all oblong-elliptic

with short obtuse and deeply emarginate acumen, membranous,

early finely pilose becoming glabrous, nitidulous, the medial petiolule

canaliculate and margined; panicles 4-5 cm. long, the 1-2 cm. long

branchlets mostly 3-flowered, the pedicels 3-4 cm. long; calyx 1.5-2

mm. long, cupulate, sparsely pilose with short obtusish teeth; corolla

ashy pubescent, the tube 1.5 cm. long, 1 mm. in diameter, the lobes

5 mm. long, 1.5 mm. wide at base; carpels 5, oblong, obtuse, trans-

versely sulcate, obtusely carinate dorsally, 1 cm. long, 7 mm. wide.—

The Ule collection from a tree 5-15 meters tall with white flowers.

Rio Acre: In woods near Porto Carlos, Ule 9499 (det. Krause).

Amazonian Brazil and Bahia to Guiana.

13. RAUIA Nees & Mart.

Resembles Galipea but the rather small flowers with linear pubes-

cent petals connate only at base, nearly valvate. Stamens 4 or 5

and only lightly affixed to petals, the connective without appendage.

Known species with simple leaves. — Probably belongs, as Bentham

and Hooker thought, to Galipea; the name is for A. Rau, a con-

temporary professor of natural history.

Rauia resinosa Nees & Mart. Nov. Act. Acad. Leop.-Carol. 11:

169. 1823; 94.

Glabrous except the floriferous branchlets, the older branches

slender, terete, the younger foliose ones angled as those of the white

puberulent corymbs; petioles 1-3 cm. long, glandular; leaves lustrous

green or later metallic, membranous, pellucid punctate, oblong-

elliptic, long attenuate-acuminate (acumen 1-1.5 cm. long, 3-4 mm.

wide), mostly 6-8 cm. long, 1.5-2 cm. wide, the pale midnerve and

laterals prominent with the reticulate venation especially beneath;

buds angled, incurved at tip; calyx black glandular, 5-angled, 3 mm.

high; petals 10-13 mm. long, scarcely 1 mm. wide, spreading or

contorted, without minutely, within long white-pilose and biseriately

glandular; carpels finally separating, 8 mm. long, 6 mm. across,

ventrally and laterally carinate, transversely deeply sulcate and

resinous punctate; seeds obovoid, tuberculate.— The small long-

campanulate flowers are in open inflorescences, their primary

branches 3-4 cm. long the ultimate 3-5-flowered; leaves thin; a low

(1-2 dm.) forest shrub with half decumbent stems (Tessmann).

Illustrated, Mart. Fl. Bras. 12, pt. 2: pi. 16.

Loreto: Mouth of the Rio Santiago, Tessmann 3989. Brazil.

FLORA OF PERU 683

Rauia Ulei Krause, Notizbl. Bot. Gart. Berlin 6: 145. 1914.

Younger branchlets, petioles and leaves beneath puberulent,

finally glabrous; petioles 3-4.5 cm. long; leaves thin-herbaceous (or

coriaceous?), nitidulous, oblanceolate-oblong, slenderly acuminate

(acumen 12-16 mm. long), long cuneate-narrowed to base, 1.5-2 dm.

long or longer, 6 cm. wide or wider, lateral nerves 13-15, slender,

prominent beneath; flowers in terminal erect pseudo-panicle, rachis

little complanate, puberulent, 12-18 cm. long; calyx campanulate,

sparsely pilose, 2.5-3 mm. long, divided to the middle or lower into

5 broadly ovate obtuse lobes; linear-lanceolate acute densely pubes-

cent lobes of the corolla 12-15 mm. long, connate below into a short

cylindric tube; upper 2 stamens fertile, shorter, lower 3 sterile, longer,

their pubescent filaments a little shorter than corolla, filaments

fertile, stamens subulate; disk shortly denticulate; ovary about 2.5

mm. long, with short subcapitate style. — Shrub or arborescent 2-12

meters high. Differs from R. resinosa Mart. & Nees by the longer

petioled larger leaves, different inflorescence (Krause). The leaves

dry coriaceous, black above, brown beneath, the flowers in narrow

panicles.

Rio Acre: Monte" Mo, Ule 9495, type.

14. CUSPARIA Humb.

Usually simple stemmed shrubs or trees with fleshy or leathery

often long-petioled leaves crowded near the tips of the stems, the

leaves simple (leaflets reduced to 1) or composed of 2-7 leaflets,

these rather large and elongate, sometimes oval, with heavy midrib

and ordinarily acute or acuminate. Flowers except rarely moderately

large, pedicellate, commonly crowded, infrequently racemose, axillary

or apparently extra-axillary, more or less irregular and with small

sometimes bilabiate calyx. Petals linear to long-spathulate, shortly

united toward base; stamens 5 (now and then 4-8), attached medially

to the petals, all fertile or only 2 or 3 with anthers, these linear and

with flattened more or less barbate filaments. Disk about cupulate,

5-dentate and more or less including the ovary. Fruit parts 1-3 and

free from each other from the first, each part 1-seeded. — Could be

united as by Bentham and Hooker with Galipea, in which they also

included Rauia, which would be more practical and probably more

natural taxonomy. The publication of the name, derived from the

native "Cuspare," dates from 1822 when validated by De Candolle,

Me"m. Mus. Hist. Nat. Paris.

684 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Among fruiting specimens not determined but possibly belonging

here mention may be made of Williams 3727 (Killip & Smith 27150}

from Iquitos, fruits rather tardily separate so maybe a Pilocarpus

but leaves 5-foliate.

Leaves 5-7-foliate C. toxicaria.

Leaves simple.

Leaves obtuse or barely acute C. ucayalina.

Leaves sharply acuminate C. acuminata.

Cusparia acuminata Pilger, Verh. Bot. Ver. Brandenb. 47:

154. 1905.

A shrub of 3-9 meters, glabrous except the spiciform inflorescence,

the membranous elliptic or somewhat obovate leaves long-cuneate

to base, obtusely long-acuminate, 2-3 dm. long, 7-11.5 cm. wide,

the nerves and reticulate veins prominent beneath; petioles 12-55

mm. long; rachis stout, puberulent-pilose, 6 cm. long; flowers white,

the cupulate dentate-incised calyces shortly tomentulose-pilose;

petals 5, lightly coherent below, obtuse, villous-tomentose both

sides, 7 mm. long; stamens 5, two fertile with filaments villous above,

2.5 mm. long, anthers to 2 mm. long, connective extended 0.75 mm.

obtusely; staminodia 3; disk annulate, truncate, scarcely more than

1 mm. high; ovary 5-celled, sparsely pilose apically, style short,

stigma little enlarged.— F.M. Neg. 12499.

Loreto: Mouth of Rio Santiago, Tessmann 4077. Brazil.

Cusparia toxicaria [Spruce] Engl. in Mart. Fl. Bras. 12, pt. 2:

114. pi. 24. 1874.

Branches a dm. or so in diameter, the younger densely red-

tomentose; petioles longitudinally sulcate, 3-4.5 dm., 3-5 mm. thick

below but apically much enlarged, early red-pilose as the stout

peduncles, pedicels and calyces; leaves membranous, green, lustrous,

with 5-7 often more or less unequal broadly lanceolate long- and

acutely-acuminate leaflets, the medial finally 3-4 dm. long, 10-13

cm. wide, with acumen 2-3 cm. long, petiolule 2-2.5 cm. long;

midnerve and reticulate venation with lateral nerves prominent;

flowers white, essentially racemose, solitary or in 2's or 3's, the

rachis 1-3 dm. long; pedicels 1-2 mm. long; calyx cupulate, the

ovate-lanceolate lobes equaling the tube, 2 mm. long, persisting and

spreading in fruit; petals 5, 1.5 cm. long, 2.5 mm. wide; filaments

1 cm. long, pilose and within, below the anthers, barbate; ovary

glabrate, the elongate style glabrous; carpels black, lustrous, carinate

FLORA OF PERU 685

ventrally, about 13 mm. long. — I am not certain that the Rio Acre

collection is correctly identified. F.M. Neg. 19222.

Rio Acre: Xapury, Ule 9496 (det. Krause). Brazil.

Gusparia ucayalina Huber, Bol. Mus. Goeldi 4: 573. 1906.

Type a low shrub with few if any branches; petioles 3 cm. long,

canaliculate above, enlarged at base and apex, ferruginous tomen-

tulose as the ultimate branchlets of the narrow panicles; leaves

oblong, cuneate at base, rounded-obtuse or acutish, 2-2.5 dm. long,

7-9 cm. wide, glabrous, except nerves beneath minutely strigillose,

subcoriaceous, drying more or less brownish; peduncles about 5 cm.

long, pseudoterminal; flowers in a narrow but obvious panicle, the

branchlets about 1 cm. long; pedicels 3 mm. long; calyx 5-parted

medially or more deeply, the rotund-ovate obtuse or apiculate lobes

gray-puberulent without; petals white, connate only at base, spathu-

late, acutish, 6 mm. long, marginally toward the base whitish

tomentulose like the 3 thick ligulate sterile stamens, the filaments of

the 2 fertile conglutinate ones puberulent; ovary glabrous, exceeded

by the disk, the style two or three times longer, stigma oblong-clavate,

5-sulcate. — Similar in inflorescence to C. paniculata Engl., 120, but

resembles C. macrophylla, 115, C. cuneifolia, 116, C. Gaudichaudiana,

116, etc., more in ovary and style (Huber). There is an unnumbered

negative of the type at Chicago Museum, accompanied by a scrap.

Illustrated, Huber, I.e. 574 (flowers).

Loreto: Pampa del Sacramento between Sarayacu and Santa

Catalina (Huber 1513, type).

15. MONNIERIAL.

Finely subappressed hispidulous and somewhat glandular hirsute

branching annual (or sometimes suffrutescent) with soft petioled

3-foliate leaves, the leaflets lance-elliptic, and small bilabiate flowers

in bifurcate somewhat scorpioid cymes. Sepals 5, very unequal, the

outermost foliate and bract-like, the next one about half as long,

the inner minute or obsolete. Stamens 5, attached to corolla tube, the

2 fertile with bearded filaments, the sterile bearded medially. Disk

overtopping ovary, the 5 carpels free except at the short style.—

Genus suggests in aspect especially in herbaria some Euphorbiaceae.

Monnieria trifolia L. Sp. PI. ed. 10. 1153. 1759.

Low and herbaceous, again definitely woody except the upper-

most branchlets and then as much as 1 meter high; petioles to about

686 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

3 cm. long; leaflets elliptic to oblong-elliptic, shortly acute both ends

or obtusish at the shortly narrowed tip, or, especially the middle

one, attenuate to the more or less elongate petiolule, to about 5.5 cm.

long, 2 cm. wide, pale green, soft membranous; flowers white, the

forked inflorescence with long peduncle to about 4.5 cm. long, the

dense flowering branches 1-2 cm. long appearing closely bracted

by the foliate outer sepals, these 5 mm. long or longer; carpels about

2 mm. long. — The sharp aromatic roots have been used in medicine.

Illustrated, Pflanzenfam. ed. 2. 19a: 292. 1931.

San Martin: Chazuta, Klug 4024- — Loreto: La Victoria, Williams

2612. Caballo-Cocha, Williams 2030; 2096; 2135. Widely distrib-

uted, Brazil to Mexico.

16. RAVENIAVell.

Peruvian species a foetid tree with large simple leaves and

terminal erect few-flowered inflorescences of glandular green some-

what bilabiate funnelform flowers. Calyx lobes very unequal,

strongly imbricate. Petals medially connate. Fertile stamens 2

adnate to the tube below the upper lip. Disk cupulate. Carpels 5.

Ravenia biramosa Ducke, Archiv. Inst. Biol. Veg. Rio Jan. 2:

48. 1935.

Small foetid tree, glabrous except the sparsely and minutely

pubescent inflorescence, the flowering portion of this typically bi-

parted, the racemiform branches finally to. 12 cm. long, alternately

subdistichiflorous; petioles 8-15 mm. long, deeply canaliculate above;

leaves obovate- rarely oblanceolate-oblong, narrowed to petiole,

long and subabruptly acuminate, 13-28 cm. long, 5-10.5 cm. wide,

membranous, little lustrous, subconcolored, distinctly pellucid punc-

tate, irregularly pinnate-nerved and reticulate, more prominently

beneath than above; inflorescence terminal, erect, long-peduncled

(peduncle 8-15 cm. long); bracts caducous, linear, gland-tipped, to

6 mm. long, pilose; pedicels 5-9 mm. long, enlarged above; flowers

green, densely glandular; sepals strongly imbricate, elliptic-orbicular,

coriaceous, scariose-margined, the much larger outer 2 strongly

convex, about 14 mm. long, glabrous or sparsely pilose, sericeous

within; corolla at an thesis about 22 mm. long, medially connate

with wide tube the throat densely pilose within, bilabiate, the lower

lip of 4 obtuse parts the 2 inner of these longer and less deeply

divided than the 2 outer, the upper lip 1 linear-cuneate longitudinally

plicate concave petal; fertile stamens 2, adnate to the tube below

FLORA OF PERU 687

the upper lip, anthers glabrous, connective basally auriculate;

staminodia 3, densely lanate as stamens at base; disk cupulate,

lightly 5-crenulate; carpels 5, fruit unknown. — R. pseudalterna Ducke,

I.e. 49, has the apparently alternate leaves opposite a stipuliform

appendage or reduced caducous leaf 6-15 mm. long, flowers usually

3-5, composed of 2-flowered cincinni. According to Ducke these

two species have the most strongly zygomorphic corolla with bilabi-

ate limb; intermediate in this character are R. amazonica Huber, Bol.

Mus. Goeldi 5: 423. 1909, with smaller well-petioled leaves, and

R. polygalaecalyx Ducke, Archiv. Jard. Bot. Rio Jan. 4: 101. 1925,

the calyx polygaloid, the leaf-axils barbate within, while the original

species of eastern Brazil, R. infelix Veil., has weakly zygomorphic

flowers, not bilabiate. — Ducke observes that the Amazonian species

differ from it further in their entirely green flowers but according to

the collector of the 3-5 meter shrub or liana of Peru the flowers were

green and white; these have become broken but appear zygomorphic

and bilabiate; vegetatively the specimens agree in general with the

diagnosis of R. biramosa. Apparently Killip & Smith 28396, with

fruits about 1.5 cm. long, long, 2.5 cm. broad, belong here but are

generically aberrant for each cell contains 2 seeds; notwithstanding

that the development of 1 or 2 seeds is traditionally used to key out

genera related here its importance is probably not basic since 2

ovules in any case may be present, sometimes at least. Standley

gave an herbarium specific name (perhaps rightly) to these collec-

tions, so imperfect, which I prefer to record as R. biramosa Ducke,

var. peru viana Macbr., var. nov. in axillis petiolorum intus parcis-

sime flavidobarbatis; foliis 12-30 cm. longis, 7-13 cm. latis, late

oblongo-ellipticis, vix vel haud obovatis; floribus 3-9 (paullo notatis)

simpliciter racemosis vel etiam racemis breviter biramosis, pedunculis

solum circa 3-4 cm. longis. — Klug 3047, type.

Loreto: Mishuyacu near Iquitos, Klug 1316. Balsapuerto, Klug

2951 ; 3047; Killip & Smith 28396. Amazonian Brazil.

17. RUTAL.

Disagreeably pungent herbs sometimes suffrutescent, with terete

branching stems, simple or compound leaves and greenish-yellow

flowers in terminal leafy bracted corymbs, panicles or cymes. Calyx

with 4-5 segments that persist and the same number of petals, these

often dentate or ciliate. Torus thick, urceolate, 8-10-glandular or

-foveolate with this number of stamens at its base, the alternate

shorter. Ovary sessile, 4-5-lobed; ovules pendulous from central

688 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

axis. Capsule 4-5-lobed, the lobes, if dehiscent, at apex, and with

many angled albuminous seeds.

Ruta chalepensis L. Mant. 69. 1767.

Unpleasantly redolent glaucous perennial herb sometimes form-

ing clumps several dm. tall and becoming woody toward or at the

base; leaves bi- or tripinnate or ternately divided, ovate-triangular,

the oblongish segments obtuse; sepals entire or crenate, the fringed

petals 7-9 mm. long; ovary lobes acute as those of the ovoid capsule,

this at maturity 7-9 mm. across. — An occasional introduction, usually

in stony places. The similar R. graveolens L. may occur; its petals

are entire or dentate and the ovary and capsule lobes are rounded.—

Illustrated, Fiori & Paol. Icon. Fl. Ital. 297.

Huanuco: Weed at Muna, 3989. — Junin: Tarma, Killip & Smith

21784.— Loreto: Rio Itaya, Williams 173. Iquitos, Williams 3552.

Introduced from Southern Europe. "Ruda."

18. DICTYOLOMA DC.

Ashy pubescent shrub with thin bark and hard wood, only faintly

bitter when freshly cut, bipinnate leaves of many entire glandular-

punctate leaflets and ample supra-axillary cymosely branched

panicles mostly near the tip of the stem. Flowers polygamous, all

parts 5, the calyx small, the petals lightly imbricate, sericeous, the

disk tumid, hirsute, 5-lobed, the filaments with large bifid hirsute

scales. Styles connate into 1 with large 5-lobed stigma; ovules 4-5

in each cell. Capsules 5, distinct, compressed, introrsely bivalved,

each valve with 3 or 4 very flat suborbicular seeds that are sur-

rounded, except for a narrow cleft, by an elegantly veined wing, the

veins mostly radiate but several concentric. The name has been

conserved. The genus was once included in the Simaroubaceae.

Dictyoloma peruvianum Planch, in Hook. Lond. Journ. Bot.

5: 583. 1846.

Small tree or shrub with simple trunk commonly 4-6 meters high;

leaves 2-4 dm. long with 20 or more pinnae the intermediate ones

with 12-15 pairs of lanceolate or narrowly ovate-lanceolate leaflets,

the lower and upper pinnae shorter; leaflets canescent puberulent

beneath, firm, 3-5 cm. long, 6-12 mm. wide; panicles 5 dm. long or

longer, ashy sericeous including the subsessile flowers, these with

deeply parted calyx, oblong, shortly mucronulate petals 5 mm. long,

FLORA OF PERU 689

2 mm. wide; ovary densely tomentose-hirsute; capsules about 12 mm.

long, half as wide. — Illustrated, Engl. & Drude, Veg. Erde 15: 117.

The leaves are used locally as a substitute for soap but more

commonly as a fish "poison"; also recorded (by Bues) as a remedy

for gonorrhoea.

Cajamarca: Valley of the Tabaconas near Ja£n, 900 meters,

Weberbauer 6161.— Huanuco: Pozuzo, Ruiz & Pav6n. Tocache and

on the Huallaga, Poeppig 1878; 2419. — Junin: Near La Merced,

Kittip & Smith 24012. — San Martin: Tarapoto, Williams 5530;

6160; Spruce 3888. Chazuta, King 4090 (det. Standley).— Ama-

zonas: Chachapoyas(?), Mathews 1657, type. — Loreto: Yurimaguas,

Williams 4296(1}', Kittip & Smith 27543; Ule 6261. Balsapuerto,

Klug 3088. — Cuzco: At 700 meters, Bues. "Huaman-samana"

(Williams), "achuhua" (Bues). "Barbasco negro" (Spruce).

SIMAROUBACEAE [DC.] Lindl. Quassia Family

Reference: Cronquist, Brittonia 5: 128-147. 1944 (with references

to other papers), and I.e. 469-470, a revised key.

Like Rutaceae but the leaves usually pinnate and eglandular at

least on the surfaces, bark often bitter, filaments sometimes append-

aged, carpels often lightly connate or distinct; Bentham remarked

that these are little "differences" but highly characteristic. The

family moreover as indicated by Engler, Pflanzenfam. ed. 2. 19a:

364. 1931, contains six sharply defined groups, including in Peru

Picramnia. Brunellia R. & P. long considered as belonging here is

now allied to or included in the Cunoniaceae, which see. Besides

the following the Chinese "Tree of Heaven," Ailanthus altissima

(Mill.) Swingle, Journ. Wash. Acad. Sci. 6: 490-498. 1916 (or possibly

but scarcely A. glandulosa Desf.; cf. Suringar, Mitt. Deutsch.

Dendr. Ges. 41: 45. 1929), commonly cultivated in warm lands,

may be grown in Peru and as is its habit elsewhere, may persist;

it is a smooth-barked tree with pinnate leaves of 13-25 sparsely

serrate lanceolate-ovate leaflets, ciliate but glabrous beneath and

small polygamous greenish flowers in large panicles followed in the

pistillate trees with ample clusters of reddish samaras 3-4 cm. long.

The South American members of the family provide bitter woods,

notably Quassia, and some yield violet dyes, particularly species of

Picramnia. Key after Bentham.

Ovary lobed, the carpels soon distinct and 5.

Stamen filaments unappendaged and as many as the petals.

1. Picrolemma.

690 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII

Stamen filaments appendaged, at least in male flowers, sometimes

only at base, and twice as many as petals except Quassia.

Flowers unisexual; stigmas separate 2. Simarouba.

Flowers perfect; stigmas united or somewhat lobed.

Leaf rachis winged; petals erect, nearly glabrous. .3. Quassia.

Leaf rachis not winged; petals somewhat spreading, usually

pubescent 4. Simaba.

Ovary entire, the carpels solid even in fruit; styles usually free;

stamens as many as petals or sepals 5. Picramnia.

Vegetative Key (Peru only)

Leaflets rounded, emarginate or very obtuse, sometimes mucronate

or exceptionally and in part acute.

Petals glabrous; flowers unisexual; stigmas distinct. 2. Simarouba.

Petals pubescent; flowers perfect; stigmas united or lobed.

4. Simaba.

Leaflets regularly acute to acuminate.

Leaf rachis winged; flowers large 3. Quassia.

Leaf rachis not winged; flowers 1-3 (5) mm. long.

Flowers to 5 mm. long in narrow irregular panicles.

1. Picrolemma.

Flowers usually smaller in racemes or spikes, apparently.

5. Picramnia.

I. PICROLEMMA Hook. f.

Reference: Cronquist, Brittonia 5: 143-144. 1944.

A perfectly smooth shrub-tree with simple fistulose trunk, the

wood extremely bitter, and imparipinnate alternate leaves with many

pairs of entire petiolulate oblong-lanceolate leaflets. Flowers dioe-

cious, small, orange, subfasciculate, pedicellate, ebracteolate, in irreg-

ular panicles shorter than the leaves. Male flowers with cupulate

4-lobed calyx, 4 oblong deciduous petals inserted with and opposite

to the 4 smooth stamens, the disk obscure; female 5-merous, the

stamens rudimentary, disk elevated, ovary 5-parted with spreading

lobes, the stout style with capitate stigma; ovules solitary near tip

of cell, pendulous. Drupes by abortion solitary, oblong, sessile,

obtuse, the mesocarp thin.

Ants often inhabit the hollow stems or branches.

FLORA OF PERU 691

Calyx teeth shorter than the tube; anthers longer than filaments.

P. Huberi.

Calyx teeth longer than tube; anthers shorter than filaments.

P. Sprucei.

Picrolemma Huberi Ducke, Archiv. Jard. Bot. Rio Jan. 4: 197.

1925.

Like P. Sprucei but leaves with 8-10 pairs of oblique obtuse or

subcordate leaflets to 15 cm. long, 6 cm. wide, the rachis not ridged

or winged; panicle few-flowered, about 5 cm. long; calyx scarcely

more than 1 mm. high, the teeth shorter than the tube; petals 6-7

mm. long, about 1 mm. wide; staminodia 5, linear, longer than the

stamens, the large anthers longer than the filaments. — Pistillate

flowers and fruit unknown. Sometimes 6 meters high. Huber noted

the flowers as very fragrant, Klug as white. F.M. Neg. 12540.

San Martin: Klug 2657. — Loreto: Quebrada Grande de Cancha-

huaya, Ucayali, Huber 1471, type.

Picrolemma Sprucei Hook. f. in Benth. & Hook. f. Gen. PI.

1: 312. 1862. P. pseudocoffea Ducke, Archiv. Jard. Bot. Rio Jan.

4: 196. 1925, fide Cronquist.

Shrub 1-several meters high, the leaves with usually 4-9 pairs of

elliptic-oblong to ovate or broadly elliptic leaflets mostly 8-18 cm.

long, 3-7 cm. wide, usually rounded or obtuse at base, abruptly

linear-caudate, the acumen 1 cm. long or longer, the rachis margined

or with 2 ridges; panicles ample, open, the staminate 1.5-3 dm. long,

pistillate little shorter; calyx about 1 mm. high, the teeth longer than

the tube; petals orange, 2.5-5 mm. long; filaments much longer than

the small anthers, the staminodia short and broad; fruit brown

or reddish, 2-3 cm. long. — Petals orange or yellow, fruit bright

orange (Killip & Smith). Illustrated, Mart. Fl. Bras. 12, pt. 2:

pi. 46. F.M. Negs. 22062; 12542 (both P. pseudocoffea).

Loreto: Iquitos, Killip & Smith 27294- San Antonio, Killip &

Smith 29482. Florida, Klug 1985. Mishuyacu, Klug 166; 442.

Amazonian Brazil. "Cafe"-rana" (Huber).

2. SIMAROUBA Aublet

The Peruvian species a large tree, the pinnate leaves with alter-

nate leaflets. Inflorescences dioecious, the staminate larger than the

pistillate; sepals 5 (4-6), united at base, the petals the same number,

distinct. Stamens appendaged only at base (squamate), 10 (8-12),

692 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

in the pistillate flowers reduced or wanting. Carpels usually 5 (4-6),

lightly united, with short common style and broad divergent stigmas,

1-ovuled, the ovule pendulous. Fruit composed of several distinct

drupes, these often with subapical tips the flesh thin. — Name orig-

inally written as above. In herbaria the leaflets are brittle and fall

readily.

Simarouba amara Aublet, PI. Guiana 2: 860. pis. 331, 332.

1775.

Leaves with or without a terminal leaflet, the glabrous leaflets

usually oblong, cuspidate, 5-10 cm. long, about 2.5-4 cm. wide,

with many very fine lateral nerves united near the margin; panicles

3 dm. long or longer, the many white or yellowish-green flowers

mostly approximate on the branches; calyx 1 mm. long or shorter,

the acute lobes ciliate; petals ovate to lanceolate, about 3.5 mm.

long; stamens about 1.5 mm. long, the scales villous (male flowers).

Cronquist, Bull. Torrey Club 71: 229. 1944, from whose work I

have compiled, has noted petals as 3.5-4.5 mm. long, anthers 0.7-1

mm. long, the leaves typically rugulose-punctate beneath, dull and

glabrous or in var. opaca Engl. (S. opaca (Engl.) Radlk.) papillate

and glaucous beneath, this not, fide Cronquist, as yet known from

Peru. The following collections from trees 10-15 meters high.

San Martin: Zepelacio, Klug 3741- Near Moyobamba, Weber-

bauer, 290. — Loreto: Pumayacu, Klug 3207. Mishuyacu, Klug

639; 323. Near Iquitos, Tessmann 5431. — Rio Acre: Mouth of Rio

Macauhan, Krukoff 5240. Brazil and Bolivia to Central America;

Antigua, West Indies. "Simarouba" (Aublet); "marouba" (Brazil).

3. QUASSIA L.

An extremely bitter shrub or tree much like Simaba but with

winged petioles and leaf-rachises and with large red tubular flowers

borne in simple or branched terminal racemes, the pedicels bracted

at base and bibracteolate at the articulate apex. Flowers 5-merous

with erect membranous contorted petals, a large columnar torus.

Filaments appendaged. Ovary deeply 5-lobed, the styles united.

This, the Quassia or one of the "bitterwoods" of commerce is

cultivated in adjacent countries, probably also in Peru.

Quassia amara L. Sp. PL ed. 2. 553. 1762.

Shrub or becoming a large glabrous tree with pinnate leaves,

the winged often red rachis bearing 5 (usually) elliptic-oblong acumi-

FLORA OF PERU 693

nate entire leaflets, narrowed to base, 7-16 cm. long, 2-6 cm. wide,

and terminal racemes of large red flowers, the contorted petals about

3 (-nearly 5) cm. long; stamens exserted, the red filaments with

white pilose basal appendage; drupes black with pale spot at base,

8-15 mm. long, free from each other on the fleshy red disk. — Illus-

trated, Pflanzenfam. ed. 2. 19a: 378.

Peru: Probably in cultivation. Brazil; Guiana. "Quassia."

4. SIMABAAublet

Reference: Cronquist, Lloydia 7: 81-92. 1944 (no descriptions).

Shrubs or trees or if suffrutescent the leaves all basal the leaflets

usually opposite and in one species only with terminal leaflet. In-

florescence a simple or mixed panicle; pedicels continuous or with

joint at base. Flowers perfect; sepals 4 or 5, more or less united, the

petals as many, distinct, pubescent and more or less spreading at

anthesis. Stamens twice as many as petals the filaments with pubes-

cent appendage which is sometimes free except at base. Carpels

4 or 5, lightly united, 1-ovuled, the style 1 with entire or slightly

lobed stigma. Fruit sometimes with only 1 mature drupe, usually

several.

Besides the following, S. cedron Planch., the widely cultivated

"Cedron," may occur at least in haciendas or parks; it is a tree with

10-15 pairs of usually gland-tipped leaflets pubescent on the mid-

nerve above; the petals are 2-3.5 cm. long, the filament appendages

1-2 cm. long, drupes about 1 dm. long, the tree sometimes 15 meters

tall.

Simaba multiflora A. Juss. Me"m. Mus. Hist. Nat. Paris 12:

pi 27. 1825; 86.

Tree or tree-like; leaves mostly 1.5-2 dm. long with 3-9 oblong-

or obovate-lanceolate leaflets, the lateral ones 4-7 cm. long, 2-2.5

cm. wide; inflorescences ample; petals cream colored, 4-7 mm. long,

puberulent both sides; filament appendages at most 1.8 mm. long,

equaling or longer than the gynophore the free portion much shorter

than the attached part; fruit stout, symmetric, 1.5-2.5 cm. long.—

The leaflets vary from obovate and rounded at tip, where often

minutely mucronate, to lanceolate and acute or rarely even emargi-

nate, the fine lateral nerves faint both sides and obsoletely or ob-

scurely reticulate. Referred with doubt by Engler, Mart. Fl. Bras.

12, pt. 2: 211, to S. guianensis (Aublet) Engl., maybe correctly,

694 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII

sens. lat. Other species are to be expected within Amazonian

Peru. Said to grow in lowlands at times inundated; Schunke speci-

men from a river bank, tree 15 meters tall with trunk circumference

of 1 meter.

Loreto: Rio Mazan, Jose Schunke 254 (det. Cronquist). To Para

and Trinidad.

Simaba paraensis Ducke, Archiv. Jard. Bot. Rio Jan. 4: 195.

1925.

Medium size tree or larger, the new branchlets angulate and

finely ashy tomentulose as the leaf-rachises and panicles, the older

glabrate ones strongly striate; leaves with petiole to 2 dm. long, the

rachis slightly depressed, angulate-dilated, with 3-6 rarely 7 pairs

of subsessile obovate-oblong or oblong leaflets, 7 (-10) cm. long,

3 (-3.5) cm. wide, more or less attenuate at base, obtuse or retuse

at apex, subcoriaceous, glabrous (often with a few trichomes on mid-

nerve beneath), somewhat lustrous, paler and finely pinnate nerved,

very finely reticulate beneath, the darker slender nerves prominent

both sides; panicles ordinarily terminal, ample, 3-4 dm. long, the

upper branches with simple oblongish bracts; pedicels to 1.5 mm.

long; calyx parted to below the middle, the segments acute, barely

1 mm. long; petals 6-7 mm. long, 2.3-3.7 mm. wide (Cronquist),

oblong, canescent both sides; stamens about 5.5 mm. long, the

villous appendage about 4 mm. long; gynophore 1.5 mm. long, longer

than ovary, the style glabrous above. — Flowers unpleasantly scented

with valerian.

Rio Acre: Tree, 30 meters high, mouth of Rio Macauhan, Krukoff

5276 (det. Cronquist). To Amazonian Brazil.

5. PICRAMNIA Swartz

Shrubs or trees, often bitter, with imparipinnate leaves, the

entire leaflets opposite or nearly so, and small glomerulate or racemu-

lose usually dioecious flowers borne from trunks, branches or opposite

the leaves in pendulous spikes or racemes that often are greatly

elongate. Flower with 3-5 segments and petals, the latter rarely

lacking, usually imbricate with inflexed apex and, in the male

flowers, opposite, 3-5 smooth inflexed filaments inserted under the

disk, this depressed, lobulate. Stamens in female flowers reduced

to linear scales, the sunken ovary 2-3-celled, the style with 2-3

recurved lobes stigmatic within or with 2-3 sessile stigmas. Ovules

FLORA OF PERU 695

2, near apex of cells. Drupe olive-shaped, 1-2-celled, the cells