FB

FLORA OF PERU

BY

J. FRANCIS MACBRIDE

CURATOR, PERUVIAN BOTANY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V, NUMBER 2

DECEMBER 9, 1960

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FLORA OF PERU

BY

J. FRANCIS MACBRIDE

CURATOR, PERUVIAN BOTANY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V, NUMBER 2

DECEMBER 9, 1960

PUBLICATION 902

Library of Congress Catalog Card Number: 36-10426

PRINTED IN THE UNITED STATES OF AMERICA

BY CHICAGO NATURAL HISTORY MUSEUM PRESS

FLORA OF PERU

J. FRANCIS MACBRIDB

BORAGINACEAE [Juss.] Lindl.

Various in habit, ligneous or herbaceous, usually pubescent, not

infrequently harshly, the perfect normally 5-merous flowers ordinar-

ily in open and branched or scorpoid inflorescences, rarely solitary.

Calyx never entire, the lobes sometimes uneven. Corolla usually

regular, often plaited, pubescent or, in the throat, somewhat ap-

pendaged. Stamens on the tube and alternate with the lobes. Ovary

superior, bicarpellate, the commonly biovulate carpels free or united,

each remaining entire, parting along a medial false partition or more

or less lobed or separated along the medial line of the partition.

Fruit dry to fleshy in some degree, retaining the composition of the

carpels or, and generally, becoming 2-4 biovulate or uniovulate nut-

lets. Style obsolete to well-developed, simple, bilobed or twice

forked, variously situated, sometimes on an upwardly extended re-

ceptacle (gynobase) ; stigma entire or slightly bilobed.

The numerous papers of I. M. Johnston — the outstanding stu-

dent of the family — are cited with comment under the genera con-

cerned; the usefulness of this synopsis stems, of course, from his

realistic and thoughtful studies. The cytogenetic studies on the

family by Donald M. Britton, Brittonia 7: 233-266. 1951, will prob-

ably help to an understanding of relationship.

Forget-me-not is probably the best known plant in this family,

certainly one of the most beloved in gardens nearly everywhere.

Otherwise the group is more interesting to taxonomists than to horti-

culturists; however, Echium, Heliotropium and Cordia are often grown

for ornament while a few have popular medicinal repute, notably

borrago (Borago officinalis L.).

Regretfully I here record the passing of my never-to-be-forgotten

friend, Ivan Johnston.

Ovary slightly lobed if at all, sometimes 2-4-grooved laterally.

Style entire, stigma 1; inflorescences or its branches scorpoid or its

flowers solitary.

Clambering shrubs (or, as T. microcalyx, erect) ; fruit pulp aque-

ous, mesocarp fleshy, nutlets 2-4, bony. ... 1. Tournefortia,

539

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

Herbs or suffrutescent; fruit dry, parting into 2-4 bony nutlets.

2. Heliotropium.

Style 2-parted or early coherent; much-branched prostrate herbs

or ligneous below 3. Coldenia.

Style twice forked, stigmas 4; inflorescence not scorpoid; fruit

drupaceous except C. alliodora.

Flowers sessile, not in foliose panicles; calyx not inflated.

4. Cordia.

Flowers pedicellate in foliose panicles; calyx soon dilated.

5. Saccettium.

Ovary deeply 4-parted, the fruit consisting of 4 (or fewer) nutlets.

Nutlets neither margined nor appendaged.

Corolla lobes contorted in bud; "no-me-olvides" ... 6. Myosotis.

Corolla lobes imbricate in bud.

Flowers ebracteate; leaves crowded toward stout perennial

base 7. Moritzia.

Flowers often bracted or plants annual.

Leaves all alternate; perennials 8. Lithospermum.

Leaves, at least lower, opposite or in a rosette; annuals or

biennials.

Corolla yellow, throat usually smooth .... 9. Amsinckia.

Corolla white at least in large part, throat appendaged.

Nutlets keeled ventrally (or also dorsally).

Leaves mostly in a basal rosette . . 10. Plagiobothrys.

Lower leaves opposite, even the earliest.

11. Allocarya.

Nutlets grooved ventrally, keeled obscurely if at all

dorsally 12. Cryptantha.

Nutlets alate-margined or dorsally appendaged.

Appendages apically barbed; leaves alternate. . . .13. Hackelia.

Appendages or teeth apically uncinate-setose; first leaves oppo-

site 14. Pectocarya.

1. TOURNEFORTIA L.

Reference: Johnston, Contr. Gray Herb. 92: 66-68. 1930.

Lianas or shrubby and then often lax or subscandent, the flowers

in scorpoid racemes or spikes disposed in dichotomous panicles.

FLORA OF PERU 541

Calyx persistent. Corolla limb spreading. Ovary 4-celled; style (or

stigma) terminal, simple, the stigma peltate or conic, usually bifid at

tip, often sessile. Fruit fleshy, obscurely if at all lobed or definitely

consisting of 1-2-seeded irregular nutlets or of 4 consimilar 1-seeded

nutlets; nutlets often with 1-2 empty cavities. Embryo straight

or curved.

The scholarly paper of Johnston, while entitled "Treatment of

Tournefortia," actually concerns only the species of the eastern half

of South America; he divided the genus into section Eutournefortia

Johnst., the obscurely or not lobed fruit bicarpellate, composed of

1-2-seeded uneven (often joined) nutlets, the embryo straight, and

section Cyphocyema Johnst., the evidently lobed fruit obscurely bi-

carpellate, consisting of 4 similar 1-seeded nutlets, the embryo curved

and the corolla lobes very narrow with involute margins. This clari-

fication of the genus is a fine contribution to its taxonomy; most

Peruvian species belong to the latter section (some are incompletely

known) ; at least T. bicolor Sw., T. cuspidata HBK., T. hirsutissima L.,

T. ovalifolia Rusby and T. Ulei Vaupel belong to the former. The

following key attempts to lead to herbarium materials often only

in flower; no doubt several names refer more correctly to variants.

Corolla segments more or less setiform, distinctly longer than wide,

often recurving at tip; calyx segments minute or narrow.

Corolla about 1 cm. long; leaves usually rounded at base; inflores-

cences ordinarily terminal T. maculata.

Corolla to about 5 mm. long; leaves somewhat acute at base; in-

florescences axillary, often also terminal.

Stems as leaves beneath strigose tomentose T. mapirensis.

Stems as leaves green or lightly appressed strigose.

Leaves merely acute at base . . . . T. breviflora, T. psilostachya.

Leaves decurrent at base T. glabra.

Corolla segments rounded to acute, sometimes narrowed above but

not distinctly longer than wide; calyx segments about 1.5 mm.

long or longer, or broad.

Leaves more or less bicolored, tomentose beneath, usually green

and sparsely (rarely densely) strigose above . . T. polystachya.

Leaves concolor, pubescence subequally developed both sides.

Corolla tube 2.5-3 mm. long; calyx minute; leaves oblong-

lanceolate to elliptic, rarely 3.5 cm. long. . . . T. microcalyx.

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

Corolla tube about 4-12 mm. long; calyx and leaves mostly

larger.

Inflorescence lax, branchlets even in flower divaricate, some

soon several cm. long or if early suberect or short, corollas

about 1 cm. long.

Corollas about 5 mm. long, rather in corymbs; style well-

developed (unless T. glabra).

Flowers sessile; leaves ample, usually drying dark.

T. Ulei, T. glabra.

Flowers pedicellate; leaves drying brownish-green.

Calyx hispidulous; stems scabrous T. ovalifolia.

Calyx glabrate; stems scurfy T. chinchensis.

Corollas 10-12 mm. long, mostly on dichotomous branches;

style short.

Leaves ample, drying black, glabrous T angustiflora.

Leaves rather small, greenish-brown, puberulent.

T. tarmensis, T. Buchtienii.

Inflorescence dense, branchlets even in age rarely 3 cm. long

unless T. hirsutissima, T. undulata, especially if corolla

is long, this usually 5-7 mm. long.

Stems notably spreading hirsute, trichomes long; corolla

about 1 cm. long T. cuspidata.

Stems usually densely, leaves evenly more or less appressed

hispid; corolla densely strigose, shorter.

T. hirsutissima.

Stems glabrous or puberulent as leaves or these rather ob-

scurely pubescent on nerves and veins.

Upper stems and inflorescence closely reddish puberulent.

T. tarmensis, T. Buchtienii.

Upper stems glabrous or glabrate, indument if present

sparse.

Leaf nerves, at least mostly, about 1 cm. distant; peti-

oles about 1 cm. long.

Leaves about twice as long as wide.

Leaves membranous T. setacea, T. bicolor.

Leaves coriaceous T. coriacea.

Leaves about three times longer than wide.

T. longifolia.

FLORA OF PERU 543

Leaf nerves, at least mostly, about 5 mm. distant.

Leaves long-decurrent, about three times longer than

wide T. ternifolia.

Leaves short-decurrent, or subsessile, about two

times longer than wide . . T. virgata, T. undulata.

Tournefortia angustiflora R. & P. Fl. Peruv. 2: 25, pi 151a.

1799.

Spreading, branching, glabrous or nearly, the larger subherba-

ceous glabrous branches about 8 mm. in diameter with ovate-acumi-

nate leaves to 2 dm. long, 8 cm. wide, those of the flowering branchlets

half as large or smaller, narrowly acuminate, minutely hispidulous

both sides; inflorescence branches often arcuate-divaricate; calyx ses-

sile, scarcely 1 mm. long, lobes ovate, subacute; corolla greenish or

yellowish-white, sparsely and obscurely puberulent, the slender tube

1 cm. long, the lobes subrotund, about 5-6 mm. across expanded; fruit

white, succulent, glabrous, 5-6 mm. in diameter. — F.M. Neg. 1038.

Hudnuco: Chicoplaya and Pueblo Nuevo, Ruiz & Pavdn, type.

Pozuzo, 4644- Prov. Huamalies, Weberbauer 8637 (det. Vaupel). —

Loreto: Yurimaguas, Killip & Smith 28064 (det. Killip). — Rio Acre:

Seringal Auristella, Ule 9708. Ecuador.

Tournefortia bicolor Swartz, Prodr. 40. 1788. T. laevigata Lam.

Tab. Encycl. 1: 416. 1791, fide Johnston, Journ. Arnold Arb. 16: 51.

1935.

To several meters tall, finally more or less scandent, its branchlets

glabrate or sparsely appressed pubescent with antrorse trichomes;

petioles 0.5-1.5 cm. long; leaves ovate to elliptic or somewhat lanceo-

late, obtuse-rounded at base, acute, 0.5-1.5 dm. long, 1.3-9 cm. wide,

slightly lustrous, dark green, often obscurely papillate and pubescent

with fragile appressed trichomes, these more abundant beneath; in-

florescence often ample, finally lax but the racemes early crowded,

1-4 cm. long; calyx sparsely strigose, usually sessile, lobes at anthe-

sis usually 1-2.5 mm. long, little accrescent; corolla strigose, tube

4-5 mm. long, limb 6 or 7 mm. across; fruit white, succulent, gla-

brous, elobate; stigmas subsessile. — Calyx lobes subfiliform, some-

times 5 mm. long; pubescence nearly lacking or more abundant (var.

calycosa Bonn. Smith, Bot. Gaz. 14: 27. 1889). Common, especially

near Iquitos, only a few collections cited.

San Martin: San Roque, Williams 7783 (det. Killip).— Junin:

La Merced, Killip & Smith 23515 (det. Killip). — Loreto: Near

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

Mazan, Mexia 6455 (det. Johnston); Jose Schunke 121; 270 (det.

Killip). Yurimaguas, Williams 4237, 4241 (det. Standley); King

2809 (det. Killip). Near Iquitos, King 735; Killip & Smith 27413;

Williams 1395; 3571. Yarina Cocha, Tessmann 5425. Florida,

Klug 1999. American tropics. "Loblobo" (Mexia).

Tournefortia breviflora DC. Prodr. 9: 520. 1845; 76. T.

xapuryensis Vaupel, Notizbl. Bot. Gart. Berlin: 186. 1914?

Resembles T. mapirensis but green, obscurely if at all strigose,

and the slender divaricately branched inflorescence usually entirely

axillary (or also terminal); stems as petioles somewhat puberulent,

the latter 1-1.5 cm. long; leaves oblong-elliptic, acute at base, rather

abruptly acuminate, to about 1 dm. long, 4 cm. wide, the slender

distant nerves prominent only beneath; inflorescence lax, 5-8 cm. long,

sometimes with a linear bract or two; calyx about 0.5 mm. long, the

puberulent-ciliate lobes lanceolate, acute; corolla 3-4 mm. long,

greenish-yellow, obscurely puberulent, the lobes subulate-subseti-

form (Vaupel). — DeCandolle's plant, ex char., may be distinct; his

diagnosis differs in petioles 6 mm. long, leaves about 6 cm. long,

2 cm. wide, corolla 3 mm. long, the short lobes acute. However, the

Vaupel plant probably occurs in Peru like the related T. mapirensis

with in part similar range; both species seem to be scarcely separable

from T. psilostachya HBK.— F.M. Neg. 1065.

Rio Acre: Seringa! San Francisco, Vie 9710 (type, T. xapuryensis).

Brazil; Paraguay.

Tournefortia Buchtienii Killip, Journ. Wash. Acad. Sci. 17:

334. 1927.

Branches subterete, ferrugineous-tomentose and scabroid; peti-

oles opposite, to 2 cm. long; leaves oblong or oblong-elliptic, rounded

or subacute at base, acuminate, 7-14 cm. long, 3-5 cm. wide, the 10

or so lateral nerves slightly impressed above, where dark green and

appressed strigillose except the nerves, rusty hirsutulous especially

beneath; peduncles 6-7 cm. long, subternately branched, the branch-

lets 2-3-dichotomous, the subterminal flowers 2.5 mm. distant; sepals

linear, 2-3 mm. long; corolla tube 7-10 mm. long, dilated just be-

low throat, rusty tomentose, the orbicular lobes minute; stamens

near throat, linear; style about 7 mm. long; fruit ovoid-conical,

7-8 mm. long, glabrescent, white. — After author, who indicated no

relationship. Leaves of Weberbauer specimen are oblong-elliptic,

FLORA OF PERU 545

to 5 cm. long, 12 mm. wide, while Eyerdam 25123 from La Paz has

leaves to 1.5 dm. long, 5 cm. wide.

Junin: Comas, 3,300 meters, Weberbauer 6607 (det. Killip, 1937,

T. ovalifolia; 1925, T. chinchensis}. Bolivia.

Tournefortia chinchensis Killip, Journ. Wash. Acad. Sci. 17:

331. 1927.

Branches soon terete, scurfy, ferrugineous-hirsutulous; petioles

5-10 mm. long; leaves opposite, ovate-oblong to narrowly oblong,

3-7 cm. long, about 1-3 cm. wide, slightly narrowed at base, acute,

entire, the 6 or 7 nerves impressed above, prominent beneath, sub-

coriaceous, glabrous or slightly hispidulous above, glabrous beneath

except for the rusty hirsutulous nerves and veins; inflorescence 3-4-

dichotomous, the ultimate branches 2-3 cm. long in flower, the

flowers about 2 mm. apart; sepals linear, 2-3 mm. long, glabrous

or a little pilosulous; corolla tube 6-9 mm. long, rusty tomentose,

greenish, the minute orbicular but apiculate lobes glabrous; stamens

1-1.5 mm. long, style 5 mm. long; fruit depressed globose, about

8 mm. in diameter, white, glabrescent. — After Killip, who contrasted

two similar species (T. andina Britton, T. ovalifolia Rusby) as follows:

Flowers sessile; corolla tube appressed hirsute; branchlets smooth.

T. andina.

Flowers pedicellate; corolla tube tomentose; branchlets rough.

Calyx lobes 2 mm. long or less; leaves alternate T. ovalifolia.

Calyx lobes longer than 2 mm.; leaves subopposite . T. chinchensis.

However, Johnston included the shrub in T. ovalifolia Rusby.

Ecuadorian specimens by Steyermark, apparently the same, have

leaves 8 cm. long, 3.5 cm. wide.

Huanuco: Hacienda Villcabamba, Rio Chinchao, 5142 (type,

T. chinchensis') . Bolivia, Ecuador.

Tournefortia coriacea Vaup. Bot. Jahrb. 54, Beibl. 119: 3. 1916.

Shrubby, the stout branches smooth; petioles 2 cm. long; leaves

oval or broadly elliptic, short-acuminate, ample, coriaceous, glabrous;

inflorescence small with few branches; flowers about 5 mm. long; calyx

lobes narrowly lanceolate; corolla tube about two times longer than

the calyx, sparsely pilosulous with minute appressed trichomes, the

lobes obtuse; anthers affixed at base of tube, narrowly lanceolate;

pistil shorter than calyx, stigma conical. — After the author, who

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

merely remarked : at once recognizable by the large leathery leaves.

The Williams specimen has petioles 1 cm. long, leaves to 12 cm.

long, 8 cm. wide, nerves 1 cm. apart, lightly impressed above;

calyx lobes subulate, 2 mm. long, corolla puberulent, tube 7 mm.

long, lobes rounded, 1 mm. long. F.M. Neg. 1042.

Loreto: Yurimaguas, Williams 78S9 (det. Killip). Ecuador.

Tournefortia cuspidata HBK. Nov. Gen. & Sp. 3: 83. 1818.

T. obscura A. DC. Prodr. 9: 517. 1845, fide Johnston, Journ. Arnold

Arb. 16: 54. 1935.

Shrub or liana, the branchlets short-pubescent and inordinately

villous-hirsute with slender spreading brown trichomes some 4 mm.

long; petioles to about 1 cm. long or somewhat longer; leaves lance-

olate to subovate, rounded or obtuse at base, acuminate, 7-15 cm.

long, 3-6 cm. wide, closely appressed strigose; spikes approximate

at ends of elongate branches, to 3 cm. long; calyx lobes subulate-

linear, 7-9 mm. long, little longer in fruit, sparsely long-pubescent

and short-strigose; corolla tube 5-8 mm. long, densely strigose,

limb 4-6 mm. across; fruit white, fleshy, somewhat verruculose;

stigma sessile. — Common on the lower Rio Huallaga, only a few col-

lections cited.

San Martin : Juanjui, Klug S776. — Loreto: Yurimaguas, Williams

3845; 4312; 4776; 4972; 7870. Yarina, Tessmann 3773. Near Iquitos,

Klug 432. Bolivia to Central America and the West Indies.

Tournefortia glabra L. Sp. PI. 141. 1753.

Large scandent or erect shrub or small tree; branchlets puberulent

or glabrate; leaves lanceolate-elliptic to ovate, 0.5-1.5 dm. long,

narrowed to both ends and often long-decurrent on the slender

petioles, membranous, early pilosulous, at maturity nearly glabrous;

spikes frequently many, slender, sometimes greatly elongate, the

inflorescence lax; calyx lobes lanceolate, about 2 mm. long, long-

acuminate as the lobes of the pale green or whitish corolla, its tube

4 mm. long; fruits white, succulent, to about 5 mm. thick, 9 mm.

long, not lobed. — Wood bright yellow when cut.

San Martin: Pongo de Cainarachi, Klug 2730 (det. Killip). To

Mexico; West Indies.

Tournefortia hirsutissima L. Sp. PI. 140. 1753. Messer-

schmidtia hirsutissima (L.) Roem. & Schult. Syst. 4: 541. 1819.

FLORA OF PERU 547

Branches subtetragonous, typically densely (not in Peru as deter-

mined) short-hirsute as the petioles and peduncles; petioles stout,

6-9 mm. long or much longer; leaves ovate-oblong or -elliptic, usually

acuminate, rounded to narrowed at base, mostly 6-9 cm. long, about

5 cm. wide to twice as large, membranous, appressed strigose beneath,

scabrous or more sparsely hispid above; inflorescence usually ample,

corymbose or dichotomously divided; spikes dense, narrow, in fruit

3-4 cm. long; calyx hirsute or hispidulous, the calyx teeth linear to

ovate-oblong, acute or acuminate; anthers linear, at middle of tube;

corolla white, at least the tube sericeous, 4-6 mm. long, the lobes

acute or apiculate; style very stout, stigma depressed-conical; fruit

globose, white, hispidulous, about 5 mm. in diameter; nutlets 4,

ovate-oblong. — Shrubby or a liana to several meters long; the indu-

ment is sometimes so short that it is inconspicuous; Killip determina-

tions.

San Martin : Juan jui, King 4321 . Tarapoto, Williams 5781 ; 5690;

6629. — Huanuco: Puente Durand, 1,900 meters, Weberbauer 9590.

Widely distributed in tropical America.

Tournefortia longifolia R. & P. Fl. Peruv. 2: 25, pi. 150b. 1799.

Glabrous or nearly, erect, the branches sulcate-angled; petioles

about 1 cm. long; leaves oblong-lanceolate or -elliptic, acute both

ends or acuminate, to 2 dm. long, 6-7 cm. wide, submembranous

but somewhat rugose-bullate above in age, much veined; inflores-

cence dichotomous but small, few-flowered; seeds trigonous. — Type

in fruit, imperfectly known. F.M. Negs. 12943; 29254.

Huanuco: Pozuzo, Ruiz & Pavdn, type.

Tournefortia macula ta Jacq. Enum. 14. 1760; Sel. Stirp. 47.

1763. T. syringaefolia Vahl, Symb. 3: 23. 1794. T. peruviana Poir.

Encycl. Suppl. 4: 425. 1816. T. volubilis L. as to R. & P. Fl. Peruv.

2: 24, pi. 148b. 1799, fide Urban and Johnston. T. paniculata Cham.

Linnaea 4: 468. 1829?

Shrubby or sometimes more or less clambering or climbing, dark

green and inconspicuously short-strigose or leaves softly pilosulous

beneath, including the lax inflorescence and growing parts; petioles

about 1-1.5 cm. long, leaves ovate or somewhat elliptic-lanceolate,

acute to rounded at base, acuminate, mostly 5-10 cm. long, 3-5 cm.

wide, ordinarily densely and finely tuberculate; calyx 1-1.5 mm. long,

scarcely accrescent but pedicels finally 1-5 mm. long and thickening;

corolla greenish-white, tube 3-4 (-8) mm. long, limb 3-5 mm. across;

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

fruit yellow, often blotched with black, the 4 finally separate nutlets

rounded dorsally, angled ventrally. — Johnston, Journ. Arnold Arb.

30: 130. 1949, identified the plant of Jacquin. F.M. Neg. 1054

(T. volubilis).

San Martin: Tarapoto, Williams 5414; 6170 (det. Killip). Zepe-

lacio, Klug 3348. — Huanuco: Pozuzo, Ruiz & Pav6n (type, T. volu-

bilis). — Junin: La Merced, 5473. — Loreto: Yurimaguas, Williams

4778; 4971; 4973; 5121. Nanta, Raimondi 9427—Cuzco: Valle de

Santa Ana, Herrera 2632. — Puno: Gay (as T. voluUlis). To Central

America and the West Indies; Brazil.

Tournefortia mapirensis Lingelsheim, Repert. Nov. Sp. 7: 244.

1909; 76. T. subrotunda Rusby, Bull. N. Y. Bot. Card. 8: 115. 1912,

fide Johnston.

Fulvous strigose-tomentose liana (except leaves above) with axil-

lary (and terminal) leafless lax inflorescences; petioles 1-2 cm. long;

leaves broadly ovate, subcordate or rounded at base, shortly acum-

inate, (4) 8-12 cm. long, (2.5) 4.5-8 cm. wide, dark green but lightly

pubescent above, paler and densely strigose-tomentose beneath, the

nerves 5 or 6 pairs; calyx 2 mm. long, accrescent in fruit; corolla

greenish-yellow, scented, the tube very narrow, pubescent, 4 mm.

long, the subsetiform acute segments 1 mm. long; fruits early yellow,

subglobose, 4 mm. thick, finally purple to black-brown. — Mostly

after author; very distinct by its tawny pubescence, definitely axil-

lary inflorescences, the flowering branchlets leafless or the few leaves

much reduced, according to Johnston, who determined the Peruvian

specimen. Inflorescences 4-6 cm. broad, branches 1.5-2.5 cm. long

(Rusby).

Loreto: Yurimaguas, (Spruce 3889). — Rio Acre: Seringal San

Francisco, Ule 9712; 9713. Bolivia to Amazonian Brazil.

Tournefortia microcalyx (R. & P.) Johnst. Journ. Arnold Arb.

37: 295. 1956. Heliotropium microcalyx R. & P. Fl. Peruv. 2: 3,

pi. 109b. 1799. Tournefortia hispidula R. & P., in herb.

Open, often stockily branched, a meter or two tall, glabrous or

essentially or the oblong-lanceolate leaves beneath and almost tiny

flowers minutely hispidulous; leaves oblong-lanceolate, attenuate to

the slender petioles, these 5-8 mm. long, moderately acuminate,

5-10 cm. long, 1-3.5 cm. wide, membranous, obscurely venose above,

lightly reticulate-veined beneath; inflorescence axillary, usually

about 3-5 mm. in overall size, the branchlets often arcuate-spreading;

FLORA OF PERU 549

calyx segments sublinear, scarcely 1.5 mm. long, not accrescent in

fruit, usually glabrate as the nearly twice as long corolla, this with

rounded lobes, expanded about 2.5 mm. across, white; fruit composed

of 4 similar nutlets. — Shrub but open and with short or longer and

floppy branchlets, white translucent fruit. Determinations by Killip.

F.M. Negs. 1048 (T. hispidula) ; 1051.

Piura: Cerro Viento, Haught 95. — Ancash: Tambo de Pariocota,

2550.— Lima : Pachacamac, Mexia 8096; 83S3. Lurin, Pennell 1 2205.

Atocongo, Pennell 14758. Prov. Huarochiri, Weberbauer 5250. Near

Lima, Nee; Gaudichaud. Chosica, Grant 7401. — Huanuco: Chico-

playa, Isern 2080. Chinchao, Ruiz & Pavdn (type, T. hispidula).

Hills of Huanuco, Ruiz & Pavdn, type.

Tournefortia ovalifolia Rusby, Bull. N. Y. Bot. Gard. 4: 414.

1907.

Branchlets sparsely scabrous; petioles about 1 cm. long, broadly

sulcate; leaves oval, slightly decurrent at base, abruptly and acutely

cuspidate, 6-10 cm. long, 3-6 cm. wide, dark green, the 10-12 pairs

of lateral nerves prominent only beneath where yellowish or purplish;

peduncles 2-3 cm. long, the branches of the dichotomous inflorescence

elongating, flowers 2-3 mm. distant; pedicels 2-3 mm. long; calyx

2 mm. long, truncate at base, the lanceolate lobes erect; corolla tube

5 mm. long, lobes rounded; style 3 mm. long, stigma 1 mm.; fruit

globose-ovoid. — Flowers green. Perhaps should be interpreted to in-

clude T. chinchensis Killip.

Peru (cf. note above). Bolivia.

Tournefortia polystachya R. & P. Fl. Peruv. 2: 24, pi. 149,

fig. a. 1799. Heliotr opium oppositifolium R. & P. I.e. 2: pi. 108, fig. b.

T. loxensis HBK. Nov. Gen. & Sp. 3: 81. 1818, at least as to Peru.

An open shrub or tree-like in age, the flowering branchlets puber-

ulent, the ample open terminal inflorescences as the leaves beneath

fulvous-canescent villous; petioles about a fourth as long as leaves,

these, at least the lower, somewhat obliquely rounded to subcordate

at base, shortly acuminate, commonly ovate or the upper sometimes

somewhat elliptic and narrowed to base, those of the flowering

branches 6-15 cm. long, usually about 4-6 cm. wide, but attaining

2.5 by 2 dm., slightly scabrous, glabrate and finally bullate above;

calyx segments sublinear, 2 mm. long, not accrescent; corolla tube

about 4-5 mm. long, lobes rotund; fruit with white translucent pulp

at least 1 cm. in diameter. — The flowers, at first greenish-yellowish,

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

are white when fully expanded and 5 mm. across, either with a dis-

agreeable odor (2404) or fragrant (4017, 4188; and Stork & Horton);

unnoted determinations by Killip. Type of H. oppositifolium R. & P.

had hispid stems and leaves asperous above but seems to belong here.

F.M. Negs. 1056; 27094 (Mathews); 1052 (T. oppositifolia).

Cajamarca: Below Ocros, Weberbauer 2716; 271 6a. — Amazonas:

Chachapoyas, Mathews. — La Libertad: Cachicadan, Stork & Horton

9953 (det. Standley, Heliotropium arborescens) . — Lima: Ambar, Stork

1 1 433.— Huanuco : Chaglla, Weberbauer 6706. Huanuco, 2404; 3098;

Sawada 82. Mufia, 4017; 4188; Ruiz & Pavon (type, H. oppositi-

folium). Mito, 3377. Piedras Grandes, Woytkowski 127. — Junfn:

Near Tarma, Huassa-huassi, Ruiz & Pavon, type. — Cuzco: Marca-

pata Valley, Weberbauer 7877 (det. Standley). To Colombia. "An-

tagra" (Ruiz & Pavon).

Tournefortia psilostachya HBK. Nov. Gen. & Sp. 3: 78. 1818.

T. floribunda HBK. I.e. 79, at least as to Peru. T. cirrhosa Vaupel,

Bot. Jahrb. 54, Beibl. 119: 3. 1916. T. volubilis L. Sp. PI. 140. 1753,

at least in lit.

Variable in leaf size, form and indument but as interpreted usually

more or less appressed strigillose, especially the younger branchlets

and leaves beneath; petioles a few mm. to 1 cm. long; leaves rather

oblong or narrowly ovate-lanceolate to broadly ovate, obliquely

rounded at acute base, acuminate, sometimes abruptly, usually

5-8 cm. long, about 1.5-2.5 cm. wide or the lower larger; peduncles

in type terminal, the filiform spikes diffuse; calyx strigose, the

lanceolate acuminate segments spreading; corolla long-exserted, seri-

ceous, the throat inflated, the lobes subulate; anthers subsessile near

top of tube, this as long as the filiform style; stigma small, subconic. —

T. floribunda HBK. ex char., type from Colombia, has leaves obtuse

or rounded at base, somewhat canescent sericeous beneath, corolla

tube about 3 mm. long, the lobes "short"; Johnston, Contr. Gray

Herb. 92: 88. 1930, suggested that it once may have had a con-

tinuous range to eastern Brazil; however, the single Peruvian collec-

tion referred to it is more probably a form of T. psilostachya, which

name, long in use, is here retained in preference to the perhaps not

identical one of Linnaeus. Finally, Vaupel's plant, type from Ecua-

dor, has ovate leaves 4-5 cm. wide and may be distinguishable; cf.

T. breviflora. Determinations by Killip. F.M. Negs. 38975; 1043

(T. cirrhosa).

FLORA OF PERU 551

Loreto: Mishuyacu, near Iquitos, Klug 1113; 1213. Nancho,

(Raimondi 3414, det. Herb. Dahlem, T. floribunda). — Puno: Gay.

Ecuador; northern South America?

Tournefortia setacea Killip, Journ. Wash. Acad. Sci. 17: 332.

1927.

A bushy-branched liana, the younger branches, leaves above and

beneath on the nerves appressed strigillose; petioles 0.5-2 cm. long;

leaves subopposite or in threes, oblanceolate or ovate, 5-14 cm. long,

3-7.5 cm. wide, subdecurrent, acuminate, entire, membranous, dark

green above, paler beneath; inflorescence to about 7 cm. wide, few-

branched, shortly peduncled; flowers sessile; sepals linear-setaceous,

4-5 mm. long, about as long as the strigillose corolla tube, the sub-

orbicular corolla-lobes 1.5 mm. long; stamens near base of tube,

anthers linear, 2.5 mm. long; fruit conical, 3-4 mm. long, strigillose,

white. — With the variation that Johnston assigned to T. bicolor Sw.

to which Killip originally referred it; this may not be distinct, but

the author wrote: "the long setaceous sepals distinguish it."

San Martin: Sinchona, Ferreyra 1127. — Huanuco: Cushi, 4829.

Rio Pozuzo, Weberbauer 6737.— Junin: La Merced, 5579, type.

Colonia Perene", Killip & Smith 24993. — Ayacucho: Aina, Killip &

Smith 22733. Estrella near Huanta, Killip & Smith 22668.— Cuzco:

Cerro Mascac, Valle de Lares, (Herrera 1779). Bolivia.

Tournefortia tarmensis (Krause) Macbr., comb. nov. Helio-

tropium tarmense Krause, Bot. Jahrb. 37: 632. 1906.

Liana or the branches clambering, these glabrous in age, early

quadrate, sparsely as leaves both sides, reddish-pilosulous; petioles

4-6 mm. long; leaves oblong- or ovate-elliptic, shortly or subacute

both ends, 3-6 cm. long, 1-3 cm. wide, often somewhat undulate,

nearly smooth; spikes at first corymbosely congested, rather lax in

full flower, 2-5 cm. long; flower limb 7 mm. long; calyx lobes linear,

hispidulous, scarcely 2 mm. long; corolla tube greenish, puberulent

only without, the crisped spreading lobes white; anthers linear, acute,

subsessile; style elongate. — Distinguished by Krause from T. micro-

calyx (R. & P.) Johnst. especially by the much longer flowers. F.M.

Neg. 1062.

Junin: Comas, Weberbauer 6607, in part? (det. Killip, T. ovali-

folia). Palca, Prov. Tarma, Weberbauer 1750, type. Carpapata,

Prov. Tarma, Soukup 3440. Bolivia.

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

Tournefortia ternifolia HBK. Nov. Gen. & Sp. 3: 84. 1818.

Branches terete, sparsely appressed pubescent; petioles 1.5-2 cm.

long; leaves ternate, oblong, narrowed to both ends, acute, about

1.5 dm. long, half as wide, membranous, reticulate-veined, glabrous

or appressed pubescent on nerves beneath; spikes terminal, geminate,

peduncled, less than 2.5 cm. long; flowers pedicellate, secund, ebracte-

ate; calyx strigose, teeth linear-lanceolate; corolla strigose, thrice as

long as calyx (after authors). — Compare T. longifolia R. & P. to

which the Killip and Smith collection, young, may belong, or pos-

sibly there is only one species.

Cajamarca: Tomependa, Bonpland, type. — Amazonas: Chacha-

poyas, Mathews 3009 (det. Killip). — Junin: Tarma, Kittip & Smith

21920 (det. Killip).

Tournefortia Ulei Vaupel, Notizbl. Bot. Gart. Berlin 6: 186.

1914.

Scandent, the younger branchlets sparsely puberulent; petioles

1 cm. long or longer; leaves broadly elliptic or ovate, shortly acumi-

nate, to 13 cm. long, 7 cm. wide, glabrous; inflorescence to 2.5 dm.

long, lax, mostly trichotomous with divaricate branches, with sessile

leaves above the first branches; flowers subsessile, crowded, ap-

pressed puberulent; calyx lobes narrowly triangular, 1-2 mm. long,

3-4 times shorter than corolla tube, this 6 mm. long, slightly inflated,

densely yellowish-brown pubescent; flowers yellow-green, about

7 mm. long; anthers below throat; style nearly as long as tube,

stigma conical; fruit glabrous, ovoid, often crowned with pistil (after

author).— T. andina Britton, Bull. Torrey Club 26: 148. 1899, of

Bolivia with sessile flowers may be comparable. The Peruvian

specimen has glabrate flowers and no leaves in the inflorescence;

compare T. glabra L.

Loreto: Florida, mouth of Rio Zubineta, King 2083 (det. Stand-

ley). Amazon headwaters (Rio Madeira).

Tournefortia undulata R. & P. Fl. Peruv. 2: 25, pi. 149b. 1799.

Rounded shrub or tree-like, to 3 meters tall, the branchlet tips

and young inflorescences reddish pilose, soon glabrous or nearly;

flowering branches densely leafy, stout; leaves broadest at or slightly

above the middle, attenuate into the short sulcate petioles, rather

abruptly acuminate, usually 8-15 cm. long, 3-6 (8) cm. wide, coria-

ceous, the many arcuate-ascending lateral nerves rather obscure

above, prominent with veins beneath, mostly 5-7 mm. distant,

FLORA OF PERU 553

rugose-bullate in age above; inflorescences and branches densely

flowered, suberect, tardily spreading, to 7 cm. long or usually shorter;

calyx segments subulate, about 3 mm. long, to 4 mm. long in fruit;

corolla tube puberulent, 4 mm. long, lobes subrotund, about 1 mm.

long; fruit not lobed, white, about 5 mm. in diameter, style appar-

ently obsolete. — Killip & Smith 21525 from the botanical garden at

Lima was about 3 meters high in 1929. F.M. Neg. 1064.

Huanuco: Mito, 1998. Pillao, Ruiz & Pavdn, type. — Arequipa:

Near Chala, Worth & Morrison 15685. Atiquipa, 1,050 meters,

Tqfalla, type; Weberbauer 7192 (det. Killip). — Puno: Cerros de Islay,

Isern 2476. Ecuador?

Tournefortia virgata R. & P. Fl. Peruv. 2: 25, pi. 150. 1799.

Similar to T. undulata and apparently not well marked, but juve-

nile pubescence more persisting, canescent; leaves smaller, subcoria-

ceous, the fewer lateral nerves less regularly developed; inflorescence

branches recurving and incurving; calyx lobes about linear, scarcely

2 mm. long, little longer in fruit; corolla tube scarcely 4 mm. long,

rather stout, canescent puberulent, the undulate-crenulate lobes

nearly 1.5 mm. broad; fruit white, not lobed, about 4 mm. in diam-

eter, style obsolete. — Description from my collection, the white

flowers (early yellowish) with a heavy, rather unpleasant fragrance;

the many leaves, as in the similar species, are strongly waved or

undulate in life so that they often crease in pressing. The Stork and

Horton specimen in flower is doubtful; corollas about 1 cm. long,

calyx scarcely 2 mm. long.

Huanuco: Mito, 1575 (det. Killip). — Junin: Huassa-haussi, Ruiz &

Pavdn, type. — Apurimac: Pincos, 2,700 meters, Stork & Horton 10695?

2. HELIOTROPIUM L.

Reference: Johnston, Contr. Gray Herb. 81: 1-73. 1928.

Shrubs, rarely herbs, with alternate leaves and white flowers often

borne in scorpoid racemes or spikes. Ovary undivided, the fruit con-

sisting of 4 nutlets or carpels, these remaining entire, biovulate when

ripe, or breaking into 2 uniovulate nutlets at maturity. Corolla

sinuses more or less plaited in bud. Anthers obtuse or acuminate,

often more or less coherent. — Johnston's excellent revision of the

South American species has supplied the following sectional key and

the species key in part, except that vegetative characters have been

emphasized, for convenience. Nidia Gangui has presented an attrac-

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

tive account with illustrations of the genus for the species of Argen-

tina, Univ. Nat. Cordova 11: 481-559. 1955.

SECTIONAL KEY TO HELIOTROPIUM (after Johnston)

Carpels not cleft, finally separate biovulate nutlets.

Carpels with several empty cells besides the 2 fertile; corolla gla-

brous within section Tiaridium (Lehm.) Griseb.

Carpels with 2 cells, both usually fertile.

Corolla villous within; fruit adorned vesicularly.

section Schobera (Scop.) Johnst.

Corolla glabrous at least within; fruit (as carpels) smooth.

Plant prostrate-ascending; carpels often 1-seeded.

section Hypsogenia Johnst.

Erect shrubs; carpels normally 2-seeded.

section Cochranea (Miers) Reiche.

Carpels cleft, finally forming 2 uniovulate nutlets.

Tall shrubs; nutlets rough; anthers ciliate toward tip.

section Heliothamnus Johnst.

Herbs or usually less than 5 dm. tall; nutlets smooth, glabrous or

pubescent; anthers, if pubescent, only at tip.

Stigmatic disk thick, narrow on more or less developed style;

usually erect herbs or shrubs. . .section Orthostachys R. Br.

Stigmatic disk very broad, rather thin, sessile; prostrate, succu-

lent herbs section Halmyrophila Johnst.

SPECIES KEY TO HELIOTROPIUM

Prostrate herbs; leaves small, rarely 2 cm. long, or narrow, succulent.

Succulent leaves soon exceeded by spikes (section Halmyrophila).

H. curassavicum.

Herbaceous leaves longer than inflorescences (section Hypsogenia).

H. microstachyum.

Erect or suberect, usually more or less ligneous; leaves never succu-

lent, usually longer than 2 cm., always exceeded by spikes.

Herbs to ligneous, low; leaves mostly or all about 3, usually several

times longer than wide; nutlets smooth; all section Ortho-

stachys except H. Krauseanum (section Cochranea), perhaps

H. lanceolatum (section Heliothamnus?).

FLORA OF PERU 555

Leaves crenulate, pale puberulent beneath, obtuse; carpels not

cleft H. Krauseanum.

Leaves entire, subconcolor, more or less acute; carpels cleft.

Flowers clearly in bractless spikes.

Corolla rather showy, 4-5 mm. across or tube well exserted;

anthers linear, apically pubescent or glandular except

H. procumbens.

Corolla lobes rounded, tube and calyx subequal.

Plant canescent as calyx within H. Lobbii.

Plant green; corolla glabrous within H. toratense.

Corolla lobes more or less acute, the long tube prominent.

Anthers glandular-puberulent; corolla lobes acute.

H. oxylobum.

Anthers crested with antrorse trichomes.

H. lanceolatum.

Corolla small, 1-3 mm. wide, tube little exserted.

Shrub; leaves subcoriaceous; calyx segments suberect;

flowers yellowish, glabrous H. polyanthellum.

Herbs or half-shrubs; leaves herbaceous unless oldest;

calyx segments soon lax; flowers white.

Anthers (as corolla within) pubescent, linear, obtuse;

leaves slightly narrowed apically, acute.

H. pilosum.

Anthers glabrous, subovate, connective extended;

leaves, at least mostly, rounded apically, mucron-

ulate H. procumbens.

Flowers scattered on unevenly foliose branchlets . .H. piurense.

Tall and (or) ligneous; leaves mostly or all about twice longer than

wide except the subovoid H. rufipilum, sometimes H. angio-

spermum and the canescent shrubs H. erianthum, H. lippi-

oides; nutlets rough, lacunose or sculptured except the only

herbaceous species, H. indicum.

Herb, sparsely setulose-hispid; leaves decurrent on long petioles;

fruit lobed, nutlets smooth (section Tiaridium).

H. indicum.

Shrubs or ligneous below; indument, at least in part, pilose or

crinkled; petioles short; nutlets rough (section Heliothamnus

except H. angiospermum) .

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

Carpels rough with obtuse vesicles, not cleft; leaves membra-

nous, sparsely pilose as stems or glabrate; corolla villous

within, small, tube and calyx subequal (section Schobera) .

H. angiospermum.

Carpels usually lacunose or sculptured, cleft; characters other

than above at least in part (section Helioihamnus).

Leaves membranous, concolored, not rugose, often shorter

than 5 cm.; corolla tube well exserted from the 1-

1.5 mm. long calyx H. adenogynum.

Leaves subcoriaceous to membranous, rugose-bullate espe-

cially when shorter than 5 cm.

Leaves not or obscurely bullate but veins more or less

obvious; corolla tube exserted.

Inflorescence lax, the branches soon several cm. long;

style shorter than stigma H . rufipilum.

Inflorescence congested, branchlets nearly 2-3 cm.

long; style longer than stigma. . . .H. arborescens.

Leaves usually firm and more or less bullate or (and)

gray pubescent beneath; corolla tube and calyx sub-

equal or leaf-character pronounced.

Indument, especially flowering branchlets, somewhat

fulvous; style shorter than stigma.

Calyx to 2 mm. long, longer in fruit.

Leaves 3-6 cm. long, oblong- or rarely subovate-

elliptic, acute or subacute . . . H. Urbanianum.

Leaves ample, mostly ovate-acuminate, often

about a dm. long or longer H. submolle.

Calyx inconspicuous, at most 1.5 mm. long in flower,

little longer in fruit H. Mandonii.

Indument, even of branchlets, canescent; style longer

than stigma.

Leaves lanceolate, 4-5 cm. long, 10-17 mm. wide.

H. erianthum.

Leaves ovate or ovate-elliptic, about twice as long

as wide H. incanum.

Leaves elliptic, 1.5-2 cm. long H. lippioides.

Heliotropium adenogynum Johnst. Contr. Gray Herb. 81 : 42.

1928.

FLORA OF PERU 557

Laxly branched, more or less villous and usually glanduliferous,

even the calyces, these globose, finally 2 mm. long, the elliptic-lanceo-

late erect or connivent lobes somewhat persisting; petioles 3-8 mm.

long; leaves herbaceous, venose, ovate or oblong-lanceolate, 2-5 cm.

long, acute or obtuse at base, acute at apex, 2-5 cm. long, about

0.5-3 cm. wide, paler beneath, entire; spikes 4-10 cm. long; corymb

lax; corolla sparsely villous only without, not rarely glanduliferous,

2-3 times longer than calyx, to 4 mm. across, lobes orbicular; anther

cells pubescent toward tip; ovary glanduliferous; style obsolete,

stigma about 0.8 mm. long; nutlets ovate, rugose, 1-1.5 mm. long,

stipitate-glandular. — Remarkably distinct by small flowers, gland-

ular ovary and fruit, scarcely accrescent more or less deciduous

calyces; most closely related to H. Mandonii Johnst. and evidently

also to H. rufipilum (Benth.) Johnst. (author), a species varying in

some if not all of these characters. Section Heliothamnus Johnst.

Lima: Chosica, 495, type; Weberbauer 5312; (Wilkes Exped.}.

Heliotropium angiospermum Murray, Prodr. Stirp. Gott. 217.

1770; 10. H. parviflorum L. Mant. 2: 201. 1771. H. synzystachyum

R. & P. Fl. Peruv. 2: 3, pi. 109a. 1799. Tournefortia synzystachya

(R. & P.) R. & P. Syst. 4: 539. 1819. Synzystachyum peruvianum Raf.

Sylva Tell. 89. 1838. H. simplex Meyen, Reise 1: 436. 1834, fide

Johnston, Journ. Arnold Arb. 18: 18. 1937.

Perennial, sometimes flowering as an annual, rather sparsely

pubescent, erect or becoming lax or prostrate, sometimes a meter

long; petioles often to 1 cm. long or longer; leaves entire, oblong-

lanceolate to elliptic, attenuate to base, acute, small or large (2-3 cm.

long, 1-1.5 cm. wide, or 1.5 dm. long, 5 cm. wide); spikes solitary or

binate, 5-10 (15) cm. long, ebracteate; calyx lobes acute; corolla

white, glabrous but barbate within, about 2 mm. wide, the tube

subincluded in the calyx; fruit depressed, 3-4 mm. across. — Section

Schobera (Scop.) Johnst. I.e. 10. Fruit didymous (2-seeded carpels)

and covered with unique vesicular structures.

Piura: Rocky slopes, Cerro Prieto, (Haught & Svenson 11602}.

Talara, Haught 38; Norton 11592. — La Libertad: Barranco near

Pacasmayo, (Forbes). — San Martin: Tarapoto, Woytkowski 35200.—

Lima: Chosica, Mexia 04087. Atocongo, Pennell 14458a. Rio Chil-

lon, Pennell 14458. Lurin, Pennell 12217. Santa Clara, Rose 18626.

Callao to Lima, 52; Watkins 396 (and many others). Chancay,

Vargas 1235; Goodspeed 33033; Ruiz & Pavdn (type, H. synzystachy-

um).— Huancavelica: Near Pampano, Weberbauer 5394 (part). — lea:

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

Above Pisco, 1,550 meters, Weberbauer 539 4. — Apurimac: Rio Pacha-

chaca, Goodspeed Exped. 10537. — Arequipa: Near Arequipa, Meyen.

Murillo Bay, (Macrae). — Tacna: Arica, (Meyen, type, H. simplex).

Bolivia to Central America, West Indies, Texas, Florida. "Alacran."

Heliotropium arborescens L. Syst. ed. 10. 913. 1759; 39.

H. peruvianum L. Sp. PI. ed. 2. 187. 1762. H. corymbosum R. & P.

Fl. Peruv. 2: 2, pi. 107a. 1799.

Shrub, sometimes 2 meters tall, usually much lower, with many

short branchlets with lavender-purple flowers; leaves typically ovate-

or oblong-elliptic, more or less decurrent on the short or rarely 1-

2 cm. long petioles, acute, ordinarily 4-8 cm. long, about half as wide

or sometimes considerably larger, membranous or nearly, paler and

venose beneath, more or less pubescent or glabrate above when

smooth or inconspicuously bullate-rugulose in age; inflorescences 3-

10 cm. across, lightly to densely strigose or villous, the short branch-

lets congested even in fruit; calyx segments subulate-linear, about

3-3.5 mm. long, not accrescent, the corolla tube about twice as long,

appressed strigulose, lobes glabrate, rounded, 4-5 mm. across at top;

anthers elongate, crested dorsally by a wavy antrorse trichome

(Johnston); style longer than stigma; carpels pitted (lacunose), ellip-

soid, medially and ventrally affixed by a definite elliptical areola

(Johnston). — Var. grisellum Johnst. I.e. 40, differs in the longer,

more canescent indument, the leaves usually rugose and more canes-

cent beneath (all the southern Peru specimens; cf. H. Mandonii).

As Johnston remarks, the Ancash specimens are not typical. An

attractive flowering shrub forming large mounds or clumps. The

native name "Vaynilla" refers to the fragrance (Ruiz & Pavon).

Section Heliothamnus Johnst. Illustrated, Bot. Mag. pi. 1609.

Ancash: Below Ocros, 2,650 meters, Weberbauer 2710?; Soukup (t).

—Lima: Matucana, 2938; Rose 18639. Ambar, Stork 11457. Canta,

Pennell 14352; 14759; Mathews, Savatier. Amancaes, Weberbauer

1637; Soukup 3125. Near Lima, Ruiz & Pavon, (type, H. corym-

bosum); Dombey. — Junin: Huancayo, Oxapampa, Soukup 2438. —

Ayacucho: Coracora, 2,900 meters, Weberbauer 5799. — Arequipa:

Posco, (Cook & Gilbert 40). Mollendo, Weberbauer 1466; Hitchcock

22394; West 8206. South of Chala, Worth & Morrison 15686. Que-

quena, Eyerdam & Beetle 22180. — Moquegua: Carumas, Weberbauer

7272, (type, var.).— Tacna: Without locality, (Shephard). "Docto"

(Beetle), "vaynilla."

FLORA OF PERU 559

Heliotropium curassavicum L. Sp. PI. 1: 130. 1753; 14. H.

curassavicum L. var. parviflorum Ball, Journ. Linn. Soc. 21 : 227. 1884.

A glabrous glaucous succulent, the branching prostrate stems

sometimes elongating to a meter or so; leaves oblanceolate or obo-

vate, often 2-4 cm. long, 0.5-1 cm. wide; spikes (bractless) often

geminate; corolla typically 1-2.5 mm. long; anthers free, sagittate,

the cells acuminate-appendaged; stigma column in Peruvian plants

less than 0.2 mm. long, fruit 4-lobate, disk prominent, stigma broad.

— An unmistakable plant of wet or low places especially when there

is salinity; the white corollas are usually yellow at throat and some-

what purplish in age as sometimes the entire plant. Section Halmy-

rophila Johnst.

Piura: Alto de la Cruz, Stork & Norton 11355. — La Libertad:

Toward Salaverry, Worth 8891. — Lambayeque: Chiclayo, Stork 11 430.

— Lima: Callao, 5881. Rio Rimac, (Ball) and many other collections,

—lea: Paracas Bay near Pisco, (Forbes). — Arequipa: Lomas, Weber-

bauer 5732. Mollendo, Weberbauer 388; Eyerdam 25165. — Tacna:

Near Tacna, Woitschach; Werdermann 711. Chile and Patagonia to

United States; West Indies.

Heliotropium erianthum Johnst. Contr. Gray Herb. 81: 41.

1928.

A canescent shrub, the branches and dense corymbs tomentose-

villous, the leaves strigose-tomentulose above, tomentose beneath;

petioles 5-8 mm. long; leaves lanceolate, acute both ends, 4-5 cm.

long, 1- about 2 cm. wide, tessellate-rugose with impressed nerves

above, obscurely crenulate, scarcely revolute; spikes 2-5 cm. long;

calyx as corolla without appressed villous, the former orange, finally

4-5 mm. long, the lobes lanceolate-subulate, the latter 2.5-3 mm.

long, limb about 3 mm. across, lobes rounded; anthers linear-oblong,

the cells antrorsely pubescent dorsally toward tips; ovary glabrous,

style 1-1.3 mm. long, stigma about 0.8 mm. long. — Resembles H. in-

canum R. & P. with harsh ascending indument, larger flowers; also

H. argenteum Lehm., 42, of southern Ecuador, with slightly shorter

style and ovate, less rugose leaves (author). With more collections

it seems possible that Lehmann's species may be found to vary in

these characters and, admitting any variation, the shrub will prob-

ably be shown to be a part of the Ruiz and Pavon species. Section

Heliothamnus Johnst.

Piura: Huancabamba, Weberbauer 6332, type.

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

Heliotropium incanum R. & P. Fl. Peniv. 2: 2, pi. 108a.

1799; 41.

Shrub, often sprawling or openly branched, a meter or so high,

with rather showy fragrant white or sometimes purplish flowers and

brick red ovate or ovate-elliptic leaves, typically soon very rugose-

bullate and green above, canescently pubescent and reticulate-venose

beneath; leaves narrowed to the scarcely obvious petioles, acute,

ordinarily 4-6 cm. long, about half as wide, thick, more or less sub-

appressed hispid above, glabrate in age, densely and persistently vil-

lous tomentose beneath; corolla about 4 mm. across expanded top

or somewhat broader; calyx lobes subulate-lanceolate, strigose; nut-

lets nearly 2 mm. long, faintly sculptured; otherwise like the similar

H. arborescens which some forms approach closely (Johnston). —

Section Heliothamnus Johnst. F.M. Neg. 17326.

Cajamarca: Chota, Stark & Horton 10041 (det. Standley). — Huan-

uco: Near Huanuco, 3237; Ruiz & Pavdn, type. Huacho, Stork &

Horton 9405. Ambo, 2423. — Junin: Cabello above Huertas, 1327.

La Oroya, Kalenborn 75. — Huancavelica: Mejorada, Stork & Horton

10916; Weberbauer 7604. — Apurimac: Oropeza Valley, Vargas 9768

(det. Standley). Pincos, Stork & Horton 10674. Rio Pampas, 2,950

meters, Weberbauer 5855. — Cuzco: Ollantaitambo, Hen era 231; Pen-

nell 13640. Vilcanota Valley, H err era 1080. Urubamba Valley,

Herrera 1367; 1533.

Heliotropium indicum L. Sp. PI. 130. 1753; 19.

Sparsely hispid-setulose annual but often vigorous, several dm. to

a meter tall, branching above; petioles alate toward leaf -blade, usu-

ally more than half as long as the ovate-elliptic to broadly lanceolate

herbaceous venose leaves, these several cm. to a dm. long or longer,

acute both ends or more or less cordate at base; spikes bractless,

solitary, scorpoid, often becoming 2-3 dm. long; calyx lobes narrow,

2-2.5 mm. long; corolla blue or violet, sometimes white, the tube

2.5-4.5 mm. long, spreading limb 2-4 mm. across; anthers free; style

short but slender; fruit glabrous, prominent lobes divergent, the

eventually free nutlets angled, 2-3 mm. long. — Section Tiaridium

(Lehm.) Griseb.; defined by its conspicuously ribbed multicellular

glabrous fruit. Usually a weed in clearings; only a few collections

cited. Illustrated, Univ. Nat. Cordova 11: pi. 8, opp. page 524.

Piura: Near Piura, (Townsend 1393}. — San Martin: San Roque,

Williams 7801. Juanjui, Klug 3436. Tarapoto, Woytkowski 33542.

FLORA OF PERU 561

— Loreto: Soledad, Tessmann 5260. Near Iquitos, Williams 1312;

139 b. Tropics and subtropics.

Heliotropium Krauseanum Fedde, Bot. Jahresb. 34, pt. 3: 72.

1908 ; 28. H. saxatile Krause, Bot. Jahrb. 37 : 633. 1906, not Brandg.,

1905.

Erect with arcuate-spreading flowering branches, to a meter tall,

the densely leafy branchlets early somewhat pilosulous; leaves sessile

but long and often narrowly attenuate to base, rounded at tip, oblong

or oblanceolate, 2-5 cm. long, 4-8 mm. wide, crenulate, subcoria-

ceous, finely reticulate-veined and glabrate above, canescent puberu-

lent-hispidulous beneath; spikes congested and densely flowered,

3-4 cm. long in fruit; calyx lobes linear, acute, minutely hispidulous,

the nearly twice as long corolla tube sparsely so; corolla lobes white,

glabrous, nearly triangular; filaments ^ery short; style subequaling

calyx. — Meritoriously named for a genial botanist of the Dahlem

herbarium. Related to H. taltalense (Phil.) Johnst. of Chile, coarser

with somewhat larger flowers, more pubescent calyces, larger fruits;

may be a variety; resinous and pleasantly scented according to

Johnston, who placed it in section Cochranea (Miers) Reiche. F.M.

Neg. 17327.

Arequipa: Mollendo, Johnston 3533; Weberbauer 1552, type; 147.

Atiquipa to the port of Chala, Weberbauer 7188; Worth & Morrison

15647. Chile.

Heliotropium lanceolatum R. & P. Fl. Peruv. 2: 4, pi. Ilia.

1799; 40.

Shrub, the type with slender strigulose and sparsely hispidulous

flowering branchlets; leaves crowded, oblong-lanceolate, narrowed to

short petioles, acute or acuminate, mostly 2-3.5 cm. long, 4-6 mm.

wide, rather densely appressed asperous hispidulous and rugulose

above, crinkly puberulent and on the prominent nerves hispidulous

beneath; flowers crowded, the inflorescence branchlets 1-2 cm. long;

calyx segments linear-subulate, about 3 mm. long, sparsely strigulose

as the half again as long corolla tube; corolla lobes rounded or sub-

acute, the expanded top about 4 mm. across. — Characterized by

small scabrous lanceolate leaves and lax corymb of long-tubed flow-

ers; type locality listed as Huanuco in Ruiz and Pavon's journal but

not found since, so perhaps that name referred to another species, as

H. incanum; the anthers, according to the sectional character (fide

Johnston) are crested dorsally by a row of crowded antrorse trichomes;

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

1 have not dissected the one flower on type material seen. Section

Heliothamnus Johnst. F.M. Neg. 17328.

Lima: Lurin, (Ortega); (Wilkes Exped.}; (Nation). Toward Ma-

tucana, Weberbauer 96. — Huanuco: Pillao and near Huanuco(?),

Ruiz & Pavdn, type.

Heliotropium lippioides Krause, Bot. Jahrb. 37: 632. 1904; 42.

Suffruticose, the few slender strict erect branches brownish and gla-

brate below, toward the tip as the sparse leaves beneath more or less

ashy tomentose; leaves sessile or essentially, elliptic, shortly narrowed

at base, apically rounded but often mucronulate, 1.5-2 cm. long,

about 8 mm. wide, coriaceous, revolute, grayish pilose above, strongly

rugose by the impressed nerves; spikes 2-3 cm. long, densely flow-

ered, alternately geminate; calyx hispid, the linear lobes somewhat

spreading in fruit; corolla tube hispidulous, white, 6 mm. long or

about 3 times longer than calyx, lobes undulate, subglabrous; fila-

ments almost 1 mm. long; style short, conical, the stigma obtuse. —

Seems to be nearest H. incanum R. & P. but leaves much smaller

(author). Type was a meter tall. Section Heliothamnus Johnst.

F.M. Neg. 17330.

Cajamarca: Santa Cruz, 1,650 meters, Weberbauer 4122, type; 189.

Heliotropium Lobbii Johnst. Contr. Gray Herb. 81: 54. 1928.

Canescently villous-hispid shrub; branches ascending, to 1.5 dm.

long; leaves elliptic-oblanceolate, narrowed to petioles (these 1-

2 mm. long), acute, 6-17 mm. long, 2.5-6.5 mm. wide, firm, con-

spicuously pale tomentose beneath, only midnerve prominent; spikes

geminate, ebracteate, 1-2 cm. long; pedicels stout, to 1 mm. long;

calyx (as corolla without) strigose, 2.5-3 mm. long, the subequal

lobes linear-lanceolate; corolla funnelform, 4-5 mm. long, sparsely

long-strigose in throat, cylindric tube 3 mm. long, limb 2.5-3 mm.

across, lobes ovate; anthers sessile, obtuse, glandular; stigma cylin-

dric, to 1 mm. long, 2-3 times longer than style. — Habit of H. pilo-

sum R. & P. with corolla smaller and notably different in form

(author). Section Orthostachys R. Br.

Amazonas: Chachapoyas, (Lobb, type).

Heliotropium Mandonii Johnst. Contr. Gray Herb. 81: 43.

1928.

Becoming a meter or two high, the laxly branched branches early

fulvous-villous (trichomes retrorse), finally glabrescent; leaves Ian-

FLORA OF PERU 563

ceolate or oblong-elliptic, reduced into villous petioles about 1 cm.

long, acute, 7-14 cm. long, 2.5-6 cm. wide, scabrous, sparsely villous-

hispid and often by the impressed veins strongly reticulate-rugose

above, paler or canescently velutinous beneath; spikes villosulous, to

a dm. long, in a terminal dichotomous corymb; calyx sessile, slightly

villous, often glandular, globose, finally 2-2.5 mm. long with the

ovate-lanceolate acuminate lobes little plicate in the ovate sinuses;

corolla white, 3-4.5 mm. long, pubescent (often glandular) only with-

out, the cylindric tube to two times longer than calyx, limb 3-5 mm.

across, lobes obtuse; anthers densely pubescent toward apex; stigma

sessile, 0.5 mm. long; ovary eglandular; nutlets 1.5 mm. long, lacu-

nose, compressed, broadly affixed. — A coarse species very distinct

from its relatives and suggesting a Tournefortia in gross aspect

(author) ; simulates H. arborescens var. grisellum Johnst. but stigma

sessile and inflorescence in age distinctive; flowers fragrant in my

collections. Type is Mandon 387 from La Paz, Bolivia. Section

Heliothamnus.

Huanuco: Yanano, 1,800 meters, 3660. Rio Santa Domingo,

1,200 meters, 4206— Cuzco: Valle de Santa Ana, (Herrera 899).

Bolivia.

Heliotropium microstachyum R. & P. Fl. Peruv. 2: 3, pi.

llOb. 1799; 12. H. brachystachyum (DC.) Griseb. Gott. Abh. 24:

271. 1879. Heliophytum brachystachyum DC. Prodr. 9: 554. 1845.

A lightly strigulose perennial developing a ligneous caudex, the

slender prostrate stems with glomerate inflorescences in the axils of

the upper branchlets or leaves, these elliptic or suboblong, attenuate

to the slender, usually short petioles, rounded or obtuse at apex,

ordinarily 1.5-2.5 cm. long, 5-10 mm. wide, sometimes smaller or

considerably longer, often glabrous or glabrate in age; nerves slender,

impressed above, rather prominent beneath; inflorescences in fruit

more or less spicate, a cm. or two long; calyx segments linear, about

2 mm. long; corolla 2-3 mm. across at top, glabrous, tube included,

lobes rounded; carpels 2-celled, frequently maturing only a single

seed (Johnston), lacunose, nearly 2 mm. long. — Johnston (who dis-

covered the identity of the above names) noted that the upper plant

figured on Ruiz and Pavon's plate 110 is by error labeled H. pilosum,

a confusion in legend occurring elsewhere in their Flora of Peru.

Section Hypsogenia Johnst. Illustrated, Univ. Nat. Cordova 11:

pi 3, opp. page 506. F.M. Neg. 17333.

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

Lima: Matucana, 202. — Huanuco: Near Huanuco, Dombey; Kane-

hira 258. Ambo, 8168. — Junin: Tarma, Weberbauer 2370 (distr. as

H. paronichioides) ; 176; Ruiz & Pavdn, type. — Huancavelica: Huay-

tara, Weberbauer 54-27, part. — lea: Above Pisco, 2,650 meters, Weber-

bauer 5427. — Cuzco: Ollantaitambo, Hen era 2254; Pennell 13660.

Near Cuzco, Vargas 654. — Arequipa: Cotahuasi, Weberbauer 6871. —

Puno: Near Puno, (Shepard 120). Chuquibambilla, Pennell 13358.

Altos de Toledo, 4,650 meters, Meyen. — Moquegua: Carumas, Weber-

bauer 7484- Ecuador to Argentina.

Heliotropium oxylobum Johnst. Contr. Gray Herb. 81: 57.

1928.

A shrub with many slender branches, sometimes 6 dm. tall,

sparsely strigose with appressed curved trichomes; leaves lanceolate-

linear or oblanceolate, narrowed to base (petioles 1 mm. long), acute,

1-2.5 cm. long, 1.5-4.5 mm. wide, firm, enervose, pale beneath;

spikes single or paired, ebracteate, 2-6 cm. long; calyx subsessile,

1-2 mm. long, the sublinear lobes subequal; corolla yellowish, densely

strigose without and slightly short-villous within above the middle,

4-5 mm. long, the cylindric tube 3-4 mm. long, lobes ovate-deltoid,

acute; anthers to 1 mm. long, obtuse, glandular adhering; ovary

glabrous or puberulent; stigma about 1 mm. long with thick disk,

to two times longer than style; nutlets 1 mm. long, convex, inner

face angulate. — Differs from H. campestre Griseb., known from La Paz

southward, in its more slender habit, smaller leaves, more appressed

indument and pubescence within the corolla. Simulates H. lance-

olatum but apparently differs in anther character. Corolla yellowish

or tube purplish (Goodspeed Exped.). Section Orthostachys R. Br.

Huancavelica: Rio Mantaro below Surcubamba, Weberbauer 6486.

— Apurimac: Bridge of Apurimac, (Pearce, type). Rio Pachachaca,

Goodspeed Exped. 10544 (det. Standley, H. pilosum). Pincos, Stork

& Horton 10654 (det. Standley, H. pilosum). — Cuzco: Mayhua, Prov.

Paruro, Vargas 890 (det. Standley) ; 9735 (det. Standley, H. pilosum).

Heliotropium pilosum R. & P. Fl. Peruv. 2: 3, pi. HOa. 1799;

56.

Prostrate or spreading-ascending, sometimes a meter high, at

least the tips canescent with more or less appressed pubescence;

caudex ligneous as the prostrate-ascending older stems and few

branches, the herbaceous younger with soft greenish lanceolate-

elliptic leaves, attenuate to short petioles, acute, often to 3 cm. long,

FLORA OF PERU 565

to about a third as wide, the older flowering branches usually with

narrowly oblong-lanceolate leaves, frequently firmer, acuminate, 2-5

cm. long, 5 mm. wide or narrower; inflorescence often composed of

2-3 spikes finally 5 cm. long or longer; calyx segments subulate-

linear, 1.5 mm. long, little longer in fruit, lax; corolla almost minute,

tube within and without strigulose, not at all or slightly exserted,

lobes rounded. — Var. albatum Johnst. is merely a form with an

abundant appressed pubescence (author). Plate 11 Ob, not a as cap-

tioned, is obviously this species; details incorrect; nutlets are 1 mm.

long, sparsely hirsute-strigose, convex, the face distinctly acute;

corolla is subsalverf orm with stout cylindric tube not surpassing calyx

lobes. Section Orthostachys R. Br. F.M. Neg. 17339.

Ancash: Lomas de La Chay, Goodspeed Exped. 9214- — Lima:

Hills of Lima, (Nation). Rio Rimac, (Ball, type, var.); Metcalf

30249. Surco, 1,950 meters, Weberbauer 5204 (var.). Atocongo,

Pennell 14786. Chosica, Weberbauer 5319. Clays near Barranca,

Weberbauer 1649; 147. Sandy seaside hills, 5922. Amancaes, Ruiz &

Pavon, type. Lurin, 5922. Sayan, Goodspeed 33043. — Arequipa:

East of Chala, Worth & Morrison (var.). Laspinas, 2,200 meters,

Eyerdam & Beetle 22159 (var.).

Heliotropium piurense Johnst. Amer. Journ. Bot. 33: 479.

1946.

Slender erect annual 1-2 dm. high, stem at base 1.5 mm. thick,

dichotomously branched above; sparsely appressed strigose with

curved antrorse trichomes; leaves oblanceolate, long-narrowed to

slender petioled (1-2 mm. long), acute, 1-1.5 cm. long, 2-3 mm. wide

above the middle, minutely strigose, enervate; spikes terminal,

slender, 5-10 cm. long or longer, leafy-bracted; pedicels 0.5-1 mm.

long; calyx sparsely strigose, the segments unequal, erect or sub-

erect, lanceolate, acute, 2-2.5 mm. long, in fruit 3-3.5 mm. long;

corolla white, cylindric, 2-4 mm. long, a little strigillose (only out-

side), lobes oblong, erect or nearly, little exceeding calyx; anthers

extended apically, cohering; stigma broadly conical, to 3 mm. high

and broad, subsessile; nutlets about 1 mm. high, ventrally angulate,

pored, well-convex dorsally and pubescent with uncinate trichomes.

—"Keys" in the author's revision, Contr. Gray Herb. 81: 49. 1928,

to the southern H. ocellatum Cham., the calyx not uncinate pubes-

cent; in this character resembles the northern H. fruticosum L. that

is less slender and with a definite style (author).

Piura: Among sparse grasses, Cerro Prieto, H aught & Svenson

11553, type. Cabo Blanco, (Haught 256).

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

Heliotropium polyanthellum Johnst. Contr. Gray Herb. 81:

56. 1928.

Type a lax light green bush about 5 dm. tall, the many slender

branches densely strigose; leaves narrowly lanceolate or oblanceolate,

to about 1 cm. long, 2-4 mm. wide, gradually attenuate into the

1 mm. long petiole, acute, subcoriaceous, not veiny but midrib strong

beneath; spikes single or geminate, slender, ebracteate, 4-6 cm. long;

calyx subsessile, finally 1.5-2 mm. long, the slender lobes unequal;

corolla yellowish, strigose only without, the cylindric tube 2-2.5 mm.

long, less than 1 mm. thick, the limb 2-2.5 mm. across, lobes ovate;

anthers linear, obtuse, sessile, lightly coherent by the glandulosity;

ovary sparsely villosulous; stigma cylindric, about 0.8 mm., style

obsolete; nutlets about 1 mm. long, strigose. — Relationship unknown

(author). Section Orthostachys R. Br.

Cajamarca: Between Jaen and Rio Shumba, 750 meters, Weber-

bauer 6184, type.

Heliotropium procumbens Mill. Diet. ed. 8, no. 10. 1768; 52'

H. inundatum Sw. Prodr. 40. 1788.

Flowering as an annual but sometimes more or less enduring and

ligneous toward base, green or subcanescent strigose; leaves often

1-2 cm. long, about half as wide, elliptic or somewhat obovate, nar-

rowed to the much shorter petiole, rounded and mucronate at apex;

spikes usually geminate, soon several cm. long; calyx segments sub-

ulate linear, about 1.5 mm. long, lax in fruit and 2.5 mm. long;

corolla inconspicuous, the strigose tube slightly exserted; nutlets

strigulose but smooth. — See Johnston for complete synonymy, as

elsewhere; to it he added, after seeing types (Contr. Gray Herb. 92:

89. 1930), H. americanum Mill., I.e. no. 11, H. brasilianum Roth,

Nov. PL Sp. 103. 1821, and H. rigidulum DC., Prodr. 9: 540. 1845;

he notes species apparently sometimes perennial (Bolivian plateau)

and flowers larger in southern part of range. Section Orthostachys

R. Br. Illustrated, Univ. Nat. Cordova 11: pi. 12, opp. page 538.

Piura: Near Piura, Spruce. — Loreto: Middle Rio Ucayali, Tess-

mann 3111. Argentina to United States.

Heliotropium rufipilum (Benth.) Johnst. Contr. Gray Herb.

81: 44. 1928. Tournefortia rufipila Benth. Bot. Sulph. 140. 1844.

Slender, sometimes a meter or two tall, the long stems or branches

especially above more or less fulvous hispid with often somewhat

viscid and usually spreading trichomes; leaves narrowly ovate-

FLORA OF PERU 567

elliptic, acute both ends or more commonly long-decurrent on short

or elongate petioles and acuminate at apex, frequently about a dm.

long, a third as wide or much larger, membranous, obscurely veined

except in age when somewhat rugulose above, more or less hispid-

ulous; inflorescences lax in age, the branches 4-15 cm. long; calyx

segments subulate, 2 mm. long or about 3 mm. long in fruit, the

lightly strigulose corolla tube slightly exserted; corolla lobes rounded,

the flower 3-5 mm. across at expanded top. — There is considerable

difference in leaf-form and flower-size between the Haught speci-

mens determined by Johnston and others determined by him from

Ecuador; typically with herbaceous leaves, veins little impressed,

branches lax, spikes elongate, nutlets densely long-stiped glandular;

var. anadenum Johnst., I.e., is a form or race with nutlets sparsely

if at all glandular. Compare H. submolle Klotzsch and H. Urbani-

anum Krause, which will prove, probably, to be a part of this

species; H. adenogynum Johnst. is scarcely more than a variant;

Croton, Lantana and other Andean shrubs respond to local condi-

tions similarly. Section Heliothamnus Johnst.

Piura: Talara, Haught 70. Parinas Valley, Haught 169. Cerro

Viento, Haught 77 (var.). — San Martin: San Roque, Williams 7804

(det. Standley). Zepelacio, Klug 3488 (det. Standley). — Junin:

Oxapampa, Soukup 1810 (det. Ewan). Bolivia to Mexico.

Heliotropium tora tense Johnst. Contr. Gray Herb. 81: 55.

1928.

Shrubby at base, decumbent-ascending, 1.5-2 dm. high, the many

stems to 3 dm. long, spreading villosulous below, appressed pubes-

cent to tips; leaves oblanceolate, gradually attenuate to base, acute,

3-6 cm. long, 4-8 mm. wide, herbaceous, veinless, obscurely strigil-

lose, narrowly revolute; spikes terminal, geminate or ternate, 3-7 cm.

long; calyx subsessile, 3-4 (finally 5) mm. long, sparsely strigose, the

linear-acuminate lobes unequal; corolla about 4 mm. long, tube

strigose only without, 2-3 mm. long, the limb 4.5 mm. across;

anthers subulate, acuminate, obscurely granulate; stigma 0.3-0.4

mm. long, 4-lobate, subsessile; nutlets villous, 1.7 mm. long, convex,

the inner face angulate. — Well-marked species of uncertain relation-

ship (author). Section Orthostachys R. Br.

Moquegua: Torata, Weberbauer 7407, type.

Heliotropium Urbanianum Krause, Bot. Jahrb. 37: 633. 1906;

45.

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

Flowering branches, especially toward apex and crowded inflor-

escences, conspicuously fulvous villous-hispid as the younger leaves

beneath, the older canescent; petioles 3-5 mm. long; leaves elliptic

or early ovate-elliptic, acute at base, rounded or sometimes sub-

acute at apex, usually 3.5-5 cm. long, about 2 cm. wide; inflorescences

little longer than wide even in fruit, ordinarily only a few cm. long;

calyx segments narrowly ovate-subulate, 2-2.5 mm. long, hispid-

ulous; corolla blue, including the rounded lobes, more or less pilose,

the stout tube barely exserted, the expanded flower about 4 mm.

across; anthers 2 mm. long; style elongate-conical, shorter than

stigma. — Distinguished from H. rufipilum at a glance (Johnston)

but, as monographer remarks, characters mostly relatively devel-

oped; spikes and corolla-tubes usually stouter, leaves smaller, usually

rugose and broadest at or above the middle, more pubescent, the

trichomes often bulbous based, and, especially, calyx does not be-

come globose. Section Heliothamnus Johnst. F.M. Neg. 17349.

Amazonas: Chachapoyas, (Mathews); perhaps same locality,

(Lo66). Ecuador.

3. COLDENIA L.

Tiquilia Pers. Syn. 1: 157. 1805.

References: Johnston, Contr. Gray Herb. 70: 58-61. 1924; Journ.

Arnold Arb. 16: 183-185. 1935.

Annual or perennial, sometimes more or less ligneous, often

spreading or prostrate and the veiny or evenose leaves approximate

or subtending the small flowers. Calyx 4-5-parted. Corolla funnel-

form or salverform, appendaged or smooth. Style 2-cleft or 2-parted.

Ovary entire or 4-lobed, in fruit 4, rarely 2, separate nutlets, these

attached basally, usually ventrally.

C. Nuttallii Hook, of western North America has been found in

the Andes of Argentina according to Johnston, I.e. 75: 43. 1925; it is

an annual with smooth ovoid nutlets, appendaged corollas. Most

specimens being in flower, Johnston's key, which follows, is supple-

mented.

Nutlets plano-convex; stamens exserted C. dichotoma.

Nutlets spheroid to ovoid.

Nutlets narrowly ovoid with ventral scar; petioles usually villous-

ciliate C. paronychioides.

Nutlets spherical; petioles not villous.

FLORA OF PERU 569

Corolla about 5 mm. long; stamens included C. elongata.

Corolla about 1 cm. long; stamens exserted C. conspicua.

SUPPLEMENTARY KEY

Flowers very obvious; stamens well-exserted; nutlets subspherical.

C. conspicua.

Flowers more or less obscure; stamens slightly or not exserted.

Leaves about 5-7 mm. long; nutlets narrowly ovoid.

C. paronychioides.

Leaves mostly or all about 1 cm. long or longer; nutlets sub-

spherical to plano-convex.

Stamens included; nutlets spherical C. elongata.

Stamens visible; nutlets plano-convex C. dichotoma.

Coldenia conspicua Johnst. Journ. Arnold Arb. 16: 183. 1935.

Prostrate, the laxly branched stems sometimes a dm. long or

longer, the younger finely hispidulous and more or less glanduliferous

as the setose petioles (2-9 mm. long) and hispid calyces; leaves

broadly lanceolate or elliptic, entire (or essentially), 5-13 mm. long,

2-5 mm. wide, closely and coarsely pustulate-asperous above, finely

and erectly so beneath; calyx lobes linear, subindurate at base, about

9 mm. long, in fruit to 1.5 cm. long; corolla blue, tube 9 mm. long,

limb 10-12 mm. across; throat smooth; stamens exserted; nutlets

early about the top of a subcylindrical gynobase, this finally ex-

panded, turbinate and the nutlets borne basally in the apical tissue

by means of strophiodes. — Nutlets quite similar in size, shape and

markings to those of section Sphaerocarya Johnst. Contr. Gray

Herb. 70: 57. 1924. Johnston once referred the Pennell collections

to the Chilean C. grandiflora Phil.; these appear to be annual or possi-

bly biennial; flowers pale to blue-violet (Pennell), leaves 1.5-2 cm.

long.

Arequipa: Sand flat near Mejia and Cachendo, (Guenther &

Buchtien 155, type; det. Bruns, C. dichotoma'); 156. Mollendo,

(Stafford K60); Mexia 94178. Quishuarani, 1,860 meters, Pennell

13044- Sand dunes, Tiabaya, 2,100 meters, Pennell 13059.

Coldenia dichotoma (R. & P.) Lehm. Asperif. 1: 9. 1818; 58.

Lithospermum dichotomum R. & P. Fl. Peruv. 2: 5, pi. lllc. 1799.

C. Dombeyana Juss. ace. to Buck, Index DC. Prodr. 3. 108. 1858.

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

C. pentandra Juss. ace. Steud. Nomencl. 212. 1821. Tiguilia dicho-

toma (R. & P.) Pers. Syn. 1: 157. 1805.

Slightly suffrutescent with rugose veiny leaves, bark papery, soon

exfoliating, early shortly glandular pilosulous; petioles 5-10 mm.

long; leaves hispidulous or glabrate, especially above where bullate

by the strongly impressed nerves, mostly oblong-obovate, revolute,

crenulate, about 12-15 mm. long, 5-6.5 mm. wide; corolla 3-4 mm.

wide, bluish-pink, little longer than the sublinear hispid sepals; sta-

mens well-exserted. — Nutlets 2, 1.5-2.5 mm. long, strongly convex

dorsally, finely tessellate-granulate, strongly brown-mottled, the an-

terior face flat, spreading to form circular patches sometimes 15 dm.

across (Johnston).

Piura: Talara, Haught 6; Johnston 3517; Eyerdam 9008. Parinas

valley, Haught 158. — La Libertad: Salaverry, Johnston 3522. — Lima:

On the lomas, Weberbauer 5713; 144. Lurin, Ruiz & Pawn, type;

5946. "Tiquil-tiquil" (Ruiz & Pavon).

Coldenia elongata Rusby, Descr. New Sp. S. Am. PL 107. 1920;

185. C. parviflora Phil, as to Johnston, I.e., fide Johnston, Journ.

Arnold Arb. 16: 185. 1935.

Irregularly branched, shrubby, lower stems glabrate, exfoliating,

younger more or less densely villous-hispid; petioles slender, shorter

than the oblong obtuse leaves, these with 4-5 pairs of veins impressed

above, in type to 7 mm. long, 4 mm. wide, usually larger, oval-ovate,

obtuse both ends, revolute; calyx lobes linear, about 5 mm. long,

long pilose-hispid; corolla about 5 mm. long, the tube about 1.5 mm.

long, the lobes subrotund; stamens included; nutlets 4, spherical or

nearly, attached by a small basal circular scar, smooth, lustrous,

about 0.5 mm. thick or somewhat larger, brown in type, perhaps

black at full maturity. — The elongate, usually crenate leaves, tomen-

tose petioles and calyces distinguish this from C. parviflora Phil, of

Chile (Johnston).

Arequipa: Yura, (Williams 2562, type; 2521? nutlets larger);

Karl Schmidt (det. Standley, C. dichotoma). Below Arequipa, Weber-

bauer 6837. Lomas, Prov. Camana, Weberbauer 5730; 5734. Adja-

cent Chile.

Coldenia paronychioides Phil. Cat. Itin. Tarapaca 55. 1891; 60.

Lithospermum aggregatum R. & P. Fl. Peruv. 2: 4. 1799. C. aggregata

Rusby, Descr. New Sp. S. Am. PI. 106. 1920, not based on R. & P.

type.

FLORA OF PERU 571

Prostrate, greenish or canescently appressed to spreading hispid-

ulous with minute pale blue to white flowers, early herbaceous but

soon ligneous toward base and openly sprawling, in age sometimes

several dm. in diameter; petioles 1-3 mm. long; leaves oblong-

oblanceolate, mostly about 5 mm. long, 1.5-2 mm. wide; corolla

nearly hidden in the pubescence of the linear calyx lobes, these only

about 3 mm. long; nutlets narrowly ovoid, lustrous, minutely reticu-

late; habit of C. dichotoma, with which it sometimes grows (John-

ston).

Piura: Talara, Haught S3; Pennell 12202; Goodspeed Exped. 9004;

Horton 11555; West 3572; Beetle 26204. Near Piura, Weberbauer 5932.

Negritos, (Haught & Svenson 17606). Paita, Pennell 14819; (Saf-

ford 9; Ball; Williams 2913, type, C. aggregata Rusby); Weberbauer,

154. — La Libertad: Salaverry, Johnston 3523. — Ancash: Los Zorros,

Goodspeed Exped. 9197. — Lima: Quive, Pennell 14305. Sta. Eulalia,

Goodspeed 33092. Sayan, Goodspeed 17358. — Junin: Near Tarma,

Ruiz & Pavdn (type, L. aggregatum) . — Arequipa: Tingo, 2,100 me-

ters, Pennell 13102. — Puno: Pisacoma, (Meyen, distr. as Heliotropium

humifusum, fide Johnston). Northern Chile; Bolivia. "Flor de

arena" (Stork).

4. CORDIA L.

Trees or shrubs, in Peru broad-leaved and the inflorescence vari-

ous. Calyx mostly persisting. Corolla white or yellowish, red to

orange in C. sebestena (cultivation), the lobes 5-9 (-15). Stamens

5-10. Ovary usually 4-celled, ovules 1-4. Style terminal, well-devel-

oped, 2-lobed or 2-parted, the branches each 2-lobed; stigmas 4.

Fruit usually a drupe with dry or more or less mucilaginous exocarp,

or carpoid wall, chartaceous and fibrous (section Gerascanthus) . —

Genus first described in detail anatomically by Mez, Bot. Jahrb. 12:

526-588. 1890. Johnston's "The Boraginaceae of Northeastern

South America," Journ. Arnold Arb. 16: 1-64. 1935, is an invaluable

contribution to the knowledge of this family and I hope I have

correctly compiled his painstakingly observed and recorded observa-

tions pertaining to the Peruvian species.

The red or orange flowered C. sebestena L. is probably grown in

Peru for ornament; it is native to the northern coasts of South

America and the West Indies.

Corolla persisting (marcescently) ; calyx costate.

Calyx teeth obscure; fruit enclosed.

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

Leaves lanceolate, about three times longer than wide.

C. iguaguana.

Leaves oval to ovate in some degree and usually about twice

as long as wide.

Leaves usually longer than wide.

Flowers about 1.5 cm. long C. alliodora.

Flowers about 4 cm. long C. insignis.

Leaves suborbicular C. lutea.

Calyx with long subulate teeth; fruit exserted . . . . C. varronifolia.

Corolla deciduous after drying; calyx (in Peru) not tubulate nor

strongly costate nor ovate and subulate-lobed.

Nodes inflated; stems notably long-hispid C. nodosa.

Nodes not inflated; stems glabrous to variously pubescent.

Flowers showy, in globose heads; peduncles to 1 dm. long; calyx

teeth filiform C. macrocephala.

Flowers small, sometimes capitate but calyx lobes obscure, rarely

subulate-appendaged; peduncles often short.

Branches remotely spurred or nodular by persisting petiole-

bases; flowers usually congested in spikes or heads, dis-

posed rather laxly only in C. Poeppigii; fruit often shorter

than 5 mm.

Inflorescences usually simple, in any case densely flowered

(cf. C. Poeppigii).

Leaves evenly serrate, ample, with subulate teeth to

2 mm. long, 3-5 mm. apart C. macrodonta.

Leaves entire, minutely or densely or unevenly serrate,

usually only a few cm. wide (key characters below

variable, expedient).

Inflorescences soon clearly longer than broad, spiciform.

Leaves coarsely and unevenly serrate . . C. peruviana.

Leaves entire to more or less serrate but closely or

evenly C. cylindristachya, C. spinescens.

Inflorescences usually forked or capitate, especially in

flower.

Leaves more or less acuminate and acutely, mem-

branous or firm, not or obscurely rugose unless

in age, usually glabrate or sparsely pubescent;

cymes small, early dense or capitate.

FLORA OF PERU 573

Inflorescence axillary C. patens.

Inflorescence extra-axillary C. bifurcata.

Leaves if acuminate shortly and not or scarcely

acutely, usually rounded to subacute, subcori-

aceous, reticulate-rugose, often rather softly

pubescent beneath; flowers often capitate even

in age.

Leaves subentire or remotely serrulate.

Flower heads often several on more or less leafy

branchlets.

Calyx lobes subulate-appendaged; branchlets

spreading hirsute C. buddleoides.

Calyx lobes acute; branchlets lanate.

C. mollissima.

Flower heads or small forked cymes solitary;

branches soon glabrate C. scaberrima.

Leaves obviously even if rather minutely, serrulate

or crenate.

Leaves oblong- or ovate-elliptic unless young

shoots, slightly rounded to subacute.

C. Krausiana.

Leaves rather ovate, more or less obtusely and

broadly acuminate, scabrous above.

C. scaberrima.

Leaves subrotund, hispidulous both sides.

C. lantanoides.

Inflorescences mostly compound or foliose, the branches

rather laxly flowered C. Poeppigii.

Branches smooth or in any case inflorescences soon lax or

sometimes simple but laxly flowered or open and flowers

crowded, usually not simple or not typically small; fruit

often 5 mm. long or longer.

Leaves rather ovate, often acutely acuminate; inflorescence

branches few, elongate, simple or subsimple, suberect.

C. Poeppigii.

Leaves rather elliptic, cuspidate or acute or if acuminate,

abruptly or shortly; inflorescence branches short, often

many.

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

Leaves glabrous or essentially, even beneath any indu-

ment minute and obscure; ovary glabrate or glabrous.

Calyx long-obconic, 6 mm. long in bud; corolla to

12 mm. long; ovary glabrous C. Mexiana.

Calyx campanulate or pyriform, 2.5-4 mm. long; cor-

olla to about 5 mm. long.

Petioles 2-3 cm. long, geniculate at base; calyx lobes

uneven C. collococca.

Petioles shorter or in any case straight; calyx lobes

obscure, equal C. lomatoloba.

Leaves strigose or scabrous at least in part; ovary pubes-

cent except C. hebeclada, C. Ulei, C. ucayaliensis,

sometimes C. ripicola.

Flowers much congested; leaves (as to types) clearly

pubescent beneath.

Inflorescence lightly pubescent.

C. laevior, C. ripicola.

Inflorescence reddish tomentose C. Sprucei.

Flowers not congested or inflorescence branchlets long;

leaves minutely pubescent or scabrous.

Petioles about 1 cm. long; leaves mostly or all nar-

rowed to acute base.

Branchlets puberulent; calyx about 4 mm. long.

Leaves scabrous, nearly three times longer than

wide; ovary pubescent C. scabrifolia.

Leaves pubescent both sides, about twice as long

as wide; ovary glabrous. . .C. ucayaliensis.

Branchlets hirsutulous; calyx about 5 mm. long.

C. Ulei.

Petioles about 1.5-2 cm. long; leaves broadly round-

ed or shortly acute at base or very dissimilar.

Ovary pubescent; larger leaves often acute or nar-

rowed to base C. toqueve.

Ovary glabrous; all leaves rounded or truncate at

base C. hebeclada.

Cordia alliodora (R. & P.) Oken, All. Naturgeschichte, Bot. 2:

1098. 1841; 13. Cerdana alliodora R. & P. Fl. Peruv. 2: 47, pi 184.

1799.

FLORA OF PERU 575

More or less densely stellate-pubescent with small or minute

forked trichomes, especially the ample inflorescences; petioles 1-3 cm.

long; leaves rather broadly elliptic, obtusely acute both ends or

acuminate apically, usually 1-5.5 dm. long, about half as wide, entire,

coriaceous, glabrous above where slightly marked by the impressed

nerves and veins, these rather prominent beneath; inflorescence ter-

minal, laxly branched, the flowers crowded; calyx nearly globular,

strongly 10-ribbed, canescent-puberulent, truncate, 6-7 (14) mm.

long, the teeth minute, the exserted corolla tube with oblong rounded

lobes, these 5-10 mm. long, usually about 3 mm. wide; fruit fibrous,

chartaceous, 5 mm. long, about 3 mm. thick, falling within the per-

sisting calyx tube and corolla.

A distinctive feature of the plant is its myrmecophily (cf . Wheeler,

Bull. Mus. Comp. Zool., 90: 9-41. 1942); the leafy twigs and fre-

quently the axis of the inflorescence develop as irregular swellings

which serve as ant domatia; see also C. nodosa. Small ants some-

times totally destroy the leaves, cutting them into triangular pieces

and storing them symmetrically; each piece may be six times the

size of the ant that carries it by the shortest side, apparently like a

sail (Ruiz & Pavon). See Johnston, I.e., for extra-Peruvian synon-

ymy; he remarks that there is only moderate variation in the species,

in size of flowers and degree of pubescence. In Peru sometimes

attains 20 meters or more, the trunk (bark grayish, fissured) 3 dm.

or more in diameter; timber used for construction and for vehicles

(Williams); leaves and bark used as a condiment, serving as garlic

(Ruiz & Pavon) ; flowers fragrant.

Cajamarca: Hualgayoc, Soukup 3839. — San Martin: Tarapoto,

Williams 6751. — Hudnuco: Pozuzo, Ruiz & Pavon, type. — Loreto:

Lower Itaya, Williams 318. Lower Huallaga, Williams 4306. —

Cuzco: Urubamba Valley, Weberbauer, 280. — Rio Acre: Upper Rio

Jurupary, Krukoff 5220. Seringal Auristella, Ule 9719. Bolivia to

Mexico; West Indies.

Cordia bifurcata Roem. & Schult. Syst. 4: 466. 1819. Varronia

dichotoma R. & P. Fl. Peruv. 2: 23, pi. 146. 1799, not C. dichotoma

Forst., 1786.

Slender, shrubby below, often openly branched above, the leafy

flowering branchlets minutely and sparsely appressed strigillose and

(or) scabrous as the green leaves at least above, these sometimes also

hispidulous on the veins beneath; petioles about 3 mm. long; leaves

usually ovate- or oblong-elliptic-lanceolate, obliquely acute at base,

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

acutely acuminate, 6-7 cm. long, about 2 cm. wide near the middle

or sometimes (as freely interpreted) broadly ovate, rounded at base,

3-4.5 cm. wide and only about twice as long, subentire to remotely

serrate above the middle, membranous, paler beneath; peduncles 1-2

(4) cm. long, terminal on leafy branchlets or from the leaf-axils of

branchlets, the small inflorescences capituliform in flower, obviously

and subumbellately 2-4-dichotomously branched only in fruit when

sometimes about 3 cm. across, half as long; calyx appressed strigil-

lose, glabrate and reddish in fruit, with broad almost minute lobes,

about 3 mm. long or about half as long as the fleshy usually white

corolla; fruits bright red, ovoid-elliptic, 3.5-4 mm. long. — Probably

a species variable in pubescence, leaf -form, flower color and perhaps

in position and form of inflorescence (compare C. patens, C. scaber-

rima and others). Johnston, in 1930 (Contr. Gray Herb. 92: 30-34),

under the name C. corymbosa (L.) G. Don, commented at length on

apparent variability and, as regards pubescence, proposed forma

typica, leaves finely tomentulose beneath, coarse trichomes few or

lacking; f. intonsa, coarse trichomes abundant; f. detonsa, pubescence

fine, appressed; and f. hirsuta, pubescence coarse, spreading. In 1949

(Journ. Arnold Arb. 30: 90) he wrote, using the name C. bifurcata

Roem. & Schult.: flowers minute, orange, in terminal or internodal

cymes, these usually forked even in bud. Material determined by

Johnston after 1949 had white flowers, according to the collectors.

Johnston, I.e. 30: 90 et seq. 1949, has argued, apparently correctly,

for the validity of this species, confused with C. lineata (L.) Roem.

& Schult., doubtfully in South America, and C. polycephala (Lam.)

Johnst., also known only north of Peru; both have the axillary

inflorescence of C. patens HBK.

Confusion in interpretation of the shrub's characters has been

equaled by the nomenclature; professional taxonomists apparently

ought to be reminded of the old truism that whatever name used the

rose is still a rose or words to that effect; Tournefortia syringaefolia

Vahl is a case in point, apparently with another name, T. maculata

Jacq. in spite of its retaining the same "spots;" however, if a botan-

ical compiler overlooks or misapplies a rediscovered or reinterpreted

name may heaven forgive him!

Piura: Canchaque, Prov. Huancabamba, Stork 11417 (det. John-

ston, C. macrodonta) . — San Martin : Tarapoto, Ule 6478; Woytkowski

35074; Williams 5408; 5571; 6177; 6426; 6684A— Huanuco: Cha-

cahuasi, Ruiz & Pavdn (type, V. dichotoma). Tingo Maria, Allard

20494 (det. Lyman Smith, C. polycephala). Rio Santo Domingo,

FLORA OF PERU 577

4210. Pozuzo, 4560. — Junfn: La Merced, Killip & Smith. Prov.

Tarma, Woytkowski 37003. Argentina to Costa Rica.

Cordia buddleoides Rusby, Mem. Torrey Bot. Club 6: 83. 1896;

34.

Branchlets elongate, spreading hirsute; petioles stout, very short

to nearly 1 cm. long; leaves ovate, abruptly contracted at base, acute,

7-10 cm. long, 3-5 cm. wide, obsoletely serrulate, firm-membranous,

scabrous and the 6 or so lateral nerves impressed above, strigose be-

neath, the veins strongly reticulate; inflorescence ample, leafy, in

type the flowers capitate; calyx tube 1.5 mm. long, lobes subulate-

linear, 3 mm. long, reflexed; corolla little exserted, stamens included;

ovary slightly 4-lobed, style nearly 2 mm. long, 4-cleft. — After author

except that he describes branchlets as harshly tomentose; the small

heads are on long peduncles and form a somewhat leafy open inflores-

cence. Ule 9714 from Rio Acre in Bolivia has flowers in a cylindric

spike, probably due to greater maturity than type with globose

inflorescences from La Paz, matched by Lechler; the Poeppig collec-

tion lacks the apical calyx lobe appendages (Johnston) ; this northern

shrub may rather be C. scaberrima HBK. or affine. Name originally

spelled as above.

Loreto: Maynas (Yurimaguas), (Poeppig 2471). — Puno: San Go-

van, (Lechler 2424)- Bolivia.

Cordia collococca [Sandmark] L. Amoen. Acad. 5: 377. 1759

(as C. callococca). C. glabra of authors, not Linnaeus; cf. Johnston,

Journ. Arnold Arb. 21: 345; 1940, I.e. 30: 123-124. 1949.

Essentially glabrous, the indument of the open, dichotomously

branched inflorescence and leaves beneath obsolete or extremely

minute; petioles 2-3 cm. long; leaves elliptic-obovate, obliquely

rounded or subacute at base, broadly acuminate or rounded and

apiculate at apex, submembranous, finely reticulate-veined both

sides, the 7-9 lateral nerves slender; calyx obscurely puberulent,

cupulate, about 4 mm. long, unevenly lobed; corolla about 5 mm.

long; ovary glabrous; drupes bright red. — Deciduous, dioecious, the

flowers precocious, the leaves beneath pubescent with many slender

erect bristles (Johnston); umbrella-shaped, 5 meters tall, with pen-

dent ("weeping") habit (Mexia).

Loreto: Rancho Indiana, Mexia 6403 (det. Johnston). To Cuba

and Mexico. "Camungo-tiarina" (Mexia).

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

Cordia cylindristachya (R. & P.) Roem. & Schult. Syst. 4: 459.

1819. Varronia cylindristachya R. & P. Fl. Peruv. 2: 23, pi. 147a.

1799. V. macrostachya R. & P. I.e. as to plate. C. tarmensis Krause,

Bot. Jahrb. 37: 630. 1906, fide Killip. C. subserrata Krause, I.e. 627.

Branchlets and leaves beneath more or less fulvous or cinereous

tomentose-pilose; petioles 4-10 mm. long; leaves elliptic or ovate-

elliptic, acute both ends, varying in size from about 3 cm. long, 2 cm.

wide, to a dm. or so long, 4 cm. wide or under, submembranous to

coriaceous, entire or subserrate toward tip, rugose, sparsely hispid

both sides or tomentulose beneath; spikes terminal and axillary,

usually 3-5 cm. long; calyx densely pilose, lobes ovate, acute, the

slightly longer glabrous or subglabrous corolla tube about 4 mm. long,

the lobes short; filaments affixed about medially, dilated and villous

toward base; style subequaling calyx. — According to Johnston, Journ.

Arnold Arb. 30: 121. 1949, C. curassavica (Jacq.) Roem. & Schult.,

ranging north of Colombia and similar, differs constantly in the com-

plete lack of axillary spikes.

Cajamarca: Socota to Tambillo, Stork & Horton 10178 (det.

Standley). — Huanuco: Acomayo and Pillao, Ruiz & Pavdn, type;

Woytkowski 34002. Chaglla, 3641 . — Junin : Near Palca, Weberbauer

1758; Killip & Smith 23372. — Cuzco: Marcapata, near Cachu-

pampa, Vargas 9696. — Arequipa: Mollendo, Weberbauer 1530.

Cordia hebeclada Johnst. Journ. Arnold Arb. 31: 176. 1950.

Tree to 15 meters tall with regular dichotomous branches, dimor-

phic leaves and strongly heterostyled flowers, unisexual by abortion;

branchlets and leaves beneath densely puberulent; petioles often

1-2 cm. long; leaves laxly or appressed subscabrous above, the

larger elliptic-lanceolate or -ovate, rounded at base, more or less

acuminate, 1.5-2.5 dm. long, 7-15 cm. long, the smaller suborbicular

or orbicular-ovate, frequently 8-10 cm. long, often truncate or sub-

cordate at base, rounded or obtuse at apex; inflorescence corymbose,

1-3 dm. across; male flowers with cupulate calyx 3-4.5 mm. long,

densely puberulent but strigose within, the unequal lobes acute;

corolla 6-8 mm. long, oblong recurved lobes and tube 3-4 mm. long;

filaments about 4 mm. long, villous basally; ovary rudimentary,

glabrous; female flowers similar but calyx subcylindric, 2.5-3 mm.

long, filaments glabrous or very sparsely villous; style exserted,

3.5-4 mm. long, bifid; drupe glabrous, depressed globose, strongly

rugose, to 1 cm. long, 8 mm. broad, the expanded fruiting calyx

4-5 mm. across. — Probably most closely related to C. panamensis

FLORA OF PERU 579

Riley, but differing in indument and in its regular dichotomous

branching; in C. hebeclada the indument consists of a great abundance

of minute trichomes that give a thin downy uniform very soft

cover (Johnston), scarcely soft in age; perhaps an Andean variant

of C. tetrandra Aublet in which the author included it, Contr. Gray

Herb. 92: 55. 1930.

San Martin: Morales near Tarapoto, Williams 5676. Chazuta,

Klug 4050. Bolivia to Colombia. "Tahnampa-caspi" (Williams);

"quitasol" (Little), or "jutumbe" (Tafalla) in Ecuador. "Rena-

quillo" (Klug).

Cordia iguaguana Melchior ex Johnst. Journ. Arnold Arb. 33 :

63. 1952.

Tree, 8-20 meters tall, branchlets finely fulvous tomentulose, gla-

brate in age, petioles to about 2 cm. long; leaves broadly lanceolate,

obtuse or rounded at base, acute or somewhat attenuate at apex,

1-1.5 dm. long, 4-5 cm. wide, glabrous or minutely and sparsely

appressed pubescent, the 5-7 lateral nerves and many veins lightly

impressed above, rather prominent beneath; paniculate or corymbose

inflorescences terminal on young leafy branches, about 1.5 cm. across;

calyx clavate-cylindrical, prominently 10-costate, about 11 mm. long,

3 mm. thick toward the apex, obscurely appressed-puberulent, the

3-5 often unequal obtuse lobes about 1 mm. long, the sinuses obtuse;

corolla white, 23 mm. long, limb about 2.5 cm. broad, lobes 6-8 mm.

long, 8-10 mm. wide, often emarginate, tube 8 mm. long, 1-1.5 mm.

thick, tomentulose within below the middle, the throat 10-15 mm.

across, about 5 mm. deep; filaments glabrous, affixed near apex, 1-

3 mm. long, the anthers also unequal, 1.5-3 mm. long; ovary with

style about 13 mm. long (nectary lacking), forked about 1 cm. above

the fusiform base. — Section Gerascanthus with three other species

known from western South America; C. alliodora (R. & P.) Oken has

stellate indument and is myrmecophilous, the other species not and

furthermore their indument is simple; C. Goeldiana Huber, Amazonian

Brazil to Colombia, has long oblong corolla lobes while C. macrantha

Chodat, of western Ecuador, with the broad rounded corolla lobes of

C. iguaguana, has corollas 3.5-4 cm. long, larger elliptic leaves

evidently villous aspidulose beneath, and differs otherwise according

to Johnston, who, agreeably, took up the well-chosen herbarium

name of Melchior. F.M. Neg. 975.

Cajamarca: Near Jaen, 700-800 meters, Weberbauer 6218, type.

"Iguaguana."

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

Cordia insignis Cham. Linnaea 8: 122. 1833. C. Haenkeana

Mez, Bot. Jahrb. 12: 560. 1890, fide Johnston, Journ. Arnold Arb.

16: 179. 1935.

Branchlet tips densely more or less ferrugineous tomentulose with

simple trichomes as the terminal corymbose-paniculate inflorescences,

the ultimate branches of these few-flowered; petioles to 4 mm. long;

leaves ovate or suboval, rather rounded at base, basally narrowed to

finally rounded apex, about 14 cm. long, 9 cm. wide, subglabrous and

lustrous above, strigose beneath where prominently reticulate-

veined; calyx costate, 2.5 cm. long, anthers often with few irregular

teeth; corolla to 4 cm. long, coriaceous, persisting, the 5 lobes

emarginate; stamens nearly 2 cm. long, affixed near base, joined

annulately, included; ovary glabrous. — Description after Mez, who

remarked: evidently differs from C. insignis in form of leaves and

inflorescence. I greatly doubt that the Haenke collection came from

Peru; C. insignis is known only east and south of the Amazon

Basin, from eastern Brazil to eastern Bolivia, and is certainly not

to be expected in Peru (Johnston). There are apparent exceptions

known to this range unless determinations have been wrong; C.

insignis may have ovate-oblong leaves, flowers in dense cymes.

Illustrated, Mart. Fl. Bras. 8, pt. 1, pi. 1 (C. insignis}.

Peru(?) : See note above; without locality except "Peru: Haenke"

(type, herb. Monac.).

Cordia Krausiana Killip, Journ. Wash. Acad. Sci. 17: 330. 1927.

C. pauciflora Krause, Bot. Jahrb. 37: 629. 1906, not Rusby, 1896.

Flowering branches somewhat ferrugineous tomentose as the

3-5 mm. long petioles; leaves elliptic or ovate-elliptic, acute both

ends, in type 2.5-3 cm. long, 12-16 mm. wide, apparently often

larger, subcoriaceous, rugose, hispidulous and the veins impressed

above, tomentose and the veins conspicuous beneath, clearly, even

coarsely serrate; flowers capitately congested, few in type or in

vigorous examples many; calyx 5-6-parted, the pilose subacute seg-

ments 3 mm. long; corolla tube glabrous, little exserted, the lobes

rounded; filaments affixed just above middle, short, villous at dilated

base; style slightly exceeding calyx, stigma globose. — Related to C.

scaberrima HBK. but with smaller, more strongly serrate leaves

(Krause); if the shrub is specifically distinct it seems vegetatively

at least to differ only in the serrate leaves, these in my collections

to 8 cm. long, 3 cm. wide, the flower-heads in fruit 2 cm. thick, the

calyx and drupe about 6 mm. long. On the other hand this and the

FLORA OF PERU 581

plant of HBK. are doubtfully distinct specifically from the Tarapoto

shrub here referred to C. bifurcata but not typical. F.M. Neg. 991.

Ancash: Tamanco to Huaraz, 3,000 meters, Weberbauer 3161,

type. — Lima: Matucana, 180; 288k (det. Killip, C. lantanoides) .

Gordia lantanoides Spreng. Syst. 1: 653. 1825. Varronia cre-

nata R. & P. Fl. Peruv. 2: 23, pi. 146b. 1799, not C. crenata Delile,

1813.

Branches tortuous, the many leafy flowering ones minutely hir-

sutulous; petioles slender, to about 6 mm. long; leaves subrotund to

oval-elliptic, rounded both ends, 2-3 cm. long, 1.5-2 cm. wide,

crenate-serrulate nearly to base, submembranous, spreading hispidu-

lous both sides, slightly asperous and rugulose above, reticulate

venose beneath; peduncles axillary, 2-3 cm. long, to 4-dichotomous

in fruit but the flowers early in small heads; calyx glabrate in fruit,

about 4 mm. long, the ovate lobes acute, the white corolla little

exserted; drupe red, seed ovoid, rarely 2-celled (Ruiz & Pavon). —

Apparently not found in typical state since the original collection

but possibly should be drawn to include C. Krausiana Killip. F.M.

Neg. 980. *

Huanuco: Ruiz & Pavdn, type.

Cordia laevior Johnst. Journ. Arnold Arb. 16: 180. 1935.

Branchlets soon glabrescent, only toward tips shortly pubescent

with appressed or ascending trichomes; petioles 5-10 mm. long;

leaves similar, oblong-lanceolate, acute at base, long-acuminate,

1.5-2.5 dm. long, 4-10 cm. wide, glabrous above except the hir-

sutulous midrib and 7-8 lateral nerves (tertiary obscure), paler

beneath where puberulent on the many veins; cymes in forks of

branches, the lower branches spreading; calyx in bud obovate,

4-5 mm. long, 2-3 mm. thick, rounded apically, obscurely costate,

densely puberulent, the triangular lobes somewhat uneven; corolla

tube 4 mm. long, lobes 2 mm. wide and long; style sparsely pilose,

the filaments toward base, the ovary evidently toward apex. — Less

pubescent than the related more eastern C. Sprucei Mez and cyme

more loosely branched; leaves not roughened above by the many

veinlets; distributed as C. Ulei Johnst. with glabrate ovary, coarser

pubescence (author). It seems desirable to study these species

with more material before their true status (as that of C. ripicola)

can be discovered. Type a tree 6 meters high.

San Martin: Pongo de Cainarachi, (Klug 2756, type).

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

Cordia lomatoloba Johnst. Journ. Arnold Arb. 18: 12. 1937.

Arborescent, to 25 meters tall, nearly glabrous; leaves similar,

lanceolate or elliptic-lanceolate, narrowed to petiole — this 8-15 mm.

long — acute or acuminate, 8-14 cm. long, 2-5.5 cm. wide, glabrous

or obscurely strigose, reticulate both sides, lustrous above, paler and

opaque beneath, entire; primary nerves 6-8 pairs; cymes often

somewhat brown-pubescent in the forks of the branches, these lax,

slender; calyx sessile, pyriform, 2.5 mm. long, abruptly contracted

to base, the deltoid equal teeth only marginally obvious; corolla

white, 4 mm. long, tube 1 mm. long, lobes about 1.8 mm. wide,

oblong, recurved, rounded at tip; filaments villous below; ovary

glabrous. — Related to the pubescent C. naidophila Johnst. of the

Amazon, distributed as C. ecalyculata Vahl with much narrower

leaves, while C. diospyrifolia Cham, as to type by Sellow has cori-

aceous broader leaves; the characters may prove to be variable.

San Martin: Chazuta, King 4050? — Loreto: Mouth of the Rio

Santiago, Tessmann 4454 (det. Melchior, C. diospyrifolia). — Rio Acre:

Mouth of Rio Macauhan, Krukoff 5497, type; also 5345. Brazil.

Cordia lutea Lam. 111. 1: 421. 1791. C. marchionica Drake, 111.

Fl. Ins. Pacific 240 (1892), fide Johnston, Journ. Arnold Arb. 33: 65.

1952. C. rotundifolia R. & P. Fl. Peruv. 2: 24, pi. 148a. 1799.

Flowering branchlets many, canescent-lanate to puberulent in-

cluding the conspicuously ribbed calyces of the showy pale yellow

flowers; petioles 1-2.5 cm. long; leaves subrotund to broadly ovate-

elliptic, mostly 4 or 5 cm. wide and little longer, sometimes much

larger, weakly crenulate, early canescent short-pilose beneath, harshly

scabrous above; calyx soon obconic, about 1 cm. long, teeth subulate;

corolla a clear light yellow, 3 cm. long or longer, the throat dilated,

the limb 6-8 lobed; filaments villous at base; fruit ovoid, white.—

Johnston, I.e., has presented arguments that seem irrefutable that

the common occurrence of this species in the Marquesas is due to

its introduction by the French during the nineteenth century, prob-

ably as an ornamental. A decoction is used with good results for

jaundice (Ruiz & Pavon); an attractive yellow-flowered bush along

streams. F.M. Neg. 994.

Tumbez, fide Diels. — Piura: Near La Brea, (Haught & Svenson

11596}; Weberbauer 5955. — Cajamarca: Huambos, Stork & Norton

10205. — Lambayeque: Chiclayo, West 3575. — Ancash: Tambo de

Pariacota, 2557. Talara, Horton 11516. Pabur, Stork 11422.—

Lima: Lurin, Chan cay, Ruiz & Pavon (type, C. rotundifolia}; Dom-

FLORA OF PERU 583

bey, type. — Huanuco: Maranon Valley, Weberbauer, 155. — Loreto:

Mishuyacu, near Iquitos, Klug 1482. Western Ecuador; Galapagos

Islands. "Tina" or "membrillejo" (quince); "oberal" (Weberbauer).

Cordia macrocephala (Desv.) HBK. Nov. Gen. & Sp. 3: 77.

1818. Varronia macrocephala Desv. Journ. de Bot. 1: 273. 1808.

C. polyantha Benth. PI. Hartw. 121. 1843? C. guayaquilensis DC.

Prodr. 9: 496. 1845.

Often many-stemmed, matted or erect and sometimes 5 meters

tall, the flowers in well-peduncled heads, the coarsely crenate or ob-

tusely serrate leaves greenish but more or less scabrous both sides,

conspicuously veined, to about 6 cm. long, 2.5 cm. wide, often smaller,

ovate- or oblong-elliptic, obtuse or subacute. — Johnston, Journ. Ar-

nold Arb. 30: 89. 1949, remarks that the similar C. polyantha Benth.,

PI. Hartw. 121. 1843, "of Ecuador and northern Peru," and C. macro-

cephala (Desv.) HBK., of Peru, have a relative in C. Pringlei Robins,

of more northern range. A pubescent or glabrate shrub striking by

the globose inflorescences of white flowers. Pubescence of unfolding

leaves during dry period may consist of closely packed stellate scales

while in the wet period stellate and single hairs are greatly elongate

and forked hairs are on some specimens (Svenson, Am. Journ. Bot.

33: 479. 1946). F.M. Neg. 984.

Piura: Amotape Hills, (H aught & Svenson 11523). Near Frias,

Weberbauer 6433. Serran, Stork 11373 (det. Johnston, C. polyantha).

— Lambayeque, fide Weberbauer. — Lima: Chosica, 542; Mexia 04009.

Sta. Eulalia, Goodspeed 33018; Norton 10994. Sayan, Goodspeed

33042; 33046. Rio Rimac Valley, Goodspeed, Weberbauer & Metcalf

30217.— Junin: Huacapistana, Killip & Smith 24518 (det. Killip,

C. floribunda). To Colombia.

Cordia macrodonta Killip, Journ. Wash. Acad. Sci. 17: 328.

1927.

Branchlets quadrate, ferrugineous-puberulent and finely pilosu-

lous (as leaves beneath), scabrous; petioles 5-10 mm. long; leaves

broadly ovate, narrowed to base, abruptly short-acuminate, 6-12 cm.

long, 4-6.5 (8) cm. wide, coarsely and acutely serrate except in lower

third, membranous, shortly strigillose-hispid above, lateral nerves

6-7 pairs; peduncles 4 cm. long; flowers solitary (or few) at ends of

the inflorescence branchlets; calyx globose-turbinate, 2 mm. long,

appressed rusty-strigillose, the acute teeth 0.5 mm. long; corolla tube

cylindric, 3.5 mm. long, stamens in throat; fruit ovoid-conic, 5 mm.

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

long, glabrous. — After author, who remarked: distinguished from

other members of the Corymbosae DC. by the coarsely serrate

leaves; may prove to be a part of the variable C. bifurcata Roem. &

Schult. as suggested by Stork 11417.

Piura: San Antonio, Prov. Huancabamba, Weberbauer 6015, type.

Cordia Mexiana Johnst. Journ. Arnold Arb. 18: 12. 1937.

Tree, 5-7 meters, with puberulent branches; leaves elliptic or

oblong-elliptic, abruptly contracted to petiole, this 1-2.5 cm. long,

obtusely acuminate, 1.5-4 dm. long, 8-20 cm. wide, coriaceous, retic-

ulate both sides, glabrous, lustrous and minutely puncticulate above,

subpuberulent beneath; nerves 5-7 pairs; cymes laxly much branched,

2.5 dm. across, the flexuose branchlets strigose puberulent; calyx

(bud) about 6 mm. long, 2 mm. thick, stouter above, densely brown

strigose, puberulent within above the middle, the 5 deltoid erect lobes

about 1 mm. long; corolla white, 12 mm. long, tube 1.5 mm. thick,

4 mm. long, throat obvious, 2.5-3 mm. long, gradually expanded to

about 3 mm. across, lobes oblong, 4 mm. long, recurved, rounded;

filaments exserted, villous below; ovary glabrous; style deeply forked.

— Distributed as C. colombiana Killip of Ecuador and Colombia but

distinct because of the elongate corolla with well-developed throat;

this character, and the long tube, are very unusual in the Pilicordia

section (author).

Loreto: Rio Maranon near Rancho Indiana, Mexia 6459, type.

Adjacent Colombia.

Cordia mollissima Killip, Journ. Wash. Acad. Sci. 17: 330. 1927.

Densely lanate-tomentose even the contracted or early subelliptic

few-flowered inflorescences; branches terete; petioles to 1 cm. long;

leaves ovate-lanceolate, rounded or subacute at base, acute or acumi-

nate, to 7 cm. long, 3.5 cm. wide, serrulate, reticulate-veined, dark

green above, paler beneath; calyx campanulate, about 2 mm. across,

the deltoid lobes acute; corolla 3-3.5 mm. long; stamens and style

about 1.5 mm. long, the ovate-oblong anthers scarcely exserted.—

Related to C. corymbosa (L.) Don. (C. bifurcata Roem. & Schult.)

but differing in the white indument and the closely serrulate leaves

(author).

Cajamarca: Jaen, Weberbauer 6202, type. — Amazonas: Chacha-

poyas, (Mathews 3134). — San Martin: Juanjui, King 4385 (det.

Standley, C. scaberrima} .

FLORA OF PERU 585

Cordia nodosa Lam. Tab. Encycl. 1: 422. 1791; 46 and 13. C.

hispidissima DC. Prodr. 9: 475. 1845.

Distinctive by enlarged nodes below each fork, these hollow, ordi-

narily inhabited by ants; marked also by the divaricate rufous or

brownish bristles that are usually more or less developed about the

apiculate calyx tip; petioles 2-5 mm. long; leaves usually subopposite

or whorled, somewhat dissimilar, often elliptic, 1-3 dm. long, obtuse

at base, acuminate, lustrous, more or less bullate at limb in age, the

6-10 pairs of nerves impressed, reticulate; paniculate cymes sessile or

peduncled, up to a dm. across, usually puberulent with crinkly curved

trichomes or the papyraceous scarcely ribbed calyx strigillose, 5 mm.

long in bud, unevenly lobed, ordinarily persisting and becoming

fibrose; corolla white, 4-6 mm. long; style and ovary pubescent

(filaments basally) ; fruits yellow, usually more or less bristly, about

1.5 cm. long. — Shrub or tree sometimes 10 meters high, few branches,

bark fissured with many small lenticels (Williams, Field Mus. Bot.

15: 433. 1936), nearly smooth except at nodes to densely bristly

(forma hispidissima). I. W. Bailey, Bot. Gaz. 77: 32-49, pis. 6-7.

1924, gave an account of myrmecophilism here, a curiosity occurring

among Peruvian plants, notably in Melastomaceae and Moraceae;

this intriguing development was investigated first by A. F. W.

Schimper, Bot. Mitteil. Trop. Heft 1: 53-. 1888, and by Schumann,

Pringsheim's Jahrb. 382-. 1888.

San Martin: Tarapoto, Williams 6516. Juanjui, King 3775. —

Huanuco: Tingo Maria, 700 meters, Stork & Horton 9519. Valley

of the Monzon, Prov. Huamalies, Weberbauer 3679; 285. — Junin:

Rio Pichis, Killip & Smith 26731. Hacienda Schunke, 1,400 meters,

Killip & Smith 24582. Perene" Bridge, Killip & Smith 25359 —

Loreto: Florida, King 2158; 2305. Rio Nanay, Williams 330 A;

1229. Caballo-Cocha, Williams 2334- Rio Mazan, Jose Schunke 10.

Yurimaguas, Killip & Smith 27688; 27728; Williams 4305; 4677.

Iquitos, Williams 3748; 7974; Killip & Smith 27022; King 480; 510;

803. La Victoria, Williams 2679. Rio Itaya, Killip & Smith 29602.

Bolivia to the Guianas. "Almenchillo," "analliocaspi" or "ayahuas-

co" (Williams); "ibrachoreri" (Huitoto); "pucacurocaspi" (Schunke).

Cordia patens HBK. Nov. Gen. & Sp. 3: 75. 1818.

Young branchlets and leaves beneath densely flavescent tomentu-

lose; petioles 4 mm. long; leaves ovate-oblong, rounded at base, acute

or acuminate, to 7.5 cm. long or longer, rather remotely serrulate

toward tip, firm, reticulate and strigose-scabrous above, the veins and

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

nerves prominent beneath; spikes subglobose, axillary and terminal

on slender peduncles 5 cm. long; calyx strigose-hispid with minute

subulate teeth, villous in the throat as the white twice as long corolla;

stamens exserted, exceeded by the style. — After HBK. Inflores-

cences axillary, not extra-axillary as in C. bifurcata (Johnston) ; Pe-

ruvian collection was referred to C. bifurcata (C. corymbosa (L.) Don)

by Standley and Killip and the leaves are not yellowish tomentulose

beneath as are those of the type from Venezuela, which in this char-

acter resembles Tarapoto material. F.M. Neg. 38979.

Loreto: Yurimaguas, Killip & Smith 28262; Williams 3898; 3947;

4469; 4787. Contamana, Rio Ucayali, Killip & Smith 26876. To

the Guianas.

Cordia peruviana Roem. & Schult. Syst. 4: 459. 1819. Varronia

obliqua R. & P. Fl. Peruv. 2: 24, pi. 147b. 1799, not C. obliqua Willd.,

1797.

Similar to C. spinescens but leaves coarsely and irregularly serrate,

obliquely decurrent at base, mostly 5-6 cm. long, 2.5-3 cm. wide

somewhat below the middle, submembranous, lightly scabrous and

in age rugulose above, hispidulous on the reticulate veins beneath;

spikes puberulent, long-pedunculate in the upper axils, several cm.

long in fruit; calyx lobes acute. — Determinations by Killip; seems to

have little to recommend it as a distinct species but the leaf-indenta-

tion is rather more than an extreme variation of that of C. spines-

cens L.

Ancash: Tambo de Pariacota, 2546. — Lima: Chosica, 2862. Ma-

tucana, 302. — Junin: Tarma, Ruiz & Pavdn, type.

Cordia Poeppigii DC. Prodr. 9: 492. 1845.

Open or rather compact shrub, the long flowering branches often

supported in thickets, early more or less hispidulous with spreading

subappressed or incurved trichomes; petioles 5-10 mm. long, decid-

uous above the base; leaves ovate-elliptic, more or less rounded at

base and acutely acuminate, a dm. or so long, 4-6 cm. wide, those in

the inflorescence reduced, all except type ex char, sparsely scabrous

with incurved trichomes above, hispidulous beneath especially on

the slender nerves and rather obscure veins, submembranous, not or

obscurely rugulose; inflorescence when well-developed composed of a

number of slender branchlets from upper leaf-axils, and finally 3 dm.

long or longer; calyx subpapyraceous, glabrous or nearly, at least

typically, about 2.5 mm. long, the ovate lobes acute; corolla greenish-

FLORA OF PERU 587

white, campanulate, 4.5 mm. long, the subglobose anthers barely

exserted from the subtruncate limb, and well-exceeded by the slender

forked style; fruit ovoid, 4 mm. long, finally red, the calyx not or

tardily splitting. — Type, "Central Peru, Poeppig." Determinations

by Killip. F.M. Neg. 7735.

San Martin: Near Tarapoto, Williams 6969. — Huanuco: Chico-

playa, Ruiz & Pav6n. Pozuzo, 456 1. Pampayacu, 5042. — Junin:

Colonia Perene", Killip & Smith 24990. — Loreto: Iquitos, Tessmann

3560 (det. Melchior, C. floribunda); Williams 3543; 8031; Killip &

Smith 271 27; King 1 344; Mexia 6507. Caballo-Cocha, Williams 2060

(toward C. scaberrima?). "Barillas" (Mexia).

Cordia ripicola Johnst. Journ. Arnold Arb. 16: 180. 1935.

Resembles C. laevior; branchlets scabrous with short trichomes;

petioles 2-6 mm. long, leaves uniform, more oblong or somewhat

obovate, 8-14 cm. long, 3-7 cm. wide, sparsely appressed strigillose

above, closely and appressed hispidulous-puberulent beneath; calyx

(type) strigose, subpapyraceous, unevenly rupturing into 2-5 lobes;

corolla lobes about 2.5 mm. long; style and ovary sparsely pubescent

or the latter glabrous. — Calyx character, appressed leaf pubescence

and pubescent pistil separates this from C. Ulei Johnst. and the

scattered appressed trichomes and much appressed ones beneath

from C. Sprucei Mez. (author) of southwestern Brazil. The Schunke

specimen is nearly glabrous, or only scabrous; cf. C. laevior Johnst.

San Martin: Juanjui, Klug 4231 (det. Standley, C. Sprucei). —

Loreto: Mouth of Rio Zubineta, (Klug 2262, type). Rio Mazan,

Jose Schunke 292 (det. Killip). "Chore-ey."

Cordia scaberrima HBK. Nov. Gen. & Sp. 3: 71. (55). 1818.

Branches glabrous, younger tetragonous, scabrous as the leaves

both sides, these rugose, serrate, oblong, narrowed to petioles (4 mm.

long), rigid, reticulate- veined, acuminate or acute, the nerves and

midrib strigose scabrous above, veins scabrous or pilose-hispidulous

beneath; corymbs terminal, dichotomous, the heads sometimes 2 cm.

long, 1.5 cm. thick, often tardily forked; calyx striate, hispid or more

or less hispidulous, 5 mm. long in fruit with broad ovate lobes.—

The Junin specimen has leaves in shape toward var. ecuadorica Killip

but subentire, and an ellipsoid tardily forked inflorescence. The

Williams specimen with peduncles 2 cm. long, heads globose, 7 mm.

thick, calyx 3 mm. long may be the same as one by Poeppig referred

by Johnston with doubt to C. buddleoides Rusby, the calyx without

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

the subulate tips (appendages) that are very evident on that species.

The Huanuco plant, var. ecuadorica Killip, in herb, foliis plus mi-

nusve serratis dense piloso-hirsutis, may be distinct and indeed has

been so designated under a similar name by Johnston. The pedun-

cles are leafy, the globose heads to 2.5 cm. in diameter, calyx in fruit

to 7 mm. long. F.M. Neg. 997.

Piura: Ayavaca, Bonpland, type. — Huanuco: Mito, 3328. Ya-

nano, 3789. — Junin: Huacapistana, Killip & Smith 24518 (det.

Killip, C. floribunda). — Loreto: Santa Rosa, Yurimaguas, Williams

4779 (det. Killip). Ecuador.

Cordia scabrifolia A. DC. Prodr. 9: 485. 1845; 53.

Branchlets and inflorescences minutely puberulent; leaves elliptic-

lanceolate, acute at base, acutely short-acuminate, not markedly

dissimilar, shortly appressed scabrous-strigose above or pustulate,

sometimes smooth and glabrous(?), beneath green, on the nerves

pubescent with some very short, usually inwardly directed stout tri-

chomes; calyx cupulate, about 3 mm. long, nearly as broad, the lobes

about even; corolla about 5 mm. long; anthers nearly 2 mm. long,

exserted; ovary pubescent. — No doubt occurs in Peru.

Rio Acre: Seringal Auristella, Ule 9717. Mouth of Rio Macauhan,

Krukoff 5336; 5672. Amazonian Brazil; British Guiana.

Gordia spinescens L. Mant. 2: 206. 1771. Varronia ferruginea

Lam. Tab. Encycl. 1: 418. 1791; Poir, Encycl. 4: 263. 1797. C. fer-

ruginea (Lam.) Roem. & Schult. Syst. 4: 458. 1819. C. pyrifolia

Willd. ex Roem. & Schult. I.e. 802. C. Thibaudiana DC. Prodr. 9:

489. 1845. C. pauciflora Rusby, Mem. Torr. Bot. Club 6: 83. 1896.

Petiole bases persisting spur-like on the upper branches, the

branchlets, petioles and peduncles more or less tomentulose or red-

dish-hirsute; petioles 5-10 mm. long, leaves often broadly ovate-

elliptic, 7-15 cm. long, about half as wide, acute both ends or more

or less abruptly acuminate at apex, scabrous above, more or less

villous-hispidulous beneath, entire to obsoletely to coarsely serrulate,

firm to subcoriaceous, the nerves and reticulate veins correspondingly

conspicuous; peduncles on the upper petioles or several in a terminal

inflorescence, 1-several cm. long; spikes often few-flowered, some-

times ellipsoid, calyx somewhat campanulate, to about 4 mm. long,

the ovate lobes acute, corolla 5-8 mm. long, tube glandular (Worth &

Morrison). — The persisting indurated spur-like petiole bases sug-

gested the name according to Johnston, who worked out the synon-

FLORA OF PERU 589

ymy, Journ. Arnold Arb. 30: 104. 1949; includes also the Colombian

C. laxiflora HBK. and C. riparia HBK. Nov. Gen. & Sp. 3: 72. 1818,

and 71, pi. 207 (C. riparia). But it is not clear that the breaking

down of the petiole cells does not occur irregularly and its taxonomic

significance is therefore debatable. F.M. Neg. 1020 (C. pyrifolia).

Arequipa: South of Chala, Worth & Morrison 15697. Bolivia

to Mexico.

Cordia Sprucei Mez, Bot. Jahrb. 12: 549. 1890; 53. Lithocar-

dium Sprucei (Mez) Ktze. Rev. Gen. 2: 977. 1891.

Branchlets apically strigose, subferrugineous-tomentulose, gla-

brate, terete or nearly; petioles to 7 mm. long, geniculately incurved

at base; leaves suboval or obscurely obovate, to 1.5 dm. long, 7 cm.

wide, attenuate to rounded base, acutely acuminate, subglabrous

and very lustrous above, pubescent beneath, prominently reticulate

both sides; inflorescence terminal, laxly paniculate, ferrugineous to-

mentulose, much shorter than leaves; calyx before anthesis entire,

urceolate, ecostate, irregularly much lobed, very slightly strigose

within; corolla with stamens to 8 mm. long, more than three-fourths

divided, the 5 rounded lobes finally reflexed; filaments affixed nearly

at throat, long pilose at base, anthers glandular at tip; ovary yellow-

ish pilose as style at base; stigma discoid; drupe tomentulose, to

12 mm. long, 7 mm. thick, the calyx scarcely enlarged. — After author,

who placed it in section Pilicordia; species confused by Fresenius,

in Mart. Fl. Bras. 8, pt. 1: 16, with C. umbraculifera A. DC.

Peru (perhaps). Rio Negro, Brazil.

Cordia toqueve Aublet, PI. Guian. 1: 228, pi. 90. 1775; 52.

C. heterophytta Poir. Diet. Sci. Nat. 10: 409. 1818, fide Johnston.

Branchlets, slenderly branched inflorescence and leaves beneath

softly puberulent-tomentulose; leaves strongly dissimilar, those on a

single branch suborbicular to broadly ovate, alternate or opposite;

larger, long-elliptic, narrowly ovate or lanceolate-oblong ones, often

acute or narrowed to base, rounded or acute at apex, finely pubes-

cent, especially on midrib and lateral nerves or the younger velvety

(Schunke); calyx short conic or proculiform, about 3.5 mm. long,

opening unevenly; corolla about 4 mm. long; anthers oblong, scarcely

1 mm. long. — At once distinct from all relatives with strongly hetero-

morphic foliage by the obviously pubescent ovary, fruit and leaves

(Johnston).

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

Loreto: Lower Rio Nanay, Williams 578. Mouth of Rio Santi-

ago, Tessmann 4125. Rio Mazan, Jose Schunke 360 (det. Killip).

To eastern Brazil and northern South America. "Bacuri" (Wil-

liams).

Cordia ucayaliensis Johnst. Journ. Arnold Arb. 16: 181. 1935.

C. Ulei Johnst. var. ucayaliensis Johnst. Contr. Gray Herb. 92: 57.

1930.

Resembles C. Ulei but leaves more obovate-elliptic, about 1.5 dm.

long, 8-9 cm. wide, and, especially, finely, sometimes sparsely ap-

pressed strigose and scabrous above, rather softly hispidulous to

puberulent beneath; fruiting calyx scarcely lobulate, 8 mm. across,

the ellipsoid fruits 12 mm. long, about 7 mm. thick. — The validity

of the above name may be questioned as the place of publication of

the synonym is not given; readily distinguished from C. ripicola by

the scabrous-strigose upper leaf surface (author). Type a 6-meter

tree, the half as tall trunk 2 dm. in diameter, but Williams noted it

twice as high, the short trunk tortuous.

Loreto: Flood-free forest, Yarina Cocha, Tessmann 5457, type.

Caballo-Cocha, Williams 2333. Yurimaguas, Williams 3832; 4387.

Without data, Poeppig 2196.

Cordia Ulei Johnst. Contr. Gray Herb. 92: 56. 1930.

Branchlets hirsutulous, petioles 5-12 mm. long; leaves obovate-

oblong or oblong-lanceolate, mostly broadest above the middle, be-

low this cuneately contracted or rarely abruptly obtuse or rounded

to base, at apex often rounded or obtuse and abruptly acuminate

(acumen 5-10 mm. long), appressed pubescent beneath, above gla-

brous or sparsely puberulent on the 5-6 nerves, these prominent

beneath and verrucosely joined by transverse veins; inflorescence

terminal, finally short-pedunculate, laxly paniculate, scarcely exceed-

ing the leaves; flowers sessile; calyx poculiform, lightly appressed

strigose, 4-5 mm. long, 2 mm. thick; corolla nearly 8 mm. long,

5-merous, tube cylindric; filaments exserted, villous below; ovary

quite glabrous. — Probably most closely related to C. Sprucei Mez

with pubescent ovary (author); adjacent to southeastern Peru and

doubtless occurring in the general region; a tree attaining 12 meters

(Ule).

Loreto: Iquitos, Tessmann 3613 (det. Killip, C. Sprucei}. — Rio

Acre: Seringal San Francisco, Ule 9716, type.

FLORA OF PERU 591

Cordia varronifolia Johnst. Journ. Arnold Arb. 33: 62. 1952.

Branchlets, leaves beneath and long-ribbed calyces more or less

villosulous; petioles 2-5 cm. long; leaves elliptic-oblong or ovate-

oblong, obtuse or rounded at base, subobtuse, 4-9 cm. long, 2-4 cm.

wide, often crenate-dentate except toward base, densely and minutely

areolate-bullate and with short flexible bulbous-based setae, beneath

deeply reticulate-venose; cymes few-flowered, usually terminal, early

condensed, solitary or rarely geminate, becoming 1-2 cm. long; calyx

10-costate, 1.5-2 cm. long, tube 10-12 mm. long, 4-8 mm. thick be-

low the middle, 3-4 mm. at the top, rounded at base, lobes subulate,

flexuose, 5-9 mm. long, free in bud, the sinuses broad, obtuse; fruiting

calyx to 2.8 cm. long, tube 17 mm. long, 8 or 9 mm. thick; corolla

marcescently persisting, funnelform, 3-3.5 cm. long; limb 2.5-3.5 cm.

across, the semicircular lobes 1 cm. wide, 5-7 mm. long, emarginate,

narrow sinuses strongly acute, the tube about 1 cm. long, 2-3 mm.

thick, lineately villosulous below within, the throat to 12 mm. across

at top; ovary glabrous, attenuate, 4-ovulate, obscurely annulate at

base; style villosulous, 1.5-2.5 cm. long, fork 12-20 mm. above

the base, the very narrow lobes 2-3 mm. long; fruit dry, ovoid,

enclosed by calyx and persisting corolla-tube, the endocarp bony,

4-celled, apparently 4-seeded. — The bullate leaves suggest species in

section Varronia, the long-persisting calyx and corollas members of

the section Gerascanthus; actually allied to C. par vif olio, DC. and

C. elaeagnoides DC. of Mexico, section Rhabdocalyx, the three known

species characterized by the nut-like fruit (Johnston). The widely

distributed type collection was designated as a new species in many

herbaria under the name of the collector by the author's eminent

friend Killip. Type was 2-3 meters tall.

Cajamarca: Pion, valley of the Maranon, Prov. Cutervo, Weber-

bauer 7138 (type).

5. SACCELLIUM Humb. & Bonpl.

A tall smooth tree with alternate entire or remotely dentate leaves

and terminal leafy panicle, the fruiting calyces long-pedicellate, ac-

crescent, oblanceolate-elliptic, 5-dentate, membranous, reticulate-

venose (or more or less nerved?). Flowers unknown. Drupe in-

cluded, obovoid globose, beaked by subulate style, tetragonous above,

sub-4-lobed, little fleshy, the cells 1-seeded, 4 with 2 cavities at the

base of each; seeds albuminous, the cotyledons long-plicate.

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

Saccellium lanceolatum Humb. & Bonpl. PI. Aequin. 1: 47,

pi. 13. 1806.

Branchlets hirtellous toward tip; leaves lanceolate, acute both

ends, reticulate-veined, the principal nerves prominent beneath, hir-

tellous both sides, less than 1.5 dm. long, 7.5 cm. wide, petioles

8-10 mm. long; peduncles superaxillary or terminal, branches few,

about 2 dm. long, the filiform subapical pedicel 6-10 mm. long; calyx

straw colored, pulverulent to about 2.5 cm. long, the drupe 4 or 5

times shorter, lustrous, fragile below, bony above; seed ovoid, pyri-

form, glabrous. — The flowers according to HBK. Nov. Gen. & Sp. 7:

161, were described by Bonpland : limb 6-parted, lobes equal, corolla

segments 5, style 1, stigma bifid. The distinctly lanceolate leaves

and the long foliose branches readily characterize the species (John-

ston, Journ. Arnold Arb. 16: 183. 1935). Shrub or tree to 6 meters

tall, the accrescent calyces yellowish-green (Weberbauer). Illus-

trated, Trans. Linn. Soc. Bot. 1 : 25, pi. 6.

Cajamarca: Near Tomependa, Bonpland, type. Valley of the

Rio Maranon, mouth of Rio Chinchipe, 400 meters, Weberbauer 6217;

6226. Between Jaen and Bellavista, Weberbauer 6209a. Also south-

ern Bolivia to northern Argentina.

6. MYOSOTIS [Dill.] L.

In Peru usually a semi-prostrate or ascending simple or branched

annual, sometimes more enduring, conspicuously pubescent with soft

broadly oblanceolate leaves becoming bractiform above, and small

blue flowers, corolla lobes contorted in the bud. Filaments filiform,

nutlets 4, erect, ellipsoid, lustrous with a basal areola. Stigma disci-

form. — M . virginica (L.) BSP. of North America with small (1-2 mm.

wide) flowers in a very hispid bilabiate calyx has been found in Argen-

tina (Johnston, Contr. Gray Herb. 75: 42. 1925).

The "no-me-olvides" of South American gardens is the following

species.

Myosotis azorica H. C. Wats. Bot. Mag. 70: pi. 4122. 1844; 28.

Stems becoming numerous, elongate, the hispid villous indument

retrorse especially toward the base; lower leaves often several cm.

long, 1-2 cm. wide or wider; racemes solitary or geminate, bractless,

at least above; calyx densely strigose and also with spreading unci-

nate trichomes slightly but obviously irregular, the erect lanceolate

lobes longer than the tube, the lower one the longest; corolla 5-7 mm.

FLORA OF PERU 593

broad; nutlets black, dorsally convex, the face strongly angled and

keeled. — There are several older but still obscure names that are

probably synonymous (Johnston).

Peru: Cultivated and probably escaped, at least in the vicinity

of gardens.

7. MORITZIA DC.

Reference: Johnston, Contr. Gray Herb. 78: 17-20. 1927.

Hispid to strigose perennials, the alternate leaves tufted at base,

the upper reduced, the blue flowers in bractless usually forked uni-

lateral spikes or racemes. Calyx narrowly lobed, accrescent in age,

the ribs uncinately pubescent. Corolla tube cylindric with spreading

imbricate obtuse lobes, the throat with pubescent intruded append-

ages or spots. Stamens affixed above middle, filiform, stigma bi-

lobed. Ovules 4. Nutlets solitary, erect, ovoid, abruptly contracted

below with small areola at base of the strong ventral keel, the gyno-

base flat.

Moritzia Lindenii (A. DC.) Benth. Nat. Pflanzenfam. 4, Abt. 3:

121. 1894. Meratia Lindenii A. DC. Prodr. 10: 104. 1846.

Heavy caudex clothed with dead leaf bases and topped with many

long-decurrent leaves, these greatly reduced and sessile on the one or

two arcuate-ascending stems; indument lax or more or less appressed,

pilose-hispid, dense especially on the leaves beneath, these narrowly

oblong-obovate, the basal to 2 dm. long, to 2.5 cm. wide, those of the

stems 4-6 cm. long, 0.5-2 cm. wide or the uppermost below the di-

chotomous branched inflorescences much reduced, all with anasto-

mosing veins; inflorescences in fruit to about 7 cm. long and wide;

calyces at least lower (2-7 mm. long) shortly pedicellate, hispid with

callus-based trichomes, the lobes narrowly ovate, acuminate, 2.5 mm.

long; corolla 4.5 mm. long, campanulate, the tube barely exserted,

lobes rounded, about 2 mm. broad, sparsely short-hispidulous; style

1-2 mm. long; stamens just included or visible; nutlet white, tubercu-

late-muricate.

Piura: Huancabamba, Weberbauer 6081. Colombia; Venezuela.

8. LITHOSPERMUM L.

Reference: Johnston, Contr. Gray Herb. 70: 18-31. 1924; 78:

6-11. 1927; Journ. Arnold Arb. 33: 299-366. 1952.

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

Perennial or rarely annual, the yellow-orange or white flowers

sometimes heterostyled; never spotted, the corolla throat more or

less glanduliferous, sometimes with intruding appendages, the corolla

tube rarely pubescent. Nectary weak or well developed. Style sim-

ple with two stigmas. Stamens whorled in the corolla tube, the

globose cylindric ellipsoidal more or less ovoid pollen often some-

what constricted medially, with 6-9 pores in one row at or below the

middle of the grain or near one end. Nutlets usually smooth, white,

lustrous. — After Johnston, 312, not including Arnebia species of Asia

and Africa, which he notes differ sharply only in the nature of the

pollen, this cylindric or constricted medially with a row of 4 or 5

pores at each end. In this connection it may be remarked that char-

acters of pollen unaccompanied by significant morphological distinc-

tions have not been found important in the classification of the

Convolvulaceae or Gentianaceae except by a few botanists. John-

ston in his last work gave a detailed description including much

original comment and observation; see also Brand, Repert. Sp. Nov.

28: 10-17. 1930, for a key to the American species.

Corolla greenish-yellow; calyx in fruit 5-8 mm. long.

Stem indument appressed; corolla villous, to 18 mm. long.

L. Macbridei.

Stem indument lax; corolla hispidulous, 2.5 cm. long.

L. cinerascens.

Corolla white; calyx short.

Stems prostrate, hispidulous L. peruvianum.

Stems spreading, early tufted, closely strigose L. Gayanum.

Lithospermum cinerascens (A. DC.) Johnst. Contr. Gray

Herb. 75: 40. 1925; 8. Macromeria cinerascens A. DC. Prodr. 10:

69. 1846.

Stems about 2.5 dm. long, canescent with loosely appressed or

spreading hispid indument, the crowded lanceolate acute leaves

somewhat bulbous-pustulate, 1.5-2.5 cm. long, 4-6 mm. wide, ses-

sile and rounded at base, the midrib prominent beneath, the margins

slightly involute; racemes bracteate, few-flowered; pedicels 3 mm.

long, slender; calyx lobes narrowly linear, acute, appressed hispid-

ulous, 5-8 mm. long; corolla yellowish, tubular, a little contracted

near base, hispidulous, 2.5-3.5 cm. long, the spreading lobes about

3 mm. long; nutlets ovoid, white, lustrous, sparsely punctate, 3 mm.

FLORA OF PERU 595

long; style very long, slender, subexserted. — After Johnston, from

original description and photo.

Amazonas: Chachapoyas, (Mathews, type).

Lithospermum Gayanum (Wedd.) Johnst. Contr. Gray Herb.

78: 10. 1927. Eritrichium Gayanum Wedd. Chlor. Andina 2: 88. 1859.

L. andinum Krause, Bot. Jahrb. 37: 636. 1906. L. Shepardae Johnst.

Contr. Gray Herb. 78: 10. 1927, fide Johnston.

Stout-rooted, more or less ashy strigose, the procumbent or as-

cending stems often branched, 3-12 cm. long; leaves lanceolate to

spathulate-obovate, subobtuse or acute, about 0.5-2 cm. long, 1.5-

4 mm. wide, subcoriaceous, often short-hispid or obscurely pustulate;

racemes foliose-bracteate; pedicels very short; calyx lobes oblong or

linear, the longer white corolla 3.5-6 mm. long, the tube 2.5-3 mm.

long with rounded spreading lobes 2-2.5 mm. long; throat append-

ages pubescent, oblong; nutlets 4 or fewer, ovoid, 2-2.3 mm. long,

lustrous, about as long as the style and calyx lobes. — Most closely

related to L. distichum Ortega of Mexico, but differs in its smaller

stature, focal appendages merely puberulent swellings, glands few,

none at base of filaments and the style without bilobate sterile tip

(Johnston). F.M. Neg. 17354 (L. andinum).

Junin : Near Tarma, Weberbauer 2396 (type, L. andinum) . North

of Yauli, 3,500 meters, Stork & Horton 10885 (det. Standley, L. Mac-

bridei). — Cuzco: Andes of Cusco, Gay, type. — Puno: Rocky clay puna,

Chuquibambilla, 4,000 meters, Pennell 13357; Mexia 04195. Near

Puno, 3,750 meters, Mrs. Reid S. Shepard 114 (type, L. Shepardae).

Saxaihuaman, (Herrera 2208) . Northern Bolivia.

Lithospermum Macbridei Johnst. Contr. Gray Herb. 78: 8.

1927; 355.

A little shrub with many simple erect branches 2-6 dm. high,

shortly and often densely ashy-strigose as also the leaves and calyces;

leaves narrowly lanceolate or oblanceolate, acute or obtuse, 2-5 cm.

long, 4-6 mm. wide, crowded, strict, the midnerve prominent; ra-

cemes terminal, few-flowered, bracteate; calyx 4-5 mm. long, in fruit

5-6 mm. long, the pedicels about 1 cm. long, falcate; corolla tubular,

15-18 mm. long, the attenuate cylindric tube 3-4 mm. thick, exceed-

ing the calyx by about 1 cm., glabrous within, villous-strigose with-

out, the divaricate or recurved lobes orbicular, 2.5-3 mm. long;

throat very obscurely puberulent without appendages; stamens from

above middle of tube, the filaments shorter than the linear anthers;

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

nutlets ovoid, white, lustrous, 2.8-3.2 mm. long, included; style elon-

gate, subfiliform. — Closely related to L. viride Green of Mexico but

with many smaller narrow veinless leaves and a pallid strigose indu-

ment; corolla greenish-yellow with a conspicuous subcylindric tube

10-14 mm. long and a small limb and diverging or recurving lobes;

throat with no intrusions but with many glands, especially below

each corolla lobe; nectary represented by 5-10 villulose swellings;

style finally exserted 2-5 mm. with 2 stigmas; pollen subglobose or

subspherical, 16-20 /* in diameter with rounded or slightly angulate

sides and 8 obscure pores on the equator; mature fruiting calyx

5-8 mm. long, at most only twice as long as the nutlet (Johnston,

I.e. 355, additional characters in contrast to L. viride).

Huanuco: Densely shrubby slope, Yanahuanca, 3,000 meters,

1185, type. Grassy slope, Mito, 1592. Llata, 2,100 meters, 2253.

Lithospermum peruvianum A. DC. Prodr. 10: 77. 1846; 11.

L. aequatoriale Brand, Repert. Sp. Nov. 28: 14. 1930, fide Johnston.

A hispid perennial, ligneous at base; leaves linear, acute, 1.2-

3 cm. long, 2-4 mm. wide, tuberculate, pustulate; racemes few-

flowered, leafy-bracted; calyx appressed pubescent, the linear lobes

acute; corolla 4 mm. long, appressed pilosulous without, funnelform

above the middle, the two scarcely surpassing the calyx, the lobes

ovate, obtuse, the throat with pubescent rounded appendages above

the anthers; nutlets usually solitary, ovoid, 2 mm. long, white, lus-

trous, punctate. — Stems many, slender, elongate, hispidulous; leaves

crowded, 1.5-2.5 cm. long; inflorescence not sharply defined, the

white flowers disposed toward the end of the elongate leafy stems;

corolla tube stout, 1.5-2 mm. long, scarcely or not longer than calyx,

the limb 3-4 mm. across, its broad rounded lobes nearly as long as

the tube; throat with 5 circular glanduliferous areas, the nectary

villose; style 1-2 mm. long with 2 juxtaposed terminal stigmas; pollen

cylindric to ellipsoidal, the pores usually obscure (Johnston, I.e.).

F.M. Negs. 27082; 17352 (L. aequatoriale}.

Amazonas: Chachapoyas, Mathews, type. Ecuador.

9. AMSINCKIA Lehm.

Reference: Johnston, Contr. Gray Herb. 78: 98-100. 1927; Brand,

Pflanzenreich IV. 252: 204-217. 1931.

Rather harshly pubescent erect annuals with alternate obscurely

veined leaves and merely basally bracted (if at all) much elongating

FLORA OF PERU 597

racemes of orange to yellowish flowers. Calyx lobes erect. Corolla

tubular-salverform, the cylindrical tube as throat unappendaged, the

included stamens subsessile in the tube. Nutlets 4, erect on the

well-developed gynobase, angulate-ovoid, not edged, ventrally cari-

nate; areola medial, small, carunculate. — Constitutes a subtribe by

reason of the biparted cotyledons. See Ray and Chisaki, Amer.

Journ. Bot. 44: 529-554. 1957, for a detailed discussion of the occur-

rence and nature of heterostyly in the genus and its cytogenetics and

phylogeny. Named after Wilhelm Amsinck, early nineteenth cen-

tury mayor of Hamburg, a founder of the botanical garden.

Amsinckia hispida (R. & P.) Johnst. Contr. Gray Herb. 73: 75.

1924; 99. Lithospermum hispidum R. & P. Fl. Peruv. 2: 5. 1799.

A. angustifolia Lehm. Delect. Sem. Hamb. Hort. 7. 1831. Plagiobo-

thrys aurantiacus Brand, Repert. Sp. Nov. 20: 46. 1924, fide Johnston.

A. aurantiaca Brand, Pflanzenreich IV. 252: 208. 1931.

Erect, more vigorous individuals attaining several dm., simple or

openly branched, long-hispid-hirsute, above finely villous toward tip;

lower leaves often approximate, oblanceolate, 5-15 (-18) cm. long,

5-15 (-24) mm. wide, cauline, linear-lanceolate, gradually reduced,

all acute; racemes solitary or geminate, sometimes bracted at base,

elongating and in age lax, 1-2.5 dm. long; calyx rather tawny, hirsute

and somewhat hispid-villous, finally 4-6 mm. long, the lobes acumi-

nate- or lanceolate-linear; pedicels ascending, 1-3 mm. long; corolla

3-7 mm. long, style as long as tube; nutlets triangular-ovate, angu-

late, incurved, 2-2.8 mm. long, granulate-tuberculate, commonly

with strong irregular transverse crests, dorsally obtuse and the medial

ridge frequently cristate longitudinally. — After Johnston who gives

complete synonymy. A. tessellata Gray of western North America

has been collected in Chile and Argentina, probably as a weed; the

corolla is 6.5-7.5 mm. long, the nutlets tessellate, 3-3.5 mm. long.

F.M. Negs. 17355 (A. aurantiaca); 17356 (A. Mandonii); 17357 (A.

marginata).

Lima: Rio Blanco, 729; 2968. Chicla, (Ball). Near Cheuchin,

Ruiz & Pavon (type, L. hispidum). — Huanuco: Llata, 2288. — Cuzco:

Paucartambo, Hen era 1006a. — Arequipa: Mollendo, Weberbauer 1522

(type, P. aurantiacus). Ecuador to Chile and Argentina.

10. PLAGIOBOTHRYS Fisch. & Mey.

Reference: Johnston, Contr. Gray Herb. 78: 75-97. 1927; Brand,

Pflanzenreich IV. 252: 105-114. 1931.

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

Hispid, erect, the crowded not obviously opposite lower leaves

forming a rosette, the remaining obviously alternate. Calyx rarely

parted at base. Gynobase with 4 lateral depressions, pyramidal.

Nutlet scar more or less central. — Corolla appendages or swellings

more or less obvious. See Johnston, I.e. 68: 57-80. 1923, for a dis-

cussion of the genus; he concludes that it correctly should be drawn

to include Allocarya Greene, a premise not accepted by Abrams and

others but perhaps an equally feasible disposition.

Plagiobothrys myosotoides (Lehm.) Brand, Pflanzenreich IV.

252: 108. 1931. bithospermum myosotoides Lehm. PI. Asperif. 319.

1818. P. tinctorius (R. & P.) Gray, Proc. Am. Acad. 20 : 283. 1885 ; 80.

L. tinctorium R. & P. Fl. Peruv. 2: 4, pi. 114. 1799, not L.

Erect, often a dm. or two high, the slender branching stems from

a somewhat persisting rosette of oblong-linear to oblanceolate sub-

obtuse leaves 1-4 cm. long, 1-4 mm. wide, the similar stem leaves

alternate, more or less extending as bracts into the finally laxly flow-

ered solitary or geminate racemes; calyx rather tawny, the erect or

openly connivent lanceolate lobes finally 1.5-2.5 mm. long, shortly

exceeded by the corolla tube, the corolla limb 1-2.5 mm. broad; nut-

lets broadly ovate, rounded at base, constricted below tip, 1-1.5 mm.

long, granulate, stoutly carinate dorsally and laterally crested, acutely,

sometimes reticulately ridged or also variously papillate or muricu-

late; scar inframedial in the broad transverse ventral groove slightly

lower than the well-developed ventral keel. — After Johnston, who

remarks that the Peruvian plants seem to be the same as the typical

state from central Chile. Plants suffused with a dye that stains dry-

ing specimen papers.

Lima: Matucana, Weberbauer 5724- — Arequipa: Volcan El Cha-

chani, 4,000 meters, Pennell 13279; Bailey. Chile; California.

11. ALLOCARYA Greene

References: Johnston, Contr. Gray Herb. 78: 98-101. 1927;

Brand, Pflanzenreich IV. 252: 159-182. 1931.

Similar to Plagiobothrys but more commonly prostrate, rarely

erect, and the leaves, including the more or less remote lower, clearly

opposite and never forming a rosette. Calyx divided to base. Corolla

without appendages. Gynobase conical with 4 triangular depres-

sions. Nutlets carinate entire length of inner face. — Brand, I.e., con-

sidered all the Peruvian plants forms of one species, A. humilis

FLORA OF PERU 599

(R. & P.) Greene, and indeed the apparent differences may be shown,

eventually, to be merely variations.

Conservatively and monographically, in work where the primary

purpose of classification is concerned with relationship, these little

plants, in spite of their reasonably homogeneous f acies, perhaps would

be most intelligently treated as comprising a section of Plagiobothrys,

as by Johnston, but realistically, which is to say, floristically and

more conveniently, they are distinguishable as a closely allied but

separate group as by the illustrious Abrams, Brand, and many other

able students. Brand's discussion, I.e. 17, is pertinent here, while his

quotation of Graebner is apropos for any number of similar circum-

stances in classification. Unfortunately many of Brand's judgments

are questionable in view of the unevenness of his work, especially in

drawing generic lines; here for example he considers Allocarya itself

as divisible generically instead of sectionally in order to express the

apparent specific relationships. Plagiobothrys plurisepalus (F. v.

Muell.) Johnst. I.e. 81: 75. 1928 if an Allocarya will make advisable

the conservation of the latter name against Maccoya F. v. Muell.,

1859; cf. Brand, I.e.

Perennials; nutlets rugose or ridged.

Style well-exserted from mature nutlets; flowers axillary or pseudo-

scapose A. Kunthii.

Style at most as long as nutlets; flowers in bracteate racemes.

Calyx tips reddish; leaves greenish A. humilis.

Calyx tips obscurely flavescent; leaves canescent. .A. pygmaea.

Annual; nutlets not rugulose A. Macbridei.

Allocarya humilis (R. & P.) Greene, Pittonia 1: 17. 1887; 84.

Myosotis humilis R. & P. Fl. Peruv. 2: 5. 1799. Plagiobothrys humilis

(R. & P.) Johnst. Contr. Gray Herb. 68: 75. 1923.

Glabrescent or sparsely strigose perennial (calyx appressed villos-

ulous) with prostrate or often ascending stems, usually rooting at

lower nodes; leaves linear, widened at base, rounded at apex, 2-6 cm.

long or longer, 1.5-3 mm. wide, sometimes only ciliate; racemes irreg-

ular, alternate bracted, finally very open, elongating to 9 cm.; pedi-

cels to 1 mm. long; calyx canescent, the erect lanceolate lobes tawny

pubescent above, 2.5-3.5 mm. long; corolla white, the limb 1-2 mm.

across; nutlets ovate, acute, 1-1.5 mm. long, granulate, unevenly

reticulate-ridged, definitely carinate only above the middle, scar

suprabasal; style at most equaling nutlets. — Described as a hispid

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

annual, no doubt by error. Young or poorly developed plants with

glomerate racemes among the leaves; simulate A. congestus typically

at least with canescent indument. Characteristically begins to flower

in wet often stony soils.

Ancash: Catuc, east of Huaraz, 3,000 meters, 2510. — Lima: East

of Canta, Pennell 14640. Matucana, 2,400 meters, 459. Near Chi-

cla, Ball. Banos, (Wilkes Exped.). — Huanuco: Pillao, Ruiz & Pawn,

type. Pampayacu, Sawada 94. — Junin: La Quinua, 3,600 meters,

2013. — Apurimac: Chincheros, Stork & Horton 10745 (det. Stand-

ley). Ampuy, Goodspeed Exped. 10631 (det. Standley); Vargas 515

(det. Standley, Cryptantha granulosa). — Cuzco: La Raya, 4,300 me-

ters, Pennell 13504- Marcapata, Vargas 9712 (det. Standley). Quis-

picanchis, Vargas 920 (det. Standley, Cryptantha granulosa) . Sicuani,

(Hickeri). — Moquegua: Carumas, Weberbauer 7325a. Bolivia.

Allocarya Kunthii (Walp.) Macbr. Field Mus. Bot. 4: 119. 1927.

Anchusa Kunthii Walp. Nov. Act. Caes. Leop. Nat. Cur. 19. suppl.

372. 1843. A. linifolia (Lehm.) Macbr. var. Kunthii (Walp.) Macbr.

Proc. Am. Acad. 51 : 545. 1916. Antiphytum Walpersii A. DC. Prodr.

10: 122. 1846. Eritrichium Walpersii (A. DC.) Wedd. Chlor. And. 2:

90. 1859. Plagiobothrys Kunthii (Walp.) Johnst. Contr. Gray Herb.

68: 74. 1923; 83.

Perennial, cespitose-tufted or the more or less developed repent

stems rooting at the nodes and forming mats; leaves crowded toward

base, linear, subobtuse, 1-2.5 cm. long, to 1 mm. wide, sparsely ap-

pressed hispid-villous and ciliate, thin at dilated base; flowers soli-

tary, apparently scapose in stemless plants from the congested leaves,

otherwise in the axils of the stem leaves; calyx slightly villosulous,

rounded in fruit, the narrow lobes 2-4 mm. long, the stout pedicels

3-6 mm. long; corolla white with yellowish appendages, tube 3-

3.5 mm. long, limb 4-6 mm. broad; nutlets ovate, acute, 1.4-1.8 mm.

long, coarsely reticulate-rugose at least above the middle, ridges often

spiculate, medial keel definite toward apex, areola suprabasal; style

much longer than nutlets. — This follows Johnston's interpretation

and is probably correct, but, as he remarks, type not seen.

Puno: Low pampa, Lake Titicaca, (Shepard 4)- Granja Modelo

de Chuquibambilla, Mexia 04185. Adjacent Bolivia to Ecuador.

"Sillo-sillo" (Mexia).

Allocarya Macbridei (Johnst.) Ferreyra, comb. nov. Plagiobo-

thrys Macbridei Johnst. Contr. Gray Herb. 78: 88. 1927.

FLORA OF PERU 601

Prostrate or laxly ascending annual with many slender minutely

appressed villous stems 3-several cm. long; leaves linear, little dilated

toward base, subacute, sparsely hispid-villous; racemes lax, regularly

bracted, the linear bracts 0.5-2 cm. long; calyx appressed villous,

finally 2-2.5 mm. long including the erect lanceolate lobes; corolla

subtubular, about 1 mm. wide; nutlets ovate, about 1.5 mm. long,

somewhat angled, irregularly tuberculate or papillate, very obscurely

carinate; areola suprabasal, large, more or less excavated. — Clearly

related to the Chilean A. procumbens (Colla) Greene but lacking

transverse ridges (Johnston). Doctor Ramon Ferreyra, the distin-

guished Peruvian botanist, has kindly transferred this interesting

species to which our friend assigned my name. Brand included this

in A. humilis (R. & P.) Greene, but it is probably distinct.

Lima: In short grass, Viso near Matucana, 2,700 meters, 599,

type.

Allocarya pygmaea (HBK.) Macbr. Field Mus. Bot. 4: 119.

1927. Anchusa pygmaea HBK. Nov. Gen. & Sp. 3: 92. 1818. Eri-

trichium humilis (R. & P.) A. DC., var. congestum Wedd. Chlor. And.

2: 88. 1859. Plagiobothrys congestus (Wedd.) Johnst. Contr. Gray

Herb. 68: 75. 1923.

Resembles A. humilis but not so coarse, canescently hispid-villous,

at least the leaves which usually exceed the ordinarily glomerate

racemes, these 1-3 cm. long, leafy-bracted; calyx canescent or at

most flavescent. — Amsinckia humifusa [Poeppig] Walp. Nov. Act.

Caes. Leop. Nat. Cur. 19. suppl. 371. 1843, essentially a nomen

nudum, probably belongs here in part (Johnston). F.M. Neg.

32969.

Arequipa: Chiray, 3,900 meters, Weberbauer 6889. — Puno: Chu-

quibambilla, Pennell 13397. Occa Pampa, 3,700 meters, (Shepard

107). — Moquequa: Carumas, 4,000 meters, Weberbauer 7325. To

Argentina.

12. CRYPTANTHA Lehm.

References: Johnston, Contr. Gray Herb. 74: 1-114. 1925; I.e. 78:

30-74. 1927.

Peruvian species (at least) erect slender hispidulous annuals, one

(C. parviflora) not producing cleistogamous flowers, all with opposite

basal leaves and inconspicuous white flowers in bracted or bractless

racemes. Calyx parted basally. Nutlets usually 4, laterally affixed

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

by a medially, often basally dilated groove to a more or less pyrami-

dal gynobase and somewhat enclosed in the mature calyx. — Brand

disagreed with Johnston in application of some names, especially as

to C. parviflora (Phil.) Reiche, which he restricted to Chile; the care-

ful and thoughtful later student saw the type as well as several

Chilean and Peruvian collections.

Nutlets thin-margined, dissimilar; spikes not bracted or only at base.

C. parviflora.

Nutlets obtusely if at all margined; spikes bracted except C. peruviana.

Corolla showy, to 7 mm. wide; style scarcely or not exserted.

C. varians, C. granulosa.

Corolla 0.5-1.5 (2) mm. wide; style longer than nutlets.

Calyx in fruit 3-4 mm. long, narrow lobes suberect to tips.

C. filaginea.

Calyx in fruit 5-6 mm. long, broad lobes spreading at tips.

C. limensis.

Corolla to 1.5 mm. wide; style barely longer than nutlets.

C. peruviana.

Cryptantha filaginea (Phil.) Reiche, Anal. Univ. Chile 121: 829.

1908; 46. Eritrichium fllagineum Phil. I.e. 90: 536. 1895.

A dm. or two tall, the basally or apically branched slender stems

finely strigose and hispid, the leaves linear or lanceolate, obtuse,

1-2.5 (-3) cm. long, 1-2 mm. wide, pustulate, finely short-hispid;

spikes solitary or geminate with caducous linear bracts 3-6 mm. long;

fruiting calyx oblong, 3-4 mm. long, usually crowded, sessile, readily

deciduous, the lobes finally linear, appressed short hispid below, hir-

sute on prominent midrib, above slightly pubescent; corolla white,

narrow, rarely 2 mm. broad; nutlets usually 4, nearly alike, the axial

one most persistent, tuberculate, usually obscurely granulate, 1.2-

1.8 mm. long, acute, angled, often margined, the narrow or closed

groove forked at base; style much longer than nutlets, the gynobase

a third to a fourth shorter. — An old plant, probably C. parviflora,

with the type, possibly suggested the name (Johnston).

Arequipa: Mollendo hills, Johnston 6280; Hitchcock 22384- Chile.

Cryptantha granulosa (R. & P.) Johnst. Contr. Gray Herb. 68:

54. 1923; 44. Myosotis granulosa R. & P. Fl. Peruv. 2: 5. 1799.

C. latifolia Johnst. I.e. 73: 73. 1924, fide author.

FLORA OF PERU 603

A few cm. to 3 dm. tall, the stems often laxly branched, finely

more or less appressed hispid; leaves sessile, linear to oblong, 2-

3.5 cm. long, 2-4 or rarely 5-10 mm. wide, sparsely pustulose-setose;

spikes solitary or geminate, 3-12 cm. long, the few to many bracts

3-5 (-10) mm. long; fruiting calyx subsessile, ovate, 2-4 mm. long,

mature lobes linear or lance-linear, obtuse, sparsely appressed hispid,

but short-hirsute on midrib below; corolla about 5 mm. long, 4-

4.5 mm. across at tip; nutlets 4, alike or the axial subpersistent one

slightly the largest, 1.3-1.8 mm. long, tuberculate or papillate and

somewhat glandular, rounded or obtuse at base, acute, angled, groove

closed or dilated basally; gynobase to three-fourths length of nutlets,

about equaling the style. — After Johnston, who decided his C. lati-

folia was a luxuriant phase of the species; Brand described the throat

glands as oculiform. Illustrated, Pflanzenreich IV. 252: 66, fig. 4-

Lima: Chancay, Ruiz & Pavon, type; Stork & Vargas 9350. Mt.

San Augustin near Lima, Weberbauer 5695. Chorillos, near Lima,

5861 (type, C. latifolia). Near Mala, Vargas 9320. — Arequipa: Mol-

lendo, (Hitchcock 22857); Johnston 3532(?); Mexia 04181; Worth &

Morrison 15765 and east of Chala, 15611. Near Atiquipa, Worth

& Morrison 15658.

Cryptantha limensis (A. DC.) Johnst. Contr. Gray Herb. 70:

46. 1924; 48. Eritrichium limense A. DC. Prodr. 10: 133. 1846.

C. Macbridei Johnst. I.e. 73: 74. 1924. C. Woitschachii Brand, Repert.

Sp. Nov. 20: 317. 1924.

Finely appressed hispid, sometimes a dm. or so tall, the oblong-

lanceolate subobtuse leaves more or less pustulate, 1-3 cm. long,

2-4 mm. wide; spikes solitary or geminate, 3-5 cm. long; fruiting

calyx subsessile, ovate-oblong, 5-6 mm. long, the finally linear to

oblanceolate lobes very hirsute below on midrib, the herbaceous

spreading upper part merely hispid; corolla subtubular, about a mm.

wide; nutlets 4, alike or nearly, truncate at base, acute, angled, tuber-

culate, 1.5-1.7 mm. long, the groove dilating into a suprabasal tri-

angular areola; axial nutlet the most firmly set, now and then the

largest, the gynobase about three-fifths as high, the style longer. —

After Johnston (as determinations) who concluded that both recently

proposed species are clearly synonyms.

Lima: Stony slopes, Chorillos, 5869 (type, C. Macbridei). San

Augustin base, Weberbauer 5693. Barranca, Worth & Morrison 9107;

Weberbauer 5700 (this and preceding det. Brand C. granulosa; col-

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

lections may be mixed). Near Lima, Abadia, type. — Arequipa: In

and on edge of fertile belt, Mollendo, Johnston 3531 . North of Mol-

lendo, Worth & Morrison 15474- Without data, (Woitschach, type,

C. Woitschachii).

Cryptantha parviflora (Phil.) Reiche, Anal. Univ. Chile 121:

821. 1908; 39. Eritrichium parviflorum Phil. Fl. Atac. 39. 1860.

C. piscoensis Brand, Repert. Sp. Nov. 20: 49. 1924. C. umbelliformis

Brand, I.e. 317, fide Johnston. C. Seleri Brand, I.e.

Slender, usually with a single stem laxly branched from near base,

sometimes about 2 dm. tall, more or less appressed canescent hispid

and often also with few short-hispid trichomes; leaves linear, mostly

1-4 cm. long, 1-2 mm. wide, appressed hispid or spreading short hir-

sute, somewhat pustulate beneath, the upper early rather obscurely;

solitary or geminate spikes, these many, 1-5 cm. long, bractless or

with one bract at base; fruiting calyx ovate or oblong-ovate, 1.5-

2 mm. long, sessile, readily deciduous, the mature calyx lobes lanceo-

late-oblong, connivent, densely short hirsute; corolla 0.3-0.5 mm.

broad; nutlets usually 4, clearly different, ovate or triangular-ovate,

tuberculate or muricate, acute, rounded at base, dorsally convex,

the edges sharply margined, the groove at least below dilated into a

shallow triangular areola; odd nutlets abaxial, largest and most per-

sistent about 1 mm. long, the consimilar ones 0.6-0.8 mm. long, read-

ily deciduous; gynobase about two-thirds height of odd nutlet, the

style much surpassing the latter. — Johnston saw no authentic mate-

rial of C. Seleri but assumed it to be the hispid plant collected by

Pennell and C. umbelliformis to be a young state of this, while C.

piscoensis with appressed pubescence is a somewhat coarser form

collected at Quive by Pennell. He remarked the close relationship

of this species to the North American C. angustifolia (Torr.) Greene

and C. Grayi (V. & R.) Macbr., resembling the former in nutlets,

the latter in aspect and measurements.

Lima: Quive, Pennell 14300. Near Chosica, Weberbauer 5337

(type, C. umbelliformis}. — lea: Above Pisco, 1,200 meters, Weber-

bauer 5375 (type, C. piscoensis). — Arequipa: Tingo, Pennell 13112.

Above Arequipa, Pennell 13177. Yura, (Born). — Moquegua: South-

west of Moquegua, Weberbauer 7398a. Chile.

Cryptantha peruviana Johnst. Contr. Gray Herb. 73: 74. 1924;

50. C. Weberbaueri Brand, Repert. Sp. Nov. 20: 318. 1924 and

C. cajabambensis Brand, I.e. 319, fide Johnston. C. debilis (Phil.)

FLORA OF PERU 605

Reiche var. peruviana (Johnst.) Brand, Pflanzenreich IV. 252: 49.

1931.

Short-hispid, sometimes long-branched, slender, often a dm. or

two tall, the many obscurely pustulate linear leaves 1-3 (-6) cm.

long, 1-2 mm. wide; spikes solitary or geminate, early congested,

finally to 5 cm. long, bractless; fruiting calyx subsessile, ovate, 2.5-

3 mm. long, the narrowly lanceolate lobes erect, appressed hispid-

villous, somewhat hirsute along the midrib; corolla sub tubular, the

limb 0.8-1.4 mm. wide; nutlets 4 or fewer, alike, the subpersistent

axial one always present, all lanceolate-ovate, acute, 1.5-1.8 mm.

long, coarsely muricate or also rugose, granulate, acute at edges, the

groove closed or basally dilated; style barely exceeding nutlets, the

gynobase about three-fifths as long. — Closely related to C. globulifera

(Clos) Reiche of Chile and Argentina with fruiting calyces 3-4 mm.

long, the lobes connivent above with somewhat spreading tips (John-

ston) ; stems stouter, leaves a little longer, var. peruviana according

to Brand. I have not seen C. debilis, described as decumbent-

ascending, nutlets 4, transverse-rugose — data contrary to Brand's

conclusion.

Ancash: Hacienda Cajabamba between Samanco and Huaraz,

(Weberbauer 3041, type, C. cajabambensis) . Rocky places, 3,200

meters, Ocros, (Weberbauer 2658, type, C. Weberbaueri) . — Lima:

Rocky slope, 3,600 meters, Rio Blanco, 674, type. — Arequipa: In

gravel along streams, base of El Misti, Pennell 13235. El Chachani,

Hinckley 77. — Moquegua: Open mixed formation, Tocata, 2,200

meters, Weberbauer 7398. Chile?

Cryptantha varians Brand, Repert. Sp. Nov. 24: 57. 1927.

Erect, slender, branched and lightly hispid with rather long some-

what spreading trichomes; leaves linear, 2-4 cm. long, 2-3 mm. wide;

terminal spikes elongating, sometimes sparsely bracted by reduced

leaves; sepals oblong, obtuse, 2 mm. long; corolla to 3.5 mm. long,

4 mm. across at tip; throat glands semilunate; gynobase subulate,

little shorter than the fruiting setose-hispid calyx, this 2.5 mm. long;

nutlets 1-3, ovoid, acute, 1.25 mm. long, brown, densely whitish

granulate, the groove at base divaricately forked. — Referred by

Johnston to C. granulosa (R. & P.) Johnst. but perhaps distinguished

by the smaller flowers, more pubescent calyx, and fewer nutlets —

differences of course that may be shown to be variable by study of

sufficient collections.

Lima: Rocky outcrop north of Chosica, Weberbauer 5840, type.

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

13. HAGKELIA Opiz.

References: Johnston, Contr. Gray Herb. 68: 43^48. 1923; Brand,

Pflanzenreich IV. 252. 1931.

Herbs, ordinarily biennial or perennial with broad veiny alternate

leaves and flowers in paniculately disposed racemes, inconspicuously

or not bracted, the pedicels slender, finally recurving. Calyx basally

parted, the divisions spreading. Corolla lobes rounded, imbricate,

the throat with trapeziform intruded glanduliform appendages. Sta-

mens medially affixed, included, the short filaments slender, the an-

thers slender to elliptic. Style slender, scarcely or not surpassing the

4 nutlets, these erect, ovate, affixed ventrally to a pyramidal gyno-

base by a broad medial areola, marginally glochidiate with frequently

basally confluent appendages, dorsally smooth. — Commemorates a

German professor, Josef Hackel, of the early nineteenth century,

who shared the lively interest of his botanical friends.

KEY (after Brand)

Nutlets similar; throat glands subquadrate; calyx 3, corolla 6 mm.

long, blue to violet H. andicola.

Nutlets dissimilar; throat glands semilunate; corolla white or pale.

Corolla inconspicuous, tinted to white; glands trifid at base.

H. parviflora.

Corolla showy, white; glands subentire H. revoluta.

Hackelia andicola (Krause) Brand, Pflanzenreich IV. 252: 120.

1931. Cynoglossum andicolum Krause, Bot. Jahrb. 37: 635. 1906.

Stems ascending-erect, more or less white hispid as the long peti-

oles of the lower lanceolate leaves, these 6-8 cm. long, to 12 mm.

wide, puberulent-hirsutulous but green, herbaceous, reduced upward,

finally sessile, nearly oblong; racemes short, forming a narrow pan-

icle; pedicels about 1 mm. long; calyx white hispidulous and ciliate,

3 mm. long; corolla subrotate with short tube, 6 mm. long, 8.5 mm.

wide; throat glands yellow, papillose, subquadrate; nutlets similar,

the body as the appendages about 3 mm. long; the scar triangular,

small. — In part after Brand; the group needs revision with much

material; if the flower and fruiting characters are constant it may

be distinct. H. mexicana (Cham. & Schlecht.) Johnst. seems to be

similar but the nutlets are said to be dissimilar, the aculei long and

FLORA OF PERU 607

short as those of H. revoluta; it has been found as near as Ecuador.

F.M. Neg. 49606.

Cajamarca: Toward Hualgayoc, 4,100 meters, (Weberbauer 4230,

type).

Hackelia parviflora (Krause) Brand, Pflanzenreich IV. 252: 122.

1931. Cynoglossum parviflorum Krause, Bot. Jahrb. 37: 634. 1906.

Type slender, weakly erect, puberulent hispidulous-pilose; basal

or lower leaves elliptic or ovate-elliptic, petioles to 5 mm. long, the

gradually reduced upper ones sessile, subovate, all acute, the largest

5 cm. long, 2 cm. wide; racemes short, terminal, lax; pedicels 1-

1.5 mm. long; calyx densely strigose, 2 mm. long, lobes lanceolate,

slightly longer than the glabrous corolla tube, this shorter than the

rounded corolla lobes; throat glands puberulent, unevenly curved or

bilobate at base; nutlets dissimilar, the largest 3 mm. long without

the appendages, 2 minute (Brand). — See remark under H. andicola.

Illustrated, Brand, I.e. 121, fig. 13 (opened flower). F.M. Neg. 17358.

Junin : Between Tarma and La Oroya, 3,700 meters, ( Weberbauer

2520, type).

Hackelia revoluta (R. & P.) Johnst. Contr. Gray Herb. n. ser. 68:

45. 1923. Cynoglossum rewlutum R. & P. Fl. Peruv. 2: 6. 1799. Lap-

pula revoluta (R. & P.) Brand, Repert. Sp. Nov. 14: 148. 1915.

C. ovatifolium Griseb. Goetting. Abhandl. 24: 271. 1879, fide Brand.

C.Fiebrigii Krause, Bot. Jahrb. 37: 634. 1906, fide Brand.

Becoming perennial from a rhizome, the 1-few rather stout stems

often several dm. tall, branched above, more or less hispidulous-

pilosulous as the petioles and calyces; leaves typically oblong- or

ovate-elliptic (petioles of lower to 1.5 dm. long), not greatly reduced

upward, all acute, green but appressed puberulent-hispid, those on

the stems mostly 5-10 cm. long, 2-4 cm. wide except the sessile

uppermost on the lax flowering branchlets; racemes short; recurving

fruiting pedicels nearly equaling the calyces, these with oblong seg-

ments 3 mm. long; corolla campanulate, about 5 mm. long, lobes and

tube subequal; throat glands large, subquadrate; nutlets dissimilar,

2 long-glochidiate, 2.5 mm. long, 2 very shortly appendaged (Brand).

—F.M. Neg. 17359.

Johnston, I.e., included here the proposed Bolivian species of

Krause but apparently with some reservation on the ground of in-

sufficient materials; the problem seems to concern the constancy and

significance of the apparent differences, which are certainly slight;

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

actually there may be only one species — variable — extending from

Mexico to Argentina. The following specimens are referred here as

determined, without consideration of Brand's characters. Forma

ovatifolia (Griseb.) Brand and f . Fiebrigii (Krause) Brand refer to the

obvious intangible variation in leaf-form, the latter including indi-

viduals with more lanceolate leaves, mostly 5-15 mm. wide.

Lima: San Juan de Chicla toward Puente de Anchi, (Weberbauer

207, det. Brand; 169). — Junin: Huassahuassi, Ruiz & Pavdn, type.

Near Yauli, 3,500 meters, Stork & Horton 10880 (det. Standley).

Near Huancayo, Soukup 3990. — Apurimac: Chincheros, Stork &

Horton 10744 (det. Standley). Prov. Canas, 3,725 meters, Vargas

11038 (det. Standley) ; 50 (det. Brand). Saxaihuaman, Herrera 2203.

Hacienda Ayusbamba, Vargas 901 (det. Standley). Prov. Anta,

Herrera 3630; 3631 . Northward in the Andes from Argentina.

14. PECTOCARYA DC.

Reference: Johnston, Contr. Gray Herb. 70: 34-39. 1924.

Ascending to erect annual herbs with linear and opposite lower

leaves. Flowers leafy-bracted, borne on most of the stems or branches.

Calyx basally divided, accrescent. Corolla tubular, the small imbri-

cate lobes ascending, the throat with intruded appendages. Stamens

affixed at middle of tube, the anthers oblong or elliptical. Nutlets 4,

obovate to linear, divaricate, geminate, uncinately pubescent, affixed

apically to a low broadly pyramidal gynobase. — After Johnston who

remarks: a very distinct genus.

Nutlets subulately fringed dorsally as marginally P. gracilis.

Nutlets smooth dorsally P. lateriflora.

Pectocarya gracilis Johnst. Contr. Gray Herb. 70: 36. 1924; 115.

Stems finely strigose, erect or ascending, leaves filiform, linear,

1-3 cm. long, 0.7-1 mm. wide, appressed hispid-villous; flowers all

chasmogamic; corolla short, salverform; nutlets densely pubescent,

the short soft trichomes uncinate; body of nutlet obovate-oblong,

1.5-1.7 mm. long, about 7 mm. broad without a proper margin but

the back and upper sides with subulate subterete appendages 0.6-

1 mm. long, these ending in uncinately tipped bristles. — Apparently

related to P. lateriflora but distinctive by the peculiar nutlike ap-

pendages (Johnston).

Arequipa: Sandy pampa, El Chachani, 2,400 meters, Hinckley 41,

type. Above Arequipa, Pennell 13165. "Estrella gatevdora."

FLORA OF PERU 609

Pectocarya lateriflora (Lam.) DC. Prodr. 10: 120. 1846; 117.

Cynoglossum lateriflorum Lam. Encycl. 2: 239. 1786. C. pilosum

R. &P. Fl. Peruv. 2:6,pJ. m&. 1799. C.lineareR.&P.lc. P.line-

aris (R. & P.) DC. Prodr. 10: 120. 1846.

Stems finely appressed, hispid-villous or strigose; leaves filiform

to linear-lanceolate, 1-5 mm. long, 1-2.5 mm. wide, usually atten-

uate both ends, ordinarily somewhat canescently appressed hispid-

villous; flowers all chasmogamic; corolla short salverform; nutlets

usually not identical, the abaxial one usually more pubescent and

roughened, the others commonly smooth and glabrate or sparsely

pubescent; nutlet body obovate or slightly oblong, 2-2.5 mm. long,

1-1.8 mm. broad with a definite medial dorsal ridge and finally a

well-developed spreading margin, pectinately and lacerately cleft or

divided into subulate or triangular teeth that terminate in uncinately

tipped bristles. — After Johnston (as other descriptions), who re-

marks: characterized by its obovate definitely margined nutlets and

rather coarse stems and leaves; a plant of the lomas and desert margins.

Lima: Huara, Ruiz & Pavdn. Amancaes, 200 meters, Pennell

14806. Atocongo, 400 meters, Pennell 14791. Canta, 2,800 meters,

Pennell 14583. Obragillo, Wilkes Exped. Rio Blanco, 3,600 meters,

675. San Geronimo, 150 meters, 5907. Near Chosica, Weberbauer

5330. — Arequipa: Atiquipa, Worth & Morrison 15654- Mollendo,

Johnston 3534; Hitchcock 22382. — Moquegua: Open mixed formation,

Weberbauer 7396. Ecuador.

VERBENACEAE Endl. Verbena Family

Avicenniaceae included expediently; Moldenke, Phytologia7: 123

et seq. 1960.

Herbs, shrubs or trees commonly with opposite or verticillate

(rarely alternate) leaves, simple except Vitex (in Peru), and variously

disposed but usually hermaphrodite gamopetalous flowers, the calyx

persisting, rarely with separate segments, the corolla sometimes more

or less zygomorphic, the tube ordinarily well developed. Stamens 4,

in 2 pairs, or 2, sometimes unequal in length; again, only 2 fertile.

Ovary superior, not lobed or parted, 2-4 (10) -celled or in Duranta

4-carpellate, 8-celled, becoming drupaceous or subcapsular (in Avi-

cennia capsular), finally often separating into achene-like nutlets or

pyrenes. Style terminal, single, entire or the stigma 2 (4-5) -parted.

In so far as this account of the species found to date (1955) in

Peru is complete, thanks are due to the extraordinary industry of

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

Harold Norman Moldenke as indicated by the records published by

him since 1934. But it has been difficult to compile intelligently from

his descriptive work because usually it lacks indication of characters

peculiar to the plant described or its apparent relationship; some-

times discussions of the latter have been promised in monographs in

preparation; Aegiphila, Petrea, Vitex and a few small genera have

been monographed by him to date (1959). Furthermore, individual

peculiarities often are obscured in the monotony of detailed diagnosis

of characters common to related forms. This bespeaks the almost in-

finite patience of the observer but questionably advances knowledge

or favors its dissemination in reviews like this, for identification.

The family obviously is closely knit or reticulate in living char-

acter-development; the generic key therefore is difficult to use and,

like Moldenke's, is after former students, especially the most recent

one, Briquet. A simpler key could be devised if number of stamens

and number of nutlets were found not sound indicators of related

groups. Basistemon Turcz. once referred here belongs to the Scroph-

ulariaceae (see Moldenke, Pennell, Bull. Torrey Club 63: 348. 1936).

Avicenniaceae Endl. may properly be a family according to the in-

formative and well-presented data assembled by Moldenke, Phyto-

logia 7: 135-139. 1960.

Teak (Tectona grandis L. f.) of southern Asia is of course the out-

standing member of the family; Moldenke, Phytologia 1: 154-164.

1935; 5: 112-120, 134-142. 1955, compiled an encyclopedic account

of this beautiful and valuable tree. Species that have added to man's

esthetic well-being include the colorful Verbena mats and Lantana

hedges, many Peruvian.

Inflorescence simple or narrowly paniculate, sometimes short or capi-

tate or condensed spikes, sometimes extended, branched or pan-

icled; plants (Peru) often semiherbaceous and (or) low.

Fruit splitting into 4 (rarely fewer) nutlets (pyrenes).

Stamens 4; calyx not or little accrescent 1. Verbena.

Stamens 2; calyx in fruit basally wider 2. Hierobotana.

Fruit splitting into 2 nutlets.

Nutlets 1-celled, 1-seeded; calyx not inflated.

Calyx not or little longer than wide; stamens (fertile) 4.

Calyx truncate; teeth more or less obscure; fruit somewhat

soft 3. Lantana.

Calyx 2-4-cleft or dentate; fruit dry, firm 4. Lippia.

FLORA OF PERU 611

Calyx clearly longer than wide, narrow, costate.

Stamens 4 5. Bouchea.

Stamens 2 6. Stachytarpheta.

Nutlets 2-celled, 2-seeded; calyx inflated or accrescent.

Nutlets echinate; plants pilose 7. Priva.

Nutlets rugulose; plants glabrescent 8. Castelia.

Inflorescence more or less compound (sometimes reduced), or simple

and (or) axillary; plants commonly at least in age shrubby,

lianas or arborescent.

Flowers variously disposed but not characteristically cymose.

Fruit with 2 united 1-seeded nutlets; flowers racemose; calyx

showy 9. Petrea.

Fruit with 2 or 4 nutlets, these 2-several-seeded; calyx small.

Calyx distinct from fruit; racemes often few-flowered.

Nutlets 2 10. Citharexylum.

Nutlets 4 11. Duranta.

Calyx merged with fruit; racemes few-flowered.

12. Rhaphithamnus.

Flowers cymose or cymose-paniculate at least characteristically

(inflorescence rarely reduced, axillary).

Flowers more or less zygomorphic; stamens 4, or 2 fertile.

Drupe with 4 unparted nutlets; bracts showy. .13. Amasonia.

Drupe with 1 nutlet.

Leaves simple; fertile stamens 2 14. Cornutia.

Leaves normally compound; fertile stamens 4. . .15. Vitex.

Drupe 4-parted with 4 or fewer nutlets; stamens 4, long-

exserted 16. Clerodendrum.

Flowers regular; stamens alike, 4 or 5.

Calyx subentire to lobed; drupe (often dry) with 2-4 nutlets.

Calyx not accrescent 17. Callicarpa.

Calyx accrescent 18. Aegiphila.

Calyx with nearly free imbricate segments; capsule tomentose,

1-seeded 19. Avicennia.

1. VERBENA L.

Glandularia Gmelin, Syst. 2: 920. 1791. Junellia Mold. Lilloa 5:

392. 1940.

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

References: Schauer in DC. Prodr. 11: 522-700. 1847; Perry, Ann.

Mo. Bot. Gard. 20: 239-362. 1933; Schnack & Covas, Darwiniana 6:

469-476. 1944.

Herbs or more or less suffrutescent, generally diffuse or even de-

cumbent, matted or tufted, rarely glabrous. Leaves seldom verticil-

late, sometimes imbricate, small or rarely scale-like, alternate, entire

or variously incised. Flowers in elongate to capitate spikes or these

rarely reduced, 1-few-flowered, usually terminal; bracts ordinarily

narrow; corollas mostly small but sometimes colorful, funnelform to

salverform, sub-bilabiate. Calyx 5-costate, slightly if at all accres-

cent. Stamens 4, didynamous, the anthers now and then append-

aged, sometimes vestigially, the cells rarely slightly diverging. Ovary

4-celled, ovules solitary, lateral just above base. Fruit dry with 4

finally separating nutlets.

For confirmation of my impression that Junellia Mold., a group

name for several small alpine more or less depressed ligneous species,

is correctly a part of Verbena, representing merely an ecological con-

dition, see Troncoso, Darwiniana 8: 487. 1949, who showed that

habit varies and that anthers here may be appendaged; compare, too,

Phyla under Lippia. On the other hand there may be a natural rela-

tionship within the scope of Glandularia as defined by Schnack and

Covas, I.e., but since their conclusion was based on relatively few

species and neither they nor apparently other students have followed

up their work after more than a decade the soundness of the segrega-

tion is certainly open to question; Schauer was probably right in

grouping the glandular-tipped anthered species as a section. How-

ever, here may be recorded Schnack and Covas observations: Ver-

bena restricted to species with funnelform corollas, various tones of

purple, rarely white; style at most three times longer than ovary;

anther cells equal to or smaller than connective, the appendages

glandular; inflorescence nearly always a terminal panicle of spikes;

stem parenchyma in 4 interrupted bands; chromosomes small with

basic number 7 or multiple. Glandularia was defined : corollas salver-

form, colors various; style much longer than ovary; anther cells ex-

ceeding connective, the 2 upper anthers with prominent, vestigial

gland or none; inflorescence simple or branched generally once; corti-

cal parenchyma continuous, not interrupted at the angles (below

inflorescence); chromosomes large with basic number n=5, yn=I5.

These characters were illustrated, I.e., for V. officinalis L. and for

G. laciniata (L.) Schnack & Covas. Perry, I.e. pi. 13, showed trans-

verse sections of the fruit of V. Wrightii Gray and V. bracteata Lag.

FLORA OF PERU 613

& Rodr. to illustrate the subtrigonous shape of the latter in contrast

to the subcylindric form of V. Wrightii, this representing section

Glandularia. It may be noted that in other families as in the related

Boraginaceae chromosome count alone may not be taxonomically

conclusive. Incidentally, Schnack and Covas have merely implied

transfers to Glandularia, as source of specific publication has not

been indicated.

Lily M. Perry's revision of the North American species, while

applicable directly to only a few Peruvian plants, has been most

helpful because of her concise and clearly contrasting presentation

of salient as well as less important characteristics, these not lost or

concealed in irrelevant verbosity or repetitive phraseology.

Finally may be mentioned Moldenke's Hybridity in the Verbena-

ceae, Am. Midi. Nat. 59: 333-370. 1958 with bibliography; possibly

in Peru occurs among other hybrids X V. Bealei Mold. (I.e. 342),

and Phytologia 2: 129-151. 1946 (V. hispida X V. litoralis).

Leaves or segments subulate or minute, usually pungent, rigid; de-

pressed shrubs except V. juniperina.

Plants forming compact polsters; leaves minute V. minima.

Plants not polster-forming or at least not as above.

Subpulvinate shrub; minute leaves obtuse V. aretioides.

Laxly or intricately branched shrub; leaves or segments pungent.

V. juniperina.

Leaves not subulate nor rigid nor pungent; not depressed shrubs or

not compactly, unless V. villifolia, V. Weberbaueri.

Leaves 3-divided nearly to base, rarely in part entire or bifurcate.

Leaves closely canescent strigillose-hispidulous.

Segments entire or bifurcate V. calcicola.

Segments dissected V. lucanensis.

Leaves more or less canescent hirsute-villous-strigose.

V. fasciculata, V. clavata.

Leaves lightly hispidulous, greenish.

Branches villous; shrubby in age V. fissa.

Branches puberulent, rather herbaceous.

Leaf segments hirsute, incised V. Ferreyrae.

Leaf segments entire or 1-3-lobed V. variabilis.

Leaves more or less trifid but obviously cuneate-based or petio-

lately narrowed, the segments variously serrate to pinnate or

614 FIELD MUSEUM OP NATURAL HISTORY — BOTANY, VOL. XIII

dissected, rather entire (cf. V. lucanensis) . See third con-

trast below.

Compact, branches short, ligneous; leaves shortly lobed, villous.

V. villifolia.

Diffuse, decumbent ascending, often ligneous toward base.

Primary leaf segments often closely pinnately lobed or dis-

sected and narrowly in varying degree.

Leaves mostly or all shorter than 1 cm V. microphylla.

Leaves mostly or all larger.

V. tenuisecta, V. dissecta, V. laciniata.

Primary leaf segments typically sparsely pinnately incised to

coarsely serrate.

Leaf segments broad, very cuneate-based.

V. trifida, V. cuneifolia.

Leaf segments narrow, alate petiolate or subsessile.

V. Matthewsii, V. pogonostoma.

Primary leaf segments entire, bifid or trifid V. occulta.

Leaves variously cleft or serrate to entire, sometimes 3-lobed.

Leaves at most to about 3 times longer than wide, more or less

cuneate-based, unevenly incised or lobulate, rarely entire.

Leaves entire.

Bracts subulate; corolla yellow or tube red; calyx teeth

unequal V. aspera.

Bracts rhomboid; corolla pale azure; calyx teeth equal.

V. Echegarayi.

Leaves never entire.

Flowers conspicuous, yellow, scarlet or in part red-purple,

except V. Tessmannii.

Corolla not lilac.

Leaves coarsely serrate-crenate V. peruviana.

Leaves somewhat pinnate or trifid or only incised.

V. Berterii.

Corolla lilac V. Tessmannii.

Flowers often lilac, purple or white, scarcely showy (3-

7 mm. long) unless V. occulta.

Leaves not at all connate (ex char.).

Spreading or depressed, suffruticose, matted in age.

Spikes often subfastigiate; leaves incised. V. Hayekii.

FLORA OF PERU 615

Spikes cylindric; leaves in part 3-lobulate.

V. Weberbaueri.

Erect or arcuate-ascending perennial or weedy; stems

4-angled V. parvula, V. officinalis.

Leaves slightly connate basally.

Leaves entire below, medially 3-lobed V. occulta.

Leaves, at least typically, unevenly serrate or cleft.

V. hispida.

Leaves usually much longer than wide, oblong to lanceolate but

often attenuate to base.

Leaves at least slightly connate.

Stems subterete; spikes often 2-several cm. long, hispid,

glandular V. hispida.

Stems acutely angled; spikes to about 1 cm. long, hispid-

ulous V. bonariensis.

Leaves attenuate to subsessile or petiolar base; stems angled.

Spikes little elongating but sometimes open below in fruit,

cymose or subpanicled V. brasiliensis.

Spikes soon elongating and open at least below, often simple.

Leaves coarsely crenate-serrate.

Plants more or less pubescent . . V. litoralis, V. Carolina.

Plants glabrate or essentially glabrous . . . . V. glabrata.

Leaves pinnately incised, cleft or lobulate . . V. officinalis.

Verbena aretioides Hayek, Bot. Jahrb. 42: 163. 1908, not R. E.

Fries, Nov. Act. Soc. Sci. Upsala ser. 4, 1, no. 1: 109. 1905. Junellia

Hayekii Mold. Lilloa 5: 396. 1940.

A small subpulvinate shrub, the short branches prostrate, the

ligneous trunk about 5 mm. thick, the young branchlets densely

short-pubescent as the minute opposite basally connate imbricate

leaves, these cuneate-oblong, in part entire, in part 3-lobed, obtuse,

carinate; heads terminal, 1-3-flowered, the oblong pubescent bracts

little shorter than the calyx, this strigose, angled, 3 mm. long; corolla

glabrous, the tube scarcely exserted; anthers not appendaged. —

Extraordinary in the partly entire, partly 3-lobed leaves; flowers

larger than those of the related V. Philippiana Ktze. and similar

species (author) ; on the other hand it is the same as V. calcicola Walp.

according to Johnston. F.M. Neg. 17402.

Puno: Tacora to Pomarape, Stuebel lOOb, type.

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

Verbena aspera Gill. & Hook, in Hook. Bot. Misc. 1: 163. 1829;

544. Junellia aspera (Gill. & Hook.) Mold. Lilloa 5: 393. 1940.

Shrubby, the branches rigid or virgate, the younger spreading,

subherbaceous, pilose-hirtellous; leaves alternate or often proliferous-

fasciculate, oblong-elliptic or subspathulate, acute, somewhat sub-

strigose-hirtellous both sides; spikes 2, densely, soon laxly imbricate,

a dm. long or longer, sessile, pubescent, the subulate bracts nearly as

long as the tubular calyx, 5 mm. long with 4 very unequal subulate-

acuminate teeth; corolla glabrous, the slender curved tube well-

exserted, the limb 5-parted; nutlets 4, at first united. — The elongate

corolla is subcernuous, yellow (Pennell) or lobes yellow, tube reddish

(Eyerdam).

Ancash: (Moldenke). — Ayacucho: Huacata, 3,000 meters, Weber-

bauer 5779 (det. Johnston). — Arequipa: Near Arequipa, Pennell

13187; (Meyen); Eyerdam & Beetle 22121. Chachani, (Hinckley 54).

Chile; Argentina.

Verbena Berterii (Meisn.) Schauer in DC. Prodr. 11: 551. 1847.

Shuttleworthia Berterii Meisn. Gen. PI. Comment. 198. 1839.

Suffruticose, the procumbent tetragonal branches ascending, ca-

nescent-hirsutulous as the calyces, these to 8 mm. long; leaves strigose-

hispidulous, pinnatifid or trifid, the intermediate division tridentate,

entire at attenuate base, the linear-lanceolate segments subacute,

revolute; spikes capitate, many-flowered; bracts linear-lanceolate,

about 3 mm. long; corolla 2.5-4 cm. long, the tube more or less

pubescent above, limb large, throat villous; connective appendages

dorsal, scarcely exserted or exceeding anther tips, continuous to base;

calyx contorted over the fruit of 4 nutlets, these striatulate, reticu-

late-rugose apically. — Differs from the Brazilian V. tenera Spreng.

and the Chilean Glandularia sulfurea (D. Don) Schnack & Covas in

the pubescent corolla, also from the former in leaves, from the latter

in sessile appendages. Annual or perennial herb, corolla red-purple,

limb bluish-lavender (Metcalf).

Junin: (Moldenke). — Huancavelica: Near Cordova, Metcalf 30267.

— Ayacucho: Puquio to Lucanas, 3,200 meters, Metcalf 30322. Chile.

Verbena bonariensis L. Sp. PL 1: 20. 1753; 541; (Perry, 254).

Tall, acutely tetragonal stems asperous-hispidulous; leaves more

or less subconnate and subcordate, the lower cuneate-oblong, lanceo-

late, coarsely serrate, scabrous-hirsute both sides, nerves prominent

FLORA OF PERU 617

beneath, spikes short, subcylindrical by the crowded flowers, long-

peduncled in dense terminal cymes; bractlets subulate-lanceolate,

acuminate, hirtellous and ciliate, slightly if at all longer than the

hispidulous-angled calyx, this 3 mm. long, cleft after closing over

the short fruit; corolla slightly exserted, blue, violet or purple, finely

villous; nutlets about 2 mm. long, subreticulate apically, muricate-

scabrous. — The related V. rigida Spreng. or var. rigida (Spreng.)

Ktze., to be expected, has glandular inflorescences, bractlets and

corolla tube longer than calyx; probably more correctly classified as

a variant as at least the glandular character varies in other specific

entities. Said to be an introduction in Peru.

Lima: Botanical Garden, Soukup 2911. — Arequipa: Near Are-

quipa, Raimondi. North of Paraguay, Argentina, Brazil.

Verbena brasiliensis Veil. Fl. Flum. 17. 1825; Icones 1: pi. 40.

1827; 542. V. litoralis HBK. var. pycnostachya Schauer in DC. Prodr.

11: 542. 1847; var. brasiliensis (Veil.) Briq. Ann. Cons. Jard. Bot.

Geneve 7-8: 292. 1904. V. approximate, Briq. I.e., fide Moldenke;

cf. V. litoralis HBK. below, recognized by Perry.

Vigorous erect perennial to 2 meters tall, conspicuously 4-angled;

leaves distant, cuneate at base, lower obovate, upper lanceolate, acu-

minate, acutely incised-serrate; cyme open, compound, the long

branches linear-lanceolate, foliate; spikes fascicled at the tips, in

fruit 2-5 cm. long; calyx pubescent; corolla small, purple; nutlets

trigonous, about 2 mm. long, striate, apically reticulate.

Cajamarca, Libertad, Huanuco, Junin, Cuzco (Moldenke). To

Paraguay.

Verbena calcicola Walp. Nov. Act. Acad. Nat. Cur. 19, suppl. 1:

378. 1843; 550. V. gynobasis Wedd. Chlor. And. 2: 156. 1857, fide

Johnston.

Herbaceous or scarcely suffruticose, canescent hirtellous (or strigil-

lose) all over, the stems decumbent, the branches ascending; leaves

sessile, usually parted to base into 3 obtuse linear revolute segments

or entire, rarely bifurcate, 1-1.5 cm. long; spikes terminal, shortly

peduncled, after anthesis ovate, in fruit to 1.5 cm. long; bracts linear,

about as long as the 5-cleft calyx, its lobes linear, 4-5 mm. long;

corolla reddish, half again longer than calyx, limb to 1 cm. across,

the rounded lobes subemargmate; anther connective unappendaged,

ovary 4-lobulate even to middle; one stigma horned and smooth,

other branches obtuse and papillose. — Included by Schauer in V. cla-

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

vata R. & P. and conceivably it may be an ecological condition.

F.M. Neg. 17409.

Cuzco: (Gay). — Arequipa: Near Arequipa, Meyen, type. Mol-

lendo, Weberbauer 1455. — Moquegua: Torata, Weberbauer 7464 (det.

Johnston). Tacora, on road from Tacora to La Paz, (Weddell, type,

V. gynobasis).

Verbena Carolina L. Syst. ed. 10, 852. 1758-1759; (Perry 268).

V. polystachya HBK. Nov. Gen. & Sp. 2: 274. 1818, fide Perry,

y. caroliniana L. (sic) forma vel var. polystachya (HBK.) Loes. Repert.

Sp. Nov. 9: 362. 1911.

Stems usually solitary and erect, branched, hirsute; leaves sub-

sessile or attenuate into a short petiole, lanceolate-oblong or some-

what elliptic-oblong, to 8 cm. long or sometimes longer, subobtuse

or acute, coarsely crenate-serrate, strigose-pustulate above, hirsute-

strigose, especially in the venation, beneath; spikes in panicles, slen-

der, open in fruit; bracts ovate, acuminate-subulate, ciliate, to about

half as long as the hispidulous calyx, this in fruit 2 mm. long, the

obtuse mucronate lobes connivent; corolla tube scarcely exserted,

the limb about 2 mm. across; nutlets trigonous, dorsally convex,

striatulate, the commissural faces nearly smooth, meeting at right

angles.

Lima: Atocongo, Pennell 14770 (det. Johnston). — Junin: Huan-

cayo, Soukup 3182. To Arizona.

Verbena clavata R. & P. Fl. Peruv. 1: 21, pi. 33, fig. b. 1798; 550.

Canescently strigose-hirsute all over, suffruticose, the many

branches ascending; leaves sessile, 3-parted nearly to base, 6-8 mm.

long, the divaricate oblong-clavate segments quasi-verticillate, en-

tire, revolute or 2-3-cleft; spikes sessile in the uppermost leaves,

capitate, many-flowered; bracts lanceolate, subequaling the calyx,

this 6 mm. long, unequally 5 plicate-subulate, dentate, both cinere-

ous hirsute; corolla purplish, tube strigose above, about 9 mm. long,

the throat pubescent, the linear-oblong lobes emarginate; anther ap-

pendages exserted, clavate, stipitate, arcuate-divergent. — Var. cas-

mensis Mold., Phyt. 4: 58. 1952, has scarcely revolute leaves mostly

drying blackish, about 3 mm. wide, with short unevenly spreading

twisted trichomes both sides; flowers lilac, fragrant. Leaves resem-

ble those of V. trifida; spreading pubescence differs from both plants.

Forma albiflora Mold, was first noted formally from Ferreyra 2507.

FLORA OF PERU 619

Ancash: Near Huaraz, Sandeman 4611. Lomas de Casma, (Fer-

reyra 8081, type var.). Lomas de Mongon, Goodspeed 9184- —

Arequipa: Near Islay, Isern 2477. Mollendo, West 8202; Eyerdam

25159; Mexia 04170; Goodspeed Exped. 15719. — Moquegua: Tacora,

Isern 2478. — Tacna: Candarave, Metcalf 30374. Ruiz & Pavdn,

(Tafalla), type. Chile. "Tacpa" (Mexia).

Verbena cuneifolia R. & P. Fl. Peruv. 1: 22, pi 32. 1798; 539.

Stems erect, tetragonal, hispid, subsimple or corymbose above;

leaves broadly ovate, at base cuneate, sessile, strigose-hispid on both

sides, 5 cm. long, nearly as wide, deeply 3-cleft and laciniate, the

lanceolate segments acutely incised pinnatifid or coarsely serrate;

spikes subternate, terminal, oblong, finally cylindric, about 2.5 cm.

long, canescent and glandular; bracts lanceolate, about 5 mm. long;

calyx subulately 5-lobed, becoming constricted over the fruit, nearly

8 mm. long; corolla blue, the tube somewhat funnelform, about

10 mm. long, pubescent above; nutlets reticulate- tuberculate above.

-F.M. Neg. 17411.

La Libertad: (Moldenke). — Ancash: Cayabamba, Weberbauer 3060.

Pomabamba, Huaraz, Raimondi. — San Martin: Moyobamba, Ruiz

& Pavdn, type. — Huanuco: Yanahuanca, 1210. — Junin: Acopalca,

Stork 1 0920; Soukup 3537. Huancayo, near Huancayo, Soukup 3970;

Raimondi. — Huancavelica: Tayacaya, Raimondi. — Ayacucho: (Mol-

denke).— Cuzco: Valledel Paucartambo, Herreral409. Chile. "China-

verbena."

Verbena dissecta Willd. ex Schauer in DC. Prodr. 11: 552. 1847.

Glandularia dissecta (Willd.) Schnack & Covas, Darwiniana 6: 475.

1944.

Early decumbent canescent hirsutulous annual, the subterete

branchlets erect; leaves cuneately decurrent into petioles, 3-parted,

pinnatifid or laciniate, the linear subobtuse segments entire or den-

tate, subrevolute, strigose above, hirsute beneath; spikes terminal

and axillary, fastigiate, the flowers spreading, cymose-paniculate;

bracts oblong, acute, much shorter than calyx, this 8 mm. long, the

unequal teeth subulate-aristate, hirsute and shortly stipitate gland-

ular, glabrate in the intervals; corolla violet, subexserted as the an-

ther appendages. — Compare V. laciniata (L.) Briq., of which, perhaps,

it is correctly a part.

Puno: Pisacoma, (Meyen). To Uruguay. Chile; Bolivia; Brazil.

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

Verbena Echegarayi Hieron. Bol. Acad. Nut. C6rdoba 4: 66.

1881. Junellia Echegarayi (Hieron.) Mold. Lilloa 5: 395. 1940.

Similar to V. aspera; spreading older branches prominently no-

dose, the younger hirtellous and with alternate leaves and short

axillary branchlets of fasciculate ones, the lower leaves oblong-lan-

ceolate or subspathulate, subobtuse, the upper cordate-lanceolate,

mucronulate, mostly 5-8 mm. long, 1.5-3 mm. wide, sparsely hirtel-

lous; spikes 2-7 cm. long; bracts rhomboid, subacuminate-mucronate,

4.5 mm. long, 1.5 mm. wide, ciliate, glabrate or sparsely hirtellous;

calyx slightly hirtellous, 5 mm. long, the 4 teeth equal; corolla pale

azure, the tube 8 mm. long, the segments subdenticulate or emargi-

nate. — Otherwise like the related species, ex char, for which I am

indebted (as often previously) to the librarian of Arnold-Gray Her-

barium, Mrs. L. Schwarten.

Arequipa: Airampul, Arequipa to Puno, Weberbauer 1396 (det.

Moldenke). Argentina.

Verbena fasciculata Benth. Bot. Voy. Sulphur 153. 1844.

Apparently suffrutescent, ascending-erect, very hirsute, to several

dm. tall; leaves sessile, 3-5-parted to base, the younger fascicled in

axils, the lanceolate or linear segments entire or 3-5-parted, the

larger divisions to 5 cm. long, 4 mm. wide, the others smaller and

much narrower, all acute; spikes sessile or shortly peduncled, dense,

the lanceolate-subulate bracts equaling the costate hirsute calyx, this

with setaceous teeth and little exceeded by the corolla tube; anther

cells distinct; upper style lobe very short, thickly pulvinate-stigma-

tose, the slightly longer lower lobe naked, incurved-acute; nutlets

rugose. — Type collected by Hinds at "Huamantango," probably

Huamantamba near Canta, Lima, which is in the scope of Cuming's

brief visit.

Peru: On the coast, (Cuming 954, fide Bentham).

Verbena Ferreyrae Mold. Phytologia 3: 279. 1950.

Herb, the ascending-erect obtusely angled stems and branches

more or less pubescent as the leaves (especially beneath) ; nodes an-

nulate; leaves sessile, almost basally 3-parted, 3-6 cm. long and wide,

the segments pinnately parted and incised, usually gradually wid-

ened toward base where often 8 mm. wide, revolute, the lobes acute,

somewhat pilosulous above, glabrescent in age; inflorescence termi-

nal, capitate in anthesis, 4 cm. long or longer later, the principal

spikes usually ternate; peduncles 1.5-4 cm. long, retrorsely short-

FLORA OF PERU 621

pubescent; bractlets oblong-linear, 6-8 mm. long; calyx tubular,

short-pubescent, 5-costate-apiculate, 6 mm. long; corolla sky-blue,

tube 8 mm. long, limb about 9 mm. across, densely pilose in throat.

— Author suggested no relationship in spite of the implied honor to

a colleague.

Ayacucho: Puquio, Prov. Lucanas, 3,200 meters, (Ferreyra 5491,

type).

Verbena fissa Hayek, Bot. Jahrb. 42: 165. 1908.

Small, shrubby, the branches densely and canescently villous;

leaves opposite, 3-parted to base, the linear divisions about 2 mm.

wide, obtuse, revolute, strigose-setose above, subsetose-tomentose

beneath; spikes subcapitate, finally little elongate, the linear-

lanceolate setose bracts subequaling calyx, this 5-angled, strigose,

much longer than corolla-tube; corolla 1 mm. long, limb 4-6 mm.

broad, lilac, fragrant. — Near V. thymoides Cham, but more depressed,

more densely pubescent, larger flowers; as V. calcicola Walp., cited

as a synonym in Index Kewensis, may also prove to be a variant of

V. clavata R. & P. as suggested by Johnston. About 3 dm. tall,

flowers white or pink, on middle and lower green slopes (Johnston).

F.M. Neg. 17416.

Ancash: Below Ocros, Weberbauer 2768, type; 169. Hualas,

Raimondi. — Arequipa: Posco, Cachendo, (Guenther & Buchtien 183;

133a}. Mollendo, Weberbauer 389; 143, 147; Johnston 3560. Near

Chala, Ferreyra 5554 (det. Moldenke, V. clavata}. — Tacna: Canda-

rave, Weberbauer 7385 (det. Johnston, V. clavata}; Metcalf 30374-

Verbena glabrata HBK. Nov. Gen. & Sp. 2: 276. 1818; 542.

Glabrous or essentially except the hispidulous lanceolate-

acuminate bracts and calyx; branches tetragonous; leaves oblong-

lanceolate, cuneate to short petioles, incised serrate, prominently

reticulate- veined beneath, about 5 cm. long or longer, 2 cm. wide;

spikes ternate, filiform cylindric, the lateral long-peduncled, 2.5 cm.

long, the intermediate longer; corolla roseate, tube little exserted. —

To 1 meter tall, corolla tube red, blue-violet above, fruit red-maroon

(Metcalf). Schauer says this is more compact, more glabrate than

V. litoralis HBK. var. pycnostachya Schauer; compare V. bonariensis

L., V. brasiliensis Veil.

Lima: (Moldenke). — Ayacucho: Puquio to Quebrada de San An-

tonio, 2,800 meters, Metcalf 30294. — Cuzco: (Moldenke). Venezuela;

Ecuador.

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

Verbena Hayekii Mold. Phytologia 2: 148. 1946. Junellia

Hayekii Mold. Lilloa 5: 396. 1940. V. aretioides Hayek, Bot. Jahrb.

42: 163. 1908, not R. E. Fr. 1905. V. procumbens Hayek, Bot.

Jahrb. 42: 163. 1908, not Forsk.

Suffruticose, the elongate prostrate rooting branches glabrous,

the erect short branchlets sparsely setose; leaves sessile, oblong-

obovate, cuneate to base, coarsely incised dentate, strigose-hirsute

both sides; spikes subfastigiate, often branched at base, finally cylin-

dric, the ovate-lanceolate acute setose bracts subequaling the calyx,

this angled, 5-dentate, 3 mm. long; corolla lilac, 6 or 7 mm. long;

upper anthers unappendaged. — Near V. Weberbaueri but differs in

the long rooting branches, longer flowers, different inflorescence

(Hayek). Corolla white with lilac or pink lobes, pubescent in center

(Stork & Horton). F.M. Neg. 17442.

Junin: La Oroya, 3,700 meters, Weberbauer 2573, type, 180;

Kalenborn 36; 176. — Huancavelica: Near Huancavelica, Stork &

Horton 10820.— Cuzco: (Moldenke).— Puno: (Moldenke). Bolivia?

Verbena hispida R. & P. Fl. Peruv. 1: 22, pi. 34, fig. a. 1798; 542.

Spreading-erect or ascending, the rigid branches pilose-hispid;

leaves semiamplexicaul, oblong or lanceolate, cuneate to base, acum-

inate, coarsely and unevenly serrate or incised, subentire or bilobu-

late, lineate-rugose, strigose hispid on nerves both sides and also

hirtellous beneath; spikes hispid and glandular, terminal, peduncled,

often ternate, dense, cylindric, finally about 7 cm. long; bracts lance-

olate, at least as long as the subulate-dentate calyx, this scarcely

more than 2 mm. long; corolla tube about 4 mm. long. — Determina-

tions mostly by Johnston; Raimondi, by Herb. Dahlem. F.M.

Negs. 17555; 24686; 29686.

Cajamarca: Cascas, Raimondi. — San Martin: (Moldenke). — La

Libertad: (Moldenke). — Lima: Canta, Pennell 14582. Near Lima,

Raimondi. — Huanuco: Near Huanuco, Ruiz & Pavon, type. Mito,

1704.— Junin: Tarma, 1006; Killip & Smith 21855; 21925.— Huanca-

velica, Stork & Horton 10820 (det. Standley). — Loreto: (Moldenke).

— Arequipa: Isern 165. — Cuzco: Sicuani, (Hickeri). Prov. de Cer-

cado, Herrera. Cerro de Colquipata, Pennell 141 84- Yucay, Soukup

259. — Puno: Chuquibambilla, Pennell 13409. — Moquegua: Tacora,

Isern 166. Ecuador; Chile and Argentina.

Verbena juniperina Lag. Gen. & Sp. Nov. 19. 1816; 544. Junel-

lia juniperina (Lag.) Mold. Lilloa 5: 396. 1940; var. grisea (Johnst.)

FLORA OF PERU 623

Mold. I.e. 397. V. juniperina Gill & Hook, in Hook. Bot. Misc. 1:

163. 1830.

Much branched pubescent shrub typically a meter or so high, the

opposite persisting leaves deeply 3-parted, the segments acerose or

spinose in age and then very rigid; heads terminal, the subulate

bracts longer than the calyx, this about 6 mm. long, somewhat in-

flated, about half as long as the corolla tube. — New leaves spring

from the axils of the old, the latter thus simulating stipular spines

(Gillies and Hooker). The variant grisea Johnst., Contr. Gray

Herb. 81 : 96. 1928, was only 3-5 dm. high, canescent, the branchlets

mostly represented by leaf -fascicles, rarely longer than 1 cm. A low

very dense shrub of open rocky slopes, the flowers deep lavender

(Pennell); purple with yellow spots (Metcalf).

Huancavelica: Cordova, Metcalf 30268. — Ayacucho: Parinaco-

chos, Raimondi. Lucanas, Raimondi. Rio de Lomas, 2,900 meters,

Weberbauer 5771. — Arequipa: Misti, Weberbauer 1392; 130. Nevado

Chachani, Pennell 13259; (Hinckley 76, type, var.)- — Moquegua:

Torata, 3,200 meters, Weberbauer 7461 (var.). — Tacna: Near Can-

darave, Metcalf 30388 (var.). Argentina. "Pomera."

Verbena laciniata (L.) Briq. Ann. Cons. Jard. Bot. Geneve

7-8: 296. 1904; 552. Erinus laciniatus L. Sp. PI. 630. 1753. V. eri-

noides Lam. Tabl. Encycl. 1: 57. 1791. V. multifida R. & P. Fl.

Peruv. 1: 21, pi. 33. 1798. Glandularia laciniata (L.) Schnack &

Covas, Darwiniana 6: 475. 1944.

Much branched, decumbent, the ascending branches rooting

toward base, more or less strigose-hirsute especially on the nerves

of the leaves beneath, the terminal or lateral pedunculate spikes

canescent; leaves decurrent, cuneate at base, 3-parted, pinnately

lobed or laciniate, spreading stellate-hirsutulous, the laciniae lance-

olate-linear or widely subacute, entire or subdentate and subrevolute;

spikes soon elongating and lax; bracts lanceolate, acuminate, spread-

ing, often as long as calyx, this angulate, hirtellous and with some

stiped glands, the subulate-aristate teeth finally connivent, to 8 mm.

long, the corolla shortly exserted; anther appendages minute, sub-

barbate. — Type from Peru by Feuille'e. Stems wiry, prostrate, form-

ing loose mats to a meter or more wide, the odorless, clear, mauve

to violet flowers in flattened heads to 5 cm. across (Balls) ; as inter-

preted, leaves trifid or much divided, the Linnaean plant was said

to have scarlet flowers (Hooker); red-violet (Schauer). Illustrated,

Darwiniana, I.e. 472, fig. 1 (flower parts).

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

La Libertad: Rio Negro, West 8115. — Lima: Rio Blanco, 678 (det.

Johnston); Raimondi 10703; Killip & Smith 21754 (det. Standley);

Ferreyra 6502 (det. Moldenke). — Huanuco: Chinche near Yana-

huanca, 1260. — Junin: (Moldenke). — Cuzco: Ollantaitambo, Hen era

8450; 3442; Pennell 13677. Sicuani, (Hickeri) . Near Cuzco, (Herrera

18); Vargas 241 (det. Standley). To Chile; Argentina; Brazil.

Verbena litoralis HBK. Nov. Gen. & Sp. 2: 276, pi. 137. 1818;

542. y. bonariensis L. var. litoralis (HBK.) Hook. Bot. Misc. 1:

166. 1830.

Rather similar to V. bonariensis but lightly strigillose; leaves

scabrous above and lanceolate-oblong, attenuate to subsessile base;

spikes more or less cymose or subpanicled; bracts ovate-lanceolate,

at most about as long as the calyx, this 2-2.5 mm. long, subtruncate

with minute teeth. — Hooker and Schauer added a "t" to the name;

the former remarked, I.e.: seems to be a variety with shorter spikes

than usual. Perry, Ann. Mo. Bot. Gard. 20: 257. 1933, retained it

as distinct. Actually, most material referred here seems doubtfully

distinct from y. Carolina L. A common weed, only a few of many

collections cited; authors gave Truxillo, Santa and Lima specimens

as type collections.

Cajamarca: Truxillo, Bonpland, type; Osgood & Anderson 36. —

Amazonas: Chachapoyas, Williams 7534. — San Martin: Tarapoto,

Williams 5911. San Roque, Williams 7087. — Ancash, fide Moldenke.

— Lima: Near Lima, 54. Rio Blanco, Killip & Smith 21541.—

Huanuco: Mito, 1705. Pillao, Woytkowski 34175. — Junin: Chan-

chamayo Valley, Schunke 1724- — Loreto: Iquitos, Williams 7894- Rio

Nanay, Williams 1297. Yurimaguas, Williams 4367. — Arequipa:

Posco, Cachendo, (Guenther & Buchtien 132, det. Bruns). — Cuzco:

Prov. Cercado, 3,350 meters, Herrera 1505. Valle del Urubamba,

Herrera 1664. Valle del Paucartambo, Herrera 3362. — Tacna: (Mol-

denke). Uruguay and Chile to Central America. "Verbena del

campo."

Verbena lucanensis Mold. Phytologia 3: 279. 1950.

Ex char, rather similar to V. Ferreyrae; branches as peduncles

(4-15 mm. long) densely and divaricately short-pubescent; leaves

1-4 cm. long and wide, deeply 3-parted, leaf-segments densely puber-

ulent both sides, deeply and irregularly dissected, to 2.5 mm. wide;

inflorescence terminal and in the upper axils, to 3.5 cm. long; bract-

lets lanceolate, about 3 mm. long; calyx narrowly 5-dentate, 3-4 mm.

FLORA OF PERU 625

long; corolla purple, the tube 6-7 mm. long, puberulous above, the

limb 6 mm. broad; calyx not inflated in fruit but 5-parted, the 4

cocci 2 mm. long, glabrous, lustrous, scrobiculate-ridged dorsally,

the commisural surface white-papillose below the upper third.

Ayacucho: Nazca to Puquio, 1,500-2,000 meters, Prov. Lucanas,

(Ferreyra 5493, type).

Verbena Matthewsii Briq. Ann. Cons. Jard. Bot. Geneve 10:

104. 1907.

Tall, the stout erect stem obtusely 4-angled (branches in type

few), spreading pilose (some shorter-stiped glandular trichomes inter-

mixed) especially the leaves, these alate-petiolate or subsessile, deeply

3-parted, about 4 cm. long, the incised-crenate segments scarcely

0.5 mm. wide, the lobules obtuse; spikes terminal, at anthesis 2.5

cm. long, 1.5 cm. broad, elongating, the peduncles pubescent; bracts

narrowly lanceolate, acuminate, 5-7 mm. long; calyx oblong, tube

5-angled, 5 mm. long, indument in part glandular, teeth lanceolate,

acuminate, 2 mm. long; corolla tube exserted, 2-3 mm. long, the

limb 5 mm. across, puberulent without, the obovate lobes sometimes

subemarginate; gland of anthers small. — Neighbor of V. Berterii

(Meisn.) Schauer and V. clavata R. & P. with procumbent branches,

the former too with shorter bracts, the latter with 3-divided leaves,

the segments pseudoverticillate (author). — Type was distributed as

V. laciniata (L.) Briq. and doubtfully a distinct species but as here

listed leaf-segments broader, in type seen acute but Lima and

Huanuco specimens with more obtuse segments could be placed here.

Species name here spelled with two t's, as it was sometimes spelled

by the collector. F.M. Neg. 24691.

Cajamarca: Chota, Stork & Horton 10032 (det. Standley, V.

laciniata). — Ancash: Chacchan, 2539 (det. Johnston, V. laciniata). —

Lima: Above Lima, Mathews 498, type. Lomas de Grenades, Stork

& Vargas 9341- — Junin: Chamiseria to Acopalca, Soukup 3537(1}.

— Loreto: (Moldenke). — Ayacucho: Prov. Lucanas, 3,000 meters,

Ferreyra 5470 (det. Moldenke).

Verbena microphylla HBK. Nov. Gen. & Sp. 2: 272, pi. 133.

1818; 551.

Low or creeping, much branched, strigose-hispidulous or hirtel-

lous; leaves subsessile, cuneate at base, 6-10 mm. long, 3-parted, the

segments obovate-oblong, obtuse, revolute, entire or the medial

trifid, the lateral bifid; spikes capitate, subsessile terminal and lateral,

many-flowered; bracts lanceolate, shorter than the calyx, this 6 mm.

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

long, the short ovate teeth acute; corolla glabrous except pilose

throat, the tube about 1 cm. long, the lobes emarginate; anther

appendages black, corniculiform, equaling the throat; fruit half as

long as calyx, the nutlets finely tuberculate dorsally. — Scarcely more

than a part of V. laciniata (L.) Briq. but perhaps recognizable and

apparently a high Andean state; listed by Ball as "V. diffusa Willd."

Ancash: Above Samanco, at Cajamba, 3,700 meters, Weber-

bauer 3042; 171. — Lima: Prov. Chancay, Ferreyra 3531 (det. Mold-

enke). Above Lima, at Chicla, 3,420 meters, Ball; Weberbauer 226;

169. — Ayacucho: Puquio, Prov. Lucanas, Ferreyra 5509 (det. Mold-

enke). — Cuzco: Sicuani, Vargas 9827 (det. Standley). — Puno: Sal-

cedo, Soukup 70. Orurillo, Soukup 870. Chuquibambilla, Pennell

13359 (det. Johnston). Sta. Lucia, J. Sharpe 58. "Salvia de la

Virgen" (Soukup). To Uruguay and Ecuador.

Verbena minima Meyen, Reise um die Erde 1: 451. 1834.

Junellia minima (Meyen) Mold. Lilloa 5: 398. 1940.

Pulvinate-cespitose, much branched, the rooting branches more

or less elongating, the leafy branchlets very short; leaves entire, im-

bricate, linear-subulate, pungent, spreading-recurved, glabrous ex-

cept minutely ciliolate toward the shortly connate base, 3-5 mm.

long; flowers 2-4 at tip of axillary branchlets, sessile; bracts ovate,

acute, ciliolate, equaling the calyx; calyx 4-lobed (5-lobed, Meyen),

the equal lobes very obtuse, laciniate-ciliolate, otherwise, as tube,

glabrous or puberulent; corolla white or purple- tinted, scarcely as

long as calyx tube, the 5 subequal lobes spreading, the throat shortly

barbate or hispidulous; stigma lobes minute, obscure, equal (Wed-

dell). — The specific name, sens, lat., may be properly V. seriphioides

Gill & Hook, in Hook. Bot. Misc. 1: 164. 1830, as apparently the

plants are similar, possibly identical.

H. Francia Chisaki, a member of the University of California

herbarium staff, kindly noted for me that Meyen in his now rare

work gave a quite adequate description as a footnote; the citation,

Nov. Act. Acad. Nat. Cur. 19, suppl. 1: 379. 1843, is therefore a

repetition of publication.

Forms flat cushions to one-half meter broad (Weberbauer) . Illus-

trated, Weddell, Chlor. And. pi. 62B. F.M. Neg. 17431.

Cuzco: Prov. del Cercado, Gay. — Arequipa: Chivay, 4,200 meters,

Weberbauer 6895. — Puno: Grass steppes, Lake Titicaca, Weberbauer

441; 85. Mesa alta de Muno, Raimondi (det. Werderman, V.

seriphioides}. Near Lake Titicaca, Meyen, type; (Shepard 26}.

FLORA OF PERU 627

Salcedo, Soukup 43. — Moquegua: Torata, Weberbauer 7472 (det.

Johnston). Bolivia; Chile. "Koletta."

Verbena occulta Mold. Phytologia 3: 280. 1950.

Erect-ascending obtusely angled branches and branchlets ap-

pressed-pilosulous or early spreading pubescent as the short (to

2 cm.) or obsolete terminal peduncles; leaves somewhat amplexicaul,

about medially 3-parted, the divisions often few-dentate or if short-

lobed the lobes subacute, 1-nerved beneath, dark green and densely

subappressed pubescent both sides with different length trichomes;

spikes 1.5-4 cm. long; bractlets conspicuous, lanceolate, acuminate,

8-10 mm. long, appressed pubescent as calyx, this 5 mm. long,

5-lobed; corolla purple, the tube 8-10 mm. long, limb 4-5 mm. wide.

— A curious species according to the author, who has listed a forma

alba Mold, from Ancash and Junin.

Cajamarca: (Moldenke). — La Libertad: Near Nevado Cajamar-

quillo, (Ferreyra 1298, type). — Ancash: (Moldenke). — Junin: Cerro

de Pasco, 3,800 meters, Ferreyra 6586 (det. Moldenke).

Verbena officinalis L. Sp. PI. 20. 1753.

Ascending to erect, the stems glabrous or nearly glabrous, the

more or less ovate leaves strigillose both sides, all except the upper

petiolar narrowed to base, 1-2-pinnate or 3-5-cleft, the pinnae or

lobes incised; spikes ternate or paniculate, slender, elongate; bracts

usually about half as long as the glandular calyces, these 2-2.5 mm.,

subtruncate, minutely dentate; corolla tube slightly exserted, the

limb about 4 mm. broad, lobes subrotund; nutlets trigonous, scarcely

2 mm. long, strongly striate, somewhat reticulate above, the faces

muricate.

Peru (probably). Widely introduced from Europe.

Verbena parvula Hayek, Bot. Jahrb. 42: 162. 1908.

Perennial from a stout multicipital root, the subsimple arcuate-

ascending to erect stem 6-10 cm. tall (type), sulcate-quadrate, ap-

pressed setose-scabrous as the leaves both sides; leaves oval or obo-

vate, contracted into petiole, subobtuse, 10-14 mm. long (type), un-

evenly incised-crenate-serrate or nearly pinnately lobate, lineate-

nerved beneath; spikes terminal, simple or few-branched, early nar-

rowly conic, finally elongate; bracts ovate, acute, setose-ciliate, sub-

equaling the calyx, this 1.5 mm. long, minutely setulose; corolla lilac,

3 mm. long, subpuberulent. — A small delicate plant with the habit

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

of a dwarf V. officinalis L. but with completely different leaves

(author). Has been found also in Junin and Tacna (Moldenke).

Variety gigas Mold., Phytologia 7: 85. 1959, has erect stems to 9 cm.

tall, elongate internodes, elliptic-oblanceolate leaves to 7.5 cm. long,

2.5 cm. wide; aspect of V. litoralis, inflorescence of V. parvula (au-

thor) ; Junin, Tarma, Killip & Smith 21925, type. F.M. Neg. 17436.

Cuzco: Paucartambo, Balls B6784- Bolivia.

Verbena peruviana (L.) Britton, Ann. N. Y. Acad. Sci. 7: 197.

1893; 537. Erinus peruvianus L. Sp. PI. 630. 1753. V. chamaedry-

folia Juss. Ann. Mus. Paris 7: 73. 1806, fide Moldenke. V. melindres

Gillies, Bot. Reg. pi. 1184- 1828. Glandularia peruviana (L.) Small,

Man. S. E. Flora 1139. 1933.

Stems filiform, procumbent, pilose-hirsute, furcately branched;

leaves oblong-lanceolate or -ovate, broadly cuneate at base, coarsely

serrate or crenate, strigose above, hispid especially on nerves be-

neath; peduncles long, solitary; spikes early congested-capitate,

some elongating, becoming lax; bracts lanceolate-subulate, ciliate,

much shorter than the canescent hirsute calyces. — Type from Para-

guay!; based on a plant of Abbe" Feuille'e, Per. 3: 36, pi. 25. 1725, as

noted by Briquet, Ann. Cons. Jard. Bot. Geneve 7: 290. 1904; as he

pointed out, Britton's transfer (now acceptable) was prompted on

the basis of a misdetermination and the plant of Jussieu, as that of

Linnaeus, is unknown in Peru unless in cultivation. Flowers brilliant

scarlet; there is a forma rosea Mold, in Uruguay and probably else-

where. Illustrated, Bot. Reg. pi. 1184.

Peru: In cultivation. To Uruguay. "Melindres Colorado,"

"pampas."

Verbena pogonostoma Klotzch, ex Walp. Repert. 4: 31. 1844-

1848; 550.

Diffuse, subcanescent, the branches hirtellous, the leaves above

and beneath on the nerves strigose-hispidulous; leaves sessile, cuneate

to base, 3-parted, about 4 cm. long, the narrow laciniae pinnatifid-

incised and dentate above, lower entire, 2 mm. wide, the ultimate

divisions linear-lanceolate, acute, revolute; spikes shortly peduncu-

late, terminal and lateral, fastigiate, much elongating before anthesis,

finally 3 dm. long, dense, strigose-hirtellous and glandular; bracts

lanceolate-subulate, as the unequal teeth of the longer calyx, this

6 mm. long, colored; corolla funnelform, the yellowish tube scarcely

exserted, gradually ampliated from base to barbate throat, the blue

FLORA OF PERU 629

limb with short rounded lobes; fruit included, the nutlets reticulate

dorsally. — Type from garden material of Berlin-Dahlem, origin un-

certain; seems, with V. Matthewsii, to be a part of V. laciniata.

F.M. Neg. 17440.

Ancash: Ocros, Weberbauer 2723 (det. Hayek, V. trifida). — Lima:

Near Chosica, Weberbauer 5338 (det. Dahlem). Paraguay?

Verbena tenuisecta Briq. Ann. Cons. Gard. Bot. Geneve 7-8:

294. 1904. Glandularia tenuisecta (Briq.) Small, Man. S. E. Flora

1139, 1508. 1933.

Decumbent-ascending, 2-3 dm. high, the tetragonal branches

early sparsely pilose; leaves triangular, 3-parted-pinnatifid, 2-3.5 cm.

long, 2-3 cm. wide, the linear subobtuse or acute (the superior) seg-

ments 1-3 mm. wide, lightly substrigose especially on the nerves

beneath, glabrescent; spikes terminal, solitary, fastigiate, early ovate-

capitate, 1.5 cm. long, finally 3-4 cm. long, relaxing; bracts ovate-

lanceolate, 2-3 mm. long, puberulent; calyx tube 7 mm. long, teeth

1-2 mm. long, densely substrigose all over and with a few dark

glandular trichomes, the teeth setose at tip; corolla blue, the glabrous

tube exserted 3-5 mm., limb about 1 cm. across, the broad emarginate

lobes pilose at throat; anther gland scarcely exserted; nutlets about

3 mm. long, strongly reticulate. — Differs from V. tenera Spreng. of

Brazil in its larger leaves, pubescent calyx and evenly puberulent

bracts, from V. dissecta Willd. in its merely fastigiate inflorescence

as well as by the strigose calyx (author); these differences are the

sort, one can imagine, that may exist in varying degrees. Naturalized

plants have been referred to V. tenera Spreng. and to V. laciniata

(L.) Briq. (V. erinoides Lam.), and Moldenke, Fl. Texas, 3, pt. 1: 41.

1942, has suggested that it may prove referable to one of several

older names, as V. aristigira S. Moore and V. mendocina Phil.

Commonly cultivated and, in southeastern United States, naturalized

(Moldenke); whether these plants centering around V. laciniata

are species, genetically, must await results of modern methods of

investigation. In any case Briquet's plant as to type is scarcely in

Peru. F.M. Neg. 17451.

Cuzco: (Moldenke). Paraguay; Brazil.

Verbena Tessmannii Mold. Phytologia 3: 45. 1948.

Herb, the acutely tetragonal stems, elongate branches, petioles

(4-6 mm. long), and peduncles (2-5 cm. long) hirtellous with divar-

icate trichomes; leaves lanceolate, attenuate at base, acute, 2-4 cm.

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

long, 7-15 mm. wide, firm-chartaceous, unevenly and coarsely den-

tate, harshly appressed strigose above with bulbous-based trichomes,

shortly pubescent beneath especially on the 4 pairs of slender lateral

nerves; spikes elongating to 4.5 cm., dense; bractlets lanceolate,

6 mm. long, subulate, puberulent and long-ciliate; calyx tubular,

about 13 mm. long, 5-costate, puberulent and with a few black

capitate glands, the 4 subulate tipped teeth 2 mm. long, often

recurving; corolla lilac, the slender tube nearly 2 cm. long, unevenly

pulverulent, the limb about 8 mm. across. — Type from campos,

Parana, formations of shrubs and tall herbs (Tessmann).

Peru: (Moldenke). Brazil.

Verbena trifida HBK. Nov. Gen. & Sp. 2: 273, pi. 134. 1818; 539.

Canescent, the tetragonal branches and subcapitate apical spikes

hirsutulous; leaves sessile, crowded, mostly fasciculate in axil of

branches, basally cuneate, 12-25 mm. long, generally trifid, the

intermediate segment largest, sometimes one or the other part bifid

or entire, the lanceolate laciniae revolute, midnerve prominent, lan-

ceolate, acuminate; bracts lanceolate, about equaling the tubular

calyx, this 6 mm. long, nervose, the conspicuous subulate teeth sub-

equal; corolla tube little exceeding calyx, the rounded lobes emar-

ginate. — Not seen by Perry, nor placed. Weberbauer 2723 was re-

ferred here by Hayek perhaps correctly, but this, typically at least,

has leaves with only three narrow lobes; the type came from Mexico.

Peru (doubtful; cf. note above). Colombia (Moldenke); to

Mexico.

Verbena variabilis Mold. Bot. Gaz. 106: 164. 1944.

Perennial herb, to 1 meter tall; stems tough, substrigillose above

as the very slender branchlets; leaves sessile, decussate-opposite or

approximate, mostly 3-parted with broad segments, each segment

entire or with 1-3 large lobes, the leaves on branchlets 3-parted but

segments linear and pinnatifid, obtuse, revolute, green but strigillose

both sides, 2-3 cm. long, 1-2 cm. wide; peduncles strigillose, to 9.5

cm. long, with 1 or 2 pairs of 3-parted bracts near tip and medial;

spikes 2-5 cm. long, dense; bracts long-acuminate, 5 mm. long,

glandular with erect spreading trichomes; rachis densely spreading

puberulent; calyx about 5 mm. long, glandular; corolla pale purple,

the tube somewhat exserted. — Roots "used against fleas," according

to the author, who suggested no relationship.

Huancavelica: Cordova, Metcalf 30255, type.

FLORA OF PERU 631

Verbena villifolia Hayek, Bot. Jahrb. 42: 164. 1908.

Small shrub, the branches prostrate, the younger densely villous

as the opposite shortly petioled cuneate-based trifid leaves, these

with cuneate middle lobe 3-lobed, the central lobes bilobed; primary

nerves prominent beneath; spikes terminal, subsessile, early fasti-

giate, finally short-cylindric, the rachis villous; bracts ovate-lanceo-

late, acute, longer than the calyx, this appressed pilose, 4 mm.

long, accrescent in fruit, the roseate corolla scarcely longer; upper

anthers unappendaged; fruits 3 mm. long, scrobiculate-rugose. —

My collection (det. Johnston) was prostrate in stones along trail, the

flowers pink, the villosity yellowish. F.M. Neg. 17458.

Junin: La Oroya, 942; Weberbauer 2582, type; 180.

Verbena Weberbaueri Hayek, Bot. Jahrb. 42: 163. 1908.

A little depressed shrub, the simple (or nearly) branches only

5-10 cm. long; leaves ovate, 1-1.5 cm. long, obtusely incised-crenate-

serrate, densely appressed setose both sides; spikes densely cylindric,

setose-hispid, 2-3 cm. long, the ovate-lanceolate acute bracts shorter

than the shortly 5-dentate calyx, this 2-3 mm. long; corolla lilac,

3-4 mm. long, setose-puberulent without. — With V. Hayekii belongs

to section Verbenaca, subsection Pachystachyae Schauer, DC. Prodr.

11: 539 (Hayek). F.M. Neg. 17459.

Cuzco: (Moldenke). — Puno: Lake Titicaca, 3,700 meters, Weber-

bauer 440, type; 185. Bolivia.

2. HIEROBOTANA Briquet

Like Verbena but with only 2 fertile stamens and the calyx slightly

accrescent in fruit. The genus is therefore academic, as the occur-

rence of 2 or 4 stamens, or 4 and only 2 fertile, is traditionally ac-

cepted as a generic character within the family. The character,

however, as the number of nutlets, probably results in an arbitrary

delimitation of groups that naturally are interrelated and should be

merged, with sectional names indicating probable origins. Parallel

situations occur in the Solanaceae.

Hierobotana inflata (HBK.) Briq. Pflanzenfam. 4, Abt. 3a: 148.

1897. V. inflata HBK. Nov. Gen. & Sp. 2: 273, pi. 135. 1818; 550.

A canescent strigose-hispidulous procumbent shrub with sessile

leaves, 3-parted nearly to base and terminal early sessile and oblong

spikes, slightly peduncled and lax in fruit; leaves about 2.5 cm. long,

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

the lateral segments 2-3-parted, the intermediate 3- or 5-parted, the

laciniae linear, subobtuse, revolute; bracts lanceolate-subulate,

shorter than the calyx, this 4 mm. long, herbaceous, the nerves ending

in 5 subequal subulate mucronate teeth; corolla tube not exserted,

the small limb with rounded emarginate lobes, the throat obscurely

or scarcely villous; fruit obovoid, the calyx enlarged but only slightly

inflated; nutlets 4, reticulate-areolate dorsally. — Strongly apart by

the fruiting calyx widened at base, the stamens only 2 (Briquet);

expediently, however, it could be retained among the species it re-

sembles otherwise, but it is more clearly defined and probably more

remotely derived than the traditionally separated Lippia; however,

the only basic difference is the stamen reduction, the calyx character

appearing in other species, as in V. juniperina Lag.

Ayacucho: Rio de Lomas, 2,500 meters, Weberbauer 5749. To

Colombia.

3. LANTANA L.

Shrubs, rarely tall herbs, often sprawling or subscandent and

sometimes forming thickets alone or with other plants, usually

harshly pubescent with simple trichomes. Flowers often brightly

colored orange or red tints and not infrequently changing the tone

in age, ordinarily in capitate, less usually cylindric axillary peduncled

spikes. Corollas solitary, bracted, sessile, the spreading limb ob-

scurely if at all oblique, the 4 or 5 broad lobes obtuse or retuse.

Stamens 4, didynamous, medially affixed, included; anther cells

parallel. Ovary 2-celled, the solitary ovules basal and erect or sub-

basal and lateral. Style often short, the stigma oblique or sublateral.

Drupes more or less fleshy with 2-celled nutlet or finally two 1-celled

nutlets, smooth or rugulose dorsally. — Species apparently not stable

and may be fewer than described; the key here as elsewhere is thus

devised expediently, and is scarcely usable without much material;

there are probably about half as many genetic species (or even fewer)

than names.

Leaves ovate or ovate-lanceolate, rarely somewhat oblong-elliptic,

ordinarily acuminate, usually acutely, cordate to somewhat de-

current at base, mostly several to many cm. long and drying

dark (unless L. Sprucei, L. Svensonii); branches often angled,

erect or clambering.

Flower heads soon spiciform; corolla lilac-rose or throat yellowish.

Leaves hirtellous to glabrate beneath L. trifolia.

FLORA OF PERU 633

Leaves pilose-tomentose beneath L. Fiebrigii, L.fucata.

Flower heads little if at all longer in fruit; corolla orange-red

except as noted.

Peduncles shorter or little longer than leaves.

Branchlets partly white hispidulous, not aculeate.

L. Weberbaueri.

Branchlets glandular or aculeate or both.

Glandulosity, aculei often obvious.

L. foetida, L. armata, L. Moritziana, L. Camara.

Glandulosity, aculei lacking or at least one obscure; corolla

not orange-red (ex char.).

Corolla red, well-exserted; leaves early plane.

L. cujabensis.

Corolla whitish; leaves rugose, cordate (type).

L. Chamissonis.

Corolla white; leaves plane, long-decurrent . .L. canescens.

Corolla (known) rose-lilac or yellow; leaves rugose.

L. rugulosa.

Peduncles even in flower mostly longer than leaf (types, ex-

pedient).

Flowers yellow, turning red; indument partly viscid.

L. tiliaefolia.

Flowers white or tinted; indument eglandular.

Branchlets and peduncles glabrescent L. Sprucei.

Branchlets and peduncles early long-hirsute. . .L. Svensonii.

Leaves (unless last two species above) mostly or all at most about

4 cm. long, shortly or not acuminate, often subobtuse, often dry-

ing gray-green; branches subterete, prostrate-ascending or low.

Leaf indument various but not velvety beneath (all may be phases

of one).

Corolla yellow to orange, rarely whitish; peduncles mostly to

3 times longer than leaf.

Leaves firm, soon strongly rugose.!/, reptans, L. scabiosaeflora.

Leaves chartaceous, rather faintly rugulose L. limensis.

Corolla lilac, purple or white (unknown, L. Pavonii] ; peduncles

mostly 2 times longer than leaf.

Corolla purplish.

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

Leaves acute at base, about ovate.

L. Ferreyrae, L. angustibracteata.

Leaves attenuate (decurrent?) to base L. Pavonii.

Corolla white L. Haughtii.

Leaf indument compactly tomentose beneath; bracts all broad;

corolla lilac or white L. Haughtii, L. Zahlbruckneri.

Lantana angustibracteata Hayek, Bot. Jahrb. 42: 168. 1908.

Branches and leaves beneath appressed setose-villous; leaves

ovate-oblong or ovate, subobtuse, the upper in type to about 3 cm.

long, half as wide, crenate-serrate, deeply nervose-rugose, appressed

setose above, as the erect peduncles (these longer) and the lanceolate

acute bracts that are shorter than the flowers, the outer slightly

recurved; heads ovoid-globose; corolla nearly 1 cm. long, puberulent,

lilac. — Similar to L. velutina Mart. & Gal. but eglandular and bracts

small as in L. canescens HBK. and L. trifolia with entirely different

leaves (author). F.M. Neg. 17461.

San Martin: Pacasmayo to Moyobamba, (Stuebel 35, type). —

Ancash: (Moldenke).

Lantana armata Schauer in DC. Prodr. 11: 597. 1847.

Tetragonal branches and peduncles densely hirsute-aculeate;

leaves broadly ovate, the rounded base abruptly decurrent into

petiole 12-18 mm. long, attenuate acuminate to apex, 5-10 cm.

long, nearly 4-6.5 cm. wide, coarsely crenate, lineately rugose, as-

perous above, subcanescently hirtellous beneath; heads hemispheric,

the involucral strigose hirsute, outer bracts linear-lanceolate, as long

as lower flowers, the inner half as long, subulate-lanceolate; corolla

tube 8 mm. long.

Lima: (Moldenke). Colombia.

Lantana Camara L. Sp. PI. 627. 1753; 598. L. aculeata L. I.e.

974, fide Schauer.

Branches often spreading, tetragonal, glabrate or sens. lat. pubes-

cent or hirsute-pilose, more or less aculeate or merely scabrous, the

younger usually puberulent as the rigid peduncles, these about as

long as or longer than the leaves, and the capitate inflorescences;

petioles short, somewhat or scarcely merging into the cuneate or

rounded base of the ovate-oblong shortly acuminate leaves, these

serrate-crenate, reticulate rugose, very scabrous above, often canes-

FLORA OF PERU 635

cently short-villous beneath; heads subumbellate, the oblong-lance-

olate acute to subulate bracts 4-7 mm. long, appressed strigose; tube

of the (usually) early yellow, or orange, or changing orange to red

corolla puberulent, about 1 cm. long, limb 6-8 mm. wide; calyx mem-

branous, 3 mm. long; drupes black, 3 mm. thick. — Highly variable

vegetatively, including the flowers that may be white or yellow or

more often change, besides reds, to blues or purples; many given

varietal names (!); most worthy of this taxonomic attention is var.

aculeata (L.) Mold. Torreya 34: 9. 1934, prickles or spines conspicu-

ous, often recurved, the yellow-orange flowers finally pink or roseate

and the var. mista (L.) L. H. Bailey, Cycl. Am. Hort. 884. 1900,

young branchlets spreading hirsute, outer flowers soon saffron or

brick red, inner orange; it has been collected in Piura, Lima and

Tacna (Moldenke).

An infusion of the leaves is used medicinally (Herrera). Only a

few of the many collections referred here are cited; probably variable

and should include several named sorts.

San Martin: Tarapoto, Goodspeed Exped. 35021; 35041; Williams

5763. Zepelacio, Klug 3385. — Lima : Hacienda Tarate, Soukup 2894-

— Huanuco: Near Huanuco, 2038. Pozuzo, 4719; 4567; 4762. Rio

Chinchao, 5052. — Junfn: La Merced, 5320. Chanchamayo Valley,

Schunke 1577. Huancayo, Oxapampa, Soukup 2315. — Apurimac:

Pincos, Stork & Horton 10665. — Loreto: Yurimaguas, Williams 3804;

3940; 4941- Iquitos, Williams 3580; 1426; 7911—Cuzco: Prov. de

Paruro, (Raimondi). Valle de Santa Ana, Herrera 964- Valle del

Marcapata, Herrera 1150. Valle del Urubamba, Herrera 3183; 3218.

South and Central America; widely naturalized. "Aya-manchana,"

' 'lauraimana' ' (Herrera) .

Lantana canescens HBK. Nov. Gen. & Sp. 2: 259. 1818; 607.

Canescently serrico-strigose and glandular-punctate, sometimes

more strigose-hirtellous and scabrous; branches strict, tetragonal,

sulcate; leaves opposite, ovate-oblong or oblong-lanceolate, attenu-

ate and entire to petioles, crenate-serrate toward acuminate apex,

lineate-rugose; peduncles subequaling or shorter than leaves; heads

more ovoid, finally ovoid-oblong, the laxly imbricate spreading bracts

ovate-oblong or lanceolate, acuminate, the larger outer involucral,

the inner half as long as the white corolla; fruit nearly dry, the nut-

lets costate-rugose. — May not be in Peru, but following, at least

sens, lat., apparently belong to it; but compare L. Svensonii. F.M.

Neg. 39489.

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

- Apurimac: Pincos, Stork & Horton 10666 (det. Standley, L. rugu-

losa). Prov. Abancay, Goodspeed Exped. 10585; Vargas 9804. —

Cuzco: Valle de Lares, Herrera 787. Valle del Urubamba, Herrera

1549. To Mexico; Venezuela and Brazil. "Canirca" (Herrera).

Lantana Chamissonis (D. Dietr.) Benth. & Hook, ex Jackson

in Ind. Kew. 2: 28. 1895, or Briq. Nat. Pflanzenfam. 4, Abt. 3a: 151.

1895. Lippia Chamissonis D. Dietr. Syn. PI. 3: 598. 1843; 580.

Stems herbaceous, erect, divaricately branched, the branches

quadrate, hirsute; petioles about 1.5 cm. long; leaves cordate-ovate,

acuminate, to about 7 cm. long, serrate, venose-rugose, early strigose,

then scabrous above, canescent-tomentose beneath; peduncles simple

or geminate, shorter than the leaf; heads subglobose; bracts herba-

ceous, ovate, shortly acuminate, squarrose, or in fruit reflexed, stri-

gose-hirsute as membranous calyx, exceeded by corolla tube; corolla

white, throat flavescent; capsule exposed, lustrous. — Some specimens

resemble L. fucata Lindl. and some herbarium material (det. Mol-

denke) has leaves cuneately narrowed at base!

Junin: (Moldenke). Paraguay; Brazil.

Lantana cujabensis Schauer, in DC. Prodr. 11: 599. 1847.

Branches spreading, elongate, subherbaceous, obtusely 4-angled,

unarmed or muricate as the finely strigillose peduncles; leaves gland-

ular punctate beneath, glabrate or subtomentose; heads hemispheric,

bracts scabrous hirtellous, the outer broadly ovate, acuminate, early

involucral, the inner oblong-lanceolate, acute, to about a third shorter

than corolla. — Otherwise very much like L. Camara L. (author);

may, at least in Peru, largely supplant L. Camara but doubtfully

distinct. Collected, according to Moldenke, also in Tumbez, Junin

and Loreto.

Peru: Without locality, Poeppig 1405. — Loreto: Boqueron Padre

Abad, Woytkowski 34400. Brazil; Bolivia.

Lantana Ferreyrae Mold. Amer. Journ. Bot. 38: 326. 1951.

Prostrate leafy closely branched shrub, canescent-strigose includ-

ing the ovate acute bracts without, the antrorse pubescence especially

dense on the leaf -venation, petioles (3-5 mm. long), and peduncles,

these 3.5-5.5 cm. long; leaves ovate or ovate-elliptic, acute at base,

rounded to subacute at apex, 1.5-3 cm. long, 7-18 mm. wide, scabrous

and bullate above by the 4-5 slender impressed lateral nerves and

veins; heads hemispheric, a cm. or two wide; bracts 6 mm. long,

FLORA OF PERU 637

2-4 mm. wide, often slightly resinous glandular, as the tube (8 mm.

long) of the white corolla, the limb 4-5 mm. across. — The author

records the collector as his good friend and colleague but curiously

does not compliment him by justifying the proposal of this as a

new species.

Amazonas: Road to Leimebamba, near Chachapoyas, 1,800 me-

ters, (Ferreyra 7130, type).

Lantana Fiebrigii Hayek, Bot. Jahrb. 42: 169. 1908.

Small shrub with virgate 6-angled strigose branches and opposite

or ternate ovate or elliptic leaves contracted into the short petioles;

leaves appressed crenate-serrate, rugulose, scabrous above with bulb-

ous-based setae, canescent-tomentose and finely glandular-punctate

beneath, 5-8 cm. long, 4-5 cm. wide, the longer strigillose peduncles

erect; heads soon elongating, finally cylindric, the ovate long-acumi-

nate densely pubescent bracts shorter than corolla tube, the little

larger outer ones involucrate; corolla lilac-rose with yellowish throat,

the densely puberulent tube 1 cm. long, limb 6-7 mm. broad. — Very

near L. trifolia L. but leaves tomentose beneath, flowers nearly twice

as large (author); however, Standley referred the following to the

Linnaean species, perhaps correctly, as a variant. Much other ma-

terial, if this is distinct, belongs here rather than to L. trifolia. Also

in Ancash and Junin (Moldenke).

Cajamarca: Prov. Cutervo, Stork & Horton 10156. — Huanuco:

Chinchao, Stork & Horton 9579. — Cuzco: Marcapata, Vargas 9686.

Prov. Convention, Vargas 817. To Argentina.

Lantana foetida Rusby, Bull. N. Y. Bot. Card. 4: 431. 1907.

Glandular with spreading trichomes; branches divaricate, acutely

quadrate; petioles stout, to 1.5 cm. long; leaves ovate, abruptly acute

or truncate or subcordate at base, acute or subobtuse at apex, the

many teeth mostly obtuse, about 5-10 cm. long, 3-7 cm. wide, rugose,

paler with yellowish veins beneath; peduncles to 1 dm. long; heads

1.5 cm. thick or in fruit 2-2.5 cm.; bracts ovate, acute, 2-4 mm. long;

calyx 2 mm. long, truncate, corolla pubescent, 1 cm. long, curved,

2 mm. thick at ventricose middle, lobes to 3 mm. wide, spreading;

longest stamens reaching to middle of tube; fruit 7 mm. long, 5 mm.

thick, rounded, black, lustrous. — Key character after a hort. spec.

Dudley Herb., det. Moldenke; Bang 2034 pt., Bolivia, type.

Huanuco: Acomayo, Woytkowski 34-250. — Junin: San Ramon,

Constance & Tovar 2223. — Apurimac: Pincos, Stork & Horton 10665.

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

— Cuzco: Machu-Picchu, Herrera 3193 (distr. as L. Camara and

L. trifolia). To Patagonia.

Lantana fucata Lindl. Bot. Reg. 10: pi. 798. 1824. L. lilacina

Desf. Cat. Hort. Paris ed. 3. 392. 1829; Schauer in DC. Prodr. 11:

604. 1847.

Erect glandular punctate shrub, the tetragonal divaricate branches

as the peduncles and bracts strigose-hirtellous; leaves crenate, ovate,

oblong or lanceolate-triangular, acute or obtuse, subcuneately pro-

duced into petiole at the rounded or slightly cordate base, impressed

lineate and reticulate-rugose, above green but hirtellous-scabrous,

beneath pubescent or canescently tomentose; heads hemispheric,

soon spiciform, the membranous, pale, ovate or oblong acuminate

bracts squarrose, the inner half as long as the corolla, the smaller

outer subcordate-ovate; calyx minute; corolla rose-lilac, the tube

6-8 mm. long. — The leaf shape at base appears to be distinctive.

Peru: (Moldenke). To Brazil; Venezuela.

Lantana Haughtii Mold. Phytologia 1: 463. 1940.

Branches slender, 4-angled, early densely appressed-puberulent;

leaf scars large, corky, divergent; petioles very slender, 4-8 mm. long,

short whitish pubescent; leaves lanceolate or elliptic-ovate, acute or

acuminate at base, obtuse, 1-3.5 cm. long, 0.7-2 cm. wide, firm-

chartaceous, dark green but bullate and closely appressed-strigose

above, gray-green tomentulose beneath, the 4-6 secondary nerves

deeply impressed above; inflorescences axillary, capitate, 2 peduncles

per node, 3-6 cm. long; heads canescent pubescent, 1-1.5 cm. in diam-

eter; bractlets broadly elliptic or ovate, acute, 9 mm. long, 4 mm.

wide, canescent-pubescent or substrigose; corolla white, the tube

7 mm. long, the limb about 3 mm. wide.

Piura: Amotape Hills, Haught 139, type; 42. — Cajamarca: (Mol-

denke).— Amazonas: (Moldenke). — Junin: La Oroya, Kalenborn 73

(det. Moldenke).

Lantana limensis Hayek, Bot. Jahrb. 42: 166. 1908.

Procumbent, suffruticose, the subterete branches minutely ap-

pressed scabrous as the slender peduncles that twice exceed the ovate

leaves; leaves shortly acuminate, somewhat produced into petiole,

4-6 cm. long, 2-4 cm. wide, serrate, rugose, early appressed setulose,

densely soft-pubescent beneath; heads umbellate-fastigiate, not elon-

gating, 2.5-3 cm. broad, the inner linear obtuse little ciliate bracts

FLORA OF PERU 639

half as long as corolla tube, the outer subspathulate, about equaling

the corolla, this puberulent, about 1 cm. long, orange. — Resembles

in habit L. scabiosaeflora but fruit unknown (author); perhaps a

coastal race. F.M. Negs. 7870 (as L. scabiosaeflora) ; 17473.

Piura: Serran, Stork 11367.— Lima: Canta, Pennell 14615 (det.

Johnston). Barranco, limestone area, (Weberbauer 1654, type). San

Bartolome", Weberbauer 5264. Trailing in lomas along the sea, 5959.

Chosica to Matucana, Goodspeed 11319. — Junin: (Moldenke).

Lantana Moritziana Otto & Dietr. Allg. Gartenzeit. 9: 369.

1841, and ex Schauer in DC. Prodr. 11: 599. 1847.

Ex char, resembles greatly L. Camara and L. cujabensis; prickles

lacking or scattered and minute, the branches and peduncles hirtel-

lous, these shorter than leaves; leaves scabrous also beneath; bracts

strigose, subulate, much shorter than corolla, the outer lanceolate,

early involucral, the lower subequaling the flowers; calyx 2 mm. long,

scabrous; corolla tube 8 mm. long, limb 4 mm. broad, the smaller

lateral lobes truncate, others subquadrate, entire. — Like the variable

L. Camara L. unless for the narrow foliaceous outer bracts (Schauer).

Has been collected in Junin, Huanuco, Loreto, Apurimac, Madre De

Dios (Moldenke).

San Martin: Tarapoto, Woytkowski 35021 (det. Cuatrecasas,

L. Camara}; 35182 (det. Standley, L. tiliaefolia). Chazuta, Klug

4074 (det. Standley, L. tiliaefolia'). Venezuela.

Lantana Pavonii Mold. Phytologia 1: 464. 1940.

Slender margined branchlets, petioles — these 2-5 mm. long —

leaves above and peduncles (2-3 cm. long) more or less sparsely hir-

sute or hirsutulous; leaves elliptic or subrotund, acuminate at base,

acute or obtuse, 1-3 cm. long, 3-15 mm. wide, crenate-serrulate,

chartaceous, in age scabrous and bullate above, rather densely hir-

sute beneath; inflorescences solitary, the heads to 8 mm. long, 19 mm.

wide. — A perplexing species (author) ! Seemingly referable here is a

Madrid specimen with an herbarium name suggesting a decumbent

habit, from the Ruiz and Pavon expedition. F.M. Neg. 29684.

Peru(?): Without data, Ruiz & Pavdn, type (Delessert Herb.,

Geneva) .

Lantana reptans Hayek, Bot. Jahrb. 42: 167. 1908.

A shrub with long procumbent-ascending appressed strigose

branches; leaves broadly cuneate-ovate, subentire below, serrate

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

above, 1.5-2 cm. long, 1-1.5 cm. wide, reticulate-rugose, the nerves

impressed above where sericeous-scabrous, prominent beneath where

strigose-tomentose; peduncles several times longer than leaves, ap-

pressed setose; heads hemispheric, 1.5-2 cm. broad, the ovate-lance-

olate acute bracts nervose, appressed sericeous-strigose, the outer

little shorter than the corolla- tube, the involucrate outer equaling it;

corolla subpuberulent, 6-7 mm. long, lemon-colored. — By habit and

indument easily distinguished from L. scabiosaeflora (author). F.M.

Neg. 17484.

Cajamarca: Below Santa Cruz, 1,300-2,000 meters, (Weberbauer

4143, type). — La Libertad: (Moldenke). — Ancash: (Moldenke). —

Huancavelica: (Moldenke). — Ayacucho: (Moldenke).

Lantana rugulosa HBK. Nov. Gen. & Sp. 2: 261. 1818; 602.

Branchlets quadrate, pilose hirsute, or the petioles (8-12 mm.)

and peduncles, these solitary, equaling or exceeding the leaf; leaves

ovate, acute at base, acuminate, 7-10 cm. long, 3.5-5 cm. wide,

coarsely mucronate-crenate, strongly reticulate-rugose, hirtellous

above, canescently lanate tomentose; heads hemispheric, scarcely

longer after an thesis, involucrate, the lanceolate acuminate bracts

strigose as the very short calyx, the inner half as long as the flowers,

the outer foliaceous, 3-5-nerved, equaling the head; corolla villosu-

lous without, tube 8 mm. long, curved and dilated medially; drupe

dark violet. — May be a part of L. Camara. — The Cajamarca plant

is var. parvipedunculata Mold. Said to be a robust shrub a meter

tall or taller.

Cajamarca: Chota, Stork & Horton 10087; 10074; Raimondi 309.

— Huanuco: Pillao, Woytkowski 34045. Pozuzo, 4769. Muiia, 4022;

4062; 4268. Mito, 1523. — Junin: Agua Dulce, Woytkowski 35443.—

Ayacucho: Yanamonte, Weberbauer 5615. Pampalca, Killip & Smith

22244-— Apurimac: Huancarana, West 3763. To Colombia. "Aia"

or "aya-rosa."

Lantana scabiosaeflora HBK. Nov. Gen. & Sp. 2: 261. 1818;

602.

Branches procumbent, slender, subangulate, pilose as the short

petioles and filiform peduncles, these 3.5-10 cm. long; leaves sub-

rhombic-ovate, cuneate to entire base, less than 2.5 cm. long, coarsely

crenate-serrate toward acute tip, strigose scabrous above, softly pilose

beneath; heads small, hemispheric; bracts herbaceous, all lanceolate,

acute, hirtellous strigose and ciliate both sides, lineate-nerved, the

FLORA OF PERU 641

inner about as long as corolla tube, the longer outer involucral; calyx

about 1 mm. long, bifid, pilose, ciliate; corolla yellow or white with

yellow center, finely sericeous, the slightly curved tube oblong-ventri-

cose medially. — There is a f. albida Mold, in Ecuador and probably

in Peru. F.M. Neg. 39488.

Piura: Huancabamba, (Bonpland, type). — Cajamarca: Socota,

Stork & Horton 10106. — Ancash: Casma, Goodspeed Exped. 9180.

Chacchan, 2535. Banos de Chancos, Sandeman 4607 (det. Standley,

L. limensis). — Lima: East of Saydn, Prov. Chancay, Goodspeed

(33040). Matucana, 80; Goodspeed Exped. 11333.— Junin: Huari-

aca, 3103. — Huancavelica: Mantaro, Stork & Horton 10400. — Huan-

uco: Near Huanuco, 2086; 3220. — Apurimac: Curahuasi, Vargas

9609. Pincos, Stork & Horton 10659; 10666. Abancay, Vargas 741.

— Arequipa: Chala, Worth & Morrison 15606. — Moquegua: Torata,

Weberbauer 7404; 7406. Ecuador.

Lantana Sprucei Hayek, Repert. Sp. Nov. 2: 162. 1906.

Branchlets subtetragonal, sparsely appressed pilose; leaves oval

and obtuse or the upper rhomboid-lanceolate and acute, long-decur-

rent, coarsely crenate-serrate, plane, minutely setulose scabrous both

sides and also shortly hirtellous beneath; peduncles glabrescent, to

three times larger than the subtending leaf; heads fastigiately umbel-

late, not elongating, the larger outer bracts involucral, the inner

ovate, subacute, slightly puberulent, about as long as the subgla-

brous corolla, this 11-15 mm. long; style 2 mm. long. — Affine L. his-

pida HBK. of Mexico but glabrescent, leaves plane, peduncles longer,

flowers larger, bracts less pilose (author) ; suberect, prostrate or rarely

erect shrub common on rocky slopes, dunes and sea cliffs, the flowers

white to pale purplish with yellow center, leaves acute to obtuse with

appressed to spreading pubescence (Svenson). Type from Chanduy,

near Peru. F.M. Neg. 34336.

Piura: Cerro Prieto, (Haught & Svenson 11619; 11575). Amotape

Hills, (Haught & Svenson 11536). Ecuador.

Lantana Svensonii Mold. Geogr. Distrib. Verb. 78. 1942.

Low shrub, the subquadrate branchlets, often subulate petioles

(5-10 mm. long) and solitary peduncles — these to 1.5 dm. long —

early spreading hirsute, more or less subappressed pubescent to gla-

brate in age; leaves ovate or ovate-elliptic, crenate-serrate almost to

the narrowly long-attenuate petioles, acute, 2.5-7.5 cm. long, 1-

3.5 cm. wide, chartaceous or membranous, densely substrigose or

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

subvelutinous above, shortly pubescent or tomentulose beneath;

heads ovate-cylindric, to 2.3 cm. long, 2 cm. wide, at base with folia-

ceous ovate acute strigose-puberulent bracts (often ciliate) to 13 mm.

long, 9 mm. wide; corolla hypocrateriform, white or pale lilac, yellow

at throat, the slender puberulent tube about 5 mm. long, limb 3 mm.

wide. — Compare L. Chamissonis in herb, not as to type, ex char.

Soukup 4199 was a bush 6-7 dm. tall, forma albiflora Mold. Phyto-

logia 5: 12. 1954.

Tumbez: Hacienda Chicama, Weberbauer 7676; 7660; 7686. —

Piura: Parinas Valley, Haught 127. Negritos, Haught 12. La Brea,

Horton 11579. — Cajamarca: (Moldenke). — Lambayeque: Olmos,

Soukup 4199. — Arequipa: (Moldenke). Ecuador.

Lantana tiliaefolia Cham. Linnaea 7: 122. 1832; 600.

Branches spreading, tetragonal, unarmed or finely aculeate, vil-

lous-pubescent or villous as peduncles (these at least as long as leaves),

the indument now entirely viscid, now partly glandular; leaves sub-

rotund or ovate, cordate, truncate or cuneate into short petioles,

acuminate, serrate-crenate, reticulate-rugose, villous both sides or

strigose scabrous above, shortly villous or tomentose beneath; heads

hemispheric, bracts strigose-hirtellous, the oblong inner 2-3 times

shorter than corolla tube, the outer lanceolate or subspathulate, two

times longer, flowers orange or red. — None of the following is prob-

ably distinct from L. rugulosa. F.M. Neg. 34340.

San Martin: Tarapoto, Woytkowski 85182 (det. Cuatrecasas) ;

Williams 5387 (det. Standley). Chazuta, King ^07^.— Loreto :

Iquitos, Williams 1365. — Cuzco: Valle de Lares, Diehl. Quilla-

bamba, Soukup 153. Valle del Apurimac, Herrera. Brazil. "Aye

manchana."

Lantana trifolia L. Sp. PI. 626. 1753; 606.

Suffruticose, the stems strict, hexagonal, the peduncles quadran-

gular, both strigillose, hirtellous or hirsute; leaves 3-4-verticillate, or

opposite, ovate or oblong, attenuate at base, acute or acuminate,

several cm. to a dm. long, crenate-serrulate, often reticulate-rugose,

above scabrous or strigose, beneath resinous, punctate and hirtellous-

tomentose; peduncles solitary in the axils, finally subequaling the

leaves; spikes soon oblong, much elongating, 1-1.5 cm. thick; bracts

herbaceous, ovate, cuspidate-acuminate, 5-nerved, strigose, laxly im-

bricate, the lower to 1 cm. long, 3 mm. wide, the upper about as long

FLORA OP PERU 643

as corolla tube or 5-6 mm. long; lower lip of corolla short, middle

lobe sessile at throat. — Variable in indument; corolla small, lilac, the

throat yellow (Schauer); drupes purple or paler, 2-3 mm. thick.

There is forma hirsuta Mold, in Colombia and of course elsewhere

and a forma oppositifolia Mold, in San Martin. L. Fiebrigii Hayek,

Bot. Jahrb. 42: 169. 1908, of Bolivia, is very near but has leaves

tomentose beneath, flowers 1 cm. long, 6-7 mm. wide (author).

San Martin: Tarapoto, Goodspeed Exped. 35014; 35020; 35184;

Williams 5400; 7053. Zepelacio, King 3310. — Lima: Hacienda Villa,

Soukup 2915. — Huanuco: Piedras Grandes, 3675. Rio Chinchao,

4975. — Junin: Raimondi 7564; 12864. La Merced, 5323. Hua-

capistana, Killip & Smith 24112. — Loreto: Puerto Arturo, Killip &

Smith 27715. Yurimaguas, Williams 4224; 4982—Cuzco: Valle del

Urubamba, Herrera 1748 (det. Werdermann); 8523. Convention,

Herrera 169. Machu-Picchu, Herrera 3193. Warmer America.

Lantana Weberbaueri Hayek, Bot. Jahrb. 42: 166. 1908.

Low procumbent shrub with unarmed branches densely stipitate-

glandular, spreading, setulose and hispid-setose; leaves opposite,

regularly serrate, ovate, cuspidate-acuminate, rounded basally and

minutely produced into petiole, 6-8 cm. long, 3.5-4.5 cm. wide, line-

ate-rugose, densely appressed setose above, setose-villous on nerves

beneath; peduncles pubescent as the branches, erect-spreading, little

enlarged toward apex, slightly exceeding the leaves, the hemispheric

heads 2.5-3 cm. broad, the much larger outer bracts (subequaling

the flowers) involucrate, the inner (much shorter) ovate-lanceolate,

cuspidate, densely setose-villous; corolla 12-15 mm. long, shortly

pubescent, orange with red edges; heads globose in fruit, the pyrenes

2-3 mm. long. — Equally resembles L. robusta Schauer and L. viscosa

Schauer; from the first differs in long-setose stem and peduncles;

from the second in the shorter fewer glandular trichomes and the

not elongating heads (author). F.M. Neg. 17493.

Huanuco: (Moldenke). — Junin: Toward Palca above Huacapis-

tana, 1,900 meters, (Weberbauer 2017, type).

Lantana Zahlbruckneri Hayek, Bot. Jahrb. 42: 167. 1908.

Shrubby, the terete branches finely puberulent as the slender

peduncles, these about as long as the leaves; leaves crenate, ovate-

oblong, rounded or broadly cuneate at base, obtuse, moderately re-

ticulate-rugose, minutely but velvety puberulent above, canescently

appressed tomentose beneath; heads hemispheric, little elongating,

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

about 2 cm. broad; bracts ovate, acute, entire, tomentulose, sub-

equaling corolla-tube, the involucrate outer little larger; calyx to-

mentose, scarcely 1 mm. long; corolla lightly puberulent, the narrow

tube 6 mm. long, pale lilac with yellowish throat; drupes 3 mm. long.

— Habit of L. odorata L. but appressed tomentulose, corolla tube

narrow, heads not greatly elongating; from L. lilacina Desf. (i.e.

L. fucata Lindl.) and L. velutina Mart. & Gal. differs in the small

crenate leaves and soft indument (author); and in reference to the

last character Zahlbruckner had given it an herbarium name. Occurs

also in Lima according to Moldenke. Could be an extreme state of

L. canescens HBK. and the latter could include H. Haughtii Mold.

F.M. Neg. 17494.

Piura: Huancabamba, Weberbauer 6068. — Cajamarca: Shumba

to Jaen, Weberbauer 6181. — Amazonas: Tupen, 800-1,000 meters,

(Weberbauer 4778, type). — Loreto: Salinas de Pilluana on the Rio

Huallaga, (Ule 6822). — Arequipa: Atiquipa, Worth & Morrison 15643.

4. LIPPIA L.

Phyla Lour. Fl. Cochinch. 66. 1790. Aloysia Ortega ex Pers. Syn.

PI. 2: 139. 1807.

Much like Lantana but often less harshly pubescent if at all, the

trichomes in some herbs malpighiaceous, the spikes often elongate

(Aloysia) or subcapitate and rather few-flowered, less frequently sub-

capitate, compact. Bracts not or more or less 4-ranked, persisting

to deciduous, the calyx either compressed, bicarinate or bialate

(bracts persisting) or 2-4-angled or cleft or 4 dentate (former, Phyla;

latter, Aloysia). Corolla limb oblique, often sub-bilabiate. Fruit at

least typically included, dry, the nutlets rarely free. — Has been united

with Lantana but the mature fruit and often the calyx provide identi-

fication. In Peru the inflorescence is solid (Phyla) or if capitate more

or less open, otherwise spicate. L. aprica Phil., once accredited to

Peru, is Chilean. L. valerianoides Walp., Nov. Act. Acad. Leopold

Carol. Nat. Cur. 19. suppl. 1: 377. 1843, listed by Moldenke as a

Peruvian species, was referred in Herb. Dahlem (Berlin) to Valeri-

ana; curiously, however, it was not mentioned in that genus by

Killip. Named for A. Lippi, an Italian botanist.

Svenson, Amer. Journ. Bot. 33: 480. 1946, noted that 2-armed

trichomes in themselves do not indicate generic character (for ex-

ample, Phyla) since they are present in many families. DeCandolle,

Prodr. 17: 296. 1873, referred Phyla Lour, to L. nodiflora L. How-

FLORA OF PERU 645

ever, Moldenke (Fl. Suriname) retains Phyla (at least for one species)

on herbaceous or subherbaceous state and habit, cuneate-obovate

bracts not 4-ranked; Abrams recognizes Phyla Lour, and Aloysia

Ortega without reference, however, to Lippia, from which Aloysia

is the weaker segregate. A. ayacuchensis Mold, is omitted from key.

The Lemon Scented, Sweet Verbena or Yerba Luisa of gardens

is L. triphylla (L'He"r.) Ktze. of Chile with verticillate linear-lanceo-

late fragrant (gland-dotted) leaves and small bluish flowers in slender

spikes; also known as Aloysia triphylla (L. He"r.) Britton, the latter

group name as the Spanish, in remembrance of Maria Louisa, wife

of Charles IV of Spain.

Peduncles except L. alba ordinarily as long as or longer than the soli-

tary or geminate compact globose-ellipsoid or oblong-cylindric

inflorescence; plants erect, strict or rooting nodally; corolla often

obscure or tube included.

Leaves commonly distinctly or somewhat longer than wide, often

obovate or ovate.

Plants trailing L. subterranea, L. nodiflora.

Plants erect, virgate.

Leaves more or less villosulous beneath L. alba.

Leaves sparsely strigose L. americana.

Leaves rhombic-ovate, not much longer than wide.

L. betulaefolia.

Peduncles very short or inflorescence clearly longer than thick or not

rigidly compact or corolla tube exserted; shrubs, sometimes vir-

gately branched.

Inflorescence subcapitate but laxly, few-flowered.

Leaves flabellate-subrotund L. antaica.

Leaves cuneate-obovate to oblong-lanceolate.

Leaves entire or few-dentate, slightly or not obovate, bright

green A. Herrerae, L. Fiebrigii.

Leaves crenate-serrulate at least above base, more or less obo-

vate and canescent or pale.

Indument minute or obsolete on leaves above, granular

beneath L. ferruginea.

Indument asperous-hirsutulous above.

Leaves appressed puberulous beneath L. turbinata.

Leaves pilosulous beneath L. tayacajana.

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

Inflorescence spicate, clearly longer than thick at anthesis, often

lax, many-flowered.

Leaves rarely ternate; calyx teeth obvious to conspicuous (spe-

cies apparently in part weak but key in some cases only

from char.).

Leaves subconcolor, granular at least beneath, rugulose if at

all only in age.

Indument in part pilosulous; leaves membranous.

L. aloysioides.

Indument a puberulence; leaves subcoriaceous.

A. minthiosa.

Leaves more or less canescent beneath, early finely reticulate-

rugose unless L. peruviana, L. virgata.

Calyx teeth ovate, acute L. peruviana.

Calyx teeth subulate or ovate-subulate.

Leaves membranous, cordate; indument in part hirsutu-

lous or crisply pilose L. leptophylla.

Leaves firm, early rugulose, rounded to subobtuse; indu-

ment not or obscurely hirsute.

L. spathulata, L. scorodonioides.

Calyx teeth subulate-filiform; leaves ample, ovate, mem-

branous, subobtusely acuminate L. virgata.

Leaves 3-4-verticillate; calyx teeth minute L. triphylla.

Lippia alba (Mill.) N. E. Br. in Brit. & Wils. Sci. Survey Puerto

Rico 6: 141. 1925. Lantana alba Mill. Gard. Diet. ed. 8, no. 8: 1768.

L. geminata HBK. Nov. Gen. & Sp. 2: 266. 1818, fide Briquet.

Strict, the virgate tetragonal branches as the peduncles and leaves

above densely hirtellous-strigose-puberulent; leaves opposite or ter-

nate, elliptic to roundish-ovate, narrowed to 3-8 mm. long petiole,

acute or obtuse, crenate, more or less rugose, canescent tomentose-

villous beneath; peduncles short, axillary, enlarged above; heads

early subglobose, shortly oblong or oblong in fruit, 8-12 mm. long;

bracts often broadly ovate-acuminate, strigose, the inner about as

long as corolla tube (4-5 mm.), the outer spreading; calyx about

2 mm. long, bidentate; corolla some purple tone or white; drupes

dark violet. — A meter or two tall, aromatic.

Piura: Prov. Ayavaca, Weberbauer 6346. Huancabamba, Weber-

bauer 6013. — La Libertad: (Moldenke). — San Martin: Tarapoto,

FLORA OF PERU 647

Williams 5581. San Roque, Williams 7325; 7359—Loreto: Yuri-

maguas, Williams 4475; 4997. Caballo-cocha, Williams 2212. Rio

Itaya, Williams 183. Nanay, Williams 344- Leticia, San Salvador

and La Victoria, Williams 3055; 2636; 1559. Warm America; widely

introduced or cultivated. "Pampa-oregana," "oregano," "orega,"

"cidraero," "sideraera," "pan poregano."

Lippia aloysioides Loes. in herb, ex Mold. Phytologia 2: 10.

1941. Aloysia aloysioides Loes. & Mold, ex Mold. Phytologia 2: 9.

1941.

Shrubby, the rather slender glabrous branches with corky leaf-

scars 2-3 mm. high, the short branchlets, petioles (1-2 mm. long),

peduncles (2-3 cm. long) and rachis of the spikes, these 5-8.5 cm.

long, about 1 cm. thick in flower, sparsely (the rachis densely) spread-

ing pilose; leaves ovate, truncate or subtruncate at base, obtuse or

rounded at apex, 1.5-5 cm. long, 1.5-2.5 cm. wide, in type mem-

branous, light green, coarsely dentate nearly from base, the teeth

rounded, scabrous above, more or less pilose beneath, the nerves and

veins prominent both sides; inflorescences axillary; bractlets lanceo-

late, long-acuminate, 2-3 mm. long, calyx about 1.5 mm. long and

wide, strongly villous; corolla tube glabrous without, 5 mm. long,

the limb about 3 mm. wide. — Leaves sometimes or finally firm, rugu-

lose and obscurely cordate; compare L. virgata. As the well-chosen

name indicates, this species resembles the subgroup of species once

regarded as generically distinct when few entities had been discov-

ered; Loesener, in herbaria, included it in Lippia, and Moldenke

(while presuming to share publication) fails to say that Loesener

even recognized Aloysia! To 1.5 meters tall, the white flowers rose-

tinted (Goodspeed).

Lima: Above Chosica, Ferreyra 759 (det. Moldenke); Horton

10993 (det. L. virgata by Johnston). Surco, Soukup 3741; Weber-

bauer 5206. Near Sayan, Prov. Chancay, Goodspeed 33022; 33025.

Lippia americana L. Sp. PL 633. 1753; 579.

Sparsely strigose or glabrate; branchlets tetragonal, glandular

punctate; leaves spreading or reflexed, ovate- or oblong-lanceolate,

acuminate, crenate-serrate above the middle, the lower 7 cm. long,

half as wide, gradually reduced, repand, membranous, pale green,

scabrous above, sparsely strigose in the axils; heads pyramidately

subglobose; bracts cuneate-obovate, abruptly cuspidate, subequal-

ing the corolla, the lower little larger, all multinerved, little narrower

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

than the villous calyx, this about 1.5 mm. long, acutely bifid. — The

Ecuadorian f. hyptoides (Benth.) Mold., Phytologia 4: 292. 1953

(L. hyptoides Benth. PL Hartw. 122. 1843), is possibly the Peruvian

plant.

Piura: Huancabamba, Weberbauer 6013. Prov. Ayavaca, Weber-

bauer 6346. — Cajamarca: (Moldenke). To Mexico.

Lippia antaica Loes. & Mold, ex Mold. Phytologia 2: 19. 1941.

Branches acutely tetragonal, appressed-puberulent; petioles to

2 mm. long, often obscure, densely pubescent; leaves flabellate-

subrotund, more or less truncate at base, rounded at apex, about

2 cm. long and wide at maturity (many smaller in the axils), char-

taceous-subcoriaceous, evenly crenate revolute, scabrous and bullate

above, puberulent on the prominent veins and densely pubescent

beneath; peduncles erect, 5-10 mm. long, pubescent, 2 at each node,

the capitate inflorescence about 8 mm. long, 1 cm. wide; bracts ovate,

closely short-pubescent, 2.5-3 mm. long; corolla 4-5 mm. long, the

limb about 3 mm. wide. — The locality is not certain.

Cuzco: Anta(?), Weberbauer 5918, type.

Aloysia ayacuchensis Mold. Phytologia 6: 323. 1958.

Shrub with slender tetragonous costate glabrate branches, florif-

erous twigs many, 3-8 cm. long; leaves crowded, opposite, oblanceo-

late, 6-10 mm. long, 1-3 mm. wide, membranous-chartaceous, entire,

glabrous, pustulate above, resinous-punctate beneath, secondary

nerves apparent only beneath; inflorescences 1-2 cm. long, puberu-

lent; calyx 2 mm. long, corolla tube about 4 mm. long, exserted part

white-puberulent, limb 3 mm. wide. — Type of this very distinctive

species is in herbarium of Padre J. Soukup, and at the New York

Botanic Garden. The author does not explain the species (?) dis-

tinctiveness.

Ayacucho: At 2,407 meters, (Padre Loy 4187).

Lippia betulaefolia HBK. Nov. Gen. & Sp. 2: 264. 1818. 583.

Phyla betulaefolia (HBK.) Greene, Pittonia 4: 48. 1899.

Procumbent stems herbaceous, rooting toward base, the ascending

tetragonal branches as the entire plant usually canescent appressed

strigillose with bicuspidate eglandular trichomes; leaves rhombic-

ovate, cuneately long-attenuate into the half to nearly as long peti-

oles, acute, 3.5-7.5 cm. long, coarsely and acuminately serrate,

FLORA OF PERU 649

impressed pinnate-nerved and rugose; peduncles short, 1-3 in the

axils; heads small, ovoid, finally oblong, the many almost minute

flowers closely imbricated, little exceeded by the ciliate cuneate api-

cally dilated and abruptly cuspidate bracts; calyx scarcely 0.7 mm.

long; corolla pale rose, 1 mm. long. — Weed in cleared lands.

Loreto: Caballo-cocha, Williams 2285; 2475 (det. Standley).

La Victoria, Williams 2970 (det. Standley). — Arequipa: (Moldenke).

To Guiana; Paraguay.

Lippia ferruginea HBK. Nov. Gen. & Sp. 2: 265. 1818. 588.

More or less puberulent and ferrugineo-puberulent all over, also

somewhat resinous glandular, even to the tubular compressed calyx

— this 2 mm. long — and the 4 mm. long corolla; petioles 2-4 mm.

long; leaves oblong-obovate, cuneate to 3-nerved base, 12-18 mm.

long, half as wide, marginally reflexed, crenulate toward obtuse tip,

lineate-rugose, slightly scabrous both sides on nerves as the younger

sulcate branchlets; peduncles axillary, little incrassate above, shorter

than the reflexing leaf, the heads at most 6-flowered; bracts strongly

imbricate, subrotund-ovate, subacute, 1 mm. long; corolla puberu-

lent without, glabrous in the throat, the stamens inserted there. —

Perhaps the Vargas specimen is L. Fiebrigii but the leaves are cren-

ulate. F.M. Neg. 17503.

Piura: Huancabamba, Bonpland, type; Weberbauer 6066; Rai-

mondi. — Amazonas: (Moldenke). — Junin: (Moldenke). — Apurimac;

Prov. Abancay, Vargas 1462.

Lippia Fiebrigii Hayek, Bot. Jahrb. 42: 165. 1908. Aloysia

Fiebrigii (Hayek) Mold. Rev. Sudam. Bot. 4: 15. 1937.

Branches 4-angled, subpuberulent; petioles short; leaves little

revolute, lanceolate, acute, 1.5-2 cm. long, 3-6 mm. wide, ap-

pressed scabrous setulose above, glandular punctate beneath; heads

ovate, few-flowered, rather lax, foliose-bracted and racemosely ap-

proximate at tip of branches; ovate acute bracts and calyx subequal,

the latter 2 mm. long; corolla more than twice longer than the calyx

and, like the calyx and bracts, densely puberulent. — Belongs with

L. satureiaefolia Mart. & Schauer, L. boliviana Rusby, and L. turbi-

nata Griseb.; differs from the first in acute scabrous leaves, egland-

ular and entire bracts, from the third in entire leaves and differently

pubescent bracts, and from the second, which it resembles most, in

its smaller leaves and looser heads and in the entire not erose bracts

(author) . — May merge with L. ferruginea HBK. Type a meter shrub.

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

Apurimac: Oropeza Valley, Vargas 9777. — Ayacucho: (Moldenke).

— Cuzco: Valle del Urubamba, along river, Hen era 2230; Weberbauer

4910 (det. Hayek); 175. Calca, (Vargas 248). Bolivia.

Aloysia Herrerae Mold. Phytologia 2: 10. 1941.

Tetragonal rather slender branchlets glabrous and lustrous in age,

obscurely puberulent at tip; nodes annulate; petioles 1-2 mm. long

or obsolete; leaves entire, oblong or oblong-elliptic, acute both ends,

about 1-4.5 cm. long, 3-10 mm. wide, chartaceous, bright green and

very scabrous above with short bulbous-based trichomes, the tri-

chomes shorter and also densely resinous-punctate beneath, the re-

ticulation prominent only beneath; inflorescence axillary and terminal,

the spikes 7-12 mm. long on slender peduncles 1-2 mm. long, these

puberulent as the 2 mm. long calyces with flaring ovate teeth; corolla

tube densely pubescent, 4 mm. long, the limb about 3 mm. wide. —

Possibly a robust specimen of L. Fiebrigii Hayek and relatives with

short peduncles. The type number distributed as L. scorodonioides

HBK. or as L. boliviana Rusby; a remarkable species (author).

Cuzco: Valle del Urubamba, 3,000 meters, H err era 1534, type

(Field Museum). "Cedronsillo."

Lippia leptophylla Loes. in herb. Aloysia leptophylla Loes. &

Mold. Phytologia 2: 11. 1941.

Tetragonal medullose branches, leaves both sides, peduncles (2-

3 cm. long), axillary and terminal inflorescence more or less hirsute;

leaves somewhat amplexicaul, ovate, cordate at base, rounded at

apex, 2-5 cm. long, 1.5-4 cm. wide, membranous, coarsely dentate,

the indument beneath in part a puberulence, the trichomes above

weak, bulbous-based; spikes 5-8 cm. long, often subtended by a pair

of dentate bracts 1 cm. long, 7 mm. wide; bractlets conspicuous,

attenuate both ends, 5-6 mm. long, 1-1.2 mm. wide; calyx 3 mm.

long, the teeth subulate-attenuate; corolla tube glabrous, 5-6 mm.

long, the limb about 4 mm. across.

lea: Valle Rio de Pisco, Weberbauer 5374, type.

Aloysia minthiosa Mold. Phytologia 2: 12. 1941.

Slender branches and branchlets as well as subsessile subcoria-

ceous leaves (both sides) often obscurely but densely pulverulent-

puberulent and resinous glandular especially when young, the many

dense spikes (4-13 cm. long) including the calyces and corolla tube

within puberulent; leaf-scars very prominent; leaves elliptic-ovate,

FLORA OF PERU 651

7-19 mm. long, 5-12 mm. wide, evenly and bluntly serrulate almost

from base to obtuse apex, venation impressed only above; peduncles

slender, to about 1.5 cm. long; bractlets 1-1.5 mm. long; calyx about

2, corolla 5 mm. long, the former unequally, the latter equally 4-

lobed, its short tube puberulent within; stigma lobes minute. — Type

from a small shrub in crevice of a cliff, the euphonious name referring

to the minty fragrance of the white flowers. The original diagnosis

is here reduced by half.

Ancash: Yautan, 2564, type. — Lima: Without locality, fide author.

Lippia nodiflora (L.) Michx. Fl. Bor. Am. 2: 15. 1803; 585.

Verbena nodiflora L. Sp. PI. 20. 1753. L. canescens HBK. Nov. Gen.

& Sp. 2: 263. 1818. L. reptans HBK. I.e. Phyla canescens (HBK.)

Greene, Pittonia 4: 48. 1899. P. nodiflora (L.) Greene, var. canescens

(HBK.) Mold. Phytologia 1: 98. 1934, var. reptans (HBK.) Mold.

Alph. List Inval. Sci. Names Verb. 31. 1942.

Stems prostrate, more or less ligneous, intricately branched, mi-

nutely canescent-strigillose or scabriusculous as the leaves, these

cuneate-oblanceolate or -obovate, sessile or shortly petioled, rarely

2 cm. long, entire or usually serrate toward the apex, sometimes

obscurely nerved, ordinarily much exceeded by the nearly filiform

peduncles; flowering heads oblong-ovoid, 3-7 mm. in diameter, cylin-

dric in age; calyx often villous-compressed, with 2 ovate teeth placed

laterally, entire or emarginate; corollas white, 1.5 mm. long, about

3 mm. wide; mature fruit black-punctulate, early globose, finally

consisting of 2 parts. — Variable in pubescence and measurements, as

probably most species will be found to be when correctly interpreted.

The var. reptans (HBK.) Ktze., in Phyla (HBK.) Mold., refers to

plants with thinner, often larger, elliptic, coarsely serrate leaves, the

venation more prominent. Var. rosea (D. Don) Macbr., comb. nov.

— Zapania nodiflora (L.) Lam. var. rosea D. Don in Sweet, Brit. Fl.

Gard. ser. 2, 3: pi. 225. 1834, in Phyla (D. Don) Mold.— is usually

densely matted and canescent-strigose all over, the small leaves

cuneiform. Var. strigulosa (Mart. & Gal.) Macbr., comb, nov.—

L. strigulosa Mart. & Gal. Bull. Acad. Brux. 11: 319. 1844; 584;

P. strigulosa (Mart. & Gal.) Mold. — is the strigillose phase, the pin-

nate nerves strongly developed. In the Mexican type may be a local

species, in Peru more or less similar and found (Moldenke) in Caja-

marca, La Libertad and Lima.

The species sometimes forms mats several dm. in diameter often

on seaside dunes and cliffs.

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

Piura: Negritos, Haught 34- Talara, Haught 49. Truxillo, (Bon-

pland, type, L. canescens). — Cajamarca: Nanch6, Raimondi. — Huan-

uco : Near Huanuco, 2336. — Lima : Huara, Soukup 1 000. Near Lima,

Raimondi; Ruiz & Pavdn; 50; 51. Callao, 5880 (var. strigulosa) . —

Ayacucho: Raimondi, (var. reptans). — Arequipa: Posco, Mejia, (Gun-

ther & Buchtien 131; 131 a). Mollendo, Raimondi. Generally warm

areas.

Lippia peruviana Turcz. Bull. Soc. Mosc. 36, pt. 2: 200. 1863.

Aloysia peruviana (Turcz.) Mold. Rev. Sudam. Bot. 4: 15. 1937.

Tetragonous stems and branches finely pubescent; leaves atten-

uate to short petioles or subrounded at base, ovate, obtuse, crenate,

punctate and minutely scabrous above, more pubescent and canes-

cent beneath; spikes little exceeding the leaves, long-peduncled; calyx

hirsute all over, the triangular acute teeth subequaling the tube, the

corolla tube about three times longer than calyx. — After author, who

noted that L. scorodonioides HBK. has smaller subrotund-elliptic

leaves, acuminate calyx teeth, shorter flowers, and compared it also

with several extra-Peruvian species.

Peru: Without data on type (Mathews 585); apparently also col-

lected by Dombey (author).

Lippia scorodonioides HBK. Nov. Gen. & Sp. 2: 269. 1818; 573.

Aloysia scorodonioides (HBK.) Cham. Linnaea 7: 234. 1832. L. scoro-

donioides HBK. var. hypoleuca Briq. Bull. Herb. Boiss. 4: 338. 1896,

fide author.

Branches elongate, minutely pubescent or puberulent; leaves

sometimes ternate, elliptic, often broadly, evenly crenate, shortly

decurrent, quite obtuse, 1.5-3 cm. long, 12-18 mm. wide, rugose,

pubescent but obscurely or not (typically) scabrous above, white

tomentose beneath; peduncles solitary, axillary, subequaling the leaf;

spikes dense, early 1.5-2.5 cm. long, becoming often much longer and

lax; bracts linear, minute; calyx pilose-hirsute often all over, the

teeth more or less subulate; corolla twice as long as the calyx, pale

rose-colored, hirtellous, the throat villous. — The leaves are often

marked with a beautiful fine rugose-venation and close crenulation;

some of the following may represent the variant L. spathulata Hayek

with smaller leaves. Near Lima occur the variants Mathewsii Briq.

and detonsa Briq. I.e. 339 (A. scorodonioides var. Mathewsii (Briq.)

Mold. Phytologia 1: 95. 1934, and var. detonsa (Briq.) Mold. I.e.),

both with leaves scabrous above, lightly pubescent beneath, some-

FLORA OF PERU 653

what unevenly serrate, the first with spikes to 8 cm. long, calyx teeth

not white pilose, the second with spikes 4-5 cm. long, calyx long-

pilose all over. F.M. Negs. 24668 (var. hypoleuca); 24669 (var.

Mathewsii) .

Amazonas: Mathews 3160 (type, var.). — Ancash: (Moldenke). —

Lima: Matucana, 133; 357; Ferreyra 5420 (det. Moldenke); Weber-

bauer 5399 (var.). Prov. Huarochiri, 2250 meters, Goodspeed &

Metcalf 30237 (var. Mathewsii). Surco, Soukup 2718. Rio Chillon,

Pennell 14438 (var. Mathewsii, det. Moldenke). — Ayacucho: (Mol-

denke).— Apurimac: Oropeza Valley, Vargas 9763 (det. Standley,

L. virgata). Pincos, Stork & Horton 10660 (det. Standley). Rio

Pachapacha, Goodspeed Exped. 10541 (det. Standley). — Cuzco: Valle

del Urubamba, H err era 3417; 1535; Vargas 11062. Calca, Vargas

160. Prov. Paruro, Vargas 9732 (det. Standley). Anta, Vargas 3692.

Yucay, Soukup 717. — Puno: (Moldenke). Colombia to Argentina.

"Cedroncillo" (Vargas), "lauraimana" (Herrera).

Lippia spathulata Hayek, Bot. Jahrb. 42: 165. 1908. Aloysia

spathulata (Hayek) Mold. Phytologia 1: 95. 1934.

Branches quadrate, early sparsely puberulent; leaves spathulate-

orbicular, obtuse, briefly narrowed into the short petiole, 1-1.5 cm.

long, crenate, in type nervose-rugose and minutely scabrous-setulose

above, puberulent- tomentose beneath; racemes erect, slender, finally

lax, the rachis densely puberulent; bracts linear-lanceolate, little

longer than calyx, this 2.5 mm. long, somewhat inflated in fruit,

densely short-setose; corolla purple, subglabrous without, well-ex-

serted. — Closely related to L. scorodonioides HBK. but distinguished

by the small roundish leaves and especially by the sparser shorter

pubescence of calyx (Hayek); could be an ecological phase; when I

saw it at Dahlem I noted it as a small-leaved variant and collections

(verified by Moldenke) have suboblong-elliptic glabrate not at all

rugose leaves, calyces long-setose toward base, as Pennell 13134- A

1-3 meter shrub of rocky slopes at about 2,000 meters; flowers purple

or tube lilac, lobes white (Pennell). F.M. Neg. 17547.

Cuzco: Valle del Urubamba, Weberbauer 4911, type; 175. Sillque

to Calca, Herrera 1670. — Arequipa: Tiabaya, Pennell 13079 (det.

Johnston). Tingo, Pennell 13134 (det. Johnston). Near Arequipa,

Weberbauer 6835; Goodspeed Exped. 22129.

Lippia subterranea Rusby, New Sp. S. Am. Pis. 108. 1920.

Phyla subterranea (Rusby) Mold. Rev. Sudam. Bot. 5: 2. 1938.

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

Suffrutescent, gray with appressed trichomes, creeping and root-

ing (fibrous roots many), the branchlets 1-3 cm. long, erect; leaves

closely imbricate, oblanceolate, attenuate to sessile base, about 3

teeth each side toward obtuse tip, 1.5-2 cm. long, 3-6 mm. wide;

heads terminal, finally a cm. long or longer and half as wide; bracts

3 mm. long and about as wide, base nearly half as long, rounded apex

ciliate, mucronate; calyx campanulate, compressed; corolla 3 mm.

long, tube abruptly contracted at base, upper lip nearly twice as long

as lower, the very broad upper lip dentate, the lower with 3 spread-

ing lobes. — In dense mats in sand, the flowers blue (collector).

Arequipa: Mollendo, (R. S. Williams 2933, type).

Lippia tayacajana Mold. Phytologia 2: 21. 1941.

Striate slender branches shortly pubescent toward tips, soon gla-

brous, in age the bark somewhat shredding; leaves crowded, oblance-

olate, strongly narrowed to base (obscurely petiolate), rounded or

subacute, about 1-2 cm. long, 4-8 mm. wide, firm, minutely and

closely canescently scabrous-strigillose above, densely short-pilose or

somewhat tomentulose beneath, vein-reticulation prominent above,

secondaries conspicuous beneath, serrulate from about the middle,

slightly revolute; peduncles solitary, about a cm. long, the heads

scarcely 5 mm. long and broad; bractlets lanceolate, acute, appressed

strigose, about 4 mm. long; corolla 5 mm. long, the tube 2 mm. wide.

— Related to L. ferruginea HBK. (author); type about 1 meter tall.

Huancavelica: Near Pampas, Rio Mantaro, Prov. Tayacaja,

Weberbauer 6510, type. — Cajamarca: Weberbauer 7125.

Lippia triphylla (L'He"r.) Ktze. Rev. Gen. PL 3, pt. 2: 253. 1898;

574. Verbena triphylla L'He*r. Stirp. Nov. 21. pi. 11. 1784. Zapania

citr(i)odora Lam. Tabl. Encycl. 1: 59. 1791. L. citr(i)odora (Lam.)

HBK. Nov. Gen. & Sp. 2: 269. 1818. Aloysia triphylla (L'Her.)

Britton, in Britt. & Wils. Sci. Survey Puerto Rico 6: 140. 1925.

Glabrous or nearly except lightly scabrous on the striate branch-

lets, the 3-4-verticillate leaves above and the axillary verticillate

spikes somewhat pubescent; leaves lanceolate, often narrow, entire

or medially serrate, acute, densely glandular punctate beneath;

spikes slender, sometimes in a terminal panicle; calyx tubular,

slightly bilabiate; corolla white or bluish. — Lemon-scented or sweet

verbena of gardens, commonly cultivated as a substitute for the

beverage tea (Thea sinensis).

FLORA OF PERU 655

Ancash: Huaraz, Baiios de Chancos, Sandeman 4^12. — Lima:

Near Lima, (Soukup 1774). — Huanuco: Mito, 1641. — Ayacucho:

Stork & Horton 10797. "Cedron." Chile.

Lippia turbinata Griseb. PI. Lorentz. 195. 1874.

Rigidly branched shrub, the subtetragonal younger branches early

asperous, estriate; leaves ternate or opposite, with a fascicle of much

reduced leaves in each axil, lanceolate, attenuate to the short petiole,

very acute, 2-3 cm. long, 3-6 mm. wide, rigid coriaceous (twisted or

flexible in drying), dentate only above the base, asperous hirsute

above (trichomes callus-based), appressed pubescent beneath on the

rather prominent nerves, otherwise subglabrous; heads solitary, 3.5-

5 mm. long, 4-6 mm. thick, turbinate in fruit and to 1 cm. long, the

peduncles 2-4 mm. long; bracts multiseriate, imbricate, outer lanceo-

late, inner lanceolate-subulate to oval, rigid, medially pubescent,

subglabrous apically, medial nerve conspicuous; calyx more or less

2-cleft, densely pubescent, the segments linear-lanceolate, 1.5-2 mm.

long; corolla white, the tube 2.5-2.8 mm. long, scarcely exceeding the

bracts, pubescent and glandular punctate on the upper third, the

lips 3-lobulate, largest medial lower, to 1.5 mm. long; filaments from

middle of tube, style less than 1 mm. long; fruit 2 mm. long, seed

obovoid. — After N. S. Troncoso, Darwiniana 10: 80. 1952, who notes

that leaves may vary to obovate, bracts smaller, ovate and apiculate,

heads sometimes geminate, but the significance of these variations is

not yet known. Illustrated, Troncoso, I.e. 82 (branch, flower and

fruit in detail).

Tacna: (Moldenke). Chile; Argentina.

Lippia virgata (R. & P.) Juss. Ann. Mus. Paris 7: 73. 1806; 573.

Verbena virgata R. & P. Fl. Peruv. 1 : 20, pi. 32b. 1798. Aloysia vir-

gata (R. & P.) Juss. Ann. Mus. Paris 7: 73. 1806. A. urticoides Cham.

Linnaea 7: 238. 1832.

Branches strict, lenticellate, subterete, glabrous; leaves usually

ovate or oblong, to about 7 cm. long, half as wide, attenuate to peti-

ole, acute or acuminate, crenate or serrate, pinnate-nerved-rugulose,

membranous, subcallous-asperous above, cinereous-tomentose be-

neath; racemes at base of axillary branchlets or paniculate, scarcely

peduncled, elongate; calyx tubular, pilose-hirsute below, semi-4-cleft,

the subulate-cuspidate teeth hispid-scabrous. — The variant elliptica

Briq. (A. virgata, var. elliptica (Briq.) Mold.), leaves obtuse or

rounded, decurrent and entire at base, has been found in Cuzco,

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

fide Moldenke. My collection from a stream-bank shrub-tree 4 me-

ters tall, the white flowers fragrant; known to attain 8 meters, the

crown spreading, trunk columnar, bark pale yellow-brown with long

coarse fissures, flowers white or with a violet hue, wood used only for

fuel (Williams, Field Mus. Bot. 15: 438. 1936). F.M. Neg. 24658

(an ined. name after Pavon).

San Martin: Tarapoto, Williams 5565 (det. Standley). — Huan-

uco: Pozuzo, Ruiz & Pavdn, type. — Junin: La Merced, 5305 (det.

Johnston). — Apurimac: Oropeza Valley, Vargas 9763 (det. Stand-

ley). — Cuzco: Machu-Picchu, 2,400 meters, Herrera 3258. Alto Rio

Urubamba, Diehl 2425 (det. Standley). Prov. del Cercado, 3,450

meters, Herrera 1925. To Brazil and Argentina. "Chicharra caspi"

(Williams).

5. BOUCHEA Cham.

References: Moldenke, Repert. Sp. Nov. 48: 16-29. 1940; 49:

91-139. 1940; Grenzebach, Ann. Mo. Bot. Gard. 13: 71-100. 1926.

Herbs or more or less ligneous, the leaves rarely entire, the ter-

minal spikes or racemes rarely short. Flowers obscurely if at all

pedicellate, usually borne laxly and the bracts small. Calyx narrowly

tubular, the 5 prominent ribs more or less extended. Corolla tube

slender, the oblique limb with 5 frequently unequal or subequal lobes.

Stamens 4, didynamous, inserted medially or higher, the anthers with

parallel cells. Ovules usually solitary, erect from base, the cells 2.

Style filiform, the clavate stigma with a posterior tooth or tubercle.

Fruit dry, shortly or not exserted, oblong or linear, separable into 2

cocci. — Named meritoriously for Peter Carl and Carl David Bouche",

father and son, eminent horticulturists at the noted Schoneberg

Garden, Berlin, in the nineteenth century. Name conserved. Mol-

denke, I.e. 119, quotes from correspondence between St. Hilaire and

Kunth which shows, interestingly, that the observing and extremely

able French botanist was deterred by his friend from proposing a

new genus based on his V. pseudogervao, six years before Chamisso's

segregation was published.

Bouchea fluminensis (Veil.) Mold. Repert. Sp. Nov. 49: 117.

1940. Verbena fluminensis Veil. Fl. Flum. 1: 17. 1825; Icones, pi. 38.

1827; 117.

Herb or woody below with purple to white flowers, sometimes a

meter high, glabrous or slightly pubescent particularly the racemes,

these 1-3 dm. long, leaves ovate to elliptic-oblong, coarsely mucro-

FLORA OF PERU 657

nate-dentate down to the cuneate base, acuminate, pale beneath;

flowers subsessile; bracts linear-lanceolate, about 5 mm. long, the

bractlets about a third as long; calyx 10-13 mm. long, finely pubes-

cent; cocci basally coherent, subcylindric, 12-13.7 mm. long, 1.5-

2 mm. wide, a little exserted, beak (2.5 mm. long) short or obscure,

dorsally ridged base to apex, the commissural surface smooth, plane

or somewhat concave. — The similar B. prismatica (L.) Ktze., 97,

probably extending into Peru from northern South America, has

calyx 5-9 (10) mm. long, the 2 cocci separating, about 7.5 mm. long,

beak 1-1.5 (3) mm. long, dorsal surface often smooth, commissural

often rough. Too many collections have been made of this weedy

plant. Illustrated, Grenzebach, I.e. pis. 9-11, figs. 11-24 (fruits).

San Martin: Tarapoto, Goodspeed Exped. 35004; 35105; Williams

5469; 5561; 5808; Woytkowski 35004. Juanjui, King 3904; 4206 —

Junin: La Merced, 5304. Puerto Bermudez, Killip & Smith 26625.

— Loreto: Balsapuerto, Klug 2839. — Cuzco: Valle Lares, Diehl 2429.

Prov. Convention, Soukup 800. Ecuador and Brazil. "Pacunga,"

' 'pakungua' ' (Woytkowski) .

6. STACHYTARPHETA Vahl

Similar vegetatively to Bouchea, to which students have allied it,

but with only 2 perfect stamens and the anther cells divaricate, the

stigma orbicular-subcapitate and the ovules laterally affixed near

the base. Leaves sometimes alternate. Pubescence villous, pilose

or lacking. — St. Hilaire, as quoted by Moldenke (see Bouchea) as-

serted his disbelief in the separation of this genus; and no wonder

in view of his experience. The conserved name of Vahl supplanted

Valerianodes [Boerh.] Medic.

Spikes filiform or slender, the rachis and grooves subequal in width.

Corolla tube about 1 cm. long.

Leaves more or less decurrent; corolla tube more or less exserted.

S. cayenensis.

Leaves long-decurrent; corolla tube well-exserted . .S. peruviana.

Corolla tube about 2 cm. long S. quirosana.

Spikes to 2 cm. thick; rachis incrassate.

Leaves ovate or subrotund, coarsely serrate S. canescens.

Leaves oblong-elliptic, serrulate, more or less attenuate to base.

S. Weberbaueri.

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

Stachytarpheta canescens HBK. Nov. Gen. & Sp. 2: 281. 1818;

565.

Subdichotomous, the stems and leaves somewhat viscid tomen-

tose-lanate or hirsute-villous, the branches quadrate; leaves sub-

sessile, ovate or subrotund, about 3.5 cm. long, 2-2.5 cm. wide,

coarsely acuminate-serrate, rugose; spikes slender, finally incrassate,

elongate, the flowers recurved-spreading, the calyx in fruit appressed,

6 mm. long; bracts firm, scarious-margined, subulate, subaristate, as

long as or shorter than the acutely 4-dentate calyx, the longer corolla

blue. — Variable in abundance of indument, especially in spike, some-

times sparsely glandular; S. Weberbaueri may be found to be a part

of this species.

Cajamarca: Near Sarta (Truxillo), Bonpland, type. Brazil.

Stachytarpheta cayenensis (Rich.) Vahl, Enum. 1: 208. 1804;

562. Verbena cayenensis Rich. Act. Soc. Hist. Nat. Paris 1 : 105. 1792.

Verbena dichotoma R. & P. Fl. Peruv. 1 : 23, pi. 34- 1798. S. dichotoma

(R. & P.) Vahl, I.e. 207, fide Moldenke (at some time). S. umbrosa

HBK. fide Moldenke (at some time).

More or less dichotomous, the branches early, especially at nodes,

lanate; leaves somewhat decurrent into petiole, ovate or ovate-

oblong, acute or acuminate, coarsely serrate, or subincised, 2.5-

10 cm. long, about half as wide, glabrous to scabrous-hirtellous or

strigillose especially on the nerves beneath; spikes elongate, slender,

the flowers erect in age; bracts subulate, aristate, scarious-margined;

calyx 4-costate-plicate, about 4-6 mm. long; corolla blue, scarcely or

somewhat exserted, the limb small. — Rachis and grooves each about

the same width; in the similar and widely distributed S. jamaicensis

(L.) Vahl the latter are narrower than the former. Richard wrote

"cayennesis."

San Martin: Tarapoto, Goodspeed Exped. 35017 (det. Moldenke).

Lamas, Williams 6475. — Huanuco: Cuchero and Chinchao, Ruiz &

Pavdn (type, V. dichotoma). — Loreto: Rio Itaya, Williams 35. —

Cuzco: Valle del Urubamba, Herrera 3480. Widely distributed in

warm America.

Stachytarpheta peruviana Mold. Phytologia 1: 473. 1940.

Perennial herb, the type about 1 meter high, obtusely angled,

glabrous or essentially, the slender branches and thin-membranous

bright green leaves lustrous, the spikes to 3 dm. long and very slen-

der; leaves long-attenuate into alate indistinctly defined petiole

(about 1 cm. long), ovate, acute, 8-10 cm. long, 4-5.5 cm. wide,

FLORA OF PERU 659

the expanded portion coarsely serrate with broad subacute minutely

apiculate teeth; secondaries slender, 5-7 each side; peduncles very

short or obsolete; rachis glabrous; bractlets lanceolate, 4 mm. long,

long-acuminate, scarious-margined near base; fruiting calyx rather

deeply impressed; corolla pinkish-blue, the slender tube 7 mm. long,

the limb about 4 mm. wide. — A distinctive species (author); cf.

S. cayenensis (Rich.) Vahl.

Ayacucho: Edge of forest, along beach, Rio Apurimac Valley,

Killip & Smith 22853, type (det. Standley, S. cayenensis).

Stachytarpheta quirosana Mold. Phytologia 1: 457. 1940.

Shrubby, the slender branches sparsely subappressed pilose as the

thin-chartaceous dark green leaves above; nodes annulate; petioles

subobsolete or to 5 mm. long; leaves ternately fasciculate, usually

also with several smaller ones on greatly abbreviated twigs in their

axils, elliptic or obovate, long-cuneate at base, acute, 2.5-7.5 cm.

long, 1-3.2 cm. wide, densely short-pubescent beneath, not scabrous

above, regularly serrate at least from about the middle; secondary

nerves very slender, 5 or 6 on each side; spikes terminal, solitary,

about 1.5-5 dm. long, the flowers imbricate above; rachis short,

slender, 2-5 mm. thick, appressed pilose, deeply excavated; bractlets

lanceolate, attenuate-filiform; corolla tube 2 cm. long, the limb IL-

LS cm. in diameter.

Piura: Valley of the Quiros, Weberbauer 6343, type.

Stachytarpheta Weberbaueri Mold. Phytologia 1: 477. 1940.

Shrubby with acutely (even subalate) tetragonous densely puber-

ulent branches and peduncles, these to 2 cm. long, annulate nodes,

chartaceous subsessile regularly serrulate elliptic leaves, tomentulose

on both sides and shortly pubescent spikes very densely flowered for

7-25 cm., the rachis deeply excavated, densely short pilose; leaves

long-attenuate to petiolar base, acute or subacute, about 2-4 cm.

long, half as wide, the 5-7 secondary nerves subprominent beneath,

the veinlet reticulation abundant; bractlets oblong-elongate, about

1 cm. long; corolla about 2 cm. long. — Spikes are to 2 cm. thick.

Cajamarca: Between Tabacamos and Maranon valleys, 1,200 me-

ters, Weberbauer 6178, type. Sucse River, Stork & Horton 10092.

7. PRIVA Adans.

Reference: Moldenke, Rep. Sp. Nov. 41: 1-76. 1936; Phytologia

5: 73-80. 1954.

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

Decumbent to erect often pubescent herbs marked by a tubular

5-costate, 5-dentate calyx strongly accrescent and usually dilated in

fruit and contracted at throat. Spiciform racemes terminal or axil-

lary, the soon laxly borne small subbilabiate flowers early often essen-

tially sessile. Stamens 4, didynamous, medially affixed; anther cells

slightly or not divergent. Ovary cells 4 (Moldenke) as basal erect

ovules or by abortion 2-celled. Style filiform, unequally 2-lobed, the

minute stigma at tip of larger lobe (Moldenke). Fruit included, with

two 2-celled echinate nutlets or sometimes 1-celled. — Kobuski, Ann.

Mo. Bot. Gard. 13: 1-34. 1926, gave a concise revision with many

fine figures; Moldenke, as indicated, has revised or reinterpreted

several observations of earlier students.

Stems as petioles pilose; cocci quadrangular P. lappulacea.

Stems as petioles puberulent; cocci subspheroid P. peruviana.

Priva lappulacea (L.) Pers. Syn. PI. 2: 139. 1806; 24. Verbena

lappulacea L. Sp. PI. 19. 1753.

Glabrous or somewhat pilosulous except toward base; petioles to

3 cm. long; leaves ovate, subtruncate or subcordate at base, acute

or acuminate, coarsely subappressed serrate, mostly several-many

(14.5) cm. long, nearly half as wide, membranous, sparsely pilose-

strigose above, pale and scattered pilose-puberulent beneath; racemes

to 2 dm. long, linear-setaceous bractlets (prophylla) 1-2.5 mm. long;

calyx 2-3 mm. long, in fruit 5-7 mm. long, the trichomes uncinate;

corolla variously tinted, tube about 3.5 mm. long, largest lobes

1.8 mm. long; fruit quadrate, the united ligneous cocci 3 mm. long,

2 mm. wide, inner (commissural) surface flat or nearly, plane, not

margined, dorsally with 2 rows of acute or spinose tubercles and

obscurely roughened between them. — Illustrated, Kobuski, I.e. pi. 4,

fig. 10 (fruit); pi. 5, fig. 19 (calyx).

A weedy plant decumbent-ascending, to even a meter tall, doubt-

less growing in other than the following departments.

Tumbez: Ricaplaya, Weberbauer 7737. — Cajamarca: Raimondi.

— Lima: Esposto. — Huanuco: (Asplund 12096}. — Junin: La Merced,

5296. — Loreto: Rio Nanay, Williams 397. Rio Itaya, Williams 291.

Yurimaguas, Williams 4428. Iquitos, King 1238. Most of warmer

America.

Priva peruviana Mold. Repert. Sp. Nov. 41: 23. 1936.

Stems, branches, petioles (4-10 mm. long) and leaves beneath

more or less puberulent, glabrate in age; leaves ovate, uniformly

FLORA OF PERU 661

appressed serrulate, broadly rounded to acute base, acute, about

2-4 (4.3) cm. long, 7-2 cm. wide, strigillose above; racemes very

lax, 3-19 cm. long, the 4-22 flowers often pseud o-secund; pedicels

1-2 mm. long; calyx about 5.5 mm. long, to 7 mm. long in fruit,

uncinate-puberulent; corolla tube 5.2 mm. long, largest lobe about

1.8 mm. long; fruit obovoid-spheroid, long-attenuate at base, each

nutlet finally 5-5.5 mm. long, 2 mm. wide, dorsally echinate in 2

series (or 1 row reduced), also reticulate-rugulose, the inner surface

excavated and clearly margined. — Suggests P. portoricensis Urban,

but different (author) ; otherwise, presumably, the new name would

not have been published.

Cajamarca: Raimondi 396. — Amazonas: Chachapoyas, (Mathews

3158, type).

8. CASTELIA Cav.

Priva Adans., section Castelia (Cav.) Briq. Pflanzenfam. 4, Abt.

3A: 155. 1897.

Reference: Moldenke, Phytologia 6: 232-241. 1958.

Differs from Priva especially in the thick hard merely rugulose

pyrenes (Briquet) and in the (in part) verticillate flowers, tuberous

roots (Moldenke) . — Kobuski, following Briquet and Rusby, included

it in Priva, which classification indicates its closest living affinity,

but in floristic work it conveniently may be considered a separate

entity; moreover, Moldenke listed (after Miers and others) 12 con-

trasting characters, notably, the calyx not globosely dilated in fruit,

corolla-tube veins straight, staminode present, nutlets joined in

fruit. The name recalls Juan de Dios Castel, Spanish companion

of Loefling on his trip up the Orinoco.

Castelia cuneato-ovata Cav. Anal. Sci. Nat. 3: 134. 1801;

Icones 6, pi. 583. 1801. Priva cuneato-ovata (Cav.) Rusby, Bull.

Torrey Club 27 : 80. 1900. P. laevis Juss. Ann. Mus. Paris 7 : 70. 1806.

Somewhat succulent, a few dm. high, little branched, the broadly

ovate leaves, often with small ones in their axils, narrowed to short

petioles, obtuse or acute, crenate-serrate, 3-8 cm. long, 1-4 (5) cm.

wide, glabrous or slightly puberulent; bracts (prophylla) oblong-

lanceolate, 4-7 mm. long; flowers scattered or mostly verticillate,

sessile or subsessile; calyx purplish-striped, finally about 1 cm. long,

2-lipped, the acuminate teeth involute and contorted over fruit;

corolla more or less bilabiate, purple to reddish, lilac or nearly white,

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

the tube curved; anthers cordate or sagittate; stigma simple, unci-

nate; fruit drupaceous, indehiscent, the 2 cells covered by a dorsal

rather corky mesocarp surrounded by a thick pith, the 2 osseous

nutlets (pyrenes) about half immersed, convex dorsally, 2-sulcate

within (after Moldenke who as usual describes carefully and com-

pletely in detail). — Tuber edible, white (Eyerdam) at the end of a

long string of fibrous roots, the flowers fragrant (Balls) ; corolla white,

the tube purplish (Metcalf). Illustrated, Kobuski, Ann. Mo. Bot.

Card. 13: pi. 4, fig- 12 (fruit); pi. 5, fig. 21 (calyx).

lea: Scolnik 1082. — Tacna: Road to Calientes, Metcalf 30350. In

fields near Tacna, Eyerdam 24646; (Raimondi 1813; Rusby 2531;

Shepard 269). Chile; to Argentina.

9. PETREA [Houstoun] L.

Reference: Moldenke, Repert. Sp. Nov. 43: 1-48, 161-221. 1938.

Scandent or arborescent, glabrous or more or less scabrous with

somewhat lenticellate branchlets and generally coriaceous opposite

or whorled leaves, often purple flowers in terminal and upper axillary

usually elongate racemes, the frequently beautifully colored enlarged

calyx lobes most conspicuous at anthesis, rather green in age. Calyx

tube commonly 5- or 15-costate, ordinarily exceeded by the mem-

branous finally reticulate mostly equal lobes, these at base with a

thin sinuate 5-dentate or 5-cleft crown or corona. Corolla tube short,

the usually oblique rotate limb with 5 often unequal lobes. Sta-

mens 4, didynamous, medially affixed; anther cells parallel, the con-

nective generally enlarged. Ovary more or less imperfectly 2-celled,

the single ovule lateral, ascending or pendent. Calyx in fruit in-

durate, the lobes accrescent, the enclosed fruit indehiscent, usually

with two 1-seeded stones. — Camp, as reported in detail by Moldenke,

I.e. 12, determined by anatomic studies that the small calyx lobes

alternate to the large ones are appendages and not remnants (for-

merly considered incorrectly an epicalyx); Moldenke terms these

(entire, dentate or divided) the calycinal crest. Several of the spe-

cies defined by him are not (at least from his presentation) clearly

contrasted, their characters seemingly within the range of reasonable

or expected variations.

Often cultivated for its showy inflorescence; named by Houstoun

(according to Moldenke) for the eminent early eighteenth century

floriculturist of England, Petre, said to have first grown Camellia

japonica in Europe. Besides P. volubilis L., P. arborea HBK. and

FLORA OF PERU 663

P. Kobautiana Presl may be grown in Peru; both have basally

rounded to connate leaves, the former with hirsutulous, the latter

puberulent calyx, often glabrous or obscurely scabrous leaves.

KEY TO PETREA (after Moldenke)

Calyx tube hirsute or tomentose, 3-6 mm. long.

Calyx lobes finally about 1 cm. long; anther connective glabrous.

P. pubescens.

Calyx lobes about 2 cm. long; anther connective pilose.

P. wlubilis.

Calyx tube, if developed, short-pubescent or puberulent.

Calyx tube (mature) 8-9 mm. long P. macrostachya.

Calyx tube shorter or obsolete, or about 2-6 mm. long.

Calyx lobes (mature) to 36 mm. long, 17 mm. wide.

P. peruviana.

Calyx lobes about 1.5-2 cm. long, 5-10 mm. wide.

Leaves coriaceous, smooth, lustrous above; calyx tube to

5 mm. long P. nitidula.

Leaves more or less scabrous (as to types).

Calyx tube to 2 mm. long; leaves membranous-chartaceous.

P. atrocoerulea.

Calyx tube 3-5 mm. long; leaves often subcoriaceous.

Calyx lobes subobovate; leaves finally strongly bullate-

venose P. Martiana.

Calyx lobes suboblong to obovate in age; leaves slightly

or laxly venose P. peruviana, P. maynensis.

Petrea atrocoerulea Mold. Repert. Sp. Nov. 43: 195. 1938.

Younger branchlets short-pubescent, petioles (5-12 mm. long),

peduncles (1-3.5 cm. long) and pedicels (1.5-3 cm. long) more or less

puberulent; leaves opposite, entire, broadly elliptic, about 7-17 cm.

long, 3-11 cm. wide, at maturity gray-green, firm-membranous, ob-

scurely scabrous both sides and somewhat rugulose beneath, the 7-10

pairs of lateral nerves and densely reticulate veins prominent be-

neath; inflorescence axillary and terminal, to about 4 dm. long;

bractlets 4-7 mm. long; calyx glabrous, the tube about 4 mm.

long, 5 mm. wide at top, not costate but rugulose at base, the sub-

ovate lobes about 1.5 cm. long, 4 mm. wide above, the basally cleft

crest with acute lobes 2 mm. high; corolla tube about 13.5 mm. long,

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

sparsely puberulent toward apex where 1 cm. across, densely pubes-

cent within, the largest of the puberulent obovate lobes nearly 1 cm.

long; anther connective densely long-pilose; calyx tube in fruit to

5.5 mm. long, lobes to at least 4 cm. long, the callose crest converg-

ing inward.

Peru: Without data, Ruiz & Pavdn. Colombia; Brazil.

Petrea macrostachya Benth. Ann. Nat. Hist. ser. 1, 2: 448.

1839; 204.

Resembles in general P. volubilis L. but indument entirely a pu-

berulence especially dense only on the younger branchlets and peti-

oles; leaves broadly elliptic or ovate-elliptic, entire or sinuate, rounded

or acute at base, obtuse, emarginate, sometimes apiculate, to about

2 dm. long and half as wide or smaller, ordinarily obscurely asperu-

lous or glabrate, not bullate; inflorescence axillary or terminal, about

2.5-6.5 dm. long, nutant on peduncles 1-3 cm. long (or shorter), often

with 2-5 pairs of sterile bractlets; pedicels about 1.5 cm. long, sparsely

puberulent; calyx 7-10 mm. long, subglabrous, the obovate lobes

about 1 cm. long, 4-4.5 mm. wide, the crest united for 4 mm.,

its lobes 1 mm. long, glabrous; corolla tube to 12 mm. long, rather

sparsely hirsute within, the largest lobe 4.5-11 mm. long; anthers

short pilose; fruiting calyx indurate, strongly 10-plicate-costate, to

9 mm. long, the lobes to 27.5 mm. long, the callose crest converging

inward. — Stem said to attain 1 dm. in diameter, corolla violet-purple,

calyx pale blue. F.M. Neg. 34292.

Loreto: Fide Moldenke, Phytologia 4: 451. 1953. To Guiana.

Petrea Martiana Schauer in DC. Prodr. 11: 620. 1847; 164.

Branchlets, stout petioles (mostly 1-2 cm. long), peduncles (to

about 1 dm. long), pedicels (9 mm. long) and rachis more or less

densely fulvous puberulent; leaf -scar bases 2-4 mm. long; leaves en-

tire, broadly elliptic, usually acuminate, acute or rounded at base,

about 1-2 dm. long, rather more than half as wide, sometimes larger,

firm-membranous or subcoriaceous, sometimes rather lustrous above,

densely asperulous both sides and somewhat hirsutulous on the prom-

inent venation beneath, at maturity conspicuously bullate; racemes

solitary, but approximate toward branchlet tip, about 2.5-4.5 dm.

long; calyx tube 4-5 mm. long and as broad at apex, 15-costate,

densely short-pubescent, the thin-membranous obovate rounded or

acute glabrous lobes 14-17 mm. long, 3 mm. wide at base, to 1 cm.

wide above, the crown-like crest about 1.5 mm. high, the lingulate

FLORA OF PERU 665

obtuse lobes 1 mm. long; corolla tube 5 mm. long, upper fourth

spreading puberulent without, long-pubescent within to just below

stamens, the lobes all rotund, puberulent; calyx in fruit to 7 mm.

long, lobes to 3 cm. long, callose. — Liana, the flowers violet (Ducke).

Illustrated, Mart. Fl. Bras. 9, pi 46.

Loreto: Without data, Fox 96. Amazon; Brazil.

Petrea maynensis Huber, Bol. Mus. Paraense 4: 602. 1906; 199.

Resembles most perhaps P. pubescens, the leaves similar in form

and rarely opposite but rather lightly asperulous, not at all pubes-

cent, the early pubescent branchlets soon glabrate, petioles puberu-

lent, 4-10 mm. long, pedicels (about 9 mm. long) and calyx glabrous;

leaves to about 2 dm. long, nearly half as wide, often acute both ends,

lateral nerves (especially) prominent beneath; racemes to 4.5 dm.

long; peduncles very short if apparent; sterile bractlets persisting;

calyx tube 3.5-4 mm. long, 10-ribbed, the lobes oblong, 12.5-14 mm.

long, 2.5-4 mm. wide, the crest 5-cleft to base, the acute segments

3 mm. long, half as wide; corolla tube 9 mm. long, to 7 mm. wide at

apex, short-pilose (as lobes) from base, sparsely pubescent above

within, largest lobe 7.5 mm. long; anthers glabrous, connective erect;

fruiting calyx to 5.5 mm. long, 8 mm. wide, lobes about 2.5 cm. long,

crest converging. — As in P. volubilis L. there are often axillary fas-

cicles of bractlets. P. longifolia Mold. I.e. 197, origin unknown, is

similar but leaves to 3 dm. long, short acuminate, glabrate, and an-

ther connective recurved. Calyx nearly white, corolla violet (Ducke) ;

calyx blue, corolla center white (Kuhlmann).

San Martin: Tocache, Poeppig 1970. — Huanuco: Fide Moldenke.

— Loreto: Between Contamana and Canchahuaya, (Huber 1489,

type). Rio Ucayali, (Tessman 3960). To Colombia; Brazil; Bolivia.

Petrea nitidula Mold. Repert. Sp. Nov. 43: 168. 1938.

Rather similar to P. Martiana but less pubescent, the branchlets

and petioles (5-12 mm. long) more or less minutely puberulent, the

leaves somewhat lustrous both sides, glabrous and smooth above,

rather densely scabrous and rugulose only beneath, the pedicels

(8 mm. long, 2 cm. long in fruit) puberulent; calyx 10-ribbed, the

lobes oblong, to 2 cm. long, 3-5 mm. wide, the crest cleft to base or

early weakly united, about 1.2 mm. high; corolla tube 16 mm. long,

the lobes oblong, sparsely puberulent; calyx tube in fruit 6 mm.

long, lobes to 2.8 cm. long, crest converging. — Corolla said to be

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

blue, the calyx paler; has been confused with P. Martiana Schauer

(author) ; cf ., however, smooth specimens of P. peruviana.

San Martin: (Poeppig). Amazonian Brazil.

Petrea peruviana Mold. Repert. Sp. Nov. 43: 206. 1938.

Branchlets, opposite leaves, petioles (1-1.5 cm. long), and termi-

nal lax inflorescence including the calyx tube minutely or obscurely

puberulent; leaves entire, broadly elliptic, obtuse both ends or rounded

basally, emarginate apically, 6-18 cm. long, 3-10.5 cm. wide, firm-

membranous, faintly scabrous or glabrous both sides, the 8-15 pairs

of secondary nerves prominent, especially beneath; racemes to 3 dm.

long or longer, developed after the leaves; pedicels about 2 cm. long,

often with a medial bract; calyx tube 6-6.5 mm. long, nearly as wide

at top, smooth but weakly 10-ribbed, the membranous obovate acute

lobes 16-21.5 mm. long, 6-8 mm. wide above, crest cleft to base, the

segments obtuse, 1.5 mm. long; corolla tube 15-18 mm. long, am-

pliate to about 10-14 mm. across, only the upper fourth puberulent,

long-pubescent within, the lobes broadly elliptic, the largest 8-11 mm.

long, puberulent all over; anther connective pilosulous; fruiting calyx

lobes to 3.5 cm. long, 1.5 cm. wide, the very callose crest converging.

— With P. nitidula seems doubtfully distinct from P. maynensis, con-

sidering some variability. The var. acuminata Mold, has leaves to

23 cm. long, long- or short-acuminate, usually acute at base and

calyx crest merely sinuate. Collectors noted it as a shrub or a liana,

flowers dark blue or violet; may become scandent above in age or

when support is at hand.

Loreto: Iquitos and vicinity, King 637, type; 170; (Mexia, type,

var.); Williams 8106; Mexia 6498 (var. acuminata). Rio Nanay,

Williams 647 (var.) ; 690. — Puno : Lake Titicaca, (Meyeri) . Ecuador;

Colombia. "Sanango-sacha" (Williams).

Petrea pubescens Turcz. Bull. Soc. Imp. Nat. Mosc. 36, pt. 2:

211. 1863; 170.

Slender branchlets early fulvous or yellowish hirsutulous as the

petioles — these opposite or often 3-4-verticillate, 1-5 mm. long —

and young leaves that ordinarily precede the blue flowers; leaves

elliptic or slightly obovate, entire or repand-sinuate, attenuate to

the acute to rounded or subcordate base, rounded or obtuse and

usually mucronulate, the largest 1.5 dm. long, at least about a third

as wide or even larger, asperulous both sides, finally firm membranous

and bullate, the lateral nerves 7-15 pairs; racemes many, crowded

apically, to 2 dm. long or longer, nutant; pedicels and obconic calyx

FLORA OF PERU 667

tube strongly long-pubescent, the latter 3-3.5 mm. long, the glabrous

obovate lobes 12-13 mm. long, 3-5 mm. wide above, the acute ciliate

crest lobes 1.5 mm. long; corolla tube 6-7 mm. long, puberulent

above, long-pubescent within; anthers glabrous; calyx in fruit to

6 mm. long, lobes to 22 mm. long, callose crest converging, dorsally

setose-hispid. — The var. Klugii Mold., I.e. 172, has the mature calyx

about 3 mm. long and densely divaricate-tomentose. Shrub or tree,

recorded to 20 meters tall, or sometimes said to be a liana. F.M.

Neg. 34291.

San Martin: Chazuta, Klug 4155 (type, var.; det. Standley, P. ar-

borea HBK.).— Rio Acre: Ule 9722. Colombia; Venezuela.

Petrea volubilis L. Sp. PI. 626. 1753; 32.

Liana or early erect, lenticellate subterete branches and branch-

lets subappressed or early spreading pubescent as petioles (4-13 mm.

long), peduncles (to 7 mm. long, or these puberulent), and calyces;

leaf-scars prominent, corky based; leaves elliptic, usually entire, gen-

erally acute or obtusely narrowed at base, acute or shortly acuminate,

to about 2 dm. long, half as wide or proportionately much smaller,

firm-membranous, not bullate, harshly asperulous both sides, often

obscurely pubescent or puberulent especially on the 8-17 prominent

lateral nerves beneath; racemes axillary, erect to nutant, often 1-

3 dm. long, pedicels 8 mm. long, calyx 3 mm. long, not costate, densely

hirsute or tomentose, the 5 membranous oblong lobes 13-18 mm.

long, about a third as wide, glabrous, the crest of 5 membranous

ovate acute sparsely ciliate lobes about 1 mm. long and wide; corolla

tube 6-8 mm. long, 4-5 mm. wide at apex, puberulent above and

within where villous among the stamens, the largest lobe 5-6.5 mm.

long; fruiting calyx to 4 mm. long, the rigid lobes to 22 mm. long,

7 mm. wide, the callose crest enclosing the mouth. — Has been con-

fused particularly with P. racemosa Nees of Brazil (Moldenke) with

appressed sericeous calyx. Cultivated at least at Lima and interest-

ing in contrast to the native species; there is a forma albiflora Stand-

ley. Type from Veracruz, Mexico. Illustrated, Bot. Mag. 17: pi. 628.

Lima : Near Lima, Soukup 2903. Central America; Mexico; West

Indies.

10. CITHAREXYLUM [Juss.] L.

Rauwolfia R. & P. Fl. Peruv. 2: 26, pi 152. 1799, not L.

Reference: Moldenke, Phytologia 6: 242-256 (key), 262-320, 332-

368, 383-432. 1958; I.e. 448-505. 1959; I.e. 7: 7-77. 1960.

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

Glabrous trees or shrubs, rarely scandent, or tomentose above

with simple trichomes, now and then spinose in the axils, the leaves

sometimes spinose-dentate, usually biglandular at base. Racemes or

spikes axillary and terminal, elongate or short, the bracts of the few-

many laxly borne flowers small. Calyx tubular-campanulate, trun-

cately 5-dentate or shortly and broadly 5-lobed, cupulately accrescent

or spreading beneath the fleshy drupe, this with 2 (1) usually 2-celled

pyrenes (1-celled, C. caudatum). Corolla limb 5- (rarely 4- or 6-)

parted, the lobes subequal. Stamens normally 4, didynamous, medial

or higher, anthers ovate or sagittate, the cells parallel. Ovary per-

fectly or imperfectly 4-celled, the single ovules lateral. Stigma ob-

scurely 2-cleft or lobate. — Besides the following, Weberbauer 5916,

possibly from Piura or Ayacucho, has been listed under an unpub-

lished name by Loesener (not seen by me) ; Moldenke, I.e. 256, lists a

species "peruvianum" as a species of Aloysia from Ayacucho, not

published (1958).

Moldenke has provided an artificial but useful key, and a supple-

mentary one, prepared previously only for the Peruvian species, also

was devised, obviously, expediently. So long as variation-range of

characters accepted as indicative of specific values is unknown, no

direct and simple key is possible; leaf -texture, dentation, and pubes-

cence may prove to be less stable or significant when the shrubs are

better known.

KEY TO CITHAREXYLUM (after Moldenke)

Spines present.

Branches, branchlets acutely angled, 4-margined; nodes strongly

annulate.

Leaves densely punctate beneath, mostly serrate . . C. Herrerae.

Leaves not punctate, entire C. andinum.

Branches, branchlets mostly obtusely angled, not margined; nodes

usually obscurely annulate.

Petioles 1-3 mm. long; leaves to 1 cm. wide. . .C. Weberbaueri.

Petioles 3-15 mm. long; leaves mostly much wider.

C. megacanthum, C. flexuosum.

Spines absent.

Leaves ternate.

Leaves sharply serrate, 2-3.5 cm. wide C. quercifolium.

Leaves entire, mostly much wider C. Poeppigii.

Leaves opposite or the younger approximate.

FLORA OF PERU 669

Leaves dentate, more or less coriaceous.

Racemes to 4 cm. long, mostly much less.

Leaves merely subcoriaceous.

Branchlets verruculose (densely lenticellate) ; leaves 7-

12 mm. wide, drying dark C. pachyphyllum.

Branchlets smooth; leaves 8-40 mm. wide, not discolor-

ing C. ilicifolium.

Leaves firm- or leathery-coriaceous.

Leaves rigidly heavy-coriaceous, often discolored when

dried; racemes 1-3.5 cm. long C. dentatum.

Leaves firm-coriaceous, not discoloring; racemes 1-5-

flowered.

Leaf surface deeply punctate, margins subentire.

C. punctatum.

Leaf surface beneath epunctate, margins coarsely ser-

rate C. argutedentatum.

Racemes 4-25 cm. long.

Leaves acute, not revolute C. laurifolium.

Leaves obtuse or emarginate, revolute C. caudatum.

Leaves entire.

Leaves glabrous unless puberulent-punctate or midrib pilosu-

lous.

Racemes attenuated, few-flowered.

Both leaf surfaces deeply punctate C. punctatum.

Upper leaf surface epunctate.

Mature leaves discolored when dried, reticulation ob-

scure above C. pachyphyllum.

Mature leaves usually not discoloring, reticulation ob-

vious above C. ilicifolium.

Racemes elongate, many-flowered.

Leaves thin-membranous, fragile C. chartaceum.

Leaves chartaceous to coriaceous.

Racemes 1-4.5 cm. long; leaves rarely to 7 X 4 cm.

long, petioles 2-5 mm. long C. ilicifolium.

Racemes and petioles mostly longer; leaves larger.

Vein reticulation prominent above . . . C. reticulatum.

Vein reticulation obscure or faint above.

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

Flowers sessile C. laurifolium.

Flowers distinctly pedicellate.

Nutlets 2-celled C. suberosum.

Nutlets 1-celled C. caudatum.

Leaves puberulent beneath.

Leaves ample, 7.5-12 cm. wide C. Poeppigii.

Leaves rarely 7 cm. wide.

Leaves chartaceous C. Kobuskianum.

Leaves thin-membranous C. chartaceum.

SUPPLEMENTARY KEY TO CITHAREXYLUM

Leaves (at least many of them) 5-10 cm. long or longer.

Nutlets 1-celled; slender glabrous shrub-tree except tomentose co-

rolla, this 5-8 mm. long, tube to twice as long as 3 mm. long

subtruncate calyx; racemes often 1 dm. long or longer, pedi-

cels 2 mm. long C. caudatum.

Nutlets 2-celled; characters in part not as above.

Calyx early subtruncate, 5-denticulate; leaves entire or mostly

except C. quercifolium.

Leaves membranous; basal glands none or minute.

C. chartaceum.

Leaves coriaceous; basal glands 2, large C. Poeppigii.

Leaves coriaceous or siibcoriaceous.

Basal glands none (always?).

C. quercifolium, C. reticulatum.

Basal or sub-basal glands several C. suberosum.

Calyx non-lobed; leaves in part or obscurely serrulate; glands

none C. laurifolium.

Leaves at most about 5 cm. long, mostly or all smaller.

Branches (ex. char.) espinose.

Leaves coarsely dentate below the middle . . . C. argutedentatum.

Leaves at least in part subentire or more or less dentate above

the middle.

Racemes short; leaves coriaceous or subcoriaceous.

Leaves compactly punctate both sides C. punctatum.

Leaves obscurely or not conspicuously nor closely punctate

at least above.

FLORA OF PERU 671

Corolla lobes and throat pilose; leaves subcoriaceous.

C. ilicifolium.

Corolla glabrous except throat.

Leaves rigid; calyx 5-7-dentate or -cleft. .C. dentatum.

Leaves flexible; calyx 5-denticulate . .C. pachyphyllum.

Racemes to 6.5 cm. long; leaves membranous.

C. Kobuskianum, C. megacanthum.

Branches (ex char.) more or less spinose.

Racemes 2.5-6.5 cm. long; leaves obovate, rounded at apex.

C. Kobuskianum, C. megacanthum.

Racemes short, 2-several-flowered; leaves obscurely or not obo-

vate.

Branches, branchlets subalately quadrate, glabrous.

C. Herrerae.

Branches subterete, branchlets early acutely quadrate, puber-

ulent.

Calyx teeth in types 5 C. andinum, C. flexuosum.

Calyx teeth in type 4 but obscure C. Weberbaueri.

Citharexylum andinum Mold. Repert. Sp. Nov. 37: 217. 1934;

288.

Branches, branchlets and slender shoots glabrous, lustrous, spinose,

acutely tetragonal, the younger often 4-marginate; nodes strongly

annulate; spines slender, 6-16 mm. long, with 3 sterigma-like sub-

apical scars; leaf -scars very corky, suberose; petioles slender, 3-6 mm.

long; leaves oblong or elliptic, acute at base, rounded or subemargi-

nate at apex, about 1- nearly 3 cm. long, 5-13 mm. wide, entire,

subchartaceous, glabrate, not glanduliferous, the 4-5 arcuate lateral

nerves scarcely impressed above nor prominent beneath; racemes

2-4-flowered, to 1 cm. long; peduncles 1-4 mm. long; pedicels 1-3 mm.

long, puberulent or glabrate as the calyx, this 3.3 mm. long, distinctly

5-costate, minutely 5-apiculate; corolla tube 3.6 mm. long, tomentose

only within at throat, the oblongish lobes rounded apically, 2 mm.

long; stigma sub-bilobed; calyx in fruit cupulate, to 5 mm. wide,

3 mm. long; fruit subglobose, about 8 mm. thick, drying bisulcate.

—Closely related to C. Weberbaueri Hayek and C. flexuosum (R. & P.)

D. Don, but differs from both in the acutely angled branchlets and

glabrate leaves. Has been confused, also, with Duranta Plumieri

Jacq. but the twigs are not covered with approximate sterigmata

(author); from range, the Scolnik specimen would be more usually

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

referable to the typically northern shrub of Ruiz and Pavon. F.M.

Neg. 28393.

Cajamarca: Scolnik 1302 (det. Moldenke). Bolivia.

Citharexylum argutedentatum Mold. Repert. Sp. Nov. 37:

218. 1934; 289.

Shrub, glabrous except the obscurely furfuraceous puberulent

branches and branchlets, these slender, strongly sarmentose; decur-

rent lines (sterigmata) incrassate, 1-2 mm. long; nodes indistinctly

annulate; petioles 1-2 mm. long; leaves opposite, rotund-elliptic,

abruptly attenuate at base and shortly prolonged into petioles, 12-

19 mm. long, 9-18 mm. wide, below the middle coarsely dentate

(teeth divergent, often recurved), coriaceous, lustrous both sides,

glandular-punctate beneath, the midnerve there prominent, second-

ary veins 3 or 4, reticulation obscure; racemes only 1-3-flowered,

when 1-flowered appearing axillary in upper leaves; fruiting calyx

3-4 mm. long and thick, 4- or 5-dentate or -lobate; fruit subglobose

or oblong, 3-4 mm. long, lustrous, drying black and apiculate at apex.

— Related to C. ilicifolium HBK. and to C. pachyphyllum Mold, in

its very thick-coriaceous, waxy-nitid holly-like leaves; the greatly

reduced inflorescences are remarkable (author).

Cuzco: Ollantaitambo, (Cook & Gilbert 719; 745, type). — Puno:

Rio Limbani, Goodspeed Exped. 30464; (Metcalf 30464}- "Tasta"

or "tosta."

Citharexylum caudatum L. Sp. PI. ed. 2: 872. 1763; 299.

Glabrous with terete striate branches, the leaves laterally glandular

at attenuate base, and terminal or axillary erect racemes of fragrant

flowers; leaves elliptic or oblong, subacute, obtuse or emarginate, a

dm. long or longer, about half as wide, coriaceous to membranous,

lustrous above, subrevolute; calyx 3-4.5 mm. long, subtruncate, ob-

tusely 5-dentate, the apically puberulent corolla-tube about twice as

long; fruit lustrous, tardily black, 6-12 mm. long, 5-10 mm. wide. —

Notable in the 1-celled nutlets (not noted by Moldenke, I.e. 251); the

pedicels in flower are usually 2 (-2.5) mm. long, leaves prominently

and openly reticulate-veined. Cultivated in Chile, Cuba, and Peru

(Raimondi 11906, fide Moldenke). Moldenke notes, with reason,

that the Peruvian collections must have been cultivated or persist-

ing in abandoned areas, which explains Raimondi's observation

(quoted by Moldenke, 305), "indigenous at Chanchamayo."

FLORA OF PERU 673

Junin: Pampa hermosa, Montana de Pangao, (Raimondi 12507,

det. Moldenke). Chanchamayo, (Raimondi 12203). Mexico; West

Indies.

Citharexylum chartaceum Mold. Repert. Sp. Nov. 37: 219.

1934; 310.

Glabrous or glabrescent, only a light puberulence on leaves be-

neath and peduncles, these 1.5-3 cm. long; branchlets and shoots

acutely tetragonous and marginate; nodes obscurely or not annulate;

corky elevations (sterigmata) at tip of branches thick, suberose, to

4 mm. long or wide; petioles opposite, slender, 2-13 mm. long; leaves

oblong-elliptic, broadly elliptic or subobovate, 5-15.5 cm. long, 3.2-

7.2 cm. wide, acute or acuminate to base where sometimes 1 or 2

minute glands, acute or shortly acuminate, thin-chartaceous; mid-

nerve slender, secondary veins 5-8, the reticulation fine; racemes

terminal or on axillary branchlets, mostly nutant, 9-18 cm. long;

bractlets often 2, linear, to 4 mm. long; pedicels obsolete or to 1 mm.

long; calyx translucent, about 3.1 mm. long, 2.8 mm. across, minutely

5-denticulate; corolla tube 4.4 mm. long, densely long pilose within

at throat; lobes oblong-obovate, 3.2 mm. long, 2.3 mm. wide, pubes-

cent within; stamens set 2 mm. below throat; style 1.5 mm. long,

stigma minutely bilobate. — Related to C. quitense Spreng. and re-

ferred in herbaria to C. molle HBK.; the thin leaves with delicate

but conspicuous and yet not at all elevated venation on the lower

surface, and the lanate corolla-throat, characterize it well (author).

Tree or shrub 5-8 meters tall, in deciduous brushwood.

Tumbez: East of Hacienda Chicama, 1,000 meters, Weberbauer

7633, type; 7678. Ecuador.

Citharexylum dentatum [Tafalla] D. Don, Edinb. New Phil.

Journ. 11: 237. 1831; 320, 332 (descriptions).

Much branched, unarmed, known to attain 5 meters, the more or

less acutely quadrate branchlets and short terminal racemes puber-

lent; nodes rather obscurely annulate; leaf-scars large, borne on stout

corky sterigmata 1-2.5 mm. long; petioles thick, 1-10 mm. long;

leaves crowded, obovate-oblong or elliptic, attenuate to base, ob-

tusely rounded or acute at apex, about 2.5 (-3) cm. long, 7-19 mm.

wide, entire or acutely or spinose dentate, rigid coriaceous, revolute,

punctate and somewhat scabrous both sides but not glanduliferous,

lustrous above, paler and opaque beneath ; nerves 4-6 pairs, obscure

above, rather conspicuous as sometimes veins beneath; racemes 1-

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

3.5 cm. long, few- to many-flowered, pedicels obsolete or to 4.5 mm.

long in fruit, indurate as fruiting calyx; bracts setaceous or linear, to

3 mm. long; calyx 5-7-nerved, acutely 5-7 dentate or cleft, nearly

4 mm. long, in fruit to 5 mm. long, 8 mm. wide; corolla glabrous ex-

cept for the villous throat, the tube exserted; fruits about 1 cm. in

diameter, lustrous, black even when fresh (Moldenke). — Apparently

is the correct name for at least the individuals cited under C. ilici-

folium HBK. and C. pachyphyllum Mold. A 5-meter tree, the berries

said to be used for making ink (Stork & Hortoh). Type by Ruiz and

Pavon, perhaps from near Tarma, the name by Tafalla in part of the

expedition's collections but as a species of Rauwolfia. F.M. Neg.

17592.

Junin(?) : Ruiz & Pavdn, type. — Huancavelica: Salcabamba, 3,500

meters, Stork & Horton 10336 (det. Standley). — Ayacucho: Above

Quinua, 3,400 meters, Weberbauer 5537.

Citharexylum flexuosum (R. & P.) D. Don, Edinb. New Phil.

Journ. 11: 237. 1831; 349. Rauwolfia flexuosa R. & P. Fl. Peruv. 2:

26, pi. 152, fig. a. 1799. R. macrophylla R. & P. I.e. fig. b. C. retusum

D. Don, I.e. C. spinosum HBK. Nov. Gen. & Sp. 2: 256. 1818; 610.

Scypharia tetragona Miers, Contrib. 1: 302. 1851-1861, fide Weber-

bauer, Field Mus. Bot. 8: 84. 1930.

Branches obtusely or sometimes acutely tetragonal, the short

stout branchlets modified into spines to 5 cm. long, the flexuose

younger also spinescent and as the leaves beneath more or less pubes-

cent; leaf -scars on corky sterigmata 1.5-3.5 mm. long; leaves oblong

or subelliptic, sometimes obovate, attenuate to slender petioles (3-15

mm. long), obtuse to emarginate, mucronulate, entire, (2) 3-5 (6)

cm. long, (1) 1.5-2.5 (3) cm. wide, membranous, finely reticulate-

veined, nerves slender, 5-7 pairs, finally lustrous and glabrate above,

subtomentose to glabrate beneath; racemes terminal on axillary

branchlets, few-flowered, laxly short-pubescent; pedicels filiform,

1 mm. long; calyx campanulate, nearly 4 mm. long, membranous,

5-nerved, obtusely 5-dentate, subequaling the broad tube of the

corolla, this with rounded ciliate lobes pilose as the corolla throat;

anthers sagittate; calyx in fruit 2.5-3 mm. long, 4-6.5 mm. wide,

early subtruncate rim finally splitting into rounded lobes; fruit pur-

plish-black or black, about 8 mm. long and wide, glabrate. — Often

becoming completely or essentially glabrous except the puberulent

calyx and white or paler corolla within. The var. subglabrum Mold.,

Repert. Sp. Nov. 37: 222. 1934, apparently described the mature

FLORA OF PERU 675

leaves of a form which proved to be Lycium subglabrum Mold.

(author). The greenish-cream flowers are delightfully fragrant in

the morning and the branches are used for small crosses, the spines

serving as arms (Ruiz & Pavon). F.M. Negs. 29685; 17601 (C.

spinosum).

Cajamarca: Yanaguanga, Bonpland (type, C. spinosum). — An-

cash : (Raimondi) . — Lima : Rio Blanco, Killip & Smith 21 679. Matu-

cana, 464- San Rafael, 2409. — Huanuco: Near Huanuco, 3523; Ruiz

& Pavdn, type. Llata, 2279. Ambo, 3173; 2438.— Arequipa: Mol-

lendo, 600 meters, (Guenther & Buchtien 129; 130, det. Bruns, glabrate

form). Atiquipa, Tafalla (type, R. macrophylla). Rock slides near

Chala, Worth & Morrison 15700. "Turucasa," i.e. spina obtusa

(Ruiz & Pavon); "chama" (Tafalla), "choloquillo simarron."

Citharexylum Herrerae Mansf. Notizbl. Bot. Gart. Berlin 9:

469. 1925; 401.

Branchlets distinctly and acutely quadrate or even quadrialate,

spiny, green, early somewhat puberulent as the young leaves; spines

tetragonal, often branching; sterigmata to 3 mm. long; leaves oblong-

elliptic-lanceolate, decurrent into the 1-4 mm. long petiole, rounded

to acute, mucronulate, 1-2.5 cm. long, 3-9 mm. wide, entire or serrate

below the middle, chartaceous, soon quite glabrous, plane above,

punctate beneath, the 4-6 nerves scarcely impressed; racemes axil-

lary, to 2.5 cm. long, 4 cm. in fruit; pedicels in fruit to 1 mm. long;

calyx 3.5 mm. long, shortly 5-dentate, in fruit to 6 mm. broad, sub-

entire or lobate, glabrous; corolla 5 mm. long, lobes 1 mm. long, these

within and throat pilose; filaments inserted medially; style 2 mm.

long, the stigma sub-bilobed; fruit oblong-globose, 8 mm. thick, to

11 mm. long, lustrous, red when fresh. — Distinguished from other

spinescent species by the glabrous leaves, subalate and green branch-

lets; also, axillary shoots are frequent and persistent; they even

puncture flowers and fruits (author); the uppermost branches are

modified into spike-like thorns (Moldenke).

Cultivated; the beautiful red fruiting branches used for orna-

mentation of the nacimientos (Herrera). Tree, 3-5 meters tall (Pen-

nell). F.M. Neg. 17595.

Apurimac: (Vargas 8753; 9137, both det. Moldenke). Saxaihua-

man, Herrera 677, type. — Cuzco: Cuzco Valley, 3,200 meters, Her-

rera 1464a; Soukup 322; (Cook & Gilbert 1888; Ferreyra 9856, det.

Moldenke). San Sebastian, Pennell 13606. "Huariruru."

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

Citharexylum ilicifolium HBK. Nov. Gen. & Sp. 2: 256. 1818;

411.

Similar to C. dentatum; leaves elliptic-oblong, obtuse, 2-7 cm.

long, 1-4 cm. wide, entire or spinose-dentate, glabrous and finally

lustrous above and subcoriaceous, punctate and early pubescent

beneath; nerves 7-10 pairs, reticulation rather prominent both sides;

calyx 5-nerved, minutely 5-dentate, pubescent as pedicels; corolla

tube little exserted, white, the lobes and throat pilose; fruiting calyx

(Moldenke) cupuliform, about 3 mm. long, 6 mm. wide, puberulent

or hirtellous, the 5 conspicuous lobes at least apiculate, often sub-

serrate; fruit finally black or black-purple, 6 mm. long and wide. —

Leaves in type about 3 cm. long, half as wide, the veins densely

reticulate or prominent beneath, somewhat attenuate to base; on

younger branchlets to 5.5 cm. long, often more or less repand-dentate.

Perhaps not in Peru, at least in typical form, but Moldenke refers to

it collections by Gay and by Dombey, without, however, known

localities; the similar Peruvian shrub could prove to be a part,

genetically considered; I have not seen the Weberbauer collection

in the final preparation of this account. F.M. Neg. 17593.

Ancash: Yungay, Weberbauer 3285 (det. Hayek). Ecuador to

Bolivia. "Zitac" (Ecuador).

Citharexylum Kobuskianum Mold. Phytologia 1: 441. 1940;

424.

Younger branchlets, chartaceous leaves, especially beneath, and

axillary racemiform inflorescences (these to 6.5 cm. long), including

the herbaceous calyx more or less puberulent, the leaves above very

sparsely; nodes enlarged; leaf -scars prominent, corky; petioles very

slender, 2-5 mm. long; leaves entire, oblong-elliptic, acutely or

shortly acuminate at base, acute and shortly apiculate at apex,

2.5-5 cm. long, 1.5-2.2 cm. wide, the 7-10 secondary nerves slender,

obscure above; fruiting pedicels about 3 mm. long; fruit 5 mm. long,

4 mm. thick, glabrous, fleshy. — Named for C. E. Kobuski, whose

monograph of the genus Priva is excellent (author); the collector's

name is correctly written Jelsky (Moldenke); in lit. (and on the

labels), however, as here.

Cajamarca: Chota, (Jelski, type; Raimondi 6303). — La Libertad:

(L6pez Miranda 648, det. Moldenke).

Citharexylum laurifolium Hayek, Bot. Jahrb. 42: 170. 1908;

449.

FLORA OF PERU 677

Quite glabrous lustrous shrub, the somewhat tetragonal branches

unarmed, subangled, the lanceolate eglandular (type) leaves nar-

rowed to base, acute, 5-13 cm. long, 3-5.5 cm. wide, entire or mu-

cronulate-serrate above, coriaceous, lustrous both sides, nerves slen-

der, 4-6, veins finely reticulate; racemes terminal, erect, very strict,

several cm. to 2 dm. long or longer, the rachis stout, the many often

loosely borne flowers subsessile; calyx in fruit irregularly lobed or

bilobed, incrassate, to 6 mm. long, 7 mm. wide; fruit ovoid, about

8 mm. long, 5 mm. wide, black in drying. — Corollas lacking in type

which resembles C. caudatum L. but leaves not revolute, rachis excep-

tionally thick and calyx 2-lobed (author) ; it may be conspecific with

C. reticulatum HBK. (Moldenke) to which in manuscript I had

referred it; it may, however, as so often with northern species, be a

southern development. Type a shrub about 1 meter tall, the flowers

greenish (Weberbauer). F.M. Neg. 17594.

Puno: Sandia to Cuyocuyo, 2,500 meters, Weberbauer 873, type.

Bolivia.

Citharexylum megacanthum Rusby, Bull. N. Y. Bot. Gard.

8: 115. 1912.

Glabrate, the branches and spines stout, the former sulcate, the

latter divaricate, straight, pungent; petioles 3-5 mm. long; leaves

obovate, abruptly contracted at base, rounded at apex, 1.2-2.5 cm.

long, 8-16 mm. wide, revolute, bullate above, the 4-5 pairs of nerves

prominent beneath; racemes axillary, 2.5-5 cm. long in fruit, the

short stout pedicels recurving; calyx 6 mm. broad in fruit, the lobes

unequal; fruit subglobose, deep purple, 6-7 mm. in diameter. — A

bush, 3 meters tall. This was originally included as a synonym of

C. flexuosum (R. & P.) D. Don.

Arequipa: Mollendo, (R. L. Williams 2544, type).

Citharexylum pachyphyllum Mold. Repert. Sp. Nov. 37: 232.

1934; 491.

Rather stoutly branched densely leafy essentially glabrous shrub,

the obtusely tetragonous branches and branchlets early obscurely

puberulent or glabrate, often verruculose, the nodes (1-3 cm. distant)

faintly annulate; sterigmata only 1-2 mm. long; petioles 2-3 mm.

long; leaves oblong or elliptic, often broadly, in type, or subrotund,

subacute and eglandular at base, rounded but mucronate at tip, 1.5-

2.5 cm. long, 7-15 mm. wide, coriaceous, subentire or remotely and

minutely denticulate (2-3 repand-denticulate each edge in type),

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

coriaceous, densely punctate beneath, glands lacking, lustrous both

sides, nerves 3-5 pairs, the veins obscure; racemes 1-1.5 cm. long;

pedicels to 1 mm. long; bracts none; calyx obconic, minutely 5-

dentate, to 4 mm. long, 7 mm. wide, and in fruit unevenly lobed;

corolla tube 4.3 mm. long, tomentose within at throat, the 4-5

rounded lobes 1-3 mm. below the throat, subsessile; style stout, stig-

ma lobes indistinct; ovary imperfectly 4-celled. — Description of

flowers apparently taken by Moldenke from the more oblong-leaved

Ayacucho specimen. Leaves of Metcalf specimens suborbicular,

scarcely 1 cm. long; fruit to 11 mm. long, 13 mm. wide, deep purple,

very firm. Related to C. ilicifolium HBK., C. dentatum D. Don, C.

punctatum Greenm. (author), the type a shrub to 2 meters tall. I had

referred my collection (the type) to the shrub of HBK., that perhaps,

at least typically, only in Ecuador; it is the earliest name among

these similar shrubs, if they are not specifically distinct; the Metcalf

specimen is toward C. punctatum. Both Asplund 11496 and Mathews

1021 were referred here by Moldenke but localities were not given

by him.

Junin: Shrubby canyon, about 3,000 meters, Huariaca, 3090,

type. — Ayacucho: Below Tambo, Weberbauer 5565. — Puno: South of

Limbani, Metcalf 30 W '4 (det. Moldenke).

Citharexylum Poeppigii Walp. Repert. 4: 76. 1845; 498.

Subterete branches and branchlets as the rachis of the axillary and

terminal racemes or panicle lanuginose or glabrate; leaves opposite or

ternate, entire, subovate-oblong, abruptly decurrent into 3-4 cm.

long petiole, normally acuminate, about 2 dm. long, chartaceous or

subcoriaceous, subnitidulous above, opaque and especially on nerves

lanate beneath; nerves 9-14 pairs, reticulation dense; racemes to

1.5 dm. long; flowers sessile or subsessile, fragrant, to 18 mm. long,

glabrous; calyx obconic, 3 mm. long, not nerved, obtusely dentate,

membranous, finally puberulent; corolla tube 6 mm. long, throat

lanate; drupes ovoid-oblong. — Conspicuously marked by the pair of

large glands at the base of leaves. The var. margaritaceum Poepp. ex

Mold., Repert. Sp. Nov. 37: 233. 1934, has ovate-elliptic rarely

oblong leaves 2-2.5 dm. long, 7.5- nearly 9 cm. wide, densely

puberulent-lepidote beneath, secondary nerves 12-15, petioles to

2.5 cm. long, racemes to 4.5 dm. long, peduncles to 7 cm. long,

corolla to 7 mm. long (Moldenke); authority of variety published,

incorrectly, as Poeppig and Moldenke. Recorded as a tree to 18

meters tall. Similar species (?) to be expected as collected in adjacent

FLORA OF PERU 679

areas are C. Ulei Mold. I.e. 237 (Phytologia, 66) and C. amazonicum

Mold. I.e. 216 (Phytologia, 286), the latter with petioles 5-6.5 cm.

long, leaves membranous-chartaceous, the former with mature flowers

nearly 12 mm. long, this with name "cauchillo," occurring to the base

of the Andes in Colombia (Schultes). Type from Rio Acre, Brazil,

so of course also within Peru in one form or another. F.M. Neg.

7876 (var.).

Loreto: Rio Mazan, Josb Schunke 365. Rio Paranapura, Klug

3943 (var.). Yurimaguas, Poeppig 2219 (type, var.); Williams 7183;

Tessmann 3491. Iquitos, Williams 8030. Rio Nanay, Williams 261;

413. Brazil to Colombia. "Mullahuayo," "quiniellia bianco"

(Williams).

Citharexylum punctatum Greenm. Field Mus. Bot. 2: 189.

1907; 504.

Branches terete, the ultimate branchlets subtetragonal, cinereous

puberulent as the short (to 1.5 cm. long) racemes; leaves elliptic-

oblong to subobovate, narrowed to short petiole, usually shortly

acuminate or somewhat pungent, 1-2.5 cm. long, 5-12 mm. wide,

entire or rarely with 1 or 2 teeth, lustrous, subcoriaceous, glabrous

but conspicuously punctate above, minutely so beneath; nerves 3-4

pairs, as the reticulation visible only beneath; calyx tubular-campan-

ulate, 3-4 mm. long, 5-angled, ciliolate, sinuately 5-dentate, in fruit

deeply and acutely 5-lobed, 4 mm. long, wide (Moldenke); corolla

5-6 mm. long, pubescent only within above and in throat, the lobes

oblong, about 2 mm. long; drupe 10-12 mm. long, fleshy, lustrous,

black or purplish when dry. — Distributed as C. ilicifolium HBK.;

differs in smaller entirely glabrous leaves, not spinose-dentate; the

striking impressed punctation is diagnostic (author) ; a gnarled shrub,

to 2 meters tall, of high (3,300-3,800 meters) altitudes.

Puno: Cuyocuyo, Prov. Sandia, Weberbauer 935 (det. Moldenke,

det. Hayek, C. ilicifolium). Bolivia.

Citharexylum quercifolium Hayek, Bot. Jahrb. 42: 169.

1908; 7.

Scandent with smooth subterete branches and remotely spinose

dentate leaves, these ternate (type), oblong-elliptic, narrowed to peti-

ole, obtuse or short-acuminate, (3) 5-8 cm. long, 2-3.5 cm. wide,

entire toward base, more or less coriaceous, glabrous, sublustrous

above, opaque beneath, finely reticulate- veined; racemes erect, 2.5-

0.5 cm. long, subsecund, rachis slightly puberulent; calyx campanu-

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

late, 5-denticulate; corolla fragrant, white, 8 mm. long, exserted. —

Outstanding in the large coriaceous spiny serrate leaves (author);

seems probably an extreme variant of C. reticulatum HBK. F.M.

Neg. 17598.

Cajamarca: Toward Celendin, 3,100 meters, Weberbauer 4248,

type.

Citharexylum reticulatum HBK. Nov. Gen. & Sp. 2: 257.

1817; 13.

Branches (younger) tetragonal, glabrous, striate; leaves decus-

sate-opposite, obovate-elliptic or oblong, attenuate to short petiole,

obtuse, subacuminate or retuse, entire (rarely dentate-spinose), 5 cm.

long or longer, 16-24 mm. wide, coriaceous, strongly reticulate-veined

beneath, lustrous above, puberulent only early on nerves beneath;

racemes terminal, to 4 cm. long, erect, hirtellous; calyx minutely

repand-dentate, 5-nerved; corolla white, exserted, puberulent, to-

mentose villous in throat (Peru?) ; drupe globose, black. — Type from

Rio Catamayo, southern Ecuador. F.M. Neg. 39479.

Junin: Comas, Weberbauer 6604 (det. Moldenke, C. laurifolium).

— Lima: (Ferreyra 10426, fide Moldenke). To Colombia.

Citharexylum suberosum Loes. ex Moldenke, Phytologia 7:

49. 1959.

Resembles in general C. reticulatum (pedicels 1-2 mm. long or

obsolete) but branches and branchlets acutely quadrate, petioles to

12 mm. long, leaves to 13 cm. long, half as wide (often smaller), fre-

quently with a few scattered glands at or near base beneath, the vein

reticulation rather obvious on both surfaces; racemes 7.5-20 cm. long,

rachis slender, glabrous; calyx in fruit indurated, 5.5 mm. long, 7 mm.

across, unevenly and deeply lobed; fruit about 7x6 mm., lustrous.

Apurimac: locality unknown, Weberbauer 5916, type.

Citharexylum Weberbaueri Hayek, Bot. Jahrb. 42: 169. 1908.

Branches short, the spines simple or much branched; leaves entire,

obovate, obtuse, narrowed to short petiole, scarcely 2 cm. long, plane,

early pilose above, glaucescent, densely velutinous beneath, this in-

dument partly deciduous; racemes only 3-5-flowered, the rachis and

peduncles velvety puberulent; calyx campanulate, sub truncate, 4-

denticulate, rather velvety puberulent, little exceeded by the yellow-

ish corolla, this puberulent within. — A meter tall, resembling C.

flexuosum but the leaves not rounded, the racemes few-flowered, the

FLORA OF PERU 681

blossoms small (author) ; this as C. andinum Mold, may be genetically

distinct but the problem is certainly open to question. F.M. Neg.

17603.

Huanuco: Open mixed formation, 2,750 meters, near Chuqui-

bamba, Prov. Huamalies, Weberbauer 3731, type.

11. DURANTA L.

Reference: Schauer in DC. Prodr. 11: 615-616. 1847.

Resembles Citharexylum but, especially, the drupe is nearly or

quite included and consists of four 2-celled 2-seeded pyrenes. Leaves

sometimes verticillate. — The species as proposed seem to be tenuous;

all the characters used expediently in the following key appear to be

variable, a conclusion reached by the often discerning Otto Kuntze,

Rev. Gen. 2: 507. 1891. However, Moldenke, without offering a

solution, has revived the question, which obviously will be answered

only by information other than morphological. At this date (Febru-

ary, 1959), unfortunately, he has not published a revision, so the

following account does not benefit by the detailed contributions he

has made in the case of several other genera.

Calyx, as usually the young leaves beneath, somewhat pubescent.

Corolla tube well-exserted, 1-1.5 cm. long.

Calyx obscurely apiculate, open in fruit, not beaked.

D. Dombeyana.

Calyx teeth in some degree forming a contorted beak in fruit.

Leaves pubescent beneath unless in age.

Calyx teeth after anthesis coiled; leaves typically subacute.

D. Sprucei.

Calyx shortly beaked after anthesis; leaves rounded.

D. triacantha.

Leaves soon glabrous and lustrous; calyx beak short.

D. Skottsbergiana, D. Mutisii.

Corolla tube to about 8 mm. long, little exserted; calyx beak short.

D. peruviana.

Calyx, as leaves, glabrous or glabrate (cf . D. Sprucei, rarely).

Leaves more or less coriaceous and nerves impressed above or

obscure.

Calyx about 2 mm. thick, subcampanulate in D. armata.

Leaf nerves well-impressed.

D. lineata, D. obtusifolia, D. Mutisii.

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

Leaf nerves subimpressed.

Leaves subacute, at least the larger.

Calyx subcampanulate, apiculate D. armata.

Calyx tubular, denticulate D. coriacea.

Leaves all rounded at apex, mucronulate D. rupestris,

Calyx about 3.5-4 mm. thick, more or less campanulate.

Calyx glabrate; corolla tube scarcely exserted . . D. Mandoni.

Calyx somewhat canescent; corolla tube well-exserted.

D. Skottsbergiana, D. Mutisii.

Leaves firm-membranous, the nerves rather raised above.

D. repens.

Duranta armata Mold. Bull. Torrey Club 68: 499. 1941.

Rigid notably spiny branches and branchlets early, as the young

leaves and 1-few-flowered inflorescences, more or less appressed pu-

berulent including the calyces without; spines slender to stout, rigid,

rarely opposite or ternate, 1-3 cm. long or longer; petioles obsolete

to 3 mm. long; leaves fasciculate, oblanceolate or elliptic, about

0.5-3 cm. long, 4-14 mm. wide, the larger acute, the small obtuse,

rounded or often subemarginate, entire or 2-4-dentate above, acute

or attenuate at base, glabrous in age, lustrous above, punctate

beneath; secondary nerves 2-5; peduncles acutely tetragonal, 2-

18 mm. long; pedicels 1 mm. long, to 4 mm. in fruit; bractlets 1-4 mm.

long; calyx campanulate, 3-3.5 mm. long, 5-costate, truncately 5-

apiculate; corolla tube broadly cylindrical, 7 mm. long, glabrous

except at tip, the 5-parted limb 7-10 mm. across, densely puberulent

both sides; fruiting calyx papery, the rim usually persistently puber-

ulent-pilose; fruit tetragonal, 4 mm. long and wide, distinctly 4-lobed

and sulcate in drying, glabrous, lustrous. — Flowers violet-purple to

nearly white, the fruit red (Stork & Horton).

Junin: (Moldenke). — Huancavelica: Mejorada, Stork & Horton

10896. — Ayacucho: Near Ayacucho, Weberbauer 5499; West 3643;

3644- — Apurimac: Pincos, Stork & Horton 10685. — Cuzco: Yucay,

Soukup 738; 739; H err era 1127. Near Cuzco, (Hen era 85, type,

Berlin-Dahlem). "Upa blanca" (Herrera), "tancar" (Stork &

Horton).

Duranta coriacea Hayek, Repert. Nov. Sp. 2: 88. 1906.

Branches subterete, glabrous as the opposite entire little revolute

leaves, these short-attenuate into petiole, elliptic, acute, coriaceous,

FLORA OF PERU 683

concolor, plane, lustrous above, opaque beneath; calyx tube about

5 mm. long, shortly but subulately 5-dentate, slightly tomentulose

marginally; corolla tube 12 mm. long, nearly straight, limb plane,

finely tomentulose; drupe fleshy, 8 mm. in diameter. — After author,

who remarked: differs from D. Plumieri Jacq. (i.e. D. repens L.) in

the lustrous corolla, always entire leaves, and from D. Mutisii L. f.

in the smooth plane leaves. F.M. Neg. 17569.

Cajamarca: Ninabamba, Weberbauer 1*111. Ecuador; Colombia.

Duranta Dombeyana Mold. Bull. Torrey Club 68: 500. 1941.

Strict shrub 2-4 meters tall, the obtusely angled branchlets, leaves

beneath, and inflorescences tomentose; spines axillary, opposite, 5-

7 mm. long; petioles slender, margined above, 4-8 mm. long; leaves

decussate-opposite, often with an additional pair or fascicle on short

axillary branchlet; leaf-scars corky, semicircular; leaves rounded to

acuminate both ends, about 2-9 cm. long, 1-4 cm. wide, chartaceous,

lightly short-pubescent or puberulent above, glabrate in age, rarely

irregularly serrate, the nerves 4-6; peduncles 4-13 mm. long; axillary

inflorescences 3-7 cm. long, the terminal to 2 dm. long, the flowers

secund; pedicels obsolete, to 2.5 mm. long; calyx campanulate, 3-4

mm. long, 5-costate, flavescent-tomentose, the truncate rim obscurely

5-apiculate; corolla tube 13-14 mm. long, puberulent above, limb

8-11 mm. across, puberulent both surfaces; fruiting calyx obvolute,

rostrate, completely enclosing the subglobose glabrous lustrous um-

bonate-rostrate fruit, this to 1 cm. long and wide. — If calyx character

is variable it may be D. Sprucei Briq. My collection from a strict

3-meter tree of river bank, the flowers greenish-yellow; the author

followed my suggestion as to name but made the type an Ecuador

specimen; cf. the negative. F.M. Neg. 29682.

Piura: Quiros Valley, Prov. Ayavaca, Weberbauer 6342. — Huan-

uco: Yanano, 3798. — Junin: Huassahuassi, Ruiz & Pavdn; Dombey.

Ecuador. "Sanacassa," "tantarprieto" (Ruiz & Pavon).

Duranta lineata Hayek, Bot. Jahrb. 42: 170. 1908.

Branches quadrate, spiny, glabrous as the entire coriaceous ellip-

tic or oblong leaves, these narrowed to the short petiole, obtuse or

mucronate, about 2 cm. long, lustrous and impressed lineate-nervose

above, opaque and not strongly nervose beneath; racemes terminal

or lateral, short, the rachis 4-angled, the nutant flowers shortly ped-

icellate; calyx lightly puberulent, about 8 mm. long, denticulate; cor-

olla pale lilac, puberulent without, the tube about 12 mm. long. —

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

Belongs to the group of D. triacantha Juss. but with larger leaves, the

nerves impressed and much larger flowers; D. Benthami Briq. has

larger broadly rounded leaves, and many much longer racemes

(author); may be a local condition due to environment. Type 3

meters tall, frequent in a small ravine (Weberbauer). F.M. Neg.

17570.

Ancash: Yungay, 3,300 meters, (Weberbauer 3286, type).

Duranta Mandoni Mold. Lilloa 5: 390. 1940.

Branchlets acutely 4-angled, early appressed puberulent as the

peduncle and rachis; petioles 1-3 mm. long, canaliculate, sparsely

appressed strigillose; leaves ternate or opposite, often crowded espe-

cially on short twigs, oblong-elliptic, cuneate at base, rounded or

subemarginate, 7-23 mm. long, 5-12 mm. wide, entire but strongly

revolute, subcoriaceous, glabrous above, densely punctate beneath;

venation obscure or the secondaries apparent beneath; racemes axil-

lary, 2-5 mm. long, the flowers secund; calyx about 7 mm. long,

3-4.5 mm. across, lightly puberulent; corolla tube 1 cm. long, gla-

brate, the limb and lobes puberulent. — Tree, 3-4 meters tall, corolla

light purple, fruit yellow (Metcalf), about 12 mm. in diameter.

Ancash: (Moldenke). — Cuzco: Abancay, Vargas & Santander 512

(det. Standley, D. Sprucei). Urubamba, Soukup 61. Marcapata,

3,000 meters, Metcalf 30723. Ecuador; Bolivia.

Duranta Mutisii L. f. Suppl. PI. 291. 1781; 616. D. Plumieri

Jacq. var. strigillosa Schauer in Mart. Fl. Bras. 9: 271. 1851?

Branches subangled, unarmed or spiny; leaves opposite or ternate,

ovate or oblong-elliptic, narrowed to short petiole, acuminate, acute

or obtuse, 4-10 cm. long, 2-5 cm. wide, entire or coarsely serrate from

middle, coriaceous, glabrous or reticulate-veined, rugose, very lus-

trous above, opaque beneath; racemes axillary and terminal, many-

flowered, about a dm. long, laxly pubescent, the violet flowers nutant;

pedicels to 2 mm. long; calyx truncate, subulate-mucronate, the

pubescent tube 6-8 mm. long; corolla 9-12 mm. long, the limb

puberulent. — Spikes sometimes paniculate. Fruit orange when ripe

(West), about 7-10 mm. thick; the variety may not belong here; cf.

D. repens L.

Cajamarca: Hualgayoc, Stork & Horton 10020.— La Libertad:

Angasmarca Valley, 3,300 meters, West 8165. — Junin: Huassahuassi,

Ruiz & Pavon (type, var.). — Apurimac: Pachachaca Valley, 2,000

FLORA OF PERU 685

meters, West 3793,—Cuzco: (Moldenke). Colombia. "Cantar"

(Ruiz & Pavon).

Duranta obtusifolia HBK. Nov. Gen. & Sp. 2: 254. 1818; 616.

Type unarmed, glabrous except the linear bracts, puberulent

campanulate-tubular 5-dentate calyx and the twice as long violet

corolla; leaves opposite, ovate, obtuse, decurrent into petiole 8 mm.

long, coriaceous, reticulate-veined, about 2 cm. long, half as wide;

spikes axillary and terminal; flowers remote, pedicels pendent,

scarcely 2 mm. long; corolla tube two times longer than calyx; drupe

globose-ovoid, enclosed in the calyx. — After author; referred by

Schauer to D. Mutisii L. f .

Cajamarca: Near Jaen, Bonpland, type.

Duranta peruviana Mold. Bull. Torrey Club 68: 501. 1941.

Branches and branchlets decurrently flattened at the often shortly

spiny nodes, the younger densely short pubescent; petioles 4-10 mm.

long, ashy pubescent; leaves opposite, elliptic, acute or shortly acum-

inate, about 1.5-7 cm. long, 1-4.5 cm. wide, entire or serrate, char-

taceous, closely puberulent above, sordidly velvet beneath; secondary

nerves 6-9; inflorescence axillary and terminal, racemiform or pan-

iculate, 7-14 cm. long; peduncles to 2.5 cm. long, canescently short

pubescent as pedicels, these to 3 (-8) mm. long, tubular-campanulate

calyx, this 3.5-4 mm. long, 5-costate, truncate, 5-apiculate; corolla

tube 6-8 mm. long, densely puberulent above the calyx, the 5-parted

limb to 1 cm. across, densely canescent-puberulent both surfaces. —

Var. longipedicellata Mold., I.e. 502, pedicels after anthesis to 8 mm.

long, flowers nutant, branchlets and leaves less pubescent. Fruit

orange, showy (West).

Cuzco: Quillabamba, Soukup 825, type; West 8001. Valle Lares,

Diehl 2500. Machu-Picchu, Herrera 3209 (type, var.). "Mote-mote"

(West).

Duranta repens L. Sp. PI. 637. 1753. D. Plumieri Jacq. Select.

Am. 186. 1763; 615.

Spiny or smooth, glabrous or glabrate densely leafy shrub often

several meters tall, the slender tetragonal branches usually lax or

trailing; leaves obovate or rather elliptic, cuneate basally, obtuse to

acuminate apically, 1.5-5 cm. long, entire or serrate above; racemes

also axillary, a few cm. to 1.5 dm. long, lax, often panicled; pedicels

to 5 mm. long; calyx tubular, 3-4 mm. long; corolla blue or rarely a

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

purple tone, the tube exserted, the limb 7-9 mm. across; fruit yellow,

7-11 mm. in diameter, the calyx beak curved. — In cultivated areas

(Ferreyra); very variable. Thus there is a var. alba (Masters) L. H.

Bailey; a var. microphylla (Desf.) Mold. Phytologia 1: 483. 1941; a

var. canescens Mold.; a var. grandiflora Mold.; and a var. variegata

L. H. Bailey — all forms, their single emphasized character indicated

by the name.

Piura: Near Sullana, Ferreyra 6006. Nearly all of warm America.

Duranta rupestris Hayek, Bot. Jahrb. 42: 71. 1908.

A procumbent spiny nearly glabrous shrub, the ternate spines

1-2 cm. long, the often verticillate leaves cuneate-obovate, obtuse or

shortly 3-dentate apically, 1-2 cm. long, subcoriaceous, lustrous

above, paler and opaque beneath, impressed glandular-punctate and

the mid-nerve subpilose; racemes terminal, erect, dense, the rachis

glabrous, the shortly pedicellate flowers nutant; calyx about 5 mm.

long, 5-denticulate, villous in the throat, the violet corolla 8 mm. long,

the spreading limb subpuberulent within. — Nearest D. triacantha

Juss. but well-marked by the small often verticillate leaves, long

racemes, small glabrous calyx (author). Used for fuel (Herrera).

F.M. Neg. 17572.

Junin: Below Palca near Huacapistana, Weberbauer 1759, type;

Kittip & Smith 24839; Ferreyra 3732. Chanchamayo, Isern 2421.—

Cuzco: Valle del Urubamba, Herrera 1127. "Ttancar."

Duranta Skottsbergiana Mold. Bull. Torrey Club 68: 502.

1941.

Branches mostly very spinose, glabrate, the younger branchlets

shortly and densely pubescent as the petioles, the leaves early be-

neath and the racemiform or paniculate axillary and terminal in-

florescence; spines rigid, a half to nearly 3.5 cm. long, opposite,

ternate or scattered as the subcoriaceous lustrous leaves or these

often borne in fascicles below the older spines, mostly elliptic or

lanceolate to oblanceolate, obtuse or acute, often attenuate to mar-

gined petiole (2-11 mm. long), entire or sparsely sharp serrate toward

apex, the younger slightly pilose above, the older glabrous and be-

neath impressed punctate; inflorescence 2-19 cm. long, flowers sub-

secund, bracts foliaceous, a pair at each inflorescence branch; bract-

lets to 7 mm. long; peduncles often 1-2 cm. long; calyx campanulate,

4-5 mm. long, truncate and 5-apiculate; corolla tube broadly cylin-

dric, to about 1.5 cm. long, puberulent toward the 5-parted limb,

FLORA OF PERU 687

this 7-10 mm. across, canescent puberulent both surfaces; fruiting

calyx coriaceous, glabrate, the subglobose yellow fruit about 1 cm.

in diameter. — A shrub or small tree with blue or lavender flowers,

the branches lenticellate-verruculose; named for Carl Skottsberg, the

contemporary director of the botanical garden at Goteberg, Sweden.

Huanuco : Mito, 1 484, type. Near Huanuco, 2080; Ferreyra 6675.

Panao, Woytkowski 116. — Junin: Uspachaca, 1304.

Duranta Sprucei Briq. Bull. Herb. Boiss. 4: 344. 1896.

Branches ferrugineous-puberulent, divaricate-ascending; petioles

1 cm. long; leaves elliptic or ovate-elliptic, abruptly subacute at base,

obtuse at apex, 6-8 cm. long, 3-5 cm. wide, regularly revolute, entire,

shortly and sparsely pubescent above, appressed pubescent beneath,

little rugose; lateral racemes about 1 dm. long, pubescent with stellate

and stipitate glandular trichomes intermixed; pedicels 1-2, in fruit

3-4 mm. long; calyx tubular, 5-costate, appressed sericeous, truncate,

6 mm. long, teeth 1 mm. long, in fruit urceolate, 8-9 mm. long; co-

rolla pale blue, about 13 mm. long, limb 7 mm. broad, inside as out

closely appressed pubescent. — The var. colombiensis Mold., Phytol-

ogia 2: 18. 1941, has calyx only lightly puberulent or strigillose.

F.M. Neg. 34314.

Amazonas: Mathews 3171, type; also the variety, fide author. —

Huanuco: Mufia, 3957. Carpish, Stork & Horton 9885. — Cuzco:

Marcapata, Vargas 9704. — Puno: Rio Limbani, Metcalf 30563.

Oconeque, Metcalf 30595. "Upatancar" (Vargas). To Colombia.

Duranta triacantha Juss. Ann. Mus. Paris 7: 77. 1806.

Much branched strongly spinose shrub a meter or two tall, the

axillary subulate spines about 2.5 cm. long; branches somewhat 6-

angled; leaves ternate or few, obovate-elliptic or oblong, subsessile

at attenuate base, subacute or obtuse, mucronulate, less than 2.5 cm.

long, entire or remotely serrate toward apex, coriaceous, often revo-

lute, lustrous above, paler and punctate glandular beneath, early

pubescent on nerves, glabrate; racemes reduced or elongate, axillary

and terminal, paniculate; calyx truncate, plicate, 5-dentate, in fruit

globose, the throat closed; corolla tube exserted, the limb pubescent.

— Varies in indument, especially of the branches and racemes, the

calyx often puberulent or only tomentose at apex on the plaits,

sometimes the racemes tomentulose all over (Schauer) ; if Schauer is

correct in this degree of specific variation, several of the plants pro-

posed as species are of course not genetic entities. Type by Jos. de

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

Jussieu, Paris, labeled "Peru," is unfortunately in bud but the

pubescent denticulate calyx is about 5 mm. long; it seems to be best

matched by Ecuador specimens and is doubtfully in Peru. F.M.

Neg. 39500.

Junfn: Ruiz & Pavdn, (Dombey 253}. — Cuzco: Valle del Paucart-

ambo, (Herrera 2962)1 "Upa-ttancar" (Herrera), "sanacassa,"

"tantur-prieto." Ecuador.

12. RHAPHITHAMNUS Miers

Reference: Moldenke, Repert. Sp. Nov. 42: 62-82. 1937.

More or less spiny shrubs or small trees, the spines just above the

axils of the opposite or ternate usually entire leaves and sometimes

with 1 or 2 flowers between the larger spines, or the short racemes in

the leaf-axils. Corolla lilac, limb ampliate, the unequal lobes spread-

ing. Stamens didynamous, two slightly exserted, about medial in

or above a ring of trichomes, the 2-lobed anthers divaricate at base.

Calyx urceolate, soon adnate to the 4-celled 2-carpellate ovary, each

carpel 2-celled, each cell with a single erect anatropous ovule. Drupe

globose, blue, fleshy, with 2 suborbicular plano-convex smooth stones

excavated at base of inner face. — Related to Citharexylum with

parallel anther cells and to Duranta with similar fruiting calyx but

4-stoned (8-seeded) drupes; sometimes cultivated for ornament

(Moldenke).

Rhaphithamnus spinosus (Juss.) Mold. Repert. Sp. Nov. 42:

69. 1937. Volkameria spinosa Juss. Ann. Mus. Paris 7: 76. 1806.

Citharexylum cyanocarpum Hook. & Am. Bot. Beech. Voy. 58. pi. 11.

1832, fide Moldenke.

Branchlets mostly spiny, somewhat puberulous, spines a half cm.

to nearly 4 cm. long; petioles 1-3 mm. long; leaves finally coriaceous,

lustrous, rather elliptic to subrotund, 0.5-4 cm. long, 0.4-2.5 cm.

wide, usually somewhat acute and mucronate, acute to cordate at

base, glabrous to strigose; calyx campanulate, 2-4 mm. long, 1.3-

2.6 mm. wide, apparent in fruit only by the 5 tiny teeth in the de-

pressed top of the drupe; racemes 1-5-flowered, strigose; corolla tube

12-20 mm. long, the lobes 2.6-3.3 mm. long, 1.2-2.9 mm. wide; fruit

5-12 mm. in diameter, blue or violet, lustrous when fresh; nutlets 2,

about 6 mm. long and wide, each 2-celled, 2-seeded. — This shrub

becomes covered with flowers; gradually filling with fruits, it is

beautiful (Ruiz & Pavon). Moldenke has listed this as found in

FLORA OF PERU 689

Lambayeque; Soukup, Biota 1: 29. 1954, showed that the specimen

no doubt came from Chile, possibly from Arica, since the collector

was there. Type by Dombey at Paris, without data. Illustrated

(as R. cyanocarpus) , Trans. Linn. Soc. Lond. 27, pi. 26; Bot. Mag. Ill,

pi. 6849 (as Duranta umbellata).

Tacna (probably, since perhaps collected in Arica) . Chile; Argen-

tina. "Haumun," "chaguis" (both, Ruiz & Pavon).

13. AMASONIA L. f.

Taligalea Aublet, PI. Guian. 2: 625. 1775.

Reference: Moldenke, Repert. Sp. Nov. 46: 193-228. 1939.

More or less ligneous, often branched from the base, the leaves

alternate, opposite or scattered, usually crowded below the terminal

inflorescence of many often trichotomous cymes, these somewhat

secund and leafy-bracted. Flowers yellow, large, single or in three's,

the broadly campanulate calyx subequally 5-lobed, the corolla with

elongate tube and slightly bilabiate limb. Stamens 4, didynamous,

inserted near base, the anthers bifid at base. Style exserted, stigma

parted. Ovary 4-celled, each cell with 1 lateral ovule. Calyx as

bracts often colored, scarcely accrescent, the drupe with 4 (1-3)

finally separate pyrenes. — Name published in 1781 but conserved

(unfortunately, Moldenke), probably for George Anson (lived to

1762), who traveled in America (J. H. Barnhart).

Besides the following, other Brazilian species will probably be

discovered within Peru.

Amasonia lasiocaulos Mart. & Schauer ex Schauer in DC.

Prodr. 11: 678. 1847; 208-210.

Branches, peduncles, pedicels (3-7 mm. long) more or less short-

hirsute, the red trichomes viscid; petioles to 2 cm. long, sometimes

almost lacking; leaves usually repand or subcrenate, obovate-elliptic

or oblanceolate, cuneately long-attenuate into petiole, acute or

shortly acuminate, 7-24 cm. long, about 2-9 cm. wide, membranous,

green or beneath red-purplish, sparsely puberulent, or pilose above,

no glandular disks but punctate beneath; panicles to 2 dm. long or

larger; bracts red, often ochrous beneath, oblong-lanceolate or ellip-

tic, 1-3.5 cm. long, 5-17 mm. wide, stiped; calyx chartaceous,

9-10 mm. long, 7-10 mm. wide, puberulent, the acuminate lobes 6-

7 mm. long; corolla tube to 3 cm. long, glandular pubescent only

without, the lobes 4-5 mm. long; stamens long-exserted ; calyx in

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

fruit 1-1.5 cm. long and wide, glabrous within, lobes to 9 mm. long;

fruit about 6 mm. in diameter, lustrous.

San Martin: (Allard 20522). Brazil.

14. CORNUTIA [Plumier] L.

Reference: Moldenke, Repert. Sp. Nov. 40: 153-205. 1936.

Tall often lax and somewhat tomentulose-pubescent trees or

shrubs with opposite entire leaves and many-flowered cymes usually

disposed in terminal panicles. Calyx various in shape, entire to

sinuate or minutely dentate, rarely lobed, scarcely or not accres-

cent in fruit. Corolla limb pubescent all over within, 4-parted,

3 lobes ovate, subequal, valvate. Perfect stamens 2, medially (or

higher) affixed, the ovoid anther cells finally divaricate, the con-

nective stout; staminodia filiform. Ovary pubescent, 4-celled, the

solitary ovules lateral below the cell-apex. Stigma minutely and

unequally 2-cleft. Drupes globose, fleshy. — Plunder's name (as de-

creed by modern savants, validated by Linnaeus) records the

botanical observations of an early seventeenth century Canadian

physician, Jacques Philippe Cornut.

Calyx about 2.5 mm. long C. odorata.

Calyx about 1 mm. long C. microcalycina.

Cornutia microcalycina Pavon ex Mold. Repert. Sp. Nov. 40:

173. 1936.

Resembles (ex char.) C. odorata; leaves typically short- tomentu-

lose beneath; bractlets 2-4 mm. long; calyx about 1 mm. long,

1.5 mm. across; corolla short-pubescent; filaments densely, stam-

inodes sparsely pilose; style short-pubescent; ovary densely long-

hirsute. — Some Colombian material referred here by Moldenke has

leaves merely puberulent; some Central American specimens have

villous ones; calyx size according to monographer's key is not a

constant character, C. grandifolia (Schlecht. & Cham.) Schauer, 163,

as interpreted by him, having calyces 1-2.5 mm. long and wide; this

variable species of Mexico and Central America apparently may

have a much wider distribution. The name of Pavon was in herbaria

under the synonym Hosta Jacq.; the author's name is often, on at

least some of the Ruiz and Pavon collections, cited in this work

under both names.

Junin : Kittip & Smith 261 33. Without data (Ruiz & Pavdn) . To

Colombia.

FLORA OF PERU 691

Cornutia odorata (Poepp. & Endl.) Poepp. ex Schauer in DC.

Prodr. 11: 681. 1847; 176. Hosta odorata Poepp. & Endl. Nov. Gen.

&Sp. 3:63, pi. 269. 1845.

Branchlets stout, nodulose, acutely 4-angled, the nodes annulate,

early densely puberulent as the petioles (2-7.5 cm. long), leaves

especially beneath, peduncles (3-6 cm. long), and cymes, including

the cupulate entire or minutely denticulate subsessile calyces, these

about 2.5-3.5 mm. long and wide; leaves broadly elliptic, attenuate

to base, acuminate, 1-3.5 dm. long, 6-19 cm. wide, strigillose-

puberulent above, membranous or subchartaceous, secondary nerves

slender, 8-12 pairs, not conspicuous, the veins obscured above;

inflorescence mostly thyrsoid, to 4 dm. long, consisting of 9-12 pairs

of several times bifurcate cymes; bractlets linear, 5 mm. long; co-

rolla typically spreading, villosulous, tube about 1 cm. long, the 2

larger subovate lobes about half as long; filaments sparsely, style

densely pilose, staminodes glabrous; ovary hirsute; calyx in fruit

patelliform, puberulent, the pubescent drupe about 3 mm. thick. —

Northern forms with appressed or denser or less indument have been

given taxonomic standing; the specific status of this entity is open

to review; compare note under C. microcalycina Pavon. A bush of

stream banks, to over 2 meters tall, the large panicles of pale violet

flowers with a strong lavender scent (Svenson); sometimes a tree,

8 meters high. F.M. Neg. 7885.

San Martin: Chazuta, King 4004. — Loreto: Yurimaguas, Poeppig,

type. Near Iquitos, King 394; 662; Williams 8223. Florida, Klug

2159. To Colombia. "Dona" (Huitoto), "tal" (Williams).

15. VITEX [Tourn.] L.

Reference: Moldenke, Phytologia 5: 142-176, 186-224, 257-280,

293-336. 1955; I.e. 343-393, 404-464. 1956; I.e. 465-507. 1957; I.e.

6: 13-64, 70-128. 1957; I.e. 129-192, 197-231. 1958.

Shrubs or trees characteristically with digitately compound leaves

(rarely reduced to one leaflet or entire) and irregular flowers in

variously developed or disposed cymes. Calyx often slightly un-

equally 5-dentate or 5- (rarely 3-) lobed. Corolla sub-bilabiate,

the 2 posterior lobes ordinarily shorter. Stamens 4, didynamous;

cells subparallel to divergent or arcuate. Ovary finally usually 4-

celled, the single ovules lateral, medial or higher. Stigma acutely

cleft. Drupe rarely subincluded, the calyx often accrescent. Seeds

obovoid or oblong without endosperm.

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

Moldenke lists a bibliography, I.e. 5: 142-145, 257-258, much of

it generally applicable and mostly at least repeated by himself when

a particular citation is concerned; a pertinent reference is an earlier

(1908) but still helpful re"sum£ of the Amazonian species by the able

Swiss botanist Huber, Bol. Mus. Goeldi 5: 209-222, with plates. Here,

as usual, the monographer, notwithstanding his wealth of informa-

tion, has avoided much original expression of specific lines; perhaps

his extremely detailed but often indecisive work will some time be

the more useful to local botanists for clarification.

The "Agnus-castus," or Chaste Tree, a useful ornamental, al-

though it dies back where there is frost, is said to be adventive in

some tropical lands and was in the botanical garden at Lima in 1946;

it is V. agnuscastus L., Mediterranean (Moldenke, I.e. 5: 161-176,

186-196), and is marked by usually 7 (5-9, reduced to 1-3) narrow

(central one often 5-10 cm. long, 1-2 cm. wide) unequal leaflets,

white-puberulent beneath, and narrow elongate panicles of lilac or

purple flowers about 8 mm. long.

Inflorescence usually a dm. long or longer and rather paniculate or

dichotomous-cymose.

Calyx sub-bilabiate, nearly 5.5 mm. long; cymes supra-axillary.

V. Klugii.

Calyx 5-dentate or -lobate, 2-3 mm. long; cymes axillary or also

terminal.

Peduncles about as long as or longer than petioles . V. compressa.

Peduncles much shorter than petioles.

Leaflets 5; petioles often ampliate at apex V. cymosa.

Leaflets 3-7; petioles slender, not or little ampliate.

V. pseudolea.

Inflorescence more or less simply cymose.

Cymes mostly 3-flowered; calyx tubular-funnelform, to 2 cm. long.

V. triflora.

Cymes at least mostly or often many-flowered; calyx small.

Peduncles shorter than petioles; calyx canescent.

Leaflets long-petiolulate; calyx in fruit lax, mucro-crenate.

V. pseudolea.

Leaflets subsessile or short-petiolulate; fruit enclosed in den-

tate calyx V. gigantea.

Peduncles longer than petioles.

Calyx gray, obsoletely dentate V. orinocensis.

Calyx flavescent, obviously dentate V. flavens.

FLORA OF PERU 693

Vitex compressa Turcz. Bull. Soc. Bot. Mosc. 36, pt. 2: 224.

1863; 295.

Branchlets obtusely quadrate, lenticellate, minutely puberulent,

the leaf-scars corky; petioles flattened above with apical disk to

6 mm. wide, slender, 5-12 cm. long, more or less puberulent as the

3-7 subequal petiolulate leaflets at least beneath and the axillary but

terminally crowded inflorescences, these usually 1-3 dm. long, 3-5

cm. wide, consisting of 5-8 pairs of well-peduncled dichotomous

cymes; leaflets oblong-elliptic, the central one 7-26 cm. long, 2.5-

11 cm. wide, all usually lustrous above, undulate, with 7-12 lateral

nerves and fine reticulation rather prominent, especially beneath;

pedicels 1-3 mm. long; bracts few, small, entire to 3-petiolulate;

bractlets many, lanceolate to linear, 3-10 mm. long; calyx 2-3 mm.

long and wide, the acute teeth or lobes usually about 0.7 mm. long;

corolla 12-13.5 mm. long, tube 4-5.2 mm. long, puberulent-pilosulous

only above, more or less long-pilose within, upper lip lobes 3.6 mm.

long and wide, central lobe lower lip to 6.5 mm. long, 8.5 mm. wide,

sublanate or bearded at yellow base; filaments completely pilose, the

purple anthers widely divergent; fruit 1-1.5 cm. long and thick,

glabrous, 4-seeded. — Becomes a stout tree to 30 meters tall, the

straight trunk to a meter in diameter; flowers blue to purple, fra-

grant. Moldenke says that it is cultivated in Peru and British

Guiana, but he cites only a collection by Pavon, without data.

Peru(?) : See above. To Colombia, Trinidad and Brazil.

Vitex cymosa Bert, ex Spreng. Syst. Veg. 2: 757. 1825; 307.

Branchlets with small pith, early grayish or yellowish puberulent,

leaf -scars elevated; petioles flattened and somewhat sulcate above,

often somewhat ampliate-disciform apically, to 1.5 dm. long, densely

puberulent or short-pubescent as the twigs and leaves beneath, the

trichomes at first more or less yellowish especially on the petiolules;

leaflets 3-7, usually very unequal, the central one oblong or elliptic

to obovate or oblanceolate, several cm. to nearly 2.5 dm. long and

about 2.5-7 cm. wide, normally acute to obtusely caudate-acuminate,

acute or usually attenuate to base, in age at least firm-chartaceous,

the 10-20 secondary nerves and fine reticulation rather prominent

beneath; cymose inflorescence much shorter than subtending peti-

oles, 1-4 or several times dichotomous, the branches widely divari-

cate; pedicels 1-2 mm. long, bracts often present, 1-1.5 cm. long;

bractlets linear, to 4 mm. long; calyx densely puberulent, 2-3 mm.

long, 2-5 mm. wide, the ovate teeth obtuse or acute; corolla tube

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

6-7 mm. long, puberulent above the calyx, the 4 small lobes about

4 mm. long, half as wide, obtuse, the larger lobe rounded, crisped,

about 7 mm. long, 8 mm. wide; fruiting calyx 5 mm. long, 12 mm.

across, finely strigose, coriaceous, the 4 or 5 lobes often obscurely

dentate; fruit purplish black, about 1-2 cm. long, 1-1.5 cm. thick. —

Shrub or sometimes a tree recorded to 25 meters tall, the bark fis-

sured; flowers appearing before leaves are mature (Moldenke) in

various shades of blue to violet-rose, odorless, the succulent fruits

edible. Illustrated, Bol. Mus. Goeldi 5: pi. 4-

Lima: Near Chosica, (Weberbauer 5341). To Colombia, Brazil,

Bolivia, Patagonia.

Vitex flavens HBK. Nov. Gen. & Sp. 2: 246. 1818; 336, 359.

V. Rusbyi Mold. Phytologia 1: 103-104. 1934, fide author.

Branchlets, petioles (-1 dm. long), peduncles (-3 cm. long), and

crowded axillary cymes (2.5-5.5 X 2-3.5 cm.) flavescent or ferrugin-

eous-furfuraceous or velvety tomentose, even the corolla; leaves 5-7

foliolate, the sessile (-6 mm. petiolulate) obovate-oblong to elliptic

shortly acute or acuminate leaflets acute to attenuate to base, to

7 cm. long, merely 3 cm. wide, appressed pubescent, finally glabrate

above, more or less canescent (or ferrugineous) tomentose beneath

and densely glandular punctate; calyx campanulate, 2.5-3.5 mm.

long and little wider, shortly subappressed pubescent, sub-bilabiate,

rather acutely 5-dentate; corolla tube broadly cylindric, about 5 mm.

long, nearly 2 mm. wide at glabrous base, 3.3 mm. wide at top, pilose

within, 4 lobes 2-3 mm. long and wide, the spathulate lower 6-7 mm.

long, apically expanded to about 4.5 mm. wide; stamens and style

exserted 5 mm., the filaments pilose below, the style glabrous, branch-

lets (stigmatic) 0.5 mm. long. — V. Rusbyi was based on a tree 15

meters tall, the description particularly for flowers compiled from

Moldenke who recognized considerable variation in the vegetative

characters. Origin of type perhaps Peru, without data. Known to

attain 20 meters; used for construction, at least in Ecuador. Illus-

trated, Bol. Mus. Goeldi 5: pi. 4. F.M. Neg. 39494.

Loreto: Tessmann 3247; 4492; 4587. To Colombia and Ama-

zonian Brazil. "Pechiche" (Ecuador), "taruma" (Brazil).

Vitex gigantea HBK. 2: 245. 1818; 373.

Branchlets terete, subcompressed, softly tomentose-hirsute, to

more or less puberulent; petioles to 1 dm. long, rather flat, rarely

canaliculate, pubescent; leaves 3-5-foliolate, the shortly petiolulate

FLORA OF PERU 695

or subsessile oblong-elliptic leaflets acute to acuminate both ends,

the intermediate usually 1-2 dm. long, often about half as wide, the

outer much smaller, all entire, finely reticulate-nerved, especially be-

neath, glabrous or puberulent above, paler and flavescent pubescent

beneath; peduncle axillary, pubescent, to 2.5 cm. long; cymes 3-6-

flowered; calyx campanulate, the 5 subobtuse teeth approximate;

corolla violet, tube short, cylindric, throat dilated, the lower lip

3-lobed, spreading, with a pilose spot at base, the middle lobe 3 times

longer than the others; drupe edible, surrounded by the calyx. —

Noted by authors as a beautiful tree with globose crown, the wood

very hard; originally from Guayaquil. Obviously very closely re-

lated to V. cymosa Bert, and V. flavens HBK.; the Peruvian speci-

mens with young leaves are sparsely puberulent above and Moldenke

thinks it is not certain that they are the same species. He gives a

description of the valuable wood.

Loreto: Pongo de Manseriche, Mexia 6177; (Tessmann 4723).

Ecuador. "Pechiche" (HBK.), "moconto."

Vitex Klugii Mold. Phytologia 1: 101. 1934; 422.

Slender tetragonous branchlets, petioles (flattened and canalicu-

late, 2-6 cm. long), and cymes including the calyces more or less

minutely strigose or puberulent, nodes often annulate, central peti-

olules subequal (or central slightly longer), 2-12 mm. long; leaflets 3,

elliptic to subobovate, 6-22 cm. long, 2.5-10 cm. wide, entire, acute

at base, acuminate, membranous, glabrous or sparsely strigillose es-

pecially beneath, lustrous both sides; secondary nerves 10-18, prom-

inent beneath; cymes often supra-axillary, 4.5-9 cm. long, 2-7.5 cm.

wide, dichotomous, branchlets (as peduncles) complanate apically;

pedicels reduced or filiform, 1.5-2 mm. long; bracts often 2, folia-

ceous; calyx tubular-campanulate, 4-5 mm. long, 5-lobed, bilabiate;

corolla tube 7-9 mm. long, densely strigose without, long pilose at

throat within, lobes subequal, ovate, 3-4 mm. long, acute; stamens

subequal, exserted; fruiting calyx campanulate-cupuliform, 5-6 mm.

long, 10-12 mm. across, strigose without, rather unevenly 5-lobed,

lobes cusped; fruit hard, dry, 12-17 mm. long, 6-9 mm. thick, um-

bilicate both ends, puberulent. — Type a 4-meter tree, the flowers

blue and white (Klug) ; known to attain 20 meters, upper corolla lip

white within (Moldenke) .

Loreto: Mishuyacu near Iquitos, Klug 625, type. Colombia;

Brazil.

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

Vitex orinocensis HBK. Nov. Gen. & Sp. 2: 247. 1818; 26.

Obtusely quadrate lenticellate branchlets, as petioles and pedun-

cles— both flattened, to 5 cm. long — as well as the axillary cymes

subcanescent with a fine pulveraceous indument; leaflets 1-5, very

unequal, the 2 lowest subsessile, the others more or less petiolulate,

elliptic to obovate-oblong, narrowed to base, usually caudately and

obtusely acuminate, the central 5-11 cm. long, (2) 3.5-4 cm. wide,

the others smaller, all membranous-subcoriaceous, typically glabrate

or veins pulverulent, lustrous, reticulate- veined; cymes subumbel-

lately or corymbosely dichotomous, bracts typically none; bractlets

linear, few; calyx shortly pedicellate, campanulate, less than 2 mm.

long, truncate, obsoletely or minutely dentate; corolla blue, middle

lobe of basally barbate lower lip larger, subcordate, undulate-crenate,

the ampliate tube to 7 mm. long, limb 5-10 mm. wide; fruit unknown

to Moldenke. — The Peruvian form, var. multiflora (Miq.) Huber,

Bol. Mus. Goeldi 5: 217, pi 3, figs. 19, 20. 1908 (as var. amazonica

Huber, fide Moldenke), has pedicels very short or obsolete, foliose

bracts often developed, leaflets rounded-apiculate or acute; corolla

lilac, or blue to violet; fruiting calyx patelliform, 4 mm. across, sub-

entire; drupe oblong-obovoid, 8 mm. long, 6 mm. thick, nigrescent.

After, as elsewhere, Moldenke, who notes that the leaves usually

discolor in the variant, rarely brunnescent in the type. Var. glabra

Mold., I.e. 4: 293. 1953, may occur; type from Amazonian Colombia.

F.M. Neg. 39495.

Loreto: (Kuhlmann, fide Moldenke). To Colombia and the

Guianas.

Vitex pseudolea Rusby, Mem. N. Y. Bot. Gard. 7: 341. 1927; 91.

Very slender petiole about apically disciform, somewhat shorter

(5.5-10 cm. long) than middle leaflet (7-15 cm. long), the 5 petiolules

6-12 mm. long, the smaller blades 5-10 cm. long, 1.5-4 cm. wide, the

outer smaller, all oblanceolate, attenuate to base, abruptly and shortly

acute or obtusely acuminate, membranous, finally lustrous above,

midrib tomentulose (as petiole), secondaries about 10-20 pairs, vena-

tion strongly reticulate; peduncles 1.5-3 cm. long, crowded on young

twigs; cymes lax, about 3 times dichotomous, canescent-puberulent;

pedicels slender, 1-3 mm. long; corolla blue or lilac, larger petal

darker; stamens blue, filaments translucent (Moldenke) ; calyx violet-

green, subcrateriform (in young fruit), faintly 5-nerved, reticulate,

lax, 5-mucrocrenate; fruit purple-black, size and taste of olive (Olea).

— Near V. gigantea HBK. but not ferrugineous-tomentulose, and,

FLORA OF PERU 697

especially, V. cymosa Bert., the flowers said to appear before the

leaves are expanded (always?); perhaps conspecific with V. cymosa

(Moldenke); ex char, the word "perhaps" should read "probably."

The sweet fruits are eagerly eaten by children, and the bark is

used by elders for rheumatism.

San Martin: (Ferreyra 4829, det. Moldenke). Bolivia. "Aceituno

del monte."

Vitex triflora Vahl, Eclog. Amer. 2: 49. 1798; 156. Macrostegia

Ruiziana Nees in DC. Prodr. 11: 218. 1847.

Quadrate branchlets and axillary peduncles glabrate to more or

less ferrugineo-tomentose puberulent; petioles 1.5-6 cm. long, slen-

der, flat and canaliculate above; leaflets 3 (1-5), entire, subsessile

(petiolules 1-3 mm. long), usually subequal, obovate-oblong to ellip-

tic, more or less attenuate to base, somewhat obtusely acuminate or

cuspidate, about 7-15 (20) cm. long, membranous (to subcoriaceous),

pale green, typically glabrous or early pubescent, glabrate or strigil-

lose beneath; nerves 8-15 pairs, rather prominent as veins both sides;

cymes mostly 3- (rarely several-) flowered, often bifid, the peduncles

about equaling the petioles; pedicels 1-5 (7) mm. long, puberulent;

calyx papery, tubular-funnelform, to 2 cm. long, with 5 foliaceous

lanceolate often spreading lobes, strongly accrescent in fruit, bilabi-

ate; corolla violet (or blue), cylindric tube about 2.5 cm. long, the

exserted part fulvous-sericeous, limb glabrous within, upper lip erect,

middle rotund lobe of lower lip barbate at base; stamens little ex-

serted; drupe edible but closely pubescent, to 1.5 cm. long, 1 cm.

thick, with aroma of Eau de Cologne, especially in drying (Mol-

denke).— Var. quinquefoliolata Mold., Phytologia 1: 104. 1934, was

based on a Rio Acre specimen (Krukoff 5765} ; var. floribunda Huber,

Bol. Mus. Goeldi 5: 215, pis. 2, 3. 1909, is marked by subcoriaceous

leaves as well as many flowers. Small slender tree, flowers sky blue

(Williams); drupes fleshy, edible. Illustrated, Mart. Fl. Bras. 9:

pi 49. F.M. Neg. 22779; 5885 (Macrostegia).

Loreto: Iquitos, Klug 1492; 1254,' (Ducke 7561). Yurimaguas,

Williams 4195; Klug 2791.— Rio Acre: Ule 9727 (var. floribunda,

probably in adjacent Peru). Bolivia to the Guianas. "Tahuari"

(Klug), "taruma."

16. CLERODENDRUM [Burm.] L.

References: Schauer in DC. Prodr. 11: 658-675. 1847; Moldenke,

Fl. Suriname 4, pt. 2: 313-321. 1940.

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

Allied to Vitex but the opposite or rarely verticillate leaves simple,

the flowers often showy, the calyx little accrescent, ordinarily spread-

ing beneath the 4-sulcate 4-seeded fruit. Corolla lobes subequal.

Stamens long-exserted. — Sometimes (especially in horticulture), and

more correctly, written Clerodendron, which form therefore I prefer.

Clerodendrons have long been prominent in cultivation; several, be-

sides C.fragrans Vent, and C. Thomsonae Balf. f., included in the text,

are probably grown in Peru, for instance, C. japonicum (Thunb.)

Sweet (i.e. C. fragrans Willd. Enum. Hort. Berol. 2: 659. 1809, not

Vent.) and C. Bungei Steud. (C.foetidum Bunge, not D. Don), both

with red flowers 2-3 cm. long and opposite cordate leaves, those of

the former resinous-squamulose beneath, of the latter glabrate; also,

one may expect in gardens C. indicum (L.) Ktze., with mostly ternate

acute-based glabrous leaves, white or yellow flowers.

Leaves 2-3 cm. long, rounded-obtuse or retuse C. inerme.

Leaves about 5 to many cm. long, obtuse to acuminate.

Calyx white to roseate, 2 cm. long or longer C. Thomsonae.

Calyx green or not showy, small.

Leaves subobtuse or acute both ends; calyx lobes ovate, acute.

C. molle.

Leaves acuminate or at base subcordate.

Leaves long-attenuate to base, entire or faintly repand.

C. Tessmannii.

Leaves subcordate at base, repand C. fragrans.

Clerodendrum fragrans Vent. Jard. de Malm. pi. 70. 1804.

Volkameria japonica Jacq. Hort. Schoenbr. 3 : 48, pi. 388. 1798, not

Thunb., 1784.

Branchlets obtusely tetragonal, subtomentose; leaves subcordate,

broadly ovate, subrotund or shortly acuminate, unevenly coarsely to

repand-dentate, hirtellous above, pubescent and glandular beneath

on nerves, especially at union with the long petiole; panicles terminal,

subsessile, compact, many-flowered; bractlets many, lanceolate or

oblong-elliptic, 1.5-3 cm. long, attenuate both ends, pelviform-gland-

ular as the tubular-obconic calyces, these about 12 mm. long, also

puberulent and with 5 narrowly subulate reflexed lobes to 1 cm. long;

corolla tube glabrous, about 2.5 cm. long, the subequal obovate lobes

reflexed. — Flowers pink to bright roseate or white, fragrant, in the

cultivated state known as var. pleniflorum Schauer, the stamens and

pistil modified in some degree; the species, native to China, is more

FLORA OF PERU 699

or less naturalized in tropical America. Illustrated, Bot. Mag.

pi. 1834; Bot. Reg. ser. 2, pi. 41.

San Martin: Chazuta, Klug 4032. — Loreto: Iquitos, Williams

1465. China; tropical America.

Clerodendrum inerme (L.) Gaertn. Fruct. 1: 271. 1788; 660.

Volkameria inermis L. Sp. PI. 637. 1753. Y. buxifolia Willd. Enum.

Hort. Berol. 2: 658. 1809. C. buxifolium (Willd.) Spreng. Syst. Veg.

2: 758. 1825.

Leaves entire, obovate, retuse or obtusely acute, to about 2 cm.

long, nearly as wide, lustrous; pedicels axillary, 1-3-flowered; calyx

truncate, apiculate-denticulate, about 4 mm. long; corolla white, the

tube often 18 mm. long, the lobes about 7 mm. long. — Ordinarily a

low shrub at most about 1 meter tall, the opposite or ternate leaves

sometimes 5 cm. long, especially in cultivation, the Peruvian speci-

men possibly obtained from an adventive individual. F.M. Neg.

24622.

Peru: Without locality, Ruiz & Pavdn. East Indies; Philippines.

Clerodendrum molle HBK. Nov. Gen. & Sp. 2: 244. 1818; 659.

Branches glabrous or pilose, subaculeate, the petioles ternate,

8 mm. long; leaves entire, ovate-elliptic to subrotund, acute to atten-

uate to base, subobtuse, mostly about 5 cm. long or longer, glabrous

above, softly canescent beneath, reticulate-veined; peduncles axillary

and terminal, longer than leaves; cymes 7-9-flowered; bracts linear,

pubescent as the campanulate calyx, this with 5 ovate acute spread-

ing lobes, cupulate and little accrescent in fruit; corolla rubescent,

the cylindric tube less than 2.5 cm. long, the limb with 5 oblong lobes;

stamens very long, secund; drupe baccate, globose, sulcate, with 4

nutlets. — Flowers fragrant. Type from Guayaquil. F.M. Neg. 39497.

Peru: (Moldenke). Ecuador.

Clerodendrum Tessmannii Mold. Phytologia 1: 448. 1940.

Branchlets tetragonous, stout, often flattened at nodes and there

more or less strigillose; petioles often crowded, stout in age, to 2 cm.

long; leaves elliptic, cuneate at base, short-acuminate, about 1-3 dm.

long, nearly 3-8 cm. wide, chartaceous, glabrous, the very slender

secondary nerves 8-11; inflorescences axillary or supra-axillary, the

opposite solitary cymes 8-15 cm. long, 2-6 cm. wide, 2-7-flowered;

peduncles stout, 4-9 cm. long, glabrate as the thick 5-12 mm. long

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

erect-ascending pedicels; bracts none, bractlets few, to 12 mm. long;

calyx about 1 cm. long, 8 mm. across at top; corolla tube 22 mm. long,

its limb about 2.5 cm. in diameter. — The Ruiz and Pavon specimen

at Geneva referred to C. inerme (L.) Gaertn. may rather be this spe-

cies, if there are two entities; the former may have obtusely acuminate

leaves, corolla tube 2-3 cm. long.

Ayacucho: Rio Apurimac Valley, Kittip & Smith 22998. — Loreto:

Rio Ucayali, (Tessmann 3244, type). Rio Mazan, Josi Schunke 338.

Clerodendrum Thomsonae Balf. f. Trans. Bot. Soc. Edinb. 7:

265, 580, pis. 7, 16. 1863.

Shrubby or soon twining, more or less purplish-puberulent, espe-

cially the branchlets, petioles (1-3 cm. long), and peduncles, these

in the uppermost axils, the spreading cymes laxly few-flowered; leaves

elliptic, 5-15 cm. long, about half as wide, rounded at base, shortly

acuminate, membranous, glabrate; bractlets linear; calyx about 2-

2.5 cm. long, early yellowish, pure white during anthesis, dull roseate

in age, 5-parted to base, alately 5-angled, the lobes acutely acumi-

nate; corolla bright red, about 2.5 cm. long, the spreading lobes

6-8 mm. long, acute; fruit trichomes, black, the nutlets joined by a

brilliant red aril; stamens long-exserted. — Named for the wife of a

missionary, W. C. Thomson, whose generous interest in floriculture

resulted in her sending in 1862 a living plant to the Botanic Garden

at Edinburgh. Cultivated at Lima and elsewhere for the attraction

of the white or tinted calyces and red flowers; apparently becoming

established in clearings. Illustrated, Bot. Mag. pi. 5313.

San Martin: Tarapoto, Williams 5923. — Loreto: Iquitos, Mexia

6501a; Williams 7923. Yurimaguas, Williams 4029. Pebas, Wil-

liams 1822. Tropical Africa. "Brinco-de-dama."

17. CALLICARPA L.

Reference: Moldenke, Repert. Sp. Nov. 39: 288-317. 1936; 40:

38-131. 1936.

More or less obviously stellate-farinaceous pubescent shrubs or

trees, rarely glabrate or glabrous, opposite, usually serrulate-crenu-

late leaves and with abundantly flowered cymes, these sessile or

peduncled, axillary or supra-axillary. Flowers usually polygamous.

Calyx 4-sinuate to cleft-dentate (rarely 5-dentate), not accrescent in

fruit. Corolla tube straight, ampliate, the limb 4- (5) cleft, the equal

lingulate lobes imbricate. Stamens 4 (5), equal, at or near base

FLORA OF PERU 701

(Moldenke). Anthers oblong-elliptic, the cells parallel. Style capil-

lary, stigma depressed or peltate. Ovary 4-celled, the 2 ovules affixed

medially or higher. Drupe globose or depressed, fleshy, with 4 (or

fewer) 1-seeded pyrenes. — Punctate glands are present on the flowers,

particularly on the anthers. Ovary absent or rudimentary in small

flowers.

Callicarpa acuminata HBK. Nov. Gen. & Sp. 2: 252. 1818; 38.

Branchlets, petioles (mostly 1-2.5 cm. long), and peduncles (1.5-

6.5 cm. long) early densely floccose tomentose, at least the branches

glabrate or glabrous; leaves ovate-oblong, entire to acutely serrate,

acute at base or somewhat attenuate into petiole, acute, acuminate

or often caudate-acuminate, mostly about 1-2.5 dm. long, 4-10 cm.

wide, membranous, more or Less stellate-furfuraceous above or sub-

glabrous in age, densely tomentose or floccose, but glandular punctate

beneath; cymes mostly in the upper axils, often laxly corymbose, to

2 dm. long; bracts (if present) few, to 3.5 cm. long, 1 cm. wide;

bractlets linear, 5-10 mm. long, tomentose; calyx about 1-1.5 mm.

long and wide, granulose pulverulent, erose or obscurely 4-denticu-

late; corolla white or greenish, tube to 2.5 mm. long, lobes less than

1 mm. long, rounded, crimped or barely 2-lobed; stamens exserted;

calyx in fruit patelliform, 2-3 mm. wide, truncate, entire, the dark

purple subglobose glabrous fruit 3-4.5-.mm. long. — C. longifolia Lam.,

native to Malay, reported as cultivated at Lima, is less tomentose,

has oblong-lanceolate leaves about a third asjjwide as long, smaller

pink-lavender flowers, smaller pink or white fruits. Slender shrub or

tree, sometimes said to attain 10 meters, the thin bark purplish or

pale brown (Williams).

San Martin: Chazuta, Klug 4144- Tarapoto, Williams 6168;

6275; 6817. Juanjui, Klug 3879. Bolivia to Mexico.

18. AEGIPHILA Jacq.

Reference: Moldenke, Brittonia 1: 245-477. 1934.

Shrubs or trees, sometimes lianas, with usually entire opposite

rarely ternate or approximate leaves and cymes (sometimes panicu-

late or variously modified) subsessile, peduncled in the upper leaf

axils or terminal or both of white or tinted flowers similar to those of

Callicarpa but often more or less (never truly, Moldenke) dioecious

(heterostylous). Calyx tubular, turbinate or campanulate, trun-

cately 4-5-dentate or -lobed, accrescent in fruit. Corolla segments

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

as stamens 4 or 5, latter exserted (male flowers), affixed above the

base. Ovary imperfectly or perfectly (Moldenke) 4-celled, the sin-

gle ovule lateral. Style filiform, the bifid stigma with more or less

obvious branches. Drupes somewhat exserted, normally 4-seeded,

often dry.

Like many plants of warm regions some species at least originally

found as shrubs probably become scandent when immediate environ-

ment makes this possible; naturally, too, plants flowering as shrubs

may live to become trees.

The monographer prepared two plates — Phytologia 4: 428. 1953

— that illustrate the use of his terms, particularly as these apply to

the calyx. Of course I have had to depend upon his revision in com-

piling the following account and acknowledge my indebtedness; most

descriptions are essentially his, many proposed species being based on

one collection. While the genus contains numerous well-defined en-

tities a lot of plants given equal standing seem to be obscure or they

may be just hidden in the uniformity of characterization. The author

himself has contrived only an artificial key, and mine, concerning

the Peruvian names, is no doubt as dubious as some of the described

species. Flowers are unknown for three plants given particular

names by Moldenke: A. glabrata, A. sordida, A. umbraculiformis,

and their key position (as for many others) is at most merely sug-

gestive; also, many key characters! are scarcely of taxonomic value.

Since compiling this account Julian Steyermark has kindly sent

me a series of A. mollis HBK. including the type photograph and

cotype of var. intermedia Mold, which has made me more skeptical

of the value of A. peruviana Turcz. and thereby of other specific

judgments.

Corolla, minus lobes, finally at least twice longer than calyx.

Panicles mostly axillary and subsessile or if somewhat peduncled,

all axillary.

Leaves typically subsessile, basally cordate.

A. sordida, A. cor data.

Leaves more or less petioled, basally acute to cuneate.

Calyx appressed canescent.

Bractlets obscure or none A. integrifolia.

Bractlets to 2 cm. long, linear A. bracteolosa.

Calyx glabrous or glabrate, trichomes few or not all appressed.

Leaves rounded or apiculate apically A. multiflora.

FLORA OF PERU 703

Leaves acute to acuminate.

Leaves broadly cuneate to base.

A. peruviana, A. filipes.

Leaves long attenuate to base A. Haughtii.

Panicles mostly or all terminal and peduncled or axillary but

peduncled.

Plants not hirsute.

Bracts not involucral or lacking.

Inflorescence glabrate or glabrous, often lax and large.

Glandular disks all over leaves beneath A. glabrata.

Glandular disks all or mostly along midnerve.

Leaves falcate-plicate A. glandulifera.

Leaves not falcate-plicate A. filipes.

Inflorescence puberulent or at least compact, often small.

Leaves petioled, sometimes shortly.

Calyx shorter than 1.5 mm.; corolla tube about 3 mm.

long A. Smithii.

Calyx about 4 mm. long; corolla tube to 9 mm. long.

A. sufflava.

Leaves subsessile A. umbraculiformis.

Bracts involucral.

Leaves abruptly short-caudate A. insignis.

Leaves acute or shortly acuminate A. pulcherrima.

Plants hirsute or shortly pilose-hirsutulous.

Stems spreading hirsute A. hirsuta.

Stems shortly pilose to densely puberulent.

A. peruviana, A. mollis.

Corolla, minus lobes, shorter than or to scarcely half exserted from

calyx.

Leaves clearly decurrent, but sometimes rounded to decurrent base.

Pilose-hirsute (shortly), the leaves rounded-attenuate to decur-

rent base A. mollis.

Puberulent, the leaves rounded to the shortly decurrent base.

A. peruviana.

Glabrous, the leaves long-attenuate-decurrent.

Petioles well-defined A. triflora.

Petioles obscure A. umbraculiformis.

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

Leaves subcordate to more or less acute at base, or cuneate, but

obscurely decurrent if at all.

Leaves (mature) glabrous or glabrate or the venation lightly

pubescent; calyces minutely appressed strigillose.

Petioles lacking or nearly A. umbraculiformis.

Petioles about 5 (-8) mm. long; leaves rounded to acute bas-

ally, often almost half as wide as long.

A. chrysantha, A. vitelliniflora.

Petioles mostly longer (-1.5 cm.); leaves acute at base, often

about a half to a third as wide as long.

A. longipetiolata, A. membranacea.

Leaves (mature) clearly pubescent one or both sides (expedient

but probably variable character).

Calyx lobate or merely denticulate.

Leaves subcordate A. sordida, A. cordifolia.

Leaves rounded to subacute at base.

Stems spreading hirsute A. ovata.

Stems puberulent or tomentose.

Calyx as leaves beneath appressed flavescent pilose.

A. elegans.

Calyx as leaves strigose, villous-velutinous or tomen-

tose.

Calyx 4-lobate; stamens 4.

Petioles to 5 mm. long; leaves rounded basally.

A. Pavoniana.

Petioles to 10 mm. long; leaves subacute at base.

A. velutinosa.

Calyx 5-denticulate; stamens 5 A. Mortoni.

Calyx subfiliform dentate, the 5 teeth 5 mm. long. .A. cuneata.

Aegiphila bracteolosa Mold. Repert. Sp. Nov. 33: 115. 1933;

456.

Simulates and resembles A. integrifolia in many respects but,

especially, has a well-developed terminal panicle about a dm. long

(-2 dm.), narrow, dense, with 3-7 notably bracteolate cymes as well

as solitary often small subsessile axillary ones; petioles sometimes

ternate, stout, 1-3 cm. long; leaves minutely apiculate-subcrenate;

cymes 1-6 cm. long, 1-4.5 cm. wide; bractlets linear or nearly, to

2 cm. long; calyx 7-9 mm. long; corolla tube very narrow, to 12 mm.

FLORA OF PERU 705

long, abruptly ampliate, the lobes obtuse; stigma branches 5.5 mm.

long; ovary glabrate. — A tree to 4 meters tall of non-inundated forest

that has been confused with A. arborescens (Aublet) Gmelin (author),

i.e. A. integrifolia Jacq. Illustrated, Phytologia 2: 435.

Loreto: (Tessmann 5863). To British Guiana.

Aegiphila chrysantha Hayek, Bot. Jahrb. 42: 171. 1908; 425.

Slender branches and branchlets subterete, tomentose or pilose,

the younger indument often yellowish as on the petioles (4-8 mm.

long) and the peduncles (1-2 cm. long) of the terminal panicle (this

5 cm. long, narrow); leaves entire, oblong or ovate, more or less

rounded basally and acute, acuminate or cuspidate apically, 7-

14 cm. long, 3-7.5 cm. wide, membranous, only midrib pilose above,

secondaries (10-12 pairs) and veins especially beneath; cymes about

1.5 cm. long, 1 cm. wide; pedicels short or obsolete; bractlets few,

linear, strigose, 2-5 mm. long; calyx campanulate, 3.5 mm. long,

laxly strigose if at all, the lobes 1 mm. long; corolla funnelform, yel-

low, the tube 3-5 mm. long, acute lobes 1.8 mm. long; stamens 4,

near mouth; stigma branches 4 mm. long, lanate; ovary depressed,

4-celled; fruit ovoid, flat both ends, the cupulate calyx antrorsely

strigose, repand with 4 spreading points. — The var. glabra Mold.,

Phytologia 2: 60. 1941, refers to a collection with calyces quite gla-

brous determined by Standley as A. Smithii Mold. Moldenke orig-

inally identified King 2104 and 2204 as A. vitelliniflora, Klotzsch.

Asplund noted flowers of a Loreto specimen as sordid yellow; fruit

orange-red (Camp.); a liana (Klug). P.M. Neg. 34313.

San Martin: Moyobamba, Weberbauer 1289. Juanjui, Klug 3894

(type, var.). — Loreto: Florida, Klug 2027; 2104; 2204. Yurimaguas,

Poeppig 2314, type (as indicated by Moldenke). Ecuador, Bolivia,

Brazil. "Fetoro-ey" (Huitoto).

Aegiphila cordata Poeppig in Walp. Repert. 4: 118. 1845; 443.

Branches and branchlets as the axillary and terminal inflorescences

densely villous-hirsute with ferrugineous spreading trichomes; leaves

opposite or nearly, sessile or subsessile, ovate-oblong or rather ellip-

tic, more or less cordate, shortly acuminate, 1-nearly 2 dm. long,

6-10 cm. wide, thin-membranous, ciliate as well as laxly strigose- or

setose-hirsute both sides, especially on the nerves beneath; cymes

solitary, 3-5.5 cm. long, about 3 cm. wide, more or less glaucescent,

several crowded in the terminal panicle, this subsessile, to 14 cm.

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

long; pedicels 1-3 mm. long; bractlets filiform; calyx 4-5 mm. long,

4-dentate or -lobed; corolla red-purple, the tube 8-18 mm. long, lobes

about 3.5 mm. long, 1.5 mm. wide; stigma branches about 2.5 mm.

long; ovary flattened both ends. — One of the most distinctive species

and would be beautiful in cultivation (Moldenke). A. Froesi Mold.,

Phytologia 3 : 107. 1949, of Amazonas, Brazil, may be distinguished

at once by its more conspicuous bractlets (1.5-3 cm. long, the lower-

most often expanded apically) and quite glabrous truncate calyx

(author). Poeppig specimen cited by Moldenke as 2158, Phytologia

1: 195. 1937. F.M. Neg. 34312.

Loreto : Yurimaguas, Poeppig 281 5 (21 58?, type) . Caballo-cocha,

Williams 2088; 2318. — Rio Acre: Ule 9720. Colombia; Brazil.

Aegiphila cordifolia (R. & P.) Mold. Brittonia 1 : 185. 1932; 445.

Callicarpa cordifolia R. & P. Fl. Peruv. 1: 50, pi 77a. 1798.

Branches spreading, much compressed at nodes, the lax virgate

branchlets, petioles (7-13 mm. long), leaves early both sides and

supra-axillary and terminal inflorescences densely ochreous pubes-

cent; leaves decussate, entire, ovate or oblong, rounded, truncate or

cordate basally, commonly acuminate apically, 9.5-18 cm. long, 3.5-

7 cm. wide, membranous, dark green and in age distinctly rough

above, secondaries 6-9 pairs; solitary cymes to half as long as leaves,

the terminal panicle to a dm. long, 6 cm. wide, with few congested

cymes; peduncles stout, 4-5.5 cm. long; pedicels 1-3 mm. long, in-

crassate in fruit; bractlets to 7 mm. long; calyx often constricted

above, about 6 to nearly 7 mm. long, densely tomentose-hirsute with

divaricate trichomes 1 mm. long, the 4 ovate lobes 1.3-2.6 mm. long;

corolla tube 6 mm. long, pilose only within, the obtuse lobes about

4.5 mm. long; stamens 4, 1.5 mm. below mouth; filaments pilose;

stigma branches 5.2 mm. long; fruit about 14 mm. long, 10 mm.

thick, yellow or orange, glabrate, 4-seeded, the cupulate pubescent

calyx 1 cm. long. — A medullose shrub or tree to 3.5 meters tall

(Moldenke). F.M. Neg. 28378.

Amazonas: Chachapoyas, (Mathews). — Huanuco: Muiia, 3922.

Panatahua, Ruiz & Pavdn, type.

Aegiphila cuneata Mold. Brittonia 1: 185. 1932; 292.

Branchlets subterete, rugose-striate, early shortly pilose; petioles

stout, 1-9 mm. long, long-setose or hirsute as the leaves, these typi-

cally sparsely or glabrate above, rather densely on nerves beneath,

FLORA OF PERU 707

and the slender peduncles (2-9 cm. long), filiform pedicels (5-10 mm.

long), linear bractlets (to 17 mm. long) and calyces, these with 4 sub-

filiform-caudate teeth; leaves elliptic-obovate, shortly acuminate,

narrowed to the stout petiole, mostly about 1.5-2 dm. long, 5.8 cm.

wide, chartaceous, slightly sinuate-margined; secondary nerves 8-12

pairs; inflorescence axillary and terminal, the cymes to about 1 dm.

long, 3-5 cm. wide, dense, usually subumbellate with 2 foliaceous

bracts at base; calyx campanulate, about 5 mm. long, marginally

4-angled, the filiform tipped teeth 4-6 mm. long; corolla tube about

6.5 mm. long, pilose within, the 5 oblong-lingulate lobes 5 mm. long;

stamens 5; ovary puberulent, 4-celled. — Indument spreading or lax,

densest on peduncles. The var. hirsutissima Mold., Phytologia 4:

173. 1953, is well-named; trichomes of younger leaves, branchlets, pe-

duncles 4-5 mm. long. Shrub, 9 meters tall with white flowers (Ule) .

Loreto: Balsapuerto, Killip & Smith 28379; 28386, type. Lower

Rio Huallaga, (Killip & Smith 29040, type, var.). — Rio Acre: Serin-

gal San Francisco, Ule 9859.

Aegiphila elegans Mold. Brittonia 1: 186. 1932; 458.

Subterete flowering branches and compact terminal and axillary

inflorescences including the 2-4-lobed cyathif orm calyces densely ap-

pressed flavescent (in herb, strigose or early villous-strigose) ; petioles

thick, somewhat curved, mostly 5-7 mm. long; leaves ovate or broadly

oblong-elliptic, acutely short-acuminate, rounded at base, membra-

nous, sparsely strigose above, more or less densely (and antrorsely)

so beneath, especially on the prominent (12 or 20) lateral nerves;

bracts lanceolate, to nearly 1 cm. long; panicles dense, usually shorter

than 7 cm. long, peduncles to about 3.5 cm. and pedicels 1-5 mm.

long; bractlets many, linear, to 1 cm. long; calyx 5.5 mm. long or

somewhat longer, nearly 4.5 mm. wide, not accrescent and loosely

cupulate at base of the subglobose 7 mm. thick fruit; corolla yellow-

ish, the tube glabrous, 6.2 mm. long, the 4 lingulate lobes about

4 mm. long; stamens 4; ovary glabrous, 4-celled. — A. spicata (Rusby)

Mold., 432, of northern Bolivia, has a terminal spiciform panicle,

long-acuminate leaves, calyx subsessile, campanulate, corolla purple.

A vine (Krukoff), branches subscandent, fruit orange (Asplund), that

has been distributed as Cordia pubescens Willd.; related to A. cordi-

folia (R. & P.) Mold, (author).

Junin: Puerto Yessup, Killip & Smith 26838. — Loreto: Iquitos,

Killip & Smith 27055, type. Yurimaguas, Killip & Smith 27562;

27991. Brazil.

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

Aegiphila filipes Mart. & Schauer in DC. Prodr. 11: 652. 1847;

366.

Branches'and branchlets subterete, slender, weak and early mi-

nutely puberulent as the decussate petioles, these 5-7 mm. long;

leaves oblong to elliptic, shortly or broadly cuneate to base, usually

acuminate or abruptly acute, 8-18 cm. long, 2-7.5 cm. wide, entire

to coarsely angulate-dentate, thin-chartaceous, dark green above,

puberulent both sides, finally glabrous above and merely densely

punctate and with few-many gland disks beneath, the 10-15 pairs

of secondary nerves as the veins ordinarily obscure; axillary cymes

2-3-parted, laxly spreading, the terminal panicle 5-6 cm. long, about

as wide, few-branched, puberulent or pulverulent including the co-

rolla tube; peduncles and pedicels slender or filiform, weak, the

former 1.2-5 cm. long, the latter 2-9 mm. long; bractlets few, 1-

12 mm. long; calyx membranous, lax, campanulate or broadly cyath-

iform, about 2.5-4 mm. long, attenuate into pedicel, the rim 4-mucron-

ulate or sinuate; corolla yellow or light green, glabrous, the tube

3-6 mm. long, the 4 linear to ovate-lingulate lobes 4-6 mm. long,

obtuse; filaments 4, glabrous or puberulent, affixed below the corolla

mouth; stigma branches 1.8-2.6 mm. long; ovary 4-lobed, 4-sulcate;

fruiting calyx cupulate, about 6 mm. long, 9 mm. wide, the fruit

8-9 mm. long and about as thick, flattened apically, obviously lobed,

1-4 seeded. — A 3-7-meter tall shrub or slender tree with orange fruit

(Krukoff ) ; flowers light green (Asplund) ; a weak, low shrub or tree

(to 5 meters) or subscandent (Moldenke). Williams 3146 is not this

(Moldenke) ; besides the following, this collector made many collec-

tions, particularly at La Victoria. P.M. Neg. 20350.

Loreto : Iquitos, Williams 81 90. Lower Rio Nanay, Williams 533.

Caballo-cocha, Williams 2469. La Victoria, Williams 2622; 2850;

2856; also Killip & Smith 26882; Tessmann 3705; Asplund 14111.

To Panama and Bolivia; Brazil. "Chirapa-sacha."

Aegiphila glabrata Mold. Brittonia 1: 186. 1932; 374.

Scandent, the slender fruiting branchlets glabrous or nearly except

for a few trichomes persisting on the thick petioles (4-6 mm. long),

peduncles (4 cm. long) and somewhat incrassate pedicels (4 mm.

long) ; leaves oblong or nearly, entire or somewhat undulate, rounded

or slightly acute at base, caudate-acuminate, 15-19 cm. long, 3.5-

5.5 cm. wide, membranous or chartaceous, pale green and sublustrous

both sides, abundantly and minutely glandular beneath, the mid-

FLORA OF PERU 709

nerves and slender lateral nerves (7-12 pairs) prominent beneath;

inflorescence terminal, several cm. long (in fruit), the linear bractlets

about 6 mm. long; calyx cupulate, apparently minutely 4-apiculate,

about 5 mm. thick, twice as wide at the top and closely enclosing the

oblong-cylindric red glabrous fruit, this about 1.5 cm. long, 1 cm.

thick. — Probably related to A. martinicensis Jacq. (author); consider

also A. filipes Mart. & Schauer. Asplund's collection from a shrub

5 meters tall, branches pendent, fruit (immature) orange-red.

Huanuco: (Asplund 12835). — Junin: Pichis Trail, Kittip & Smith

25503, type.

Aegiphila glandulifera Mold. Brittonia 1: 187. 1932; 364.

Glabrous or glabrate, rarely puberulent with obtusely (rarely

acutely) quadrate or subterete branchlets, the opposite mostly falcate

and plicate leaves densely punctulate both sides and ordinarily with

many black glandular disks along the stout midrib beneath; petioles

9-13 mm. long; leaves oblong, more or less acute or acuminate each

end, 12-22 cm. long, 4.5-10 cm. wide, chartaceous, nerves 10-16

pairs, veins five; peduncled axillary cymes bracteate, bracteolate,

often reduced, or many-flowered and nearly 1 dm. long, the terminal

panicle sometimes with 200 or more cymes, usually dense; peduncles

glabrous, to 3 cm. long; pedicels slender, 2 mm. long, more or less

puberulent, the many bracts foliaceous, the bractlets linear; calyx

about 1.8 mm. long, 2.3 mm. wide, lax around corolla tube, truncate

or 4-apiculate; corolla yellowish, tube 7.8 mm. long, lobes 4, oblong,

acute, 4-6 mm. long; anthers 4, about 1.3 mm. below mouth of tube;

stigma branches 4.6 mm. long; ovary 4-celled; fruit glabrous, finally

black, 8-10 mm. long, 6-9 mm. thick, the cupulate calyx 4-4.5 mm.

long, 9 mm. wide, puberulent (author, but specimen not cited). —

Var. pyramidata L. C. Rich, ex Moldenke, Phytologia 1: 203. 1937,

resembles A. laevis (Aublet) Gmelin, 394, but has longer terminal

branches, thinner leaves densely disk-glandular and lightly strigillose

beneath as calyx, the branchlets and panicles densely strigillose or

short-strigose; type in Paris Herbarium, with description by Richard

as a species; Moldenke, I.e. 4: 432. 1953, has contrasted this variety.

Small tree to 8 meters tall, with yellow or greenish flowers; it has

been confused with A. filipes (author); Haught observed trees in

Colombia as strongly aromatic, the inflorescences pendent.

Loreto: Balsapuerto, King 3016. Rio Ucayali, Tessmann 3508.

"Chirapasacha" (Klug). To Costa Rica; Brazil.

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

Aegiphila Haughtii Mold. Phytologia 2: 8. 1941.

Extended bracts all slender or the pedicels (these 5-8 mm. long)

filiform and glabrous as all of the entirely axillary few-flowered cymes;

younger branchlets, petioles (5-10 mm. long) and opposite membra-

nous entire leaves early minutely or obscurely puberulent; leaves obo-

vate to elliptic, entire, cuneately attenuate to base, long-acuminate,

about 1-2.5 dm. long, nearly 4-8.5 cm. wide, with 10 or 20 pairs of

secondary nerves, the abundant reticulation evident; cymes 5-6.5 cm.

long, 3-4 cm. wide, on peduncles 3.5 cm. long; calyx cupulate, about

2.7 mm. long, truncate, entire, black in herbarium as the salver-

shaped corolla, this with 5 oblong-lingulate lobes 4-5 mm. long,

its slender tube twice as long. — Tree, flowers white, 8 meters tall

(Schunke) ; type a 2-meter shrub, by Oscar Haught.

Loreto: Rio Mazan, Schunke 838. Ecuador.

Aegiphila hirsuta Mold. Repert. Sp. Nov. 33: 127. 1933; 440.

Ex char, resembles A. ovata but (on the basis of a single collection)

internodes 5-6 cm. long in contrast to 7.5-14.5 cm., petioles about

5 mm. long, leaves more oblong, 6.5-13.5 cm. long, 2.5-5 cm. wide,

lustrous, secondary nerves 8-11 pah's, peduncles about 4 cm. long,

inflorescence apparently only terminal, 11.5 cm. long, 4 cm. wide;

however, the inflorescence of A. ovata was imperfectly known, and

according to the author the type of A. hirsuta was far too immature

to permit accurate measurements of floral parts. Type described as

scandent. The var. colombiana Mold., Castanea 10: 44. 1945, from

Putumayo near Peru is of course more widely distributed, but the

tree may be referable to A. ovata Mold., the apparent differences

certainly within the range of expected variation.

Peru (no doubt). Bolivia.

Aegiphila insignis Mold. Brittonia 1: 188. 1932; 436.

Branchlets obtusely tetragonal, the younger parts more or less

densely yellowish puberulent especially toward the small terminal

and axillary conspicuously foliose-bracted cymes; petioles 6-12 mm.

long; leaves opposite, mostly unequally geminate, entire, oblong,

abruptly and acutely acuminate, acute at base, membranous, lus-

trous both sides, early puberulent-strigillose on the 7-8 lateral nerves

and prominently reticulate veins beneath; axillary cymes only in

uppermost pair of leaves, about 5 cm. long and wide, the terminal

panicle of cymes to a dm. long or longer; pedicels filiform, about

FLORA OF PERU 711

2 mm. long; bracts many, ovate, oblong or lanceolate (smaller ones),

to 4.5 cm. long, 2 cm. wide, truncate to acute at base, the outer sim-

ulating an involucre, glabrous or nearly; calyx campanulate, puberu-

lent-strigillose, about 4 mm. long, the 4 broadly ovate obtuse lobes

1 mm. long; corolla tube 14-18 mm. long, glabrous, the ovate-lingu-

late lobes 4 mm. long; stamens 4, 2.6 mm. below mouth; stigma

branches 9 mm. long or longer; ovary glabrous, 4-celled. — Allied to

A. fluminensis Veil, of Brazil and A. pulckerrima Mold, (author);

the latter, in all probability, is the same.

Huanuco: Chicoplaya, Ruiz & Pavdn (Tafalla) , type.

Aegiphila integrif olia Jacq. ex Hook. & Jacks, in Ind. Kew. 1 :

46. 1895; 337. Callicarpa integrif olia Jacq. Enum. Syst. PI. 12. 1760.

C. globiflora R. & P. Fl. Peruv. 1: 49, pi. 776. 1798. A. arborescens

(Aublet) Gmelin, Syst. Nat. 2: 259. 1791. Manabca arborescens Au-

blet, PI. Guian. 1:64. 1775.

Branches and branchlets acutely angled, compressed and ampli-

ate at node, very medullose, canescently puberulent-pubescent as the

decussate petioles (4-13 mm. long) and completely axillary inflores-

cence or this as the leaves especially on the 10-15 prominent second-

aries beneath more sericeous-tomentose; leaves entire or denticulate,

oblong to obovate or elliptic, attenuately decurrent, acuminate at

apex, mostly 1-3 dm. long, a half to a dm. wide, somewhat lustrous

above where glabrate in age, glandular-dotted beneath, the veins fine;

cymes solitary, dichotomous, trifid, lobate or capitately corymbose,

usually dense; peduncles 2-5 cm. long, grooved; pedicels filiform, to

5 mm. long, obsolete (female flowers), bractlets absent or few and

short; calyx mostly clavate or turbinate-tubular, 5-8 mm. long, seri-

ceous also within, acutely 4-dentate, 4-lobed or 4-parted; corolla fun-

nelform, lobes ovate- or lanceolate-lingulate, acute, 2-4 mm. long;

stamens 4, just below mouth; stigma branches about 2.5-3.5 mm.

long, ovary glabrous; fruit 8 mm. long, 7 mm. thick, apically de-

pressed, smooth, lustrous, finally orange with 2-4 ovate plano-convex

seeds, the cupulate unevenly lobed calyx enclosing its base. — Some-

times 12 meters tall, trunk 3 dm. in diameter, rough gray bark,

flowers white (Williams).

San Martin: Zepelacio, Klug 3468. — Huanuco: Chicoplaya, Ruiz

6 Pavdn (type, C. globifera). Cuchero, Spruce 1616. Pampayacu,

5053. — Junin: Puerto Yessup, Killip & Smith 26239. — Loreto: Rio

Mazdn, Jose Schunke 343. Iquitos, Klug 108. Caballo-cocha, Wil-

liams 2052(1} ; 2328. La Victoria, Williams 2583; 2795. Rio Ucayali,

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

(Castelnau). — Cuzco: (Sandeman 8676} ; (Scolnik 903). — Puno: (McCar-

roll 94; Metcalf 30667). To Colombia, Bolivia and Trinidad. "Taba-

quillo" (Williams).

Aegiphila longipetiolata Mold. Brittonia 1: 188. 1932; 417.

Similar to A. chrysantha but glabrate and, ex char., petioles 5-

15 mm. long, leaves lanceolate or oblong, acute at base, about 7-12

cm. long, 2.5-4 cm. wide, glabrous, with about 5 pairs of secondary

nerves; panicle terminal, ample (to nearly 2 dm. long), the 11-13

cymes well-peduncled; calyx about 2.5 mm. long, with 4 small teeth;

corolla tube about 3.5 mm. long. — If this glabrous variant of A.

chrysantha Hayek, in which Hayek included Weberbauer 4667, is cor-

rectly placed the other differences could conceivably also be within an

expected range of variations for any species. Some material of

A. chrysantha has oblong-lanceolate leaves and of course inflores-

cences vary in size. The types of both plants were from the same

general region, "Yurimaguas."

San Martin: Moyobamba, (Weberbauer 4667, type).

Aegiphila membranacea Turcz. Bull. Soc. Nat. Mosc. 36, pt. 2:

219. 1863; 370.

Elongate subterete branchlets, petioles (5-10 mm. long), and

peduncles (2.5-4 cm. long) at least early short-pubescent or pilose;

leaves entire, oblong or oblong-lanceolate, acute at or narrowed

to base (acuminate), about 1.5- nearly 2.5 dm. long, 4-6.5 cm.

wide, thin-chartaceous, blackening and more or less undulate

dried as the midrib and 6-10 pairs of unevenly branched slender

nerves, glabrate above except for puberulent nerves, puberulent or

short-pubescent and closely punctate; inflorescence a terminal panicle

to at least 1.5 dm. long, the lower of the many branches often de-

flexed; pedicels filiform, puberulent, 4-5 mm. long; bracts few, small,

the bractlets linear, to 5 mm. long; calyx 2.3-3.3 mm. long, puberu-

lent, entire or 4-apiculate; corolla yellowish, the glabrate tube about

4 mm. long, the oblong-lingulate lobes 4.5 mm. long to nearly 2 mm.

wide; stigma branches about 2 mm. long; fruiting calyx glabrate,

3-4 mm. long, 8-10 mm. wide, truncate, entire, the oblong glabrous

fruit 4-seeded. — Chartaceous leaves of A. filipes Mart. & Schauer,

which, however, never become undulate; has often been confused

with Clerodendrum (Moldenke); to 3 meters tall, flowers greenish-

white (Steyermark), fruit orange (Williams).

FLORA OF PERU 713

Loreto: Rio Ucayali, Tessmann 4666. To Colombia, Venezuela,

Surinam.

Aegiphila mollis HBK. Nov. Gen. & Sp. 2: 250, pi. 130. 1818;

403. A. Mutisii HBK. I.e. pi. 181 and A. salutaris HBK. I.e. 249,

fide Moldenke. Cornutia velutina Hayek, Bot. Jahrb. 42: 172. 1908.

Branches fuscous in age and subterete, the younger subangled and

densely hirsute-lanate as the weak petioles (4-11 mm. long) and

leaves, especially above, these beneath as the peduncles (1.5-5.5 mm.

long), pedicels (1-3 mm. long) and usually calyces yellowish lanate;

leaves entire or subdentate toward acuminate or acute tip, obovate-

oblong, ovate-elliptic or elliptic, acute at base or often attenuate into

petiole, 6-19 cm. long, 2-8.5 cm. wide, membranous, the lateral

nerves 6-8 pairs well-defined beneath by their denser usually diver-

gent indument; inflorescence axillary and terminal, the opposite

cymes corymbose or panicled, bracted, spreading, to half as long as

the subtending leaves, the thyrsoid terminal panicle to 17 cm. long,

10 cm. wide, often trichotomous, bracted and foliose at least below,

the geminate bracts foliaceous, about 2 cm. long, acute both ends,

sparsely lanate, the bractlets setaceous, 3-5 mm. long; calyx 2-3.3

mm. long, 3.6 mm. wide, subquadrate, lax, subtruncate, repand or

4-mucronate; corolla white to yellow, pilose or glabrate, the tube

3.7-5.3 mm. long, lobes acute, 4 mm. long, 2 mm. wide; filaments

pilose; stigma branches 1-4 mm. long; calyx in fruit 4-angulate,

entire, greatly accrescent, the orange fruit with 2-4 oblong seeds. —

The var. intermedia Mold, described from Colombia is only tomentu-

lose-puberulent and seems to be A. peruviana; shrub or small tree,

often unpleasantly odorous, not in Peru according to Moldenke, who

has suggested that all Bolivian species referred here may belong to

his A. Steinbachii, Phytologia 2: 9. 1941, with 4-lobed calyx rim.

F.M. Negs. 17568 (Cornutia velutina); 17588 (A. salutaris).

Peru (probably; cf. note above). To Panama and Bolivia?

Aegiphila Mortoni Mold. Phytologia 1: 249. 1937.

Shrub, the obtusely angled branchlets and chartaceous leaves

above shortly pubescent, the leaves beneath and axillary dense cy-

mose capitate inflorescences white tomentose; petioles 1.5 to nearly

2 cm. long, not marginal; leaves oblong or elliptic, obtuse or acute at

base but not decurrent, acute, 9.5-13 cm. long, 3.3-4.4 cm. wide,

densely barbellate beneath along midrib and the 8-12 secondaries

(otherwise tomentose), not glanduliferous; cymes solitary, opposite,

1-2 cm. long or wide, the conspicuous linear bractlets 8-10 mm. long,

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

short fulvous-pubescent, often 3 at the base of the similarly pubescent

obscure calyx, this 5-ribbed, shortly 5-dentate, 6-7 mm. long,

3.5-5 mm. wide; corolla tube about 4 mm. long (bud), the 5 lobes

2-3 mm. long, glabrous; stamens inserted about medially; style

2-3 mm. long, branches 2 mm. long; ovary glabrous. — About 5 meters

tall with large clusters of green berries (collectors) ; allied by author

to A. truncata Mold., 289, of Colombia. Meritoriously named for

C. V. Morton of the United States National Herbarium.

Cuzco: Panticalla Pass, 3,600 meters, (Cook & Gilbert 1234, type).

Aegiphila multiflora R. & P. Fl. Peruv. 1: 50, pi. 76. 1798; 295.

Clerodendrum bolivianum Britton ex Rusby, Bull. Torrey Club 27: 82.

1900, fide Moldenke.

Branches stout, 4-angulate, compressed at the granulose-tomen-

tose enlarged nodes, the branchlets early purplish tomentose or

hirsute, finally granulose-tomentose, strigillose or glabrate as the

slender or stout petioles (0.5-2 cm. long) and peduncles (to 3 mm.

long) ; leaves ovate-lanceolate or oval, shortly attenuate into petiole,

rounded or abruptly acute, about 0.5-1.5 dm. long, half as wide,

glabrate above, rather lanate or villous beneath especially on the 6

to 20 pairs of nerves and veins, these prominent; cymes all in axils

or at nodes of preceding year, solitary, often congested, sometimes

2-3-fid; pedicels 0-3 mm. long, lanate-pilose; bractlets minute; calyx

3.6-6.2 mm. long, 4-6 mm. wide, purple pubescent or tomentose, 4-

denticulate; corolla red-purple, tube 13 mm. long, puberulent only

without or the ovate-lmgulate obtuse lobes (at least 5 mm. long,

3.5 wide) ; stigma lobes 3.1 mm. long; calyx in fruit patelliform, 8 mm.

wide, verruculose pubescent, the subrotund 4-angled dark purple

fruits depressed both ends, with 4 osseous striate seeds. — Becoming

a tree, to 7 meters tall, stems medullose but wood said to be durable

and used for boats; corolla lavender; superficially resembles Clero-

dendrum (Moldenke); the flowers are dark red. Ruiz and Pavon

apparently combined their Pillao and Huassahuassi collections and

noted it growing near water with aspect of Cornus. F.M. Neg. 17587.

Huanuco: Pillao, Ruiz & Pavdn; same department, (Asplund

13586}. — Junin: Huassahuassi, Ruiz & Pavdn, type. — Puno: Sandia,

Vargas 1311; 6948; 9644. Limbani, Metcalf 30550. Bolivia. "Utcus."

Aegiphila ovata Mold. Brittonia 1: 189. 1932; 441.

Conspicuously hirsute, a shrub or the elongate (to 3.5 meters long

or longer) branches probably more or less scandent, these subterete

FLORA OF PERU 715

and with the petioles (9-12 mm. long) most densely clothed with

spreading trichomes 1-3 mm. long, these arcuate or subappressed

and more or less bulbous-based on the membranous leaves, especially

above where rather sparse; leaves ovate, rounded at base, acutely

subcaudate-acuminate, in type 12-23 cm. long, 6 (4.5) -11.5 cm.

wide, entire, ciliate (Moldenke), the slender secondary nerves about

13 pairs, prominent only beneath; inflorescence axillary (and termi-

nal?), hirsute-villous, the few cymes opposite, subcapitate, nearly

4 cm. long, 2.3 cm. wide; peduncles about 1 cm. long, pedicels

1-2 mm. long, bractlets few, linear, to 11 mm. long; calyx campanu-

late, at least 4 mm. long, the ovate lobes rounded to a short apicula-

tion; corolla tube 3.6 mm. long, lobes 2.6 mm. long, 1.5 mm. wide;

stamens 4, just below the mouth, exserted; ovary glabrous, 4-lobed,

4-celled. — Simulates A. cor data Poeppig with cordate or subcordate

shortly acuminate leaves and is apparently distinct; type described

as a shrub. Moldenke, in his key, I.e. 277, describes the leaves as not

ciliate.

Loreto: Puerto Bermudez, Kittip & Smith 26520, type. Bolivia.

Aegiphila Pavoniana Mold. Brittonia 1: 190. 1932; 460.

Lanate-tomentose, the membranous leaves both sides and the

axillary-terminal inflorescence, or this more villous; petioles opposite,

3-5 mm. long; leaves entire, ovate or ovate-oblong, rounded at base,

acuminate, 7-12 cm. long, 4-6 cm. wide, the secondary nerves (ob-

scure above) about 6 pairs; cymes few, axillary, densely subcapitate,

2.5 cm. long, 2 cm. across, their peduncles to 1 cm. long (terminal

one 2.5-4 cm.), the terminal panicle about 7 cm. long, 4 cm. wide

with 3 cymes, the lower 2 subsessile; bracts 2, foliaceous; bractlets

linear, numerous, longer than the calyces, these thin, tubular-

turbinate, 6.2 mm. long (Moldenke), densely pubescent, the rim

4-lobate; corolla tube narrow, 5 mm. long, oblong-lingulate lobes

3.6 mm. long; stamens 4, inserted at mouth; ovary glabrous, 4-celled.

— Reminds one of some forms of A. mollis HBK. but its inflorescence

is entirely different (author) ; this is more simple and less developed,

perhaps an individual condition or variation but, ex char., there

seems to be a single character that, if correct, may indicate a distinct

species, namely, the much larger 4-lobate calyx. Moldenke, Phyto-

logia 1: 256. 1937, has discussed a Pavon inscription (apparently) on

label for this species in British Museum which reads "Huayaquil";

as he suggests, the plant may have been collected at Guayaquil (and

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

probably by Tafalla). A 5-meter tree, the flowers cream-colored

(Haught).

Peru(?): Without data, (Ruiz & Pavdn, type, Herb. Boissier,

Geneva). Ecuador.

Aegiphila peruviana Turcz. Bull. Soc. Nat. Mosc. 36, pt. 2: 219.

1863; 368.

Branches and branchlets, petioles (4-17 mm. long), peduncles

(2-6 cm. long), and pedicels (1.5-2 mm. long) all slender and at least

early densely puberulent as the decussate leaves both sides and the

axillary and terminal inflorescences including the campanulate or

turbinate-cyathiform calyces, these membranous, about 2-3.5 mm.

long, 4-apiculate; leaves lanceolate to oblong, acute or acuminate

both ends, 8-15 cm. long, 2.5-4 cm. wide, membranous or charta-

ceous, slightly rough above in age, often with many gland-disks along

midrib beneath, the 6-12 pairs of secondaries prominent; cymes soli-

tary, 5-6 cm. long, 1-4 cm. wide, bracteolate, the terminal panicle

thyrsoid or loose and spreading, 5-19 cm. long, 4-10 cm. wide; bracts

few, subfoliaceous, bractlets few, linear, 2-3 mm. long; corolla tube

about 2.5-3.5 mm. long, glabrate, the 4 ovate-lingulate lobes 3.1 mm.

long; stamens 4; stigma branches 1-4.3 mm. long; ovary glabrous,

4-lobed; fruiting calyx 4 mm. long, twice as wide, shallowly capitate,

lax around the fruit, this about 8 mm. long, 6 mm. wide, flattened

at apex, 1-4-seeded. — Very variable, shrubby or scandent (Mol-

denke) ; a few of the many collections from Tarapoto and vicinity are

cited; Klug observed it in forests, about 1 meter tall with cream-

colored corollas. It suggests A. filipes Mart. & Schauer but, for one

thing, the leaves seem to be firmer or softer. F.M. Neg. 24619.

San Martin: Tarapoto, Woytkowski 35183; Spruce 4-275, type;

Williams 5383; 5479; 5588; 6291. San Roque, Williams 7334. Juan

Guerra, Williams 6836. Zepelacio, Klug 3511. Bolivia. "Huaca,"

"chirapa-sacha," "ucullucuy-sacha" (all Williams).

Aegiphila pulcherrima Mold. Brittonia 1: 191. 1932; 435.

Resembles A. insignis unless (ex char.) the following differences

are taxonomically significant: leaves rounded at base or rarely sub-

acute, acute or very short-acuminate at apex, midrib pilose both

sides as the large veins beneath; axillary cymes to 9 cm. long, half

as wide; pedicels 2-4 mm. long, nearly all subtended by oblong or

ovate bractlets, often forming a pseudo-involucre and wider than the

pair of bracts (at base of each panicle branch), these to 2 cm. long,

FLORA OF PERU 717

4 mm. wide; calyx obconic, lobes 1.3 mm. long; corolla tube 10.4 mm.

long, lobes 7.2 mm. long; stigma branches 3.3 mm. long. — Reminds

one greatly of A. fluminensis Veil, and A. macrantha Ducke (Brazil-

ian) but a liana (Schunke) with distinctly terminal panicles, different

venation and beautifully and characteristically yellow-pilose through-

out (author). Characteristic features of a true species could ques-

tionably be known from a single collection; anyway, the yellow

pilosity ex char, is only on the midrib as regards the upper leaf

surface.

Junfn: Chanchamayo Valley, Schunke 400, type; 443 (det. John-

ston, A. mtelliniflora Klotzsch).

Aegiphila Smithii Mold. Brittonia 1: 191. 1932; 419.

Suggests in many respects A. vitelliniflora, apparent or possible

differences (ex char.) being: branchlets (early), leaves beneath and

panicles except the calyces canescent strigose; leaves lanceolate, 8.5-

13.5 cm. long, 3-4 cm. wide, glabrous above with many glandular

disks along midrib, secondaries 7-11 pairs; lower opposite cymes to

8 cm. long, half as wide; peduncles much thickened at base; calyx

campanulate, herbaceous, glabrous (Moldenke), the lobes rounded;

corolla tube to 3.1 mm. long, oblong-elliptic lobes about 2.3 mm.

long; stamens just below mouth; ovary glabrous; calyx in fruit larger,

patelliform, about 1.5 mm. long, 3.5 mm. wide, sparsely hirtellous

(Moldenke), the subglobose fruit about 5 mm. long, warty and verru-

cose, bright orange (fresh), flattened apically, 2-4-seeded. — Remark-

able because of its small calyx (also in fruit), bracteate inflorescence,

distinctive leaves, small verrucose fruit; differs except in small calyx

from A. Spruceana Mold., 416, and A. parviflora Mold., 328 (author).

Liana with white flowers in cut-over weeds (Mexia); shrub with

orange fruits (Asplund) .

San Martin : Tarapoto region, King 3894. — Junin : Puerto Yessup,

Killip & Smith 26284- — Loreto: Iquitos, Kittip & Smith 26957, type;

Williams 1390; 3689; Mexia 6499. Mishuyacu, King 1460; 1490.

Lower Rio Nanay, Williams 680. Also Tessmann 3591; (Asplund

14048).

Aegiphila sordida Mold. Brittonia 1: 192. 1932; 299.

Branchlets medium-thick, subterete, light gray, densely spreading

setose or hispid as the petioles, these stout, 7-9 mm. long; leaves de-

cussate, entire, oblong, rather long-acuminate, cordate or subcordate

at base, 22-24 cm. long, about 8 cm. wide, membranous, sparsely

setose, more densely beneath, especially along the midrib and 10 or

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

20 pairs of secondary veins, the trichomes directed variously; inflores-

cences apparently axillary, glomerate, subsessile or short-peduncled,

dense; fruiting calyx indurated, to 8 mm. long, 17 mm. wide, gla-

brate or very sparsely pilose, the rim repand; fruit oblong, apparently

1.5 cm. long, 4-seeded. — Type deplorable, leaves imperfect, only one

fruit but suggests in Peru A. cuneata (author). Moldenke has deter-

mined two collections by Killip & Smith without, however, adding

to his description. A liana with bright red fruit (Killip & Smith).

Junin: (Killip & Smith 26715}. — Loreto: Puerto Arturo, Yuri-

maguas, Williams 5054, type. Also Killip & Smith 27793, fide

Moldenke.

Aegiphila sufflava Mold. Brittonia 1: 193. 1932; 461.

Branchlets soon subterete, early densely ochrous or purplish pu-

berulent to appressed pubescent as the stout petioles (8-12 mm. long),

thyrsoid panicles, branches and peduncles (3.5-4.5 cm. long); leaves

sometimes subopposite, oblong or oblong-ovate, cordate, rounded or

subtruncate, acute or acuminate, 13 cm.-about 2 dm. long, half as

wide, membranous or subcoriaceous, light green and lustrous both

sides, glabrous above except midrib, puberulent-pubescent beneath,

especially the 9-16 pairs of lateral nerves, these prominent as the

veins both sides; inflorescence axillary and terminal, the cymes

mostly a single pair about 4 cm. long, 3 cm. wide, bracted and brac-

teolate as the terminal panicle, this to 16 cm. long, with 11 or more

cymes; lowermost bracts foliaceous, the much smaller uppermost lan-

ceolate, the bractlets linear; calyx broadly obconic, 4.4 mm. long,

3.8 mm. wide, obscurely granular, unevenly 4-lobed, two lobes some-

times 1.8 mm. long on one side, two 1 mm. long, but not constantly,

the calyx slightly apiculate; corolla glabrous, the yellowish tube

9 mm. long, the bright yellow lobes 5.4 mm. long, 2 mm. wide;

stigma branches 2.8 mm. long; ovary glabrous, depressed-oblong;

calyx in fruit heavy, about 7 mm. long, 1-1.5 cm. wide, rim entire,

obscurely puberulent; fruit globose, 10-13 mm. thick. — Type, a tree

5 meters tall; the var. Klugii Mold., Phytologia 1: 295. 1938, is more

densely appressed strigose, calyces to 7 mm. long, 5.5 mm. wide, and

often with a round gland on each lobe. Apparently related to A. ra-

cemosa Veil., Brazil and northward, with scabrous leaves and other

differences (author). Liana with yellowish flowers (Klug).

Loreto: Iquitos, Killip & Smith 27439, type; Williams 8225; Tess-

mann 5155. Florida, Rio Putumayo, Klug 2076. La Victoria, Wil-

liams 3015. Mishuyacu, Klug 2511 (var. Klugii}.

FLORA OF PERU 719

Aegiphila triflora Mold. Repert. Sp. Nov. 37: 212. 1934.

Glabrous, including the axillary 3-flowered inflorescence; branches

obtusely tetragonous, the branchlets slender; petioles 11-21 mm. long,

deeply canaliculate above, strongly incrassate and enlarged toward

base; leaves entire, oblong-lanceolate, shortly acuminate, cuneate to

base, 9.5-19 cm. long, 3.5-6 cm. wide, firm-membranous, lustrous

both sides, the 7-9 slender nerves prominent beneath, the secondary

veins not at all or obscurely arcuate-confluent marginally; peduncles

2-4 cm. long, apically enlarged as the 11-19 mm. long pedicels; bract-

lets at base of pedicels 2, filiform, about 3 mm. long; calyx campan-

ulate, about 6 mm. long, the 4 ovate-triangular teeth 0.5 mm. long;

corolla hypocrateriform, the cylindric tube nearly 4 mm. long, the

5 oblong-obovate lobes about 4.6 mm. long, 2-6 mm. wide; stamens 4,

inserted about 1.5 mm. below the throat, anthers sagittate, 1 mm.

long; style 5.7 mm. long, stigma branches 0.6 mm. long, ovary 4-

lobate, 4-celled. — Seems to be related to A. cuneata Mold, with short

setose petioles and densely hirsute many-flowered inflorescences (Mol-

denke). Type from a tree 6 meters tall, the fruit pink (collectors).

Loreto: Santa Rosa, below Yurimaguas, Killip & Smith 28858,

type.

Aegiphila umbraculiformis Mold. Phytologia 5: 225. 1955.

Stem unbranched, rather densely puberulent as the peduncles

(about 7 cm. long), the slender cyme branches and the linear bract-

lets (1.5-3 mm. long); leaves crowded-verticillate, entire, obovate,

attenuate to subamplexicaul base, acuminate, about 4 dm. long, 1.5-

nearly 2 dm. wide, membranous, paler beneath, glabrous, the 12 or

so slender lateral nerves and finely reticulate veins prominent only

beneath; inflorescence terminal, capitate, cymosely branched; pedi-

cels about 5 mm. long, sparsely puberulent; calyx campanulate, pu-

berulent, 3 mm. wide, 2 mm. long, the 5 triangular very acute lobes

about 1 mm. long; corolla unknown. — Shrub, 1-1.5 meters tall; no

relationship suggested by the author!

Huanuco: Deep forest, Tingo Maria, Stork & Horton 9495, type.

Aegiphila velutinosa Mold. Brittonia 1: 194. 1932; 437.

Subterete branches, branchlets, stout petioles (about 1 cm. long),

leaves beneath, panicle branches, peduncles (1-5 cm. long), pedicels

(1^1 mm. long) and subulate bractlets densely velutinous tomentose;

leaves oblong, acute at base, short-acuminate, 10-17 cm. long, 4-

6 cm. wide, subcoriaceous or membranous, the secondary nerves 7-11

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

pairs rather prominent beneath; inflorescence axillary and terminal;

cymes solitary, capitate, to 7.6 cm. long, 2.5 cm. wide, the narrow

thyrsoid panicle about 2 dm. long, foliose; calyx campanulate, nearly

6 mm. long, 4.5 mm. wide, strigosely velutinous, deeply lobed, the

4 triangular lobes acute; corolla glabrous, the tube about 7.7 mm.

long, widely ampliate, the apically rounded lobes 3.2 mm. long,

2.1 mm. wide; stamens at 2.5 mm. below the mouth; filaments pilose

below; stigma branches about 1.3 mm. long, densely pubescent; ovary

glabrous, subglobose. — Apparently related to A. chrysantha Hayek

but densely velutinous-pubescent, and to A. cordifolia (R. & P.)

Mold, (author).

Cuzco: Lucumayo Valley, (Cook & Gilbert 1382, type).

Aegiphila vitelliniflora Klotzsch in Walp. Repert. 4: 123. 1845;

421.

Branchlets strict, subterete; petioles opposite-decussate or approx-

imate, 4.5 mm. long, strigillose; leaves entire, membranous, oblong

or oblong-lanceolate, usually acute or subacute at base (sometimes

rounded), acuminate, 7-12 cm. long, 3-5.5 cm. wide, quite glabrous

and lustrous above except for strigillose midrib, sparsely strigillose

or appressed pubescent beneath especially on midrib and larger

veins, glandular disks few, the 6-8 pairs of slender secondaries retic-

ulate near margins; inflorescence axillary and terminal, the solitary

opposite somewhat corymbose cymes to 6.5 cm. long, 3 cm. wide, the

panicle pyramidal-thyrsoid, to 1.5 dm. long, 1 dm. wide, the many

pairs of cymes bracted, the peduncles to 6 cm. long, appressed pubes-

cent; pedicels strigillose, 1-1.5 mm. long; bractlets elongate-linear;

calyx turbinate or broadly campanulate, 2.6-5.2 mm. long, strigose

or pilose, 4-lobed or split, the ovate acute lobes 1 mm. long; corolla

yellowish or white, fleshy, the narrow tube at least 5 mm. long, pubes-

cent only within, the 4 ovate- or elliptic-Ungulate lobes about 3-

4 mm. long, acute; stamens 4 (5), at 1.5-3 mm. below mouth; stigma

branches 2.6-4.1 mm. long; ovary 4-lobate, glabrate; fruit oblong,

obtuse, smooth, 1-4-seeded, the cupulate accrescent calyx indurated.

— A 4-meter shrub that in Peru has been confused with A. filipes and

A. chrysantha; it seems possible, if Moldenke's specific lines are ac-

curate, that the Klotzsch plant is not in Peru, at least typically.

A. setiformis Rusby, 433, of northern Bolivia, has leaves usually

rounded at base, corolla said to be reddish-purple, the tube 6-7 mm.

long.

Peru (cf. note above). Bolivia; Brazil.

FLORA OF PERU 721

19. AVICENNIA L.

Reference: Moldenke, Phytologia 7: 123-168, 179-232. 1960.

Shrubs or trees, the branches and branchlets strongly nodose,

articulate, the leaves opposite, entire, persisting. Inflorescence de-

terminate, cymose, spicate or congested, when axillary generally gem-

inate. Calyx segments nearly free, the corolla tube not exserted, each

flower closely subtended by 3 imbricate scaliform bractlets (involu-

cral). Stamens 4, equal or subdidynamous. Ovary with a central

subulate placenta, 4 pendent straight ovules. Fruit a tomentulose

compressed capsule but juicy, somewhat fleshy, dehiscent by 2 valves

and regularly developing only a single seed. — The genus is sometimes

considered as constituting a family, Avicenniaceae; cf. note, under

family description. The dark brown hard wood is very durable.

The name recalls Ibn Sina, erudite Persian, 980-1037.

Avicennia germinans (L.) Steam, Kew Bull. 34. 1958; 179.

Bontia germinans L. Syst. ed. 10, 2: 1122. 1759. A. nitida Jacq.

Enum. PL Carib. 25. 1760. A. tomentosa Jacq. Sel. Stirp. Amer. Hist.

178, pi. 112, jig. 2. 1763. A. tomentosa Jacq. var. guayaquilensis

HBK. Nov. Gen. & Sp. 2: 284. 1818.

Branchlets and leaves above glabrous, the latter densely punctu-

late and beneath (at least in life) canescently and commonly com-

pactly furfuraceous tomentulose, oblong-lanceolate to elliptic or

obovate, more or less narrowed to or at base, obtuse or acute, to a

dm. long, 4 cm. wide or larger; spikes usually several cm. long, the

sessile white or yellow flowers mostly opposite, 1-2 cm. long; calyx

segments ovate, glabrous only within, 3-5 mm. long, equaling or ex-

ceeding the glabrous corolla-tube, this with lobes to 2.5 mm. long,

appressed pubescent without, tomentose within; stamens little ex-

serted; fruit somewhat oblique, about 1-2 cm. long, slightly more

than half as wide. — Steam, I.e. 33-37, gave a helpful key to the main

species of "mangrove" in the West Indies and northern South America.

Tumbez: Mangrove swamps, Rio Tumbez delta, Raimondi. Warm

American coasts to Florida and Texas.

LABIATAE Juss. Mint Family

Reference: Epling, Repert. Sp. Nov. Beih. 105: 1-93. 1938.

The familiar redolence of mint, opposite leaves on usually square

stems or branches and, characteristically, bilabiate flowers constitute

of course the distinctive traits of this group of plants, so rich in pleas-

ant products, particularly the herbs supplying taste or aroma to

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

foods; technically, only the Boraginaceae and Verbenaceae are sim-

ilar; in the former the ovule micropyle is ascending and the raphe

is divaricate, in contrast to the inferior micropyle and inward raphe of

the Labiatae. While the style is usually gynobasic in the mints, it is

sometimes terminal as in the Verbenaceae and in such cases a com-

bination of characters, of course, must be observed to separate the

connecting family tribes (Lawrence).

This account of the Peruvian members is compiled often nearly

verbatim from Carl Epling's reviews (as cited under genera) of the

South American species, works commendable for conciseness, clarity,

and sometimes frank presentation of specific problems, not under-

standable from purely morphological data. However, at the time

of his publications (1935-1939) continuities of specific relation-

ship in keys and discussions were frequently restricted to political

boundaries. Fortunately, as regards this work, many Peruvian species

have counterparts in adjacent areas and are earlier in publication

date; also, as noted in my preface, especially as regards the Andean

entities, the flora of Peru is remarkably distinctive. Most determi-

nations were by Epling and, of course, accepted by me.

The historical classification of Bentham continued by Briquet and

in our time by Epling — with customary nuances of thought prompted

by new data and (or) new interpretations — is seemingly correct; Ep-

ling devised a somewhat simplified key for the genera of northern

South America and for Peru (I.e. 1-2), (avoiding for convenience

staminal data), from which no important deviation has seemed to

me practical, even as to Peru above.

Lavender is from Lavandula vera DC., native to the Mediterra-

nean, a tomentose shrub with narrow spicate bluish flowers, the in-

cluded stamens free from the lower lip ; the name itself recalls sachet

bags and linens. Delightful mint (Mentha L.) is, naturally, from

delicious Mintha, who was transformed by Ceres into the plant; her

assuaging character, in beverage or sauce, thankfully, has not been

lost even if many have forgotten her fanciful origin.

Gordon P. De Wolf, Jr., Baileya 3: 47-57. 1955, gave a key and

interesting comment on the species of Lavandula, based on Miss

D. A. Chaytor's revision, Journ. Linn. Soc. Bot. 1937. The same

author has papers on Nepeta, I.e. 98-107. 1955, and Origanum and

allies, I.e. 2: 57-66. 1954; also on Mentha, I.e. 2: 3-11. 1954.

Stamens 4, unequal, lying on lower corolla lobe.

Flowers verticillate in racemes; lower lip of subequal segments

plane or nearly 1. Ocimum.

FLORA OF PERU 723

Flowers axillary, capitate-cymose or paniculate; lower lip more or

less saccate.

Calyx deflexed in fruit, upper tooth large; flowers axillary.

2. Coleus.

Calyx not or little deflexed, lobes or teeth not strongly diverse.

Nutlets thin erose-margined, concave-cymbiform.

3. Marsypianthes.

Nutlets angled or never thin-margined, often complanate.

4. Hyptis.

Stamens 4 (or 2 but anthers 2-celled, connective not filiform), straight

and remote or diverging.

Fertile stamens 4; suffrutescent or if herbaceous more or less erect.

Calyx as corolla obscurely or not bilabiate; stamens somewhat

exserted; herbs 5. Mentha.

Calyx as corolla more or less bilabiate; stamens little or not ex-

serted.

Calyx scarcely if at all accrescent; stamen connective deltoid.

Calyx 12-veined 6. Minthostachys.

Calyx 10-veined 7. Satureja.

Calyx reticulate-venose, soon accrescent; corolla tube annu-

late or pilose 8. Sphacele.

Fertile stamens 2; prostrate herb, leaves rounded 9. Hedeoma.

Stamens 2, parallel-ascending under upper lip, sometimes long-

exserted; anthers at least by confluence 1-celled.

Filament terminating in filiform connective of linear anther; leaves

linear only in S. rosamarinus 10. Salvia.

Filaments normal to anthers; leaves linear 11. Rosmarinus.

Stamens 4, the anthers often paired under the concave or arched

upper lip.

Calyx short, obscurely lobed and with a bump above.

12. Scutellaria.

Calyx 5- or 10-dentate.

Calyx teeth 10, spinose and recurved in fruit . . 13. Marrubium.

Calyx teeth 5, sometimes spinose and more or less recurved in

age.

Flowers purple or white.

Leaves never pinnatifid.

Corolla throat ampliate; nutlets truncate. .14. Lamium.

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

Corolla throat not or slightly ampliate; nutlets obscure.

15. Stachys.

Leaves pinnatifid 16. Leonurus.

Flowers yellow or orange 17. Leonotis.

1. OCIMUM L.

Sweetly scented herbs (Peru) with entire leaves, the floral re-

duced, bract-like, the flowers usually 6 in racemosely disposed verti-

cils, each blossom on an erect pedicel recurved at tip, reflexed in fruit.

Stamens 4, distinct, declinate on lower corolla tip, this entire, flat,

scarcely longer than the 4-lobate upper. Calyx campanulate, the

much larger often margined upper tooth recurved in fruit.

Basil or Sweet Basil as it is sometimes called, is esteemed inter-

nationally by cooks and gourmets.

Plants (at least in part) pubescent.

Corolla 3-5 mm. long 0. americanum.

Corolla 7-8 mm. long 0. basilicum.

Plants glabrous 0. micranthum.

Ocimum americanum L. Amoen. Acad. 4: 276. 1755; 72.

Branches crisply pubescent; petioles to 1 cm. long; leaves oblong-

elliptic, mostly obtuse, sparsely hirtellous, 1.5-3 cm. long, about half

as wide; bracts persisting, 3-4 mm. long; pedicels 2-2.5 mm. long;

calyx in flower 3 mm. long, in fruit 4-4.5 mm. long, upper tooth com-

planate, rotund, the lowest 1.5-2 mm. long, lanceolate-subulate;

corolla 3.5 mm. long, upper lobes subequal, lower 1.5-2 mm. long;

nutlets 1.5 mm. long. — Otherwise similar to 0. basilicum. Illus-

trated, Bot. Mag. 51: pi. 2&>2 (as 0. canum).

San Martin: Tarapoto, Williams 5399. Warm America. "Shara

masha" (Williams).

Ocimum basilicum L. Sp. PI. 597. 1753; 72.

Branches hispidulous; petioles to 1.5 mm. long; leaves subentire,

ovate-lanceolate, shortly acuminate, gl abrous or sparsely pilose toward

the rounded base, 3-5 cm. long; flowers in 3's, mostly deflexed in age

as the ciliate bracts (4-6 mm. long); pedicels to 3 mm. long; calyx

in flower campanulate, 4 mm. long, the upper tooth rounded-cordate,

decurrent, the lateral ovate, obtuse, mucronate, shorter than the

FLORA OF PERU 725

pungent lower, the mature calyx hispid-ciliate, reticulate venose,

the upper tooth 5 mm. across, the incurved lower 2.5 mm. long;

corolla 7 or 8 mm. long, lower lobes about 4 mm. long, tube glabrous

within; stamens subexserted, the upper filaments dentate-append-