V OF

ILLINOIS LIBRARY

AT URBANA-CHAMPAIGN

BIOLOGY

NOV 0 6 1996

FLORA OF PERU

BY

J. FRANCIS MACBRIDE

CURATOR, PERUVIAN BOTANY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-B, NUMBER 1

MAY 11, 1962

PUBLICATION 951

FLORA OF PERU

BY

J. FRANCIS MACBRIDE

CURATOR, PERUVIAN BOTANY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-B, NUMBER 1

MAY 11, 1962

PUBLICATION 951

Library of Congress Catalog Card Number: 36-10^26

5?o. 5

H

FLORA OF PERU

J. FRANCIS MACBRIDE

SOLANACEAE (Juss.) Lindl.

Herbaceous or ligneous, occasionally arborescent but various in

habit. Leaves alternate but often geminate and then usually un-

equal, rarely pseudo-opposite «r verticillate, simple or rarely pin-

nate, or even bipinnate. Flowers solitary to many, often disposed

in cymes (one way or another modified), hermaphrodite, infrequently

zygomorphic. Sepals commonly persisting, rather often accrescent.

Corolla rotate to tubular, rarely bilabiate, ordinarily 5-merous, aesti-

vation exceptionally imbricate, generally plicate, now and then con-

volute. Hypogynous torus seldom lacking. Ovary 2 (-5) -celled,

ovules generally anatropous. Stamens on tube, alternate with lobes,

didymous in one section only. Fruit indehiscent or dehiscent, dry to

very wet, the seeds usually many.

Seven groups indicating presumed relationships according to

Bentham and Hooker f., Wettstein, and, recently, Sleumer (Lilloa 23:

118. 1950) have been defined as to basic character in the generic key.

; To avoid dependence on the not readily observed character of the

•f embryo, that of aestivation (it too not always discernible) has been

Caused as a supplementary key-aid; much remains to be done on the

^ classification, as corolla form and calyx accrescence both occur in

many degrees and are in themselves of questionable significance.

Charles Baehni in 1946 (Candollea 10: 399-494) recorded his ob-

servations on the opening of Solanaceous flower buds.

Vassobia dichotoma (Rusby) Bitter, Repert. Sp. Nov. 15: 150. 1917

(Cyphomandra dichotoma Rusby, Mem. Torrey Bot. Club 4: 231.

1895), with detailed description (type from Yungas, Bolivia, Bang

519}, and also Coranitales by Hertzog, if found is, fide Sleumer, near

Dunalia but has flowers apically fascicled ; Rusby's description notes

a terminal inflorescence, laxly fasciculate but composite, 1.5-2 dm.

long and wide, corolla 10-14 mm. long, 16 mm. wide, purple, anthers

subsessile, berry ellipsoid, 3^4 mm. long on pedicels at least 2 cm.

long; probably V. atropioides Rusby, Bull. N. Y. Bot. Card. 4: 422.

1907, is the same.

3

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

Stamens 5 and all fertile (abnormally 1 reduced).

Aestivation of corolla lobes more or less imbricate; embryo strongly

curved except Marked and Juanulloa.

Corolla small; branched, often spiny shrubs (Atropeae).

Fruit drupaceous, with 4 nutlets; leaves usually subrotund.

1. Grdbowskia.

Fruit berry-like, often 8-seeded; leaves usually narrow.

2. Lycium.

Corolla elongate, contracted or ampliate apically; subsimple

shrubs or half -shrubs, often subscandent (Cestrinae).

Corolla amplification continuous 16. Marked.

Corolla usually contracted at apex 17. Juanulloa.

Aestivation valvate or induplicate-valvate, limb sometimes sub-

equally plicate; embryo strongly curved; fruit usually a suc-

culent berry (Solaneae).

Anther dehiscence by apical pores, cells sometimes joined, ex-

tended.

Connective of anther cells slender or not gibbous.

27. Solanum.

Connective of anther cells stout, more or less gibbous dorsally.

4. Cyphomandra.

Anther dehiscence longitudinal, sometimes early by chinks,

finally to base or nearly (cf. Solanum Pennellii, S. lyco-

persicoides and section Lycopersicum) , anther cells extended.

Calyx clearly accrescent in fruit, growth even early apparent.

Calyx appressed, not inflated nor costate or obscurely.

Flowers rotate or applanate-complanate, fasciculate.

5. Withania.

Flowers funnelform, dilated.

Section lochroma in 9. Dunalia.

Calyx inflated or lax; fruit included, rarely visible between

lobes (cf. Juanulloa}.

Flowers usually fasciculate, narrowly and deeply lobed.

5. Withania.

Flowers single or geminate.

Ovary 2-celled; calyx not or little alately angled; plants

often pubescent.

Corolla rotate or applanate-campanulate.

6. Physalis.

FLORA OF PERU 5

Corolla tubular-campanulate or broadly funnelform.

7. Cacabus.

Ovary 3-5-celled; calyx alately angled; plants glabrate

(Nicandreae) 3. Nicandra.

Calyx more or less spreading; fruit exserted.

Corolla applanate-campanulate.

8. (Hebecladus) Saracha.

Corolla tubular-funnelf orm 9. Dunalia.

Calyx not at all or scarcely accrescent even below mature fruit,

in Datura caducous, partly persisting or circumscissile and

base somewhat accrescent.

Corolla tube well-developed or corolla funnelform or large,

openly campanulate; stamens often longer than tube;

leaves never imbricate.

Ovary 2-celled; calyx not elongate-tubular, 5-denticulate,

truncate or cleft.

Calyx truncate or early subevenly 5-denticulate; co-

rolla more or less tubular or funnelform.

9. Dunalia.

Calyx unevenly, rarely shortly, and medially 5-6-cleft;

corolla campanulate 10. Poecilochroma.

Calyx cleft, often nearly to base.

Leaves entire 11. Salpichroa.

Leaves not entire 12. Jaborosa.

Ovary pseud o-4-celled by placentae; calyx long-tubular,

sometimes spathaceous (Datureae) 15. Datura.

Corolla rotate or short-campanulate; stamens not or scarcely

longer or leaves imbricate (Solaneae).

Leaves simple; flowers 1-few, or rarely many and fas-

ciculate.

Leaves imbricate; stamens exserted . . 13. Witheringia.

Leaves never imbricate; stamens not or little exserted.

14. Capsicum.

Leaves compound; flowers often in cymes, corymbs or

racemes; anther dehiscence rarely or tardily longi-

tudinal, a few species, mostly section Lycopersicum.

27. Solarium.

Aestivation of corolla lobes plicate, induplicate-valvate or imbri-

cate; embryo straight or somewhat curved; (known) fruit cap-

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

sular, baccate or a berry but often nearly dry (cf. Datura),

(Cestrinae).

Corolla lobes equal, imbricate, tube ampliate; scandent or lax,

flowers few; calyx often enclosing fruit.

17. Juanulloa, 16. Marked.

Corolla lobes unequal or induplicate-valvate, tube slender; erect

or low half -shrubs or herbs; fruit exserted.

Flowers usually many in each inflorescence.

Seeds usually few; fruit indehiscent or valvate, often tar-

dily; shrubs or trees 18. Oestrum.

Seeds usually many; herbs, often in Peru tall, subligneous

or rarely ligneous 19. Nicotiana.

Flowers solitary, terminal or lateral.

Leaves minute, closely imbricate 20. Fabiana.

Leaves not minute, lax if crowded 21. Nierembergia.

Stamens 2-4 fertile, always unequal (Salpiglossidae).

Herbs or ligneous below in age; corolla bilabiate at least slightly.

Annual or becoming half-shrubs; corolla yellow or greenish or

white with yellow center.

Corolla tube ampliate 22. Salpiglossis.

Corolla tube (Peru) subfiliform 23. Schwenkia.

Annuals, evanescent; corolla blue or purple, rarely white.

24. Browallia.

Shrubs or trees.

Leaves rugose; corolla tube spiralled 25. Streptosolen.

Leaves not rugose; corolla tube straight 26. Brunfelsia.

1. GRABOWSKIA Schlecht.

Shrubby plants with stout rigid axillary spines and fleshy or firm

rotund leaves, similar in habit to some Lyciums but the bony 2-celled

fruit with 4 stones or nutlets (pyrenes), these 1-2-seeded. — Bitter,

Bot. Jahrb. 54, Beibl. 119. 17. 1916, described the fruit as a drupe

instead of a berry because of the stony concretion about the seeds

except at base, this with 3 teeth (L'He'ritier).

Grabowskia boerhaaviaefolia (L. f.) Schlecht. Linnaea 7: 71.

1832. Lycium boerhaaviaefolium L. f. Suppl. 150. 1781. L. hetero-

FLORA OF PERU 7

phyllum Murr. Comm. Goett. 6, pi. 2. 1783. Ehretia halimi/olia

L'He>. Stirp. Nov. 45, pi. 23. 1785.

A glaucous-leaved orange-berried shrub with arching branches

(Svenson), the leaves ovate to subrotund, often shortly acuminate,

1.5-3.5 (4) cm. wide; flowers in short corymbs or panicles, violet or

white with yellowish or green veins in the pubescent throat; calyx

subtruncate, acutely 5-denticulate, 4 mm. long; corolla, with reflex-

ing lobes, about 10 mm. long; stamens exserted; fruit 7-8 mm. long.

— According to Oscar Haught this curious endemic shrub (1-3 meters

high) is a common and characteristic plant of the coast of Piura;

generally in the northern desert region (Weberbauer, 106; 153; 161).

Dammer, Bot. Jahrb. 50, Beibl. 111. 52. 1913, reported Weberbauer

3783 as the first record outside of cultivation; it was grown at Paris

from seeds sent by Joseph Jussieu (Lamarck Encycl. 3: 510. 1791);

perhaps Lycium salsum R. & P. from Lima was cultivated, if, as

possible, it is the same. Illustrated, Miers, 111. S. Am. PI.

Piura: Parinas Valley, Haught 79. Near Hacienda Nomala, Web-

erbauer 591f.5; 153. Nigritos, (Haught & Svenson 11609). Talara,

Haught 15 (det. Weberbauer); Johnston 3506. — Cajamarca: Tem-

bladera, 400 meters, Weberbauer 3783. — Lima: In rocks, Chosica,

4.86; Nunez 1870; Soukup 3797. Fifth km. east of Lima, Goodspeed

33085. Galapagos. "Gabonilla."

2. LYCIUM L.

Reference: C. L. Hitchcock, Ann. Mo. Bot. Gard. 19: 179-374.

1932.

Peruvian species spiny shrubs with small, often fascicled leaves

and 1-several axillary narrowly funnelform flowers, the lobes imbri-

cate in bud. Filaments unappendaged, sometimes pubescent at base

or ciliate-glandular, usually inserted below the middle of the corolla,

this often pubescent within toward base. Ovary 2-celled, the 2 car-

pels usually undivided, the more or less berry-like fruit 1-many

seeded. Embryo curved, at least nearly half- turned.

Calyx lobes at least two-thirds as long as tube or 2 mm. long or longer,

the lobes equal.

Corolla glabrate without, but lobes usually ciliate . . . . L. fragosum.

Corolla slightly pubescent toward base L. distichum.

Calyx lobes less than two-thirds as long as tube, usually shorter than

2 mm., often unevenly cleft and somewhat 2-lipped . L. nodosum.

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

Lycium distichum Meyen, Reise 1 : 448. 1834; 263. Grabowskia

disticha (Meyen) Nees ex DC. Prodr. 13, pt. 1: 527. 1852. L. oreo-

philum Wedd. Choris And. 2: 108. 1857. L. leiostemum Wedd. I.e.

L. divaricatum Rusby, Bull. N. Y. Bot. Card. 8: 117. 1912.

An open much-branched shrub, the branches or branchlets more

or less elongate, early densely cinereous pubescent, mostly at least

the young shoots apically armed with slender sharp spines; leaves

oblong-ovate to obovate or linear-spathulate, rounded to acute, pu-

berulent and glandular-tomentulose, 4-12 mm. long, 1-3 mm. wide,

those at base of branches commonly ovate, single or geminate, the

younger upper narrower, 1-3-fasciculate, the old nodes enlarged;

pedicels solitary at nodes 3-7 mm. long; calyx campanulate, 3-4 mm.

long, sparsely pubescent, the lanceolate acute ciliolate lobes sub-

equaling the tube; corolla obconic-tubular, the tubular portion 11-

13 mm. long, about 3 mm. across at apex, 1.5-3 mm. at base, more

or less pubescent near calyx lobes, the (4) 5 rounded lobes glabrous,

about 1.5 mm. long; filaments subequal, adnate below middle of tube

and pilose for 2 mm. above the adnate portion, the corolla tube pu-

bescent also for 4 mm. below this free portion especially along the

vascular traces of the filament (Hitchcock), or filaments and corolla

tube glabrous or with 1 or 2 trichomes within (L. leiostemum) or the

stamens inserted above the middle of the corolla (L. oreophilum);

style equaling stamens, stigma nearly 1 mm. broad; berry reddish-

purple, 6-10 seeded. — Corolla white except lavender limb (Worth &

Morrison). According to Hitchcock (from whom the synonymy is

taken), I.e., Nees wrote L. (Grabowskia) distichum (Meyen) Nees,

Nov. Act. Acad. Caes. Leop. 19, Suppl. 1: 389. 1843. Hitchcock,

I.e. 265, has carefully observed the apparent differences, at least

ex char, between the types of Weddell and Meyen but the problem

can scarcely be solved until more collections are available. Illus-

trated, Hitchcock, pi 16, fig. 26. F.M. Neg. 2461.

Cuzco: Lahumarmaqui, Urubamba, Vargas 7865; Gay (type, L.

leiostemum, Paris). — Arequipa: Mollendo, Weberbauer 1520 (ex char.

L. leiostemum); Worth & Morrison 15768 (probably but calyx not

typical); Pampa La Joya, Ferreyra 6425. — Puno: Yura, (Williams

2554, type, L. divaricatum, New York). — Tacna: Cordillera de Palca,

Weddell (type, L. oreophilum, Paris). Cordillera Tacna, 2000 meters,

Meyen (type, Berlin).

Lycium fragosum Miers, Ann. Mag. Nat. Hist. 2, 14: 132. 1854;

266.

FLORA OF PERU 9

Resembles L. distichum; leaves linear to spatulate, 2-15 mm. long,

0.5-1.25 mm. wide, densely hirtellous-puberulent, 3-8-fasciculate,

frequently revolute; calyx 2.5-3.5 mm. long, the 4 lobes nearly as

long as the tube; corolla glabrous without, the 4 oval lobes 1-1.5 mm.

long; filaments adnate medially or less, pilose the first 2-3 mm. of

their free portion, the tube also slightly pubescent adjacent to and

below the free portion; style 1-2 mm. shorter than stamens. — As

suggested by the monographer himself the type of Miers should prob-

ably be included within the probable variation range of L. distichum

Meyen; it was treated as a variant of L. salsum R. & P. (that is,

L. nodosum) by Terraciano, Malpighia 4: 530. 1891. Type Cuming

948 bears data in British Museum "Cobija, Iquiqui et Arica," (Hitch-

cock) and of course possibly extends into adjacent Peru. Illustrated,

Hitchcock, 358, pi. 16, figs. 10-12 (flowers).

Peru (see note above). Chile.

Lycium nodosum Miers, Ann. Mag. Nat. Hist. 2, 14: 139.

1854; 221. L. salsum R. & P. Fl. Peruv. 2: 46, pi. 183a. 1799, not

Bartram, 1792. L. Tweedianum Griseb. Abh. Konig. Ges. Wiss. Got.

19 : 216. 1874, fide Barkley, Lilloa 26 : 206. 1953. Citharexylum flexuo-

sum D. Don, var. subglabrum Mold. Repert. Sp. Nov. 137: 222. 1937.

L. subglabrum Mold. Phytologia 4: 292. 1953; 6: 353. 1958?

Much-branched shrub with a few short spines and often with

prominent white-pilose or -puberulent nodal (and even spinal) short

shoots; stems gray, brown in age, glabrate; leaves obovate-spatulate,

or obovate and rounded-obtuse to linear-elliptic and subovate, gla-

brous, 4-40 mm. long, (2) 3-11 mm. wide, cuneate and decurrent into

petiole 4-13 mm. long; flowers 1-3 at nodes, pedicels 3-13 mm. long,

glabrous as calyx (unless lobes), this with cupulate tube 13 mm.

long, 2 lips 2-3 mm. long and lobes 0.5-1.5 mm. long or not 2-lipped,

the lobes then 0.4-1.5 mm. long, broadly triangular, subciliate to

ciliate; corolla densely pilose within, around or between stamen in-

sertion, the tube 4-7 mm. long, the lobes 2-5 mm. long, 2-2.5 mm.

broad, usually ciliate; stamens exserted, attached 2-5.5 mm. above

base of tube, filaments more or less pilose, style 8-9 mm. long; fruit

spherical, red, 3-5 mm. thick. — Description and synonyms by Bark-

ley, who decided that L. nodosum and L. Tweedianum as delimited

by Hitchcock represent the two extremes of a very variable species;

Svenson, Amer. Journ. Bot. 33: 483. 1946, noted some variation in

size of flower parts in Ecuadorian plants in contrast to those from

Argentina, both forms or variants however described as having green

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

flowers and rounded obtuse fruit while the plant from Lima ex char,

has pale reddish to deep lavender corolla, viscous elongate reflexed

lobes and very elongated and pointed fruit. Accordingly, more in-

vestigation may show that L. nodosum Miers, also of Argentina, is

not the same as L. salsum R. & P., in which case the latter apparently

will require a new name. The native name signifying a salty spine

refers to the taste of the leaves (Ruiz & Pavon) . Illustrated, Miers,

111. S. Amer. PI. 2: 115, pi. 69a; page 131, pi. 72c (L. salsum} ; Hitch-

cock, 356, figs. 16-18; figs. 1-3 (L. Tweedianum).

Lima: San Juan de Chicla, (Anderson). Rio Blanco, Killip &

Smith 21679 (det. Moldenke, L. subglabrum). Lurin, (Mathews 450);

Ruiz & Pavdn, (type, L. salsum). Matucana, 2885 (fide Moldenke

is L. subglabrum). Near Lima, Wilkes Exped.; Nation. San Augustin,

Weberbauer 5226. Argentina and Paraguay, coastal Ecuador, Colom-

bia to Venezuela (Barkley). "Cachicasa."

3. NICANDRA Adans. Apple of Peru

Calydermos R. & P. Fl. Peruv. 2: 43. 1799.

A smooth or nearly smooth annual with ovate sinuate-toothed

leaves and solitary peduncled rather large blue or lavender flowers.

Calyx enlarged and bladder-like as in Physalis but the thin-walled

yellow berry 3-5-celled and nearly or quite dry. — The often irregu-

larly serrate leaves distinguish it in flower from Saracha species in

Peru; in fruit of course the accrescent calyx differentiates it from

Physalis; in flower it is marked by the showy blossom. Wettstein

isolated this as a subgroup, showing that the ovules are borne on

unevenly lobulate placentae (Pflanzenfam. IV, Abt. 3: 11, fig. 5.

1895). Practically, in floristic work, it must be keyed in relation-

ship to Physalis L. which it so closely resembles.

Could be a poet's inspiration, in flower or fruit; indeed, named for

one, Nicander of Colophon, Ionia (Asia Minor).

Nicandra Physalodes (L.) Gaertn. Fruct. 2: 237. 1791. Atropa

physalodes L. Sp. PI. 181. 1753. Physalis datura(e)folia Lam. Diet. 2:

102. 1786. C. erosus R. & P. I.e. Physalodes peruviana [Mill.] Ktze.

Rev. Gen. 452. 1891.

A common bushy plant of dryer, usually disturbed soils (Weber-

bauer, 106; 107). Native to Peru, it has long been grown for orna-

ment and now and then persists after cultivation.

FLORA OF PERU 11

Cajamarca: San Miguel, Ferreyra 7080; 5308; Weberbauer, 188.—

La Libertad: Chicama Valley, Smyth 35; 52 (det. Killip). — Lima:

Along Rio Chillon, Pennell 1W1 . Huara, Soukup 1004- Matucana,

295. Surco, Nunez 2687. Above Lima, (Weberbauer, 166) ; (Ruiz &

Pavori). — Huanuco: 20^1; Pavon; Stork & Horton 9365. — Apurimac:

Pachachaca Valley, Goodspeed Exped. 10565. — Cuzco: Anta, Vargas

198. Valle de San Miguel, Herrera 1990. Torontoi, Cook & Gilbert

822. — Arequipa: Mejia, (Gunther & Buchtien 108}. "Capuli cimar-

ron," "ccarapamacmam," "jarrito," "orzita de pellejo," "toccoro."

The genus Solarium, which usually would be treated at this point,

has been placed at the end of the family. (Ed.)

4. CYPHOMANDRA Mart.

Pionandra Miers in Hook. Lond. Journ. Bot. 4: 358. 1845.

Cyathostyles Schott ex Meissn. Gen. Comm. 184. 1840, nomen nudum.

Reference: Dunal in DC. Prodr. 13, pt. 1: 387, 402. 1852.

Shrub or small trees with entire, lobed, or pinnately divided

leaves; distinguished from Solarium chiefly by the more or less dor-

sally thickened (gibbous) connective of the anthers. Leaves, espe-

cially when entire, usually somewhat cordate, often obliquely. Fruit

ovoid or oblong, many-seeded, sometimes large.

This segregate or expedient genus, while not always clearly

defined, usually, at least in Peru, has a distinctive facies either

because of the leaves, the subsimple or remotely branching and

elongating inflorescence, the often large fruits (edible when cooked),

or a combination of these characteristics. Solarium incurvum R.

& P. was referred here by Werdermann in Herb. Madrid.

As most groups in this family — at least in Peru as so many

others — this requires revision by a qualified taxonomist with time

and means for the prolonged research necessary to even partial

understanding of specific relationships; as usual, recently proposed

species may prove to be the same as Amazonian ones.

KEY TO CYPHOMANDRA

Leaves more or less obliquely cordate at base (if obscurely, indument

somewhat glandular), or not entire, often broadly ovate.

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

Leaves entire (herbarium material seen), obtusely or acutely acu-

minate or cuspidate.

Leaves puberulent at least veins beneath; corolla segments

ovate-lanceolate.

Leaves uniformly pulverulent or finely puberulent beneath.

C. crassifolia.

Leaves puberulent on the veins beneath (type) . . . . C. obliqua.

Leaves quite glabrous at least beneath, where papillose; corolla

segments lanceolate or sublinear.

Corolla segments lanceolate.

Anthers linear-oblong C. splendens, C. coriacea.

Anthers broader C. obliqua.

Corolla segments sublinear C. endopogon.

Leaves, lobes or divisions (these present, at least in some leaves)

acutely and narrowly caudate-acuminate or /and pubescent.

Branchlets as leaves, at least on nerves, shortly pubescent,

rarely hirsutulous, some trichomes gland-tipped unless C.

viridiflora.

Corolla purplish, more or less puberulent . C. Ulei, C. pendula.

Corolla green, lobes lanate marginally C. viridiflora.

Branchlets as often leaves long-setose; leaves (in part, type)

3-lobed C. tenuisetosa.

Leaves acute or more or less rounded or if oblique at base, rather

oblong-elliptic; indument lax, eglandular S. incurvum.

Cyphomandra crassifolia (Ortega) Kuntze, Rev. Gen. 3, pt. 2:

220. 1898; 393. Solanum crassifolium Ortega, Dec. 9. 117. 1797.

S. betacea Cav. Icon. 6: 15, pi. 524. 1801. C. betacea (Cav.) Sendt.

Flora 28: 172, pi. 6. 1845. C. betacea (Cav.) Sendt., var. wlutina

Dunal in DC. Prodr. 13, pt. 1: 394. 1852?

A small tree usually about 3 meters high with ample fleshy

cordate-ovate long-petioled (2-3 cm.) leaves, softly pubescent at

least beneath, and more or less pendulous waxy pink-tinted glabrous

flowers; cymes simple, bifid or trifid, lateral or axillary, subequaling

the often reddish-violet petioles; calyx subcampanulate, semi-parted,

the broadly ovate subacute segments thicker and accrescent in fruit;

corolla rotate-campanulate, to 12 mm. long, the lanceolate concave

segments ciliolate, apically reflexed; stamens about 6 mm. long,

FLORA OF PERU 13

short filaments terete, anthers oblong, 5 mm. long, minute apical

pores slightly inflexed, the fleshy connective linear-oblong, rounded

at base, not calloused; ovary conical, style subcylindric, somewhat

incrassate to the small glandular stigma; fruit egg-shaped, to about

5 cm. long, maturing reddish- or brownish-yellow. — Apparently

known in Peru only in cultivation. Miers, Hook. Lond. Journ. 4:

359. 1845, records finding it "in the markets of Lima where it is

commonly used for cooking in lieu of the ordinary tomate, the

flavour of which it greatly resembles." In central Peru it is not

unusual to find one or two trees near dwellings. According to

Herrera it is used in making "dulces."

Amazonas: Near Chachapoyas, Ferreyra 7109. — San Martin:

Juanjui, Klug 4192. — Huanuco: Tingo Maria, Stork & Horton 9559.

Puente Durand, Mexia 8235. Near Muiia, 3875. — Loreto: Yurima-

guas, Williams 4198. Rio Nanay, Williams 391. Contamana,

Killip & Smith 26872. La Victoria, Williams 2615; 2827; 3021 —

Ayacucho: Weberbauer 5494 (det. Bitter). — Cuzco: La Maquina,

West 8044- Pomontama, Herrera 2984- Machupicchu, Ferreyra

2708. Ollantaitambo, Cook & Gilbert 770. Torontoi, Cook &

Gilbert 1101. Without locality, (Pawn). South America. "Tomate,"

"tomate de la Paz," "pimiento," "tomate del Campo," "gallinazo

panga" (Williams); "berenjena" (Ferreyra).

Cyphomandra coriacea (Miers) Dunal in DC. Prodr. 13, pt. 1:

401. 1852. Pionandra coriacea Miers in Hook. Lond. Journ. Bot. 4:

363. 1845.

A shrub with very heavy coriaceous glabrous ovate-cordate leaves

(petioles 5 cm. long), the larger often more than 1.5 dm. long, at

least 1 dm. wide, the smaller more than half as large; petiole and

basal leaf -lobes subequal; flowers secund in short racemes, the

crowded pedicels articulate; corolla- tube short, the divisions lan-

ceolate; anthers linear, erect, the connective fleshy. — Apparently

this is the earlier name for C. splendens Dunal of Ecuador but it

was imperfectly or incompletely described.

Amazonas: (Mathews 1971, type).

Gyphomandra endopogon Bitter, Bot. Jahrb. 54, Beibl. 119:

16. 1916.

A small glabrous (except flowers) tree to 7 meters tall; leaves

alternate or geminate, unequal, lustrous both sides, entire, sub-

coriaceous, broadly or obliquely ovate-cordate, the larger 1.5 dm.

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

long or longer, about two-thirds as wide, the smaller often only

half as large or less; petioles 2-7.5 cm. long; inflorescence greatly

elongating, usually forking, many- (often 30-50) flowered, the pe-

duncle 5 cm. long or longer; calyx 2 mm. long, 5 mm. across, the

obtuse lobes marginally puberulent, the open corolla greenish (early

pale violet), rotate-stellate, 3.5-4 cm. wide, deeply lobed, the seg-

ments 16-18 mm. long, to 2 mm. wide or little wider, pilosulous

especially toward apex or marginally; stamens about 1 mm. above

the base, anthers lanceolate, about 9 mm. long, the prominent

connective densely papillose, the anthers shortly; ovary as style

glabrous, both 9 mm. long, the stigma lobed, 2.5 mm. broad.—

When C. coriacea (Miers) Dunal and C. splendens Dunal are better

known one instead of two or three species may be found, in reality.

F.M. Neg. 2932.

San Martin: San Roque, Williams 6939. Near Juanjui, Ferreyra

4407. — Huanuco: Valley of the Rio Mayro, from the Rio Palcazu,

Weberbauer 6757, type. Tingo Maria, Ferreyra 959. — Loreto: Middle

Rio Blanco, Tessmann 3506 (det. Werdermann). Yurimaguas, Killip

& Smith 28182. Near Iquitos, King 2560 (det. Standley); Killip &

Smith 29863; Williams 3716. La Victoria, Williams 2564; 2917.

Solatium incurvum R. & P. Fl. Peruv. 2: 34, pi. 164- (115?)

1799; 155. S. recurvum Poir. Encycl. Suppl. 3: 754. 1814.

Stems glabrous, angled, nodosely articulate, to 2 meters high;

leaves ovate-acuminate, rounded, acute at the little inequilateral

base, the largest seen 13 cm. long, 6 cm. wide, long-petioled (petioles

hirsute, 5-6 cm. long), shortly crisply pubescent on both sides, prob-

ably glabrate in age, venose beneath; racemes axillary, geminate,

bifid; pedicels hirsute, incurved, violet-tinged, about 1.5 cm. long,

rusty viscid-pilose as the calyx and young leaves; calyx campanulate,

truncate, minutely dentate, 3 mm. deep; corolla pale violet, three

times longer than the calyx, or the segments 8 mm. long, acutish. —

Flowering stem apparently subherbaceous in type but recent collec-

tions are shrubs, open in growth; my 4491, in young fruit except for

two flowers, is somewhat doubtful as to true character of stamens;

the inflorescences are binate, peduncles to 5 cm. long, pedicels to

3 cm. long, corolla 1 cm. long, lanate without, young berries ellipsoid,

apiculate; its habit suggests Solanum and indeed it was placed by

Bitter in his subgroup Bassovioides but its facies is that of S. hederi-

radiculum Bitter and the berries suggest S. conicum R. & P. or one

FLORA OF PERU 15

of its relatives, so I have avoided using the implied transfer of

Werdermann in herb. Madrid to Cyphomandra.

Huanuco: Muna, Ruiz & Pav6n, type. Playapampa, 4491? —

Ayacucho: Choimacota Valley, 2,800 meters, Weberbauer 7370.

Cyphomandra obliqua (R. & P.) Sendt. Flora 28: 172. 1845;

393. Solanum obliquum R. & P. Fl. Peruv. 2: 35, pi. 165, fig. a. 1799.

Pionandra obliqua (R. & P.) Miers in Hook. Lond. Journ. Bot. 4:

359. 1845.

Stems and leaf-nerves typically minutely puberulent above and

below; leaves very oblique at the cordate base, acuminate, 1.5-2 dm.

long, 8-12 cm. wide; flowering pedicels 5 mm. long; calyx poculiform,

4-6 mm. long and broad, obtusely 5-denticulate (type); flowers

8 mm. long, 2 cm. across, segments lanceolate, acute, purple-violet,

including the connivent anthers; berries "oblong," apparently small.

— Leaves strongly oblique; stigma (figure) bifid. The Schunke and

Klug specimens have quite entire calyces (suggesting those of the

Colombian C. naranjilla Pittier, Contr. U. S. Nat. Herb. 13: 117.

1910); the Klug specimens have glabrous membranous leaves; cf.

also C. splendens. F.M. Neg. 12993.

San Martin: Tarapoto, Spruce 4229? — Huanuco: Chinchao, Haci-

enda San Carlo, Stork & Horton (det. Morton, with query). — Loreto:

Rio Mazdn, Jose Schunke 357 (det. C. crassifolia?, Standley). Near

Iquitos, Klug 484-

Cyphomandra pendula (R. & P.) Sendt. Flora 28: 173, pi. 7.

1845 ; 395. Solanum pendulum R. & P. Fl. Peruv. 2 : 39, pi. 1 74, fig. a.

1799. Pionandra pendula (R. & P.) Miers in Hook. Lond. Journ.

Bot. 4: 359. 1845.

A soft woody, single-stemmed glabrate or hirsute-pubescent and

unpleasantly scented plant to 6 meters high, branched only at the

summit; petioles to 2 dm. long, 1 cm. thick; leaves in type ternate,

but also simple or 2-11-pinnatifid, often 5-8 dm. long, 5 dm. wide,

the lowest leaflets petiolulate, the ovate-cordate segments entire,

oblique at base, acuminate, densely or sparsely pubescent with sim-

ple trichomes especially beneath; peduncles from the axils of the

branches forking remotely and greatly elongating; pedicels about

1 cm. long; calyx-divisions rotund, sub truncate, medially apiculate;

corolla 1.5-2 cm. long, rather funnelform, purplish, 4 times longer

than the calyx; lobes lanceolate, acute, reflexed apically; anthers ven-

tricose, erect; fruit large, yellow, at maturity as large as an egg. —

Werdermann leaves this in Solanum in Herb. Madrid, and the con-

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

nective, while broad, is scarcely thickened; it seems to me that Mar-

tius' genus should be more narrowly restricted, if retained. A species

of low woods. F.M. Negs. 2936; 12995.

Huanuco: Puente Durand, Mexia 824-5. Huacachi, near Mufia,

4141,' Weberbauer 6720. Prov. of Panatahuarum, especially at

Muna, Ruiz & Pavdn. — Loreto: Paca, on the Ucayali, (Huber 1569).

— Cuzco: Anta, Vargas 200. "Puruma," "papaya del monte."

Cyphomandra splendens Dunal in DC. Prodr. 13, pt. 1: 395.

1852.

Nearly glabrous, the type with vivid brownish-purple half-climb-

ing branches, the distinctly and unequally cordate-based acuminate

leaves intensely green above; petioles sulcate, to about 4 cm. long;

leaves geminate, nerved from base, reticulate-veined, rather fleshy,

sparsely punctiform puberulent above, minutely papillose beneath,

the larger 12-17 cm. long, usually about half as wide, the smaller

sometimes only a third as large; racemes branched (always?), gla-

brate or pulverulent, a dm. long or longer, the basally articulate

pedicels 1.5 cm. long or longer; calyx purplish, apiculately 5-crenate-

dentate, 4-6 mm. across; corolla 5-parted, the narrowly lanceolate-

oblong acuminate segments pubescent marginally toward apex, 10-

12 mm. long; anthers 6 mm. long, connective purple, puberulent,

linear-acuminate, basally dilated, gibbous; stigma obconic. — Type

collection from Guayaquil as noted on Ruiz and Pavon (Tafalla)

specimen in Herb. Madrid; Dunal's description from sheet in Geneva

(Herb. Boissier), accredited to Peru, where it may occur under the

name C. coriacea (Miers) Dunal. F.M. Neg. 8561.

Peru (possibly). Ecuador.

Cyphomandra tenuisetosa Bitter, Repert. Sp. Nov. 17: 352.

1921.

Type a small tree 2-5 meters high with ample diverse firm sub-

lustrous leaves ovate-cordate and entire to deeply 3-lobed, the lateral

lobes lanceolate-acuminate, the terminal rounded; pubescence in part

minutely stipitate glandular but prominently on branchlets, petioles

and leaves, especially above on the veins, subpatently long-setose

(trichomes 2-3 mm. long) ; inflorescence axillary, 20-30-flowered, pe-

duncle 5.5 cm. long; pedicels about 2 cm. long (2.5 or longer in fruit),

nearly glabrous as the calyx, this broadly campanulate, 1 cm. across,

the very broadly ovate lobes unequally (2 lobes broader) apiculate;

corolla campanulatr -stellate, lilac-blue, about 17 mm. long with a

FLORA OF PERU 17

spread of 2.3 cm., the broadly lanceolate lobes pubescent only within

at the recurved tips; anthers narrowly ellipsoid, about 8 cm. long;

style 9 mm. long, capitate stigma 1.5 mm. broad. — The two collec-

tions are alike as to flowers but one has simple leaves. The author

suggests that the latter here as in other species may occur on young

branches. Peruvian collections all show only entire leaves but may

be referable here. F.M. Neg. 2937.

San Martin: Juanjui, Klug 3873. — Huanuco: San Carlos, Mexia

8209. — Junin: Puerto Bermudez, Killip & Smith 26605. — Loreto:

Yurimaguas, Williams 4193; 4370; 4687; Killip & Smith 28005;

28703; 29093.— Rio Acre: In woods at Porto Carlos, Ule 9753, type;

Cobya 9752. "Pepino del campo" (Mexia).

Cyphomandra Ulei Bitter, Repert. Sp. Nov. 17: 349. 1921.

A shrub 2-5 meters high, with simple, broadly or rounded ovate-

cordate, shortly (velvety beneath) pubescent and somewhat stipitate-

glandular firm leaves, the larger to 2 dm. long, 13 cm. wide, the

smaller about 1 dm. X 8 cm. ; inflorescence above the leaf axils, the

indument consisting mostly of minute 2-celled glandular and egland-

ular trichomes, the peduncles only about 1 cm. long; pedicels to

2.5 cm. long, a little longer in fruit; calyx cupulate, 3-4: mm. long,

5 mm. wide, the obtuse lobes early obscure; corolla stellate, about

2.5 cm. wide, the lanceolate subacute lobes about 1 cm. long, finely

pubescent without, with a few longer usually simple marginal tri-

chomes, glabrous within except for the pilose tips; filaments 2.5 mm.

long, anthers 4.5 mm. long, papillose; style 3.5 mm. long, subpeltate

stigma nearly 2 mm. wide. — Leaves gradually narrowed to acute or

subacute apex.

Loreto: Mishuyacu near Iquitos, Klug 93. — Apurimac: Pincos, in

Escallonia forest, Stork & Horton 10711. — Cuzco: Anta, Vargas 200

(distr. as C. pendula). — Rio Acre: In woods at San Francisco, (Ule

9755, type).

Cyphomandra viridiflora (R. & P.) Sendt. Flora 28: 175. 1845;

400. Solanum viridiflorum R. & P. Fl. Peruv. 2: 38, pi. 173, fig. b.

1799. Pionandra viridiflora (R. & P.) Miers in Hook. Lond. Journ.

Bot. 4:359. 1845.

Stems woody, branched, about 2 meters high, the branchlets and

ample (the larger 1.8 dm. X 8 cm., the smaller about 1 dm. X 7 cm.)

cordate-ovate simple leaves softly villous; petioles 2.5 cm. long; pe-

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

duncles nearly as long as the leaves, drooping; calyx pilose, 6 mm.

across, half as long, the lobes acute, erect, short; corolla green, about

1 cm. long and broad, deeply parted, the ovate-lanceolate divisions

reflexed, woolly-margined; style exserted, swollen at the middle; con-

nective scabrous; berry about 5 cm. long and 3 cm. thick, villous,

yellowish; seeds lentiform, yellow. — Allied to C. pendula by Ruiz

and Pavon, and to C. crassifolia by Miers; the interpretation may

be incorrect.

Huanuco: San Antonio de Playagrande, Ruiz & Pavdn.

5. WITHANIA Pauquy

Athenaea Sendt. in Mart. Fl. Bras. 6: 133. 1846. Larnax Miers,

Ann. Mag. Nat. Hist. 2, 4: 37. 1849.

More or less ligneous, sometimes flowering as herbs, the Peruvian

species with fasciculate (1-several) 5-merous campanulate-funnel-

form flowers, deeply lobed, the lobes reflexing. Calyx more or less

inflated, minutely (Peru) dentate accrescent and closely or loosely

investing the fruit.— Werdermann, Diels, Bibl. Bot. 116: 130. 1937,

described A. Bitteriana from central Ecuador with dark purple tubu-

lar-funnelform corollas, the calyx inflated; the group therefore is prob-

ably a part of Dunalia, sens. lat. The names of Pauquy (1824) and

Sendtner have both been conserved against earlier ones. The calyx

teeth may be incurved or the calyx may be lobed, the plants when

in fruit simulating Physalis L., from which it is most readily distin-

guished (arbitrarily but conveniently) by the more or less deeply

lobed corolla, the lobes somewhat reflexed. Bitter found no granules

in the fruits he examined. Incomplete or young specimens have been

confused with Capsicum L.

Corolla white; leaves glabrous W. peruviana.

Corolla yellow; leaves villosulous W. subtriflora.

Withania peruviana (Zahlbr.) Macbr., comb. nov. Athenaea

peruviana Zahlbr. Ann. Naturh. Hofm. Wien 7: 7. 1892.

Rather open in growth, the branches slender, glabrous or essen-

tially except for the ciliate corolla lobes, these 5 mm. long; leaves

oblong-elliptic-ovate, attenuate both ends, strongly oblique at base,

often 1.5-2 dm. long, about a third as wide; berry orange, 10-14 mm.

in diameter, deeply but loosely invested by the greatly enlarged

5-ribbed and plicate calyx. — The lax calyx seems to suggest that

FLORA OF PERU 19

Withania Pauquy at best is an expedient segregate of Physalis L.

F.M. Neg. 2546.

Cajamarca: Tambillo, Jelski 55; 54 (Ml #0)- — San Martin: Tara-

poto, Spruce 4443. — Loreto: Cerro de Escaler, Ule 6803 (det. Bitter).

Withania subtriflora (R. & P.) Dunal in DC. Prodr. 13: 455.

1852. Physalis subtriflora R. & P. Fl. Peruv. 2: 42, pi. 178. 1799.

Larnax subtriflora (R. & P.) Miers, Ann. Mag. Nat. Hist. 2, 4: 38.

1849.

A shortly villous herb or ligneous toward base; branches, branch-

lets sulcate, at least in drying; petioles 0.5-1.5 cm. long; leaves in

part geminate, the larger 7-10 cm. long, to 5 cm. wide, the smaller

often about half as large, all ovate, more or less obliquely subcordate

or rounded at base, acuminate, subvillous both sides at least on the

6-7 primary nerves and rather prominent reticulate veins; pedicels

1-5, filiform, 3 mm. long, nutant; early calyx campanulate, 4.5 mm.

across, membranous, 5-nerved, truncate between the short teeth, in

age suburceolate; corolla greenish-yellow, about 6 mm. long, 10-

12 mm. across, often smaller, the ovate acutely acuminate lobes re-

flexed at tip; stamens inserted at base, filaments 3 mm. long, anthers

stout, subcordate, style filiform, stigma capitate; berry yellowish,

6-8 mm. in diameter, closely enclosed in the inflated but apically

open calyx.— F.M. Negs. 2544; 2547 (ined. name); 8566.

San Martin : Tarapoto, Spruce 41 25; (5522} . Chazuta, Klug 4111.

Zepelacio, Klug 3667. — Lima: Obrajillo and San Buenaventura, Ruiz

& Pavdn, type.— Hudnuco: Tingo Maria, Allard 21846; 21870 —

Junin: Jauja, Weberbauer 6672. — Loreto: La Victoria, Williams 2847.

Pongo de Manseriche, Mexia 6340.

6. PHYSALIS L.

Reference: Waterfall, Rhodora 60: 107-114, 128-142, 152-173.

1958.

In Peru erect or spreading annual or perennial herbs usually low,

branched and sometimes woody toward the base. Leaves entire or

sinuately dentate. Peduncles solitary or rarely several on short axil-

lary branchlets. Corolla rotate or applanate-campanulate (Sleumer) ,

(limb exceptionally 5-parted), often yellow with a dark brown or pur-

ple center. Calyx in fruit accrescent and usually bladdery-inflated,

membranous, always completely and laxly enclosing the berry, the

teeth commonly connivent. — The yellow berries are called "ground-

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

cherries," or, in English horticulture, Cape Gooseberries. P. cheno-

podifolia Lam., Illus. 2: 28. 1793, a perennial, early white-tomentose

with simple trichomes, corolla spots dark violet, anthers orange, mar-

gins violet, was described from cultivated plants, seed possibly from

Peru (author); found to date only in Mexico (F.M. Neg. 29394).

P. Alkekengi L., Sp. PI. 183. 1753, of gardens, probably sometimes

persisting as an escape, is usually unbranched and has greenish-white

scarcely lobed corollas, calyx in fruit bright red, berry red, the "straw-

berry tomato" in English; yellow-fruited species are familiar as ground

cherries, sometimes as Cape Gooseberry, all edible, especially as pre-

serves, particularly P. pubescens L., the Pear Cherry (Henry Paul

Jackson).

KEY TO PHYSALIS

Flowers blue (ex char.) ; plants somewhat stellate pubescent, rarely

in part simply or nearly glabrous P. viscosa.

Flowers yellowish or yellow; indument, if present, not conspicuously

stellate, trichomes mostly or all simple.

Pilose perennial; corolla spots, anthers (3 mm. long) deep blue-

purple P. peruviana.

Puberulent, long-pilose or glabrous annuals; anthers blue, often

1.5-2.4 mm. long.

Trichomes mostly or all long, rarely none; corolla spots 5, dark;

calyx pubescent, angled P. pubescens.

Trichomes lacking or short, few; corolla spots (typical) none;

calyx not angled in fruit P. angulata.

Physalis angulata L. Sp. PL 183. 1753; 162. P. Linkiana Nees,

Linnaea 6: 471. 1831. P. lanceifolia Nees, I.e. 473.

An essentially glabrous angular- stemmed annual several dm. to

a meter high, with ovate to linear-lanceolate cuneate-based often

sharply sinuate-dentate leaves and small (4-10 mm. long) cream-

colored flowers not spotted but with dark centers; anthers bluish

(grayish-green, Killip spec.) ; calyx subangulate, usually 2-3 cm. long,

the recurving peduncle usually shorter; seeds rufescent. — A weedy

species of wide distribution in warm countries. Determinations by

Standley. The var. lanceifolia (Nees) Waterfall, I.e. 163, is distin-

guished by the oblong-lanceolate usually subentire leaves and the

peduncles, 1-2 cm. long, erect in flower, elongating, recurving and

FLORA OF PERU 21

incurving or reflexed in fruit, the fruiting calyx 2-2.5 cm. long; corolla

usually only 4-5 mm. long, calyx in flower 4 mm. long, anthers often

only 1-1.5 mm. long.

An infusion of the entire plant (except root) is taken by the

natives to cure malaria (Ferreyra).

Piura: Cana Dulce, Haught 88. Alto de La Cruz, Stork & Horton

11353. Prov. Paita, Horton 11594- — La Libertad: Salaverry, Worth

8892.— San Martin: Jepelacio, Klug 3439. San Roque, Williams

7060. Tarapoto, Williams 6220. Juanjui, Ferreyra 4537. — Lima:

Magdalena, Nee. Chancay, Ruiz & Pavon; Wilkes Exped. — Hua-

nuco: Tingo Maria, Stork & Horton 9470; Allard 21139; 22050 (det.

L. Smith). — Junin: La Merced, 5361. — Loreto: Mishuyacu, Klug

1241? (det. Killip, P. Lagascae R. & S.). Yurimaguas, Ferreyra 4918

(det. Cowan) ; Killip & Smith 28218. Caballo-cocha, Williams 2218;

2284. Pucallpa, Soukup 3032. Iquitos, Killip & Smith 27247. Rio

Nanay, Williams 336. La Victoria, Williams 2637,—Ayacucho:

Prov. Huanta, Weberbauer 5637; Killip & Smith 23085. — Cuzco:

Rio Yanamayo, Pennell 14080. Quellouno, Vargas 7556. "Bolsa

mullaca" (Williams). "Cahuli Cimarron," i.e. False Capuli. Mexico.

Physalis peruviana L. Sp. PI. ed. 2. 1670. 1763; 141. P. edulis

Sims, Bot. Mag. pi. 1068. 1807.

A tall erect often densely pilose unpleasantly scented perennial

from a creeping rootstock, the trichomes eglandular, simple; leaves

broadly ovate-cordate or subrotund, acuminate, 5-10 cm. long, nearly

as wide; corolla yellowish, dark blue-purple spotted at base, about

12 mm. long, 1.5 wide at top; filaments slender; anthers purple or

purplish-red; calyx villous, teeth acuminate, subequaling the tube,

about 7 mm. long, soon accrescent, finally to 4 cm. long in fruit.—

Frequently cultivated in warm countries for its sweet yellow berries

and so in Peru in 1725, according to Feuille'e, where "greatly es-

teemed as a preserve." Herrera states that an infusion of the flowers

is used during childbirth. C. V. Morton, U. S. National Museum,

thoughtfully sent me (with many selected specimens) a photostatic

copy of notes on the Cape Gooseberry (Year Book 1922, Madras

Agric. Dept. 1923), an interesting account of the usefulness of the

plant for its fruits; filed with it is a report on the chemistry by J. B.

Lai (Proc. Ind. Nat. Acad. Sci. 6 (4) : 309-313. 1936), and a copy of

M. Bossin's notes on P. edulis Sims and other species (Bull. Mens.

Soc. d'Acclim. se"r. 3, 2: 64-74. 1875). Bossin regarded P. peruviana

L. as a different species. Finally, there is a file of correspondence

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

regarding this last question, highly amusing as an example of diplo-

matic "passing the buck," no specialist of the time accepting the

responsibility of declaring surely the status of the two names; I fol-

low suit.

Piura: Chicama, Haught 189. Cana Dulce, Haught 178. — Caja-

marca: Monte Seco, Soukup 3888. — Lima: Prov. Huarochiri, Good-

speed 33117 (det. Killip); Isern 2461.— Huanuco: San Rafael, Fer-

reyra 1974- Divisoria, Chanchamayo Valley, Schunke 484- — Junin:

Tarma, 1041; Killip & Smith 21902. Huancayo, at 3317 meters,

Soukup 2733 (det. Standley). — Apurimac: Prov. Abancay, Good-

speed Exped. 10571 (det. Standley); Balls 6837—Cuzco: Toward

Machupicchu, Hen era 3281; West 8023 (det. Johnston) ; Soukup 126.

Pillahuata, Pennell 13952. Prov. Anta, Vargas 148 (det. Standley).

Paucartambo, Herrera 3802 (det. Killip). "Capuli," "tomate syl-

vestre," "aguaymanto" (Vargas).

Physalis pubescens L. Sp. PI. 183. 1753; 164. P. pubescens L.,

var. hygrophila (Mart.) Dunal in DC. Prodr. 13: 446. 1852. P. hy-

grophila Mart. Flora 24, Beibl. 2: 86. 1841? P. turbinata Medic. Act.

Acad. Theod. Palat. 4: 189. 1780, fide Waterfall.

A slender generally diffuse more or less viscid-pubescent or, in

var. hygrophila, hirsutulous annual with ovate slightly cordate leaves,

subentire at least below; peduncles 3-5 (12) mm. long, to 2 cm. long

in fruit; calyx teeth lanceolate, equaling the tube, the fruiting calyx

2-3 (4) cm. long, 5-angled, somewhat retuse at base; corolla yellow

with prominent ribs or spots, 6-10 mm. long; anthers 1.5-2.4 mm.

long, usually violet. — The var. glabra (Michx.) Waterfall, I.e. 165,

is at most only puberulent.

The yellow bittersweet berries, as those of P. peruviana, are

used for preserves and are said to be sweeter; cultivated, according

to Ruiz and Pavon, for the acid-sweet fruit and also for the flowers,

which, with a little amber, form a perfume; Eyerdam found the

fragrance of the fruits suggestive of ripe pears.

Piura: Cana Dulce water-course, Haught 178. Negritos, Haught

267. Prov. Piura, Weberbauer 5940; 5941. — San Martin: Tarapoto,

Williams 6047 (det. Standley). Jepelacio, Klug 3439 (glabrate, det.

Standley, P. angulata}. Boqueron Pass, Allard 21707 (det. Lyman

Smith). — Huanuco: Divisoria, Allard 21235. — Loreto: Mishuyacu

near Iquitos, Klug 322 (det. Standley). Clearing, Pongo de Man-

seriche, Mexia 6299 (det. Morton, P. hygrophila). — Cuzco: Echarate,

Goodspeed Exped. 10483 (det. Standley). Idma, Vargas 8528. "Muy-

FLORA OF PERU 23

aca" (Mexia); "bolsa mullaca" (Klug; Williams); "capuli" (Ruiz &

Pavon).

Physalis viscosa L. Sp. PI. 183. 1753; 133. Cacabus parviflorus

Rusby, Mem. Torr. Bot. Club 4: 233. 1895, fide Rydberg.

Well-marked by the dense (typically) stellate pubescence; a per-

ennial with subcordate leaves and rather large (funnelform, 1-2 cm.

long) spotted or clear greenish-yellow flowers; calyx in flower 3-

10 mm. long, in fruit 2-5 cm. long; peduncles 1-4 cm. long, lobes

unequal, ovate; anthers yellow, about 3 mm. long; berries orange or

yellow. — A coastal species as subsp. maritima (M. H. Curtis) Water-

fall, including several variants in degree (to absence), I.e. 134, and

quality of pubescence.

Peru: (Ruiz & Pavdn). South America to the southern United

States.

7. CACABUS Bernh.

Thinogeton Benth. Bot. Voy. Sulph. 142. 1844. Streplostigma

Regel, Gartenfl. 322, pi 68. 1853.

Diffuse, or often prostrate and trailing, pilose or viscid herbs with

solitary campanulate or broadly funnelform flowers and shortly

toothed calyces, enlarged in fruit. Filaments of three lengths and

attached at various levels; stigma lanceolate. Leaves usually long-

petioled, subentire to deeply sinuate-dentate. — Distinguished from

Physalis L. particularly by the habit and the often larger flowers,

apparently usually glabrous within at the base (cf . Bitter, Repert.

Sp. Nov. 17: 245. 1921); Sleumer describes the corolla of the former

as rotate or applanate-campanulate, that of Cacabus as tubular-

campanulate or broadly infundibuliform. Seems to be an expedient

or academic segregate; however, as observed by Waterfall in his

careful review of the North American species of Physalis L., generic

definition awaits properly the considered judgment of an informed

student.

Flowers yellow; some leaves subrotund, subcordate, subentire.

C. flavus.

Flowers white, purple-tinged; leaves various, rarely ever as above in

all respects.

Flowers to 1.5 cm. long; fruiting calyx 4 mm. long, the berry partly

exserted C. pusillus.

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

Flowers 2-5 cm. long; fruiting calyx 1-2 cm. long, including the

berry C. prostratus.

Cacabus flavus Johnst. Contr. Gray Herb. 85: 177. 1929.

A prostrate sparsely villous glandular annual; leaves cordate,

3-6 cm. long, subentire; petioles 3-9 cm. long; pedicels axillary,

1-2 cm. long; calyx 9-11 mm. long, the lobes 4-5 mm. long, mem-

branous, tightly investing and finally ruptured by the fruit; corolla

yellow, 2-4 cm. long. — Apparently the only yellow-flowered species,

at least in this character suggesting Physalis L. but probably allied

to C. integrifolius Phil, according to the author; the flowers are very

much larger; the data of specimens of Velarde Nunez and Hrdlicka

do not include color of flower, but from locality (at least of former)

presumably it was yellow; otherwise both are probably C. prostratus,

sens. lat.

Arequipa: Tiabaya, Pennell 13066, type. — Moquehua: Moquehua,

Weberbauer 7459. Lomas de Pongo, Nunez 1480? Valley of Rio de

Las Trancas, Hrdlicka.

Cacabus prostratus (Dombey) Bernh. Linnaea 13: 360. 1839.

Physalis prostrata Dombey ex L'HeY. Stirp. Nov. 43, pi. 22. 1784.

P. limensis Retz. Obs. 5: 22. 1791, fide Nees. Physaloides prostrata

(Dombey) Moench. Meth. Suppl. 178. 1802. Thinogeton maritima

Benth. Bot. Voy. Sulph. 142. 1844. C. maritimus (Benth.) Benth.

Hook. Gen. PI. 2: 897. 1876. Dictyocalyx Miersii Hook. f. Trans.

Linn. Soc. 20: 203. 1847, fide Svenson. T. Lobbianum Miers, Ann.

Mag. Nat. Hist. 2, 4: 360. 1849. C. nolanoides Miers, I.e. 255, and

111. S. Am. PL 2: 51. 1849-1857. C. multiflorus Damm. Repert. Sp.

Nov. 15: 366. 1919, fide Svenson.

A grayish clammy-viscid soon trailing herb, the flexuose striate

stems more or less foliose with unevenly sinuate-angulate or dentate-

lobulate to subentire long-petioled leaves, often rather ovate and

acute to attenuate at base, a few to 10 cm. long and not rarely nearly

as wide or the upper lanceolate-oblong; peduncles solitary or gemi-

nate, a few mm. to 2 cm. long or longer in fruit, when curved or re-

flexed and the calyx then accrescent, more or less 5-10 costate-angled ;

corolla somewhat pilose, 2-5 cm. long, purple or usually white at top

where about 2.5 cm. wide; berry about 1 cm. in diameter, closely in-

vested by the firm to thin-papyraceous calyx. — Svenson, Amer.

Journ. Bot. 33: 481-482. 1946, concluded, apparently correctly, that

probably only a single species of this genus is native to the South

FLORA OF PERU 25

American coast and recorded the following important observations:

Bentham and Hooker reduced Miers' six species to C. prostratus and

C. maritimus, the fruiting calyx of the former obscurely 5-angled,

that of the latter 10-angled with 5 of the angles a little more prom-

inent, but from Bernhardi's description of C. prostratus this differ-

ence does not appear to be very much; C. multiflorus Damm. as to

type has several flowers on foliose-bracted axillary branchlets and

may be distinct or at least a local variation. The elongate inflores-

cences described by Dammer are identical on well-developed Galapa-

gos specimens; Riley, Kew Bull. 227. 1925, found no glands in the

pubescence and that seems to be the case in all specimens, quartz

grains adhering to the viscid jointed trichomes simulating glands.

Calyx increases as capsule expands; this early is clearly 2-celled but

at maturity has at base a stiped ligneous 4-celled structure with in-

curving walls, to which are attached the innermost seeds; the ripe

capsule has ten slightly elevated ribs, becomes soft, dark green to

black, a little elongate, the pale brown seeds buried in a dark viscid

substance. Svenson further describes the plant as starting to flower

when very small, the flowers solitary or in pairs in the axils of the

succulent leaves but as the prostrate stem elongates (even to 2 me-

ters fide Haught) the inflorescence becomes many-flowered in a panic-

ulate manner, the petunia-like flowers white with a dull purplish

center, purple striate in throat and purplish-tinged at apex. Weber-

bauer found the plant used for fodder. Common on sandy or stony

coastal hills. Unfortunately the type collection of C. nolanoides

(Mathews 839, Kew) is without data. Illustrated, Svenson, 404,

pi. 4, fig- 2; Miers, 111. S. Amer. PI. pi. 49 (Cacabus nolanoides).

Tumbez: Zorritos to Cancas, Weberbauer 7750 (det. Johnston,

C. maritimus). Puerto Pizarro, Ferreyra 5970. — Piura: Pima to

Nomala, Weberbauer 5931 (type, C. multiflorus}; Sandeman 4251;

Ferreyra 5884; 6008. Talara, Haught 16; Beetle 26200; Johnston

3501; Horton 11595; Stork & Horton 11634. — La Libertad: Salaverry,

Johnston 3519 (det. Johnston, C. maritimus). — Ancash: Lomas de

Monzon, Goodspeed Exped. 9176. — Lima: Chosica, Weberbauer 5336;

Grant 7397. San Geronimo, 5899. Mt. San Augustin, Weberbauer

5227; 5248. Near Lima, (Cuming 972) . Lomas de Lachay, Ferreyra

3889. Quive, Pennell 14292. Chancay, Chorillos, (Maclean). Chan-

cay, Huara, Ruiz & Pavon; Dombey, type; Beetle 9105. Puente de

Verrugas to Surco, Weberbauer 5216. Santa Clara, Weberbauer 1671

(det. Bitter); Rose 18623. Santa Eulalia Valley, Goodspeed & Stork

11480. — lea: Prov. Chincha, Weberbauer 5376; (Maclean). — Huan-

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

cavelica: Pampano to Huaytara, Weberbauer 5402. Galapagos; Ecua-

dor. "Campanillas olorosas," "suruvia," "suravilla."

Cacabus pusillus Bitter, Repert. Sp. Nov. 17: 243. 1921.

Annual herb a few cm. tall, simple or a little branched at base,

green but pubescent all over including the corollas with simple many-

celled unequal gland-tipped trichomes; petioles to 2.5 cm. long, alate

toward blade, this linear to lanceolate or ovate, finally 1-3 (5) cm.

long, about a third to as wide, both ends narrowed, obtuse or sub-

acute; pedicels axillary, solitary or binate, 4-5 mm. long, in fruit

5-6 mm. long, arcuate, incrassate; calyx 3.5-4 mm. long, 2 mm.

broad, acuminate lobes 2.5-3 mm. long, in fruit subglobose, 4 mm.

across; corolla white, lilac, or white and lilac (Weberbauer), tubular-

campanulate, 1.5 cm. long, near apex 4-5 cm. across, the short plicate

lobes acuminate; free part of filaments 5-6 mm. long, pubescent;

style 8.5 mm. long, stigmas capitate; berry very obvious between the

calyx lobes, 3 mm. thick, greenish-yellow, granules none, the seeds

only about 9, obliquely reniform, reticulate. — Leaves membranous

but fleshy in life; unique in the exposed fruit. F.M. Negs. 2528;

2529 (C. Woitschachii in herb.).

Arequipa: Tingo, Pennell 13126. Arequipa, Pennell 13045 (det.

Johnston); 13178. Yura, Weberbauer 6842, type; Guenther 12242.

Slopes of Misti, Sandeman 3823 (det. Johnston). Tia Baya, Cocker-

ell. — Tacna: Candarave, Weberbauer 7388 (det. Johnston). Near

Tacna, Woitschach.

8. SARACHA R. & P.

Bellinia Roem. & Schult. Syst. 4: 687. 1819. Kukolis and Utti-

cona Raf. Sylva Tellur. 55. 1838. Hebecladus Miers in Hook. Lond.

Journ. Bot. 4: 321. 1845.

References: Dunal in DC. Prodr. 13, pt. 1: 429-433, 682-685;

Hebecladus, 468^71. 1852; Bitter, Repert. Sp. Nov. 17: 338-346.

1921; 18: 99-112. 1922; 19: 265-270. 1924.

Perennial herbs usually or finally more or less ligneous. Peduncles

axillary or lateral, commonly solitary, sometimes reduced or obsolete,

the pedicels 1-several, 1-flowered, often subumbellate. Calyx not

obviously to more or less accrescent, little or not to widely spreading.

Corolla tubular, sometimes shortly tubular- to rotate-campanulate,

sometimes broadly and deeply, ordinarily more or less flaring at apex,

often dentate between the variously developed lobes. Filaments

glabrous or slightly to densely pubescent, anthers suboblong, emar-

FLORA OF PERU 27

ginate to cordate, frequently, as style, exserted, this with clavate or

capitate stigma. Seeds compressed-reniform, foveolate-reticulate. —

P. Isidro Saracha was a Benedictine botanist who gave rare plants

to the botanic garden in Madrid.

As pointed out by Morton in his useful paper "Notes on the

Genus Saracha," Proc. Biol. Soc. Wash. 51: 75-77. 1938, Poecilo-

chroma punctata (R. & P.) Miers is the type of Ruiz and Pavon's

genus and was so accepted, too, by Miers, 111. S. Amer. PI. App. 57.

1857. Miers himself, I.e. 148-149, retracted his own action. Never-

theless the "proper" name for Miers' genus is Saracha unless the

cognomen of Ruiz and Pavon is conserved. Otherwise Bellinia Roem.

& Schult. may be the correct name for the following plants or at least

for those not segregated (as Hebecladus). But as Morton noted the

matter is not of moment, only a few names being concerned. These

are here used as available in order to avoid expression of my opinion,

certain to be futile (see Wash. Acad. Sci. 19: 247. 1929).

Bitter, Repert. Sp. Nov. 17-20. 1924, divided the group into four

sections, excluding Hebecladus Miers but indicating, as Morton, the

transition to the latter; Morton, I.e. 76, found that the value of one

of the chief distinctions of his sections, namely the presence or ab-

sence of minute sclerotic grains among the seeds of the berries, is

highly doubtful (as also in Solanum) ; another section rested on gla-

brous filaments and annular ring; this surely, as Morton remarks, is

only doubtfully a good species character; only Macrosaracha Bitter,

characterized by its much larger campanulate rather than rotate

corollas, intermediate to Poecilochroma and Hebecladus, is well-dis-

tinguished (Morton); the corolla, especially in these groups, has 5

glands at base within, conspicuous from their copious red or scarlet

exudation, visible as swollen filament bases when dry; however, these

occur, in some degree, in otherwise dissimilar species.

The size of flowers recorded from specimens may be misleading,

partly of course from contraction but also because the corolla often

enlarges considerably after early anthesis; Bitter, too, seems to have

had a tendency to over-state measurements. The group needs revi-

sion by a student who can study living plants; characters in key are

often expedient.

KEY TO SARACHA (including HEBECLADUS)

Flowers usually 2-several, pedicellate in axils or subumbellate on

obsolete or more or less apparent peduncles; corolla shorter than

2 cm. if pedicels only 2-3, or long-tubular.

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

Filaments glabrous or essentially (sometimes granular or with a

few trichomes, as S. sordideviolacea) ; plants glabrate.

Corolla tubular, the small lobes erect or suberect.

Corolla about 1-1.5 cm. long, pedicels several. .H. umbellatus.

Corolla 2-3 cm. long, pedicels (1) 2 or 3.

H. intermedius, H. bicolor.

Corolla rotate-campanulate or rarely ventricose-campanulate.

Leaves broadly ovate or subrotund, to 4 cm. long or a little

longer; peduncles obsolete or short; corolla red, yellowish

or white, not lanate; plants shrubby, at least in age.

Corolla tube 8-10 mm. long, lobes to 5 mm. long; shrub.

H. umbellatus.

Corolla tube as lobes obscure; subherbaceous perennials or

corolla yellowish (cf. also S. alata).

Corolla purplish-reddish; stems slender.

S. lobata, S. sordideviolacea.

Corolla yellowish or white; lower branches stout, ligneous.

S. dentata.

Leaves mostly or all much longer than wide, several cm. long;

peduncles soon elongate; corolla yellowish or lanate; per-

ennial herbs (cf. H. ventricosus, S. biflora).

Corolla lanate; peduncles often 1-bracteate S. diffusa.

Corolla glabrate; peduncles not bracted. . . .S. procumbens.

Filaments pubescent; plants more or less puberulent or villous (cf.

the glabrous S. alata).

Corolla tubular, 2.5 cm. long or longer.

H. Weberbaueri Damm. 1906.

Corolla rotate-campanulate.

Indument a puberulence; corolla scarcely 1 cm. long, usually

shorter; peduncles shorter than pedicels or none; shrubby,

branched (synonyms, H. Weberbaueri, S. Bitteri).

S. dentata.

Indument uneven or more defined than puberulence, some-

times sparse, sometimes more or less villous; corolla

usually at least 1 cm. long; peduncles often as long or

longer than the (1) 2-several pedicels.

Corolla yellowish to white (or bluish at base) .

FLORA OF PERU 29

Pedicels (1) 2-3 on forked peduncles; berry often solitary,

yellowish or white; stems terete.

H. ventricosus, S. biflora.

Pedicels umbellate, usually several; mature berry dark

or black.

Indument eglandular, soon sparse; stems herbaceous.

S. contorta.

Indument usually viscid or glandular villous, early

dense; stems angled, frutescent (S. sinuosa Bit-

ter not Miers) S. villosa.

Corolla purple or blue; indument in part glandular.

S. propinqua.

Flowers solitary, rarely 2 on a peduncle, campanulate, not rotate,

never long-tubular, 2 cm. long, often longer, probably always

in full anthesis, scarlet annular glands and secretion conspicuous

(species-characters intangible).

Corolla finally 2.5 cm. long, 3-4 cm. wide, little flared apically,

lobes apiculate; filaments pubescent at or toward base.

Branches, leaves soon glabrate or glabrous S. Weberbaueri.

Branches, leaves more or less pubescent.

Filaments glabrous unless adnate part S. Urbaniana.

Filaments hirsute below H. asperus.

Corolla about 2 cm. long, 3 cm. wide or smaller, lobes acutely acu-

minate, flared apically especially if shortly acuminate; fila-

ments glabrous or nearly; plants more or less puberulent.

Flowers usually solitary; trichomes at least mostly simple.

S. ciliata.

Flowers often 2; trichomes at least mostly furcate. .S. Herrerae.

Saracha alata Dunal in DC. Prodr. 13, pt. 1: 431. 1852.

Sulcate pale green branches 4-5 mm. thick, marked with sessile

globose and linear white glands and membranous subalate angles

scarcely 2 mm. wide; leaves subsessile, geminate, ovate, cuneate to

decurrent base, obtusely subcuspidate, the larger 1-1.5 dm. long,

about half as wide, the smaller less than half as large, undulate-

subrepand, the 5-6 nerves prominent beneath and minutely white-

glandular; peduncles reflexing, 12-18 mm. long; pedicels 10-12, um-

bellate, filiform, to 1.5 cm. long; calyx subrotate, 1 cm. across, the

segments broadly ovate, vix 4 mm. long; corolla rotate-campanulate,

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

1 cm. long, 12-15 mm. wide, lobes ovate, acute, 6-7 mm. long; sta-

mens in corolla tube, the cylindric filaments 3-4 mm. long, slightly

pilose at base, anthers ovate, subacute, 1.5 mm. long; ovary sub-

globose. — Ex char, perhaps the earliest name for S. lobata Bitter or

(and) S. sordideviolacea Bitter but glabrous except for the sessile

glandulosity. F.M. Neg. 29719.

Peru(?) : Ruiz & Pavon, type in Herb. Boissier (Geneva) as S. fili-

forma Pavon, according to Dunal.

Hebecladus asperus (R. & P.) Miers, Hook. Lond. Journ. Bot.

4: 322. 1845. Atropa aspera R. & P. Fl. Peruv. 2: 45. 1799. Ulticona

aspera (R. & P.) Raf. Sylva Tellur. 55. 1838.

A harshly hirsute dichotomously branched erect purplish herb, the

type to 6 dm. high; leaves petioled, geminate, ovate or oblong-lanceo-

late, entire or repand; flowers yellow-blue, nodding and solitary, sub-

axillary; corolla campanulate, sub-10-parted, 5 acute lobes alternating

with 5 emarginate ones, all spreading, the throat and hirsute fila-

ments violet; berry white, seeds pubescent. — Ex char, this seems to

be related to S. ciliata Miers, etc., and may be found to be the earliest

name for one or more of these seemingly dubious species.

Lima: In the Amancaes Hills near Lima, Ruiz & Pavdn, type.

Hebecladus bicolor (R. & P.) Miers in Hook. Lond. Journ. Bot.

4: 322. 1845; 470. Atropa bicolor R. & P. Fl. Peruv. 2: 45. 1799.

Kukolis bicolor (R. & P.) Raf. Sylva Tellur. 55. 1838. Atropa biflora

R. & P. I.e. 44. pi 181b. UUicona biflora (R. & P.) Raf. I.e. H. bi-

florus (R. & P.) Miers, I.e.

A lightly pubescent or sometimes glabrate shrub; flowers 1-4,

axillary and lateral, the more or less nutant hirtellous peduncles

mostly about 2 cm. long; branchlets somewhat angled; petioles 5-

15 mm. long; leaves early geminate, unequal, ovate, now and then

angulate or subdentate, acute, the lower 5 cm. long, 2.5-3 cm. wide

or larger, soon glabrous, pale and often granulate beneath; calyx

weakly 5-angled, the acute lobes little or not spreading, 3-5 mm.

long; corolla tubular, striate, pilosulous, slightly constricted below

the often reddish or purple-tinged limb, the tube usually yellowish-

green, finally at least 2 (-3) cm. long; filaments exserted, thus the

bluish ovate-sagittate anthers conspicuous; stigma bilobed; berry

white (apparently rarely blue), depressed-globose, subtended by the

spreading calyx; seeds lutescent. — The tubular corolla distinctly

elongates after early anthesis; in foliage the species simulates S. bi-

FLORA OF PERU 31

flora R. & P. but is glabrate; type of H. bicolor (R. & P.) Miers not

found but ex char, there is only a single species distinguishable here.

F.M. Neg. 2530 (H. biflorus).

Ancash: Chiquian, Ferreyra 7427. Samanco, (Weberbauer 171).

— Lima: Prov. Huarochiri, Surco and Matucana, (Ruiz & Pav6n,

type, H. bicolor); Hrdlicka; Weberbauer 194 (det. Werdermann) .

Surco, Raimondi (det. Werdermann). Rio Blanco, Killip & Smith

21558; 21751. Canta, Pennell 14242. San Mateo, Isern 2295; 2545;

Ferreyra 5315. Toward Casapalca, 3,500 meters, Ferreyra 5407;

6991. Viso, 571. — Huanuco: Tingo Maria, Ferreyra 1576 (?, flowers

white, fruit blue). — Junin: Tarma to Huasahuasi, Ruiz & Pavdn,

(type, H. biflora). — Huancavelica: Cordova, Metcalf 30270, fruit

green (det. Morton). Ecuador? "Suplac" (Ferreyra).

Saracha biflora R. & P. Fl. Peruv. 2: 42, pi. 179. 1799; 431.

Bellinia biflora (R. & P.) Roem. & Schult. Syst. 4: 689. 1819. With-

eringia biflora (R. & P.) Miers, 111. S. Amer. PL 2: App. 56. 1857.

Stems sparsely pubescent above, herbaceous, angled; leaves ovate,

about 10 cm. long, 6-7 cm. wide, undulately margined, rounded at

base but subdecurrent into the narrow petiole, the upper geminate;

peduncles axillary, solitary, recurving, 1.5-2 cm. long, 2- or rarely

3-flowered; pedicels 5-7 mm. long; calyx campanulate, about 1 cm.

long, the ovate acute lobes half as long; corolla yellowish-green, cam-

panulate, spreading, to 1.5 cm. wide, the lobes acute, stamens some-

times twice as long as the corolla; berry white, somewhat flattened.

— Inflorescences, including corolla, shortly pilose. Corolla in type

little exceeding calyx, perhaps young; sepals acute, 10 mm. long,

petals acute, 12-13 mm. long, pubescent. Fruiting calyx lobes rotund-

ovate, 7 mm. long and broad, the berry deeply included.

Small yellowish fruits eaten by children (Ruiz and Pavon) ; fila-

ments glabrous or granulate; corolla white or partly violet (violet,

according to Weberbauer, 7602). Probably should not include the

Cuzco and Puno plants with deeply campanulate corollas, reflexed

lobes, suggesting those of H . ventricosus Baker.

Huanuco: Cani, 3443 (det. Morton). — Junin: Carpapata, Kil-

lip & Smith 24396 (det. Morton) . Huasahuasi, Ruiz & Pavdn, type.

— Huancavelica: Mantaro Valley, Weberbauer 7602 (det. Morton).

North of Salcabamba, 3,500 meters, Stork & Horton 10317? — Apuri-

mac : Oropeza Valley, Vargas 9798 (det. Standley, S. lobata) . — Cuzco :

Anta, Vargas 201. Marcapata, Weberbauer 7789. — Puno: Acko-

pampa, Vargas 6978. Near Limbani, Vargas 9654- "Tomatillo

cimaron."

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

Saracha ciliata Miers, Ann. Mag. Nat. Hist. 2, 3: 446. 1849.

S. lacrima-virginis Bitter, Repert. Sp. Nov. 17: 341. 1921; I.e. 339.

Witheringia ciliata Miers, 111. S. Amer. PL 2: App. 56. 1857.

Suffruticose below, the upper branches angulate, decurrent-lined,

glabrate or sparsely pubescent including the corolla with many-celled

single trichomes; leaves ovate-oblong, cuneate decurrent into petiole

(this 0.5-4 cm. long), acutely or not obtusely acuminate, to about

8 cm. long, 4-5 cm. wide, mostly smaller, membranous, green, entire

or repand; flowers solitary in axils, peduncles 6-11 mm. long; pedi-

cels 5-7 mm. long, deflexed and 1-1.5 cm. long in fruit; calyx basally

connate, 6-8 mm. long, diameter about 2-2.5 cm., lobes ovate acu-

minate, 8 mm. long (10-11 mm. long in fruit), minutely stipitate-

glandular within at base; corolla rotate-campanulate, 2-2.5 cm. long,

3-3.5 cm. across (see note below), with 5 acuminate subcucullate

pubescent lobes 2-3 mm. long; filaments (4) 6-8 mm. long, glabrous

as ovary and style, the latter 7-8 mm. long, stigma capitate; berry

white, subglobose, about 12 mm. thick. — After Bitter, who found a

single sclerotic granule in the berry examined; he noted the corollas

of Weberbauer specimens as 1.5 cm. long, 2.5-2.8 cm. across, green-

ish; S. lacrima-virginis differs ex char, in calyx lobes 6 mm. long,

corolla white, 1.7-2 cm. long, 2 cm. across, glabrous within except

lobes at tips, filaments pubescent on adnate portion, calyx lobes 6-

7 mm. long in fruit, berry with 2 subapical granules; these differences

are probably individual rather than genetic. As in the similar S. Ur-

baniana and S. Weberbaueri there are 5 blood-red (or violet) glands

at base of filaments; from this character originated the fanciful native

name; the glands are present in a number of unrelated species.

Lima: San Geronimo, 5912. Mongomarca, Weberbauer 5235.

Prov. Huarochiri, Goodspeed & Metcalf 30248. Amancaes Hills,

Soukup 3124; Weberbauer 5246a. Lima Valley, (Mathews 834, type).

— Huanuco: Tambo de Vaca, 4469? — Arequipa: Posco, (Gunther &

Buchtien, as S. lacrima-virginis ace. to Bruns) ; 1587 (type, S. lacrima-

virginis}. "Lacrima de la virgin."

Saracha contorta R. & P. Fl. Peruv. 2: 43, pi 180a. 1799; 430.

Atropa contorta (R. & P.) Spreng. Syst. 1: 699. 1815. Bellinia con-

torta (R. & P.) Roem. & Schult. Syst. 4: 689. 1819. Witheringia

contorta (R. & P.) Miers, 111. S. Amer. PI. 2: App. 56. 1857. A. um-

bellata Roth, Cat. 2: 26. 1800. Saracha umbellata (Roth) DC. Cat.

Monsp. 142. 1813 (G. Don, Gen. Syst. 4: 451. 1838). A. Rothii Poir.

Encycl. Suppl. 1: 610. 1810. Bellinia umbellata (Roth) Roem. &

FLORA OF PERU 33

Schult. Syst. 4: 688. 1818. Witheringia umbellata (Roth) Miers, 111.

S. Amer. PL 2: App. 56. 1857.

An annual with entire but often angular-dentate leaves, the larger

to 8 cm. long, minutely or scarcely pubescent on both sides, decurrent

into a petiole one third as long, and with umbellate pale yellow rotate-

campanulate flowers 12 mm. wide; peduncles declined, 6-10-flowered,

somewhat contorted as also the pedicels; filaments villous at base;

berry black.— Morton, Proc. Biol. Soc. Wash. 51: 76, 77. 1938, sug-

gested that S. umbellata is referable to A. procumbens (Cav.) R. & P.

(and probably S. contorta R. & P.), which may be correct if the de-

velopment of pubescence on filaments is not a continuing (genetic)

character; I suspect that at least in the case of these similar plants

it is not, since apparently S. contorta has not been recognized in the

much-collected area of its type, this described as a procumbent fibrous

rooted annual; herbaceous stem bright violet below. Leaves useful

as a poultice and to relieve pain (Ruiz and Pavon) . F.M. Neg. 2549.

Lima: Rough places, Canta to Obrajillo, Ruiz, type.

Saracha dentata R. & P. Fl. Peruv. 2: 43, pi 179b. 1799; 432.

Atropa dentata (R. & P.) Spreng. Syst. 1: 699. 1815. Bellinia den-

tata (R. & P.) Roem. & Schult. Syst. 4: 689. 1819. Witheringia

dentata (R. & P.) Miers, 111. S. Amer. PI. 2: App. 56. 1857. Hebe-

cladus propinqua (Miers) Bitter, var. parviflorus Bitter, Repert. Sp.

Nov. 17: 250. 1921. H. Weberbaueri Bitter, I.e. 20: 372. 1924, not

Damm. Bot. Jahrb. 37: 638. 1906, nor S. Weberbaueri Damm. I.e.

S. Bitteri Macbr. Field Mus. Herb., apparently ined.

A shortly pubescent or puberulent more or less diffuse perennial

finally more or less ligneous, the many stems or branches from a thick

base, the lower 5 mm. thick or stouter; lower petioles 1.5 cm. long or

longer, the upper only a few mm. long; leaves unequal, the lowest

ovate, several cm. long, the upper geminate, the larger of each pair

about 1.5 cm. long, half as wide, the upper reduced, ovate-oblong-

lanceolate, all entire or undulate denticulate, broadest toward acute

base, somewhat attenuate to tip, usually densely puberulent as the

1-4 (or several) -flowered umbels, including the calyces; peduncles

2-6 mm. long, pedicels 3-5 mm. long (in fruit 6-10 mm. long) ; calyx

rotate, 6-8 mm. long, 5-6 mm. across, in fruit twice as wide, the lobes

acute to acuminate; corolla white, violet-veined or -tinted, green-

dotted below, rotate-campanulate, 6-10 mm. long and broad, the

short acute lobes pubescent as (in type) the filaments below, these

4-4.5 mm. long; anthers subcordate; berry copper-colored (always?),

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

about 7 mm. in diameter. — Specimens from southern Peru may be

noted as var. diversa Macbr., var. nov., corolla circa 5 mm. longa;

filamentis glabris vel minute sparseque puberulis. The development

of pubescence on filaments has been used (as in the key) for conven-

ience but it is doubtful that it is always, especially in itself, a true

indication of specific values. F.M. Negs. 2550; 2532 (H. Weberbaueri).

Leaves (combined with lard) used as a poultice for tumors (Ruiz

and Pavon).

La Libertad: Angas-marca, Weberbauer 7197 (type, H. Weber-

baueri).— Ancash: Lomas de Mongon, Goodsp. Exped. 9183. — Lima:

Obrajillo, near Canta, Ruiz & Pawn, type. Matucana, Weberbauer

5275 (distr. as H. umbellatus). Viso, Weberbauer 109 (type, var.

parviflorus). Near Matucana, Ferreyra 5^.22 (det. Cowan, H. um-

bellatus).— Ayacucho: Ocros, Stork & Horton 10799. Prov. Lucanas,

Metcalf 30317. Puquio, Ferreyra, 7181. — Apurimac: Andahuaylas,

Stork & Horton 10716. — Cuzco: Huanoquite, Vargas 2272. — Are-

quipa: Lower slopes of Misti, Sandeman 3808. Between 3,600 and

3,700 meters, Weberbauer 6853 (type, var. diversa).

Saracha diffusa Miers, Ann. Mag. Nat. Hist. 2, 3: 447. 1849;

683. Witheringia diffusa Miers, 111. S. Amer. PI. 2: App. 56. 1857.

Nearly glabrous, dichotomously branched, described as herba-

ceous; petioles slender, 12 mm. long; leaves ovate-elliptic, subob-

tusely acuminate, narrowed to base, about 7 cm. long, 4 cm. wide,

sinuose-serrate and erose-ciliate, obscurely glandular-puberulent both

sides on the reticulate veins; umbels 4-12-flowered, peduncle slender,

2-6 cm. long; pedicels filiform, glabrous, 1 cm. long or longer; calyx

rotate-angulate, 5-dentate; corolla rotate-angulate, 16 mm. wide,

lanate without and long-ciliate; filaments glabrous, about half as

long as corolla. — May be, as Morton indicated, Proc. Biol. Soc.

Wash. 51: 76. 1938, a part of S. procumbens (Cav.) R. & P. F.M.

Neg. 33016.

Lima: Hill of Puruchuco, Mathews 775.

Saracha Herrerae Morton, Rev. Univ. Cuzco 33 (87) : 96. 1944.

Branched from base, the stems to 6 dm. long, the ascending

branches often simple, pubescent toward apex with branched tri-

chomes, these found also (with simple trichomes) on the leaves be-

neath and the denticulate margins, and short (1 cm. long) peduncles,

calyx and corolla veins without, the indument (type) otherwise sim-

FLORA OF PERU 35

pie; leaves ovate, to 7 cm. long, 5.5 cm. wide, broadly or sometimes

narrowly cuneate to base, decurrent into petioles 2-2.5 cm. long,

sparsely strigose above, the trichomes few-celled; inflorescence axil-

lary, 2-flowered, pedicels shorter than peduncles, incrassate apically;

calyx green, about 2 cm. across, 5-parted, the long-triangular lobes

spreading, 8 mm. long, 5 mm. wide at base; corolla white, rotate-

campanulate, 2 cm. long, 3-3.5 cm. broad, the short lobes long-acu-

minate, glabrous except veins and margin, the intermediate lobes

minute, the tube sparsely pilose at base of filaments, these glabrous,

straight, shorter than the corolla; glands conspicuous, red; anthers

2.5 mm. long, 1.5 mm. broad; ovary subglobose, glabrous as style,

the stigma capitate. — After author, who, in allying it to S. ciliata

Miers, distinguished it by the branched pubescence. The genetic

continuity of character such as pubescence can be proven; cf. Reed

Rollins, Rhodora 60: 145-152. 1958.

Junin: Quebrada Occopilla, Soukup 3637 (det. Morton). — Huan-

cavelica: Yauli, Stork & Horton 10859 (det. Standley, S. ciliata). —

Apurimac: Saxaihuaman, Hen era, 3088 (det. Morton); 828. — Cuzco:

Paucartambo, Balls B6731 (det. Morton). Ollantaitambo, Cook &

Gilbert 417. Urubamba, Soukup 73; 740. "Ahuhaimantu" (Her-

rera); "Aguaymantu" (Vargas).

Hebecladus intermedius Miers in Hook. Lond. Journ. Bot.

4: 323. 1845; 470.

Stems glabrous; petioles scarcely 2 mm. long (or probably the

lower much longer); leaves ovate, angulate-sinuate, to 9 cm. long,

half as wide, sparsely pilosulous-puberulent both sides; umbels

(type) 3-flowered; peduncles 12 mm. long, pedicels 2 cm. long; calyx

8 mm. long; corolla-tube floccose-hirtellous, 2-2.5 cm. long, the

limb 8 mm. long, the sublinear lobes 6-8 mm. long, denticulate in

the plicae; filaments slender, glabrous, the sagittate anthers long-

exserted. — Compared with H. asperus (R. & P.) Miers but indu-

ment soft; flowers said to be similar to those of H. biflorus (R. & P.)

Miers, i.e. H. bicolor (R. & P.) Miers.

Lima: Puruchuco, (Mathews 524, type).

Saracha lobata Bitter, Repert. Sp. Nov. 18: 103. 1922.

A minutely pubescent (trichomes mostly simple, 3-7-celled), as-

cending herb; leaves shortly (3-5 mm.) petioled, broadly ovate or

rhomboid, about 3-3.5 cm. long and nearly as broad, irregularly and

sinuately 5-6-lobed; umbels 4-5-flowered, subsessile or the peduncle

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

5-6 mm. long; pedicels 6-7 mm. long, to 10 mm. in fruit; calyx stel-

late-rotate, minutely glandular within, especially near base, 5-6 mm.

wide, in fruit twice as wide; corolla pale violet, rotate, 8-10 mm.

wide, densely pubescent within at base of glabrous (1-1.2 mm. long)

filaments; anthers cordate, only 0.3 mm. long; style 2 mm. long,

stigma capitate; berry 6 mm. wide, sclerotic granules lacking. —

Probably section Eusaracha without sclerotic berry cells. F.M.

Neg. 2553.

Lima: Stony places, Huillacachi, southwest of Matucana, Weber-

bauer 206.

Saracha procumbens (Cav.) R. & P. Fl. Peruv. 2: 43, pi. 180b.

1799; 431. Atropa procumbens Cav. Icones 1: 53, pi 72. 1791. Bel-

linia procumbens (Cav.) Roem. & Schult. Syst. 4: 687. 1819. With-

eringia procumbens (Cav.) Miers, 111. S. Amer. PI. 2: App. 56. 1857.

A procumbent-ascending laxly dichotomously branched lightly

pubescent herb, the sulcate stems more or less fistulose; petioles mar-

gined by the decurrent leaves, 1-2 (-3) cm. long; leaves solitary

below, geminate above, subobtuse or acute, the larger often 5-8 cm.

long, 3-5 cm. wide, early somewhat pubescent beneath, finally lus-

trous and glabrous (typically) both sides, entire or undulate; solitary

axillary peduncles 2-4 (-6) -flowered, 8-10 mm. long; pedicels to

twice as long and nutant in fruit, pilose or glabrate as the calyces;

corolla ochroleucous, green-centered, early ovate, plicate, soon rotate-

campanulate, marginally sublanate; filaments glabrous and equal;

berry lustrous, black, edible. — Var. repando-dentata Dunal, I.e. (S. re-

pandidentata (Dunal) Bitter, Repert. Sp. Nov. 19: 269. 1924), has

densely ciliate acuminate leaves, sparsely pubescent beneath (tri-

chomes simple), peduncles to 2 cm. long, nearly glabrous as pedicels,

corolla 17 mm. wide, acute lobes to 5.5 mm. long, berry 12 mm. in

diameter (Bitter). Leaves have cleansing, healing virtues (Ruiz &

Pavon). F.M. Neg. 33015 (var.).

Lima: San Buenaventura, Ruiz & Pavdn, type. — Huanuco: Cu-

chero, Poeppig 1419 (type, var.). Rio Chinchao, 5060. — Arequipa:

Near Mejia, (Guenther & Buchtien 105; 106, fide Bruns).

Saracha propinqua Miers, Ann. Mag. Nat. Hist. 2, 3: 446. 1849;

683. Hebecladus propinqua (Miers) Bitter, Repert. Sp. Nov. 17: 246.

1921.

Type a divaricately branched slightly puberulent shrub 1 meter

tall, the often many-celled trichomes rarely branched, most abun-

FLORA OF PERU 37

dant on the flowers; petioles 1-nearly 3 cm. long; leaves ovate or

broadly lanceolate-ovate, broadly cuneate or subrotund at base,

acutely or obtusely acuminate, 3.5-6 cm. long, 2.5-4 cm. wide, sub-

entire or undulate-lobulate, firm-membranous; inflorescence axillary,

umbelliform, 3-6-flowered; peduncles 1.5-2 cm. long; pedicels 1 cm.

long or 1.5-2.5 cm. in fruit; calyx as corolla limb, the former 8-

10 mm. long (in fruit 11 mm.), the latter (typically) about 2.5 cm.

across, the spreading acute calyx lobes 3 mm. long, pubescent with-

out with patent trichomes, minutely glandular within especially be-

low; corolla pale blue, medially violet within, 2 cm. long, the glabrous

lower part shortly campanulate, little ventricose, the triangular acute

lobes 8-10 mm. long, 7 mm. wide, with 5 small teeth between them,

obscurely cucullate; filaments 11 mm. long, glabrous only toward tip;

style 16 mm. long, stigma capitate, bilobed; berry 5-6 mm. in diam-

eter; sclerotic granules 2. — After Bitter, who notes that the corolla

form is at variance to that of other species, resembling rather the

corolla of Saracha, and he concludes that these groups are closely

related. Illustrated, Miers, 111. S. Amer. PL 38. F.M. Neg. 33136.

Lima: Near Surco, Ferreyra 5429 (det. Cowan, H. umbellatus) ;

Soukup 3731. Prov. Huarachiri at Km. 81, Goodspeed & Metcalf

30242 (det. Morton). Puruchuco, Mathews 774. Viso, Goodspeed

Exped. 11530 (det. Johnston). Matucana, 339; Ferreyra 5417 (det.

Morton) ; Weberbauer 5275.

Saracha sordideviolacea Bitter, Repert. Sp. Nov. 18: 104. 1922.

A perennial herb with ascending branches, slightly pubescent, the

trichomes mostly simple, many-celled; petioles alate, 6-8 mm. long;

leaves broadly ovate or rhomboid-ovate, usually 3^4 cm. long, entire

or obscurely repand, sparsely short-pubescent; inflorescence sessile,

3-4-flowered, the pubescent pedicels 8-10 mm. long; calyx stellate-

rotate, 7 mm. wide, the acute lobes 2 mm. long; corolla sordid violet,

rotate, 1.5-2 cm. across, densely pubescent about the filaments, these

only 2-2.3 mm. long, glabrous or with a few trichomes; anthers emar-

ginate, 1 mm. long; style to 6 mm. long, stigma obtuse. — Probably

belongs to section Eusaracha without sclerotic cells (Bitter).

Lima: Rocky cliffs above Matucana, Weberbauer 5258, type; 2950.

Hebecladus umbellatus (R. & P.) Miers in Hook. Lond. Journ.

Bot. 4: 322. 1845; 469. Atropa umbellata R. & P. Fl. Peruv. 2: 44,

pi. 181, fig. a. 1799. Atropa revoluta Dietr. Nachtrage Gartn.-Lexik.

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

1: 422. fide Roem. & Schult. Syst. 4: 633. 1819. H. Turneri Miers,

I.e. 323, fide Bitter.

A rather fragile half -shrub, the minute trichomes 2-4-celled, the

ligneous lower branches to 5 mm. thick, the uppermost puberulent

as the geminate leaves and umbelliform inflorescence, this somewhat

glandular; petioles 1-2.5 (4.5) cm. long; leaves broadly ovate, the

largest lower subcordate, 7-10 cm. long, 5-7.5 cm. wide, the upper

reduced to about half the size, entire to repand-lobulate, membra-

nous, green, the 5-6 pale nerves and veins prominent beneath; pe-

duncles axillary or lateral, 5-10 mm. long, pedicels several to many,

7-9 mm. long; calyx 4 mm. long, 5-7 mm. across in flower, thrice as

broad in fruit, the ovate-lanceolate lobes 3 (-7) mm. long; corolla

tubular-campanulate, the dull violet tube 5-8 (-10) mm. long, 5-

6 mm. across at apex, the greenish finally recurved lanceolate acute

lobes 4-5 mm. long, intermediate teeth lacking; filaments little di-

lated at base, 12-13 mm. long, glabrous as ovary and style, this

finally 15 cm. long, the stigma subglobose-capitate; anthers 1.6 mm.

long, in age subsagittate; berry 7-8 mm. in diameter with 2 subapical

granules, the many reniform seeds obviously foveolate-reticulate.—

After Bitter, Repert. Sp. Nov. 20: 375. 1924; Weberbauer observed

a blood-red sap at base of filaments sometimes even filling the tube;

the odor of the foliage reminded him of Hyoscyamus species (perhaps

H. niger L. is cultivated in Peru). Apparently H. Turneri Miers,

named for A. Dawson Turner, an English horticulturist, would be

the correct name of the plant if included in Saracha, or H. sinuosus

Miers, I.e. 7: 352. 1848?

Amazonas: Chachapoyas, (Mathews, type, H. sinuosus?). — Lima:

Chancay, Lurin, Ruiz & Pavdn, type. Lomas de Lachay, Prov. De

Chancay, Ferreyra 3861. Atocongo, Nunez 212; Goodspeed Exped.

9294. (det. Johnston). Amancaes, (Mathews 722, fide Miers). Mt.

San Augustin, Weberbauer 5223 (5228).

Saracha Urbaniana Bitt. & Damm. Repert. Sp. Nov. 17: 442.

1921.

Suffruticose; pubescence of many-celled trichomes sparse and sim-

ple, mostly ciliate on the branchlets and leaves beneath, especially

on the veins; petioles 1-3 cm. long; leaves subentire, rather rhomboid,

usually 4-7 cm. long; inflorescence 1-flowered, the peduncle 5-8 mm.

long, the pedicel to 1 cm. long; calyx nearly 1 cm. connate, lobes

10-13 mm. long, within at base minutely glandular; corolla bluish-

white, broadly campanulate, 3-3.8 cm. long, about 4 cm. wide, pubes-

FLORA OF PERU 39

cent, the lobes 5-8 mm. long, within at base glabrous but with 5 violet

glands; filaments 12-15 mm. long, pubescent below, style 1.5 cm.

long; berry about 14 mm. in diameter with 2 sclerotic cells near apex.

—A beautiful species, near S. ciliata Miers but with very much larger

flowers, and related to S. Weberbaueri Damm. but with different and

pubescent foliage.

Lima: Above San Bartolome', Weberbauer 5262; 5305.

Hebecladus ventricosus Baker in Saund. Refug. Bot. 3: pi. 208.

1870.

Erect shrub, glabrous or essentially except for puberulence on

calyx, and cilia on margins of ovate leaves, these 7 cm. long or longer;

peduncles and pedicels subequal; flowers usually 2, rarely solitary,

nutant; calyx spreading, accrescent after anthesis; corolla pale yellow,

somewhat purplish at base, campanulate-ventricose (constricted at

throat), 12 mm. long and wide, the lanceolate lobes recurved, minute

teeth in the intervals; stamens and style much exserted, the latter

finally about 2 cm. long. — Apparently the filaments as corolla are

glabrous; the author remarks that corolla shape is different from

any species described by Dunal but except in corolla resembles

H. viridiflorus (HBK.) Miers, 469, of Ecuador; that, however, has

a tubular corolla; rather it seems to be a cultivated state of S.

biflora R. & P. Type by Farris (not seen), no data; flowers 1.5-2

cm. long in cultivated examples referred here.

Peru: Without locality (Farris, type).

Saracha villosa (Zucc.) G. Don, Syst. 4: 1838; 430. Atropa

villosa Zucc. in Roem. Coll. 130. 1809. S. erecta (Zucc.) Roem. &

Schult. Syst. 4: 684. 1818. Witheringia villosa (Zucc.) Miers, 111.

S. Amer. PI. 2: App. 56. 1857. S. sinuosa Bitter, Repert. Sp. Nov.

18: 101. 1922?, vix Miers, in Hook. Lond. Journ. Bot. 7: 352. 1848.

Stems more or less woody; branches and leaves, especially be-

neath, usually densely viscid-villous; petioles winged, 12-15 mm. long;

leaves entire or sinuate, cordate-ovate, acuminate, the larger about

5 cm. long; umbels 5-9-flowered, the villous peduncle 6-8 mm.

(-1.5 cm.) long, the puberulent cernuous pedicels papillose, 4-6

mm. long; calyx accrescent; corolla greenish-yellow, rotate, small

or large, somewhat pubescent. — As suggested by Dunal, probably

this and several other names are a part of S. contorta R. & P. ; appar-

ently the plant of Miers is allied to or the same as H. umbellatus.

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

Lima: North of Barranca, Morrison & Beetle 9099; Goodspeed

Exped. 9228.

Hebecladus Weberbaueri Damm. Bot. Jahrb. 37: 638. 1906.

A puberulent shrub with greenish tubular flowers in axillary

umbels of 3-5; petioles 6-10 mm. long; leaves ovate-lanceolate,

obliquely cuneate at base, 3-6 cm. long, 2-4.5 cm. wide, peduncles

1.5-4 cm. long, pedicels 1-2 cm. long; calyx 17 mm. across, lobes

ovate, acute, 5 mm. long, stellulate without; corolla tube pilose

within, floccose without, 28 mm. long (as style), the larger teeth

6 mm. long, the smaller 1 mm.; stamens little exserted, filaments

pilose, anther sagittate, 3 mm. long, stigma clavate; berry semi-

included, 12 mm. in diameter. — Apparently related to H. intermedius

Miers but ashy pubescent even to the stems. Doubtfully distinct.

Ancash: Grass-shrub formations, Ocros, Prov. of Cajatambo,

Weberbauer 2652.

Saracha Weberbaueri Damm. Bot. Jahrb. 37: 638. 1906; 344.

S. pallascana (Bitter) Macbr. Field Mus. Bot. 8: 109. 1930. S.

Weberbaueri Damm., subsp. palascana Bitter, Repert. Sp. Nov. 19:

267. 1924.

Related to S. Urbaniana Bitt. & Damm. but nearly glabrous

and herbaceous above; leaves broadly ovate, rounded to shortly

acute base, more or less long-acuminate or caudate-acuminate;

branches supported in shrubs; peduncle 0.5-1.5 cm. long; pedicels

1.5-3 cm. long; connate part of calyx 12 mm. long in fruit; corolla

violet with green veins, 3.5-4.5 cm. long, 3.5-5.5 cm. wide, lobes

5-6 mm. long, anthers 4 mm. long; style 17-22 mm. long; berry

15-16 mm. thick. — Type a perennial herb but certainly ligneous

at base (Bitter); to 2 meters tall. Dammer noted the basally

pilose filaments as unequal; Bitter remarks that this is due to

their development at different times, finally becoming equal in

length. Perhaps always a half-shrub in age; leaves sometimes only

3-4 cm. long, half as wide, peduncles even obsolete, pedicels a cm.

long or less, corollas greenish-white, bluish-green at base within,

about 3 cm. long, 3.5 cm. broad. Such characters can scarcely be

concomitant or taxonomically significant even varietally; neverthe-

less, they indicated a subspecies (pallascana) to the over-industrious

Bitter; surely my usually protective laziness failed me that I ever

recorded it as a species.

Ancash: Ocros, Prov. of Cajatambo, Weberbauer 2657, type. —

Lima: Above Supe, (Weberbauer, 170).

FLORA OF PERU 41

9. DUNALIA HBK.

Dierbachia Spreng. Syst. 1: 512, 676. 1825. Acnistus Schott,

Wien. Zeitschr. 4: 1180. 1829, fide Kuntze. lochroma Benth. Bot.

Reg. 30: pi. 20. 1845. Lycioplesium Miers in Hook. Lond. Journ.

Bot. 4: 330. 1845. Chaenesthes Miers, I.e. 336. Cleochroma Miers,

I.e. 7: 349. 1848. Codochonia Dunal in DC. Prodr. 13, pt. 1: 482. 1850.

References: Dunal in DC. Prodr. I.e. 482 et ref. 1852; Sleumer,

Lilloa 23: 117-142. 1950.

Shrubs, often spinescent and then sometimes simulating Lycium

L. but at least as to Peru with larger leaves, longer, narrower fascicu-

late-umbellate flowers. Calyx not or slightly accrescent, but more or

less supporting to rarely sub-enclosing the berry (lochroma), tubular

to subcampanulate, somewhat unequally dentate, sometimes bilobed

or laterally (especially in age) cleft. Corolla subcampanulate to

infundibuliform-tubular or tubular, often slightly curved, the lobes

plicate or more or less induplicate-valvate. Filaments dilated or

not below, smooth or minutely appendaged, glabrous or more or

less pubescent, usually affixed basally or submedially. Calyx if

accrescent (lochroma) not inflated. Seeds reniform, somewhat com-

pressed, embryo more or less arcuate. Flowers rarely 6-merous;

Bitter noted much variation in size and count (some to 50 or about

80) of stone granules. It is doubtful if there is a natural distinction

between lochroma Benth. and Dunalia HBK., the characters appar-

ent when the original species were known being, it seems, actually the

same, developed in varying degree.

Sleumer, I.e., has given a precise key to related genera (Solaninae)

and to the species of Argentina and Bolivia but he avoids placing

the variable D. arborescens (L.) Sleumer! However, he has discussed

that species and helpfully listed all names considered valid as well

as many probably referable to Dunalia; he has followed Kuntze

in including Acnistus Schott, contrary to Hitchcock, Ann. Mo. Bot.

Gard. 19: 198. 1932, the latter using calyx and form of corolla as

basic generic indications; a number of groups could quite as logically

and much more conveniently be treated together as sections under

fewer common (generic) names, as the differences are often trivial

or somewhat variable; Nicotiana L., as recently defined, exhibits

great diversity in corolla-form, Cestrum L. in stamen character.

Omitted from key is the spinescent Fregirardia Dunaliana Wedd.

with small dark violet corollas, ex char, probably a Dunalia, sens. lat.

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

KEY TO DUNALIA (including IOCHROMA)

Corolla more or less campanulate; lobes and tube subequal; flowers

few to many, often at leafless nodes D. arborescens.

Corolla tubular to funnelform; lobes clearly shorter than tube;

flowers 1-many, often with leaves.

Leaves alternate or subopposite, subequal, usually some or all

4 cm. wide or wider; calyx somewhat accrescent, sometimes

slightly or partly enclosing berry; spines always lacking.

Corolla 3-6 cm. long, orange-red, blue or violet; filaments pu-

bescent except /. grandiflorum.

Corolla orange-red, densely tomentose, about 3 cm. long.

D. peruviana.

Corolla blue-purple or violet, puberulent or pubescent, 3-6

cm. long.

Filaments puberulent or pubescent below; corolla 3-4 cm.

long, limb 1 cm. broad D. tubulosa.

Filaments glabrous below; corolla to 6 cm. long, throat

expanded, to 2.5 cm. broad /. Warscewiczii.

Corolla about 2 cm. long, greenish-yellow or reddish to purple-

red; filaments glabrous (known).

Flowers 6-merous, reddish; filaments nearly free; anthers 3-4

mm. long D. solanoides.

Flowers 5-merous, usually yellowish; filaments affixed sub-

medially; anthers 2.5-3 mm. long Z). umbellata.

Leaves at least early unequal in fascicles, the largest rarely 4 cm.

wide (unless D. lycioides, D. spinosa, both spinescent), all

often much smaller; calyx scarcely or not accrescent.

Calyx early truncate, typically 5-cusped or -apiculate; leaves

mostly or all widest submedially, to 5 cm. long; corolla

10-12 (15) mm. long, pubescent within.

D. brachystemon, D. spathulata.

Calyx soon lobate or denticulate; leaves usually somewhat obo-

vate or oblong-spatulate, rarely 5-8 cm. long; corolla (full

an thesis) at least 1.5 cm. long.

Leaves mostly or all wider than 5 mm. and longer than 1 cm.;

spines mostly lateral or lacking.

Indument known within corolla near base (types) ; filaments

obscurely to clearly appendaged (types); branchlets

usually spiny.

FLORA OF PERU 43

Leaves 1-2.5 cm. long or if to 4 cm. long, distinctly

obovate; corolla 1.5-2.5 cm. long (following may

be 2 or 3, even a single variable entity) .

Filaments (type) free nearly to base; leaves more or

less obovate, usually about 1 cm. wide or wider.

D. obovata.

Filaments about one-third adnate; leaves lance-

spatulate to elliptic-obovate, 5-12 mm. wide

(types) or wider.

Appendages short, membranous (or none?); fila-

ments glabrous D. lycioides.

Appendages 6-7 mm. long; filaments puberulent.

Leaves suboblong, undulate D. angustifolia.

Leaves elliptic-obovate, plane . . . . D. Weberbaueri.

Leaves mostly or all 3.5-8 cm. long, often spatulate;

corolla 2.5-4 cm. long.

Corolla 2.5 cm. long; filament teeth bifid, tomentose;

type espinose D. lilacina.

Corolla 3-4 cm. long; filament teeth glabrous; spines

present (always?) D. Besseri, D. spinosa.

Indument unknown inside corolla; filaments smooth (types,

unless D. lycioides); branchlets often espinose or axil-

lary, (nodal) spines 1 cm. long.

Corolla to 2 cm. long; leaves sometimes several cm. long.

D. lycioides.

Corolla to 3 cm. long; leaves 4-10 mm. wide.

D. dolichostyla.

Corolla half as long; leaves usually 12-15 mm. wide (un-

less D. horrida).

Leaves (larger) 12-15 mm. wide; spines none.

D. Dombeyana.

Leaves 4-6 mm. wide; spines stout, apical or nodal.

D. horrida.

Leaves about 5 mm. wide, 10-12 mm. long or smaller; branch-

lets often many, 2-2.5 cm. long, spine-tipped . .D. horrida.

Dunalia angustifolia Damm. Bot. Jahrb. 50, Beibl. Ill : 54. 1913.

Branchlet tips, younger leaves beneath and flowers more or less

puberulent; nodal spines 1-2 cm. long; petioles a few (to 10) mm.

44 FIELD MUSEUM OF]NATURAL HISTORY — BOTANY, VOL. XIII

long; leaves narrowly oblong-elliptic, acute at base, subobtuse, 1.5-

2.5 cm. long, 4-8 mm. wide, often strongly undulate-margined;

pedicels solitary, 1 cm. long; calyx 4-5 mm. wide and long, broadly

ovate lobes 1.5-2 mm. long; corolla violet, 2-2.5 cm. long, about

6 mm. across at top, puberulent also within at base as the unequal

filaments, these 19-24 mm. long, appendages 7-10 mm. long; anthers

4 mm. long; style 22-27 mm. long, stigma capitate. — Sleumer, I.e.

122, suggested that this and D. Besseri may be the same, a possibility

noted in manuscript by me. F.M. Neg. 2515.

Arequipa: Base of Misti, Weberbauer 4828, type.

Dunalia arborescens (L.) Sleumer, Lilloa 23: 124. 1950. Atropa

arborescens L. Amoen. Acad. 4: 307. 1759. D. campanulata (Lam.)

Macbr. Field Mus. Bot. 8: 109. 1930. Oestrum campanulatum Lam.

Encycl. 1: 688. 1789. Lycium aggregatum R. & P. Fl. Peruv. 2: 45,

pi. 182a. 1799. L. guayaquilense HBK. Nov. Gen. & Sp. 3: 50. 1818?

L. floribundum HBK. I.e. L. ovale Roem. & Schult. Syst. 4: 698.

1819? Acnistus guayaquilensis (HBK.) G. Don, Gen. Syst. 4: 461.

1838. A. floribundus (HBK.) G. Don, I.e. A. sideroxyloides G. Don,

I.e.? Pederlea aggregate, (R. & P.) Raf. Sylva Tellur. 54. 1838. A.

aggregatus (R. & P.) Miers in Hook. Lond. Journ. Bot. 4: 341. 1845.

A. confertiflorus Miers, Ann. Mag. Nat. Hist. 2, 4: 31. 1849? A.

arborescens (L.) Schlecht. Linnaea 7: 67. 1832?

A shrub or tree sometimes 10-15 meters tall, with often ample

petioled oblong to elliptic leaves, more or less floccose at least beneath

especially when young, and greenish-white or cream-colored fragrant

flowers, few to many in fascicles along the light-barked cicatrose

branchlets; pedicels about 6-15 mm. long, slender or subfiliform

except at apex or stouter when shorter; calyx glabrate to floccose,

truncate and sometimes obscurely toothed, becoming irregularly 3-5-

lobed in age; corolla campanulate-funnelform, 8-17 mm. long, pu-

bescent within only at base of filaments, glabrous to puberulent

without; stamens often barely, sometimes shortly, exserted; fruit glo-

bose, finally orange color, 4-5 mm. thick. — This may be treated

as a variant of D. arborescens (L.) Sleumer, typically of the West

Indies (Central America, Brazil, fide Sleumer), with very slender

pedicels and soon much exserted stamens; possibly it will be found

merging with forms of the widely distributed D. arborescens, which,

as noted by Sleumer, may include a number of species but most

likely not Peruvian, described under Dunalia, Acnistus, Lycioplesium

and Lycium; the Peruvian collections vary considerably even in

FLORA OF PERU 45

characters of pedicels and flowers but may be classified as D. arbo-

rescens (L.) Sleumer, var. campanulata (Lam.) Macbr., comb. nov.

Miers, I.e. 340, described A. arborescens (L.) Schlecht. as variable

in size of leaves but glabrous as pedicels, these slender, and corolla,

this very short, broader in proportion than in any other species,

8-10 mm. long, 4 mm. across, calyx smooth, membranous, stamens

far exserted. Var. campanulatum (i.e. L. aggregatum) is more or less

pubescent, stamens scarcely exserted to exserted, leaves tomentulose

beneath. Leaves of A. guayaquilensis glabrous above, pubescent

beneath, as L. floribundum. A. sideroxyloides G. Don, I.e. from

Rio Magdalena (Lima or Cajamarca) has no distinctive character.

Often found near partly dry stream or on rocky slopes.

Earthenware used on a fire of the wood, which burns noisily,

often breaks, whence the name "quiebra ollas" (Ruiz and Pavon);

the leaves with lard are used as an emollient (Ruiz and Pavon);

for rheumatism (Klug).

Cajamarca: Rio Magdalena, Bonpland (type, L. floribundum;

also A. sideroxyloides?). — San Martin: Jepelacio, Klug 3565. Rioja,

Woytkowski 27. — Ancash: Weberbauer 2648. — Lima: Chosica, 533;

Asplund 13768; Rose 18547; Ferreyra 2840; Mexia 04001 (det.

Johnston, A. umbellatus). Atocongo, Pennell 14754- Pachacamac,

Mexia 8098 (det. Morton); 04047 (det. Johnston, A. umbellatus).

Chancay, Ruiz & Pavdn (type, L. aggregatum) ; Wilkes Exped.; Eyer-

dam 9068; 9121 (det. Johnston, A. umbellatus; 9121 abnormal from

a "rust"). — Huanuco: Chulki, Sawada 59. Near Huanuco, 2313;

Ruiz & Pavdn. Muna, 4004- Huacachi, 4142- Pampayacu, Kane-

hira 229. — Loreto: Iquitos, Killip & Smith 27157 (det. Moldenke).

To the Caribbean and Central America. "Quiebra ollas" or "ma-

capaqui."

Dunalia Besseri Damm. Bot. Jahrb. 50, Beibl. Ill: 56. 1913.

Young branches more or less tomentose puberulent; leaves sub-

lanceolate, 3.5-5 cm. long, 5-8 mm. broad; calyx-teeth ovate-

triangular, about 2 mm. long; corolla puberulent, tubular, about

3 cm. long, 5 mm. across at apex, within at base pilose-puberulent,

the teeth 1.5 mm. long; filaments puberulent, the entire (?) append-

ages about 6 mm. long. — Perhaps only a form of D. spinosa (Meyen)

Damm.; the scrap type shows no spines; the large leaves, long-

cuneate to base and slightly narrowed to obtuse tip, simulate those

of D. spinosa; Bitter found granules not solidified, so no count.

F.M. Neg. 2516.

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

Peru(?) : von Besser, type.

Dunalia brachystemon A. Br. & Bouche*, Ind. Sem. Hort.

Berol. App. 6. 1861.

Near D. spathulata (R. & P.) A. Br. & Aschers. but leaves mostly

2.5-5 cm. long, and, especially, filaments broadly bidentate, the free

portion scarcely longer than the anthers. — Probably a form; the type

from hedges and vicinity.

Huanuco: Chulki, Sawada 52. In serpentine, north of Huanuco,

Stork & Horton 9402 (det. Standley).

Dunalia dolichostyla (Bitter) Macbr. Field Mus. Bot. 8: 109.

1930. Acnistus dolichostylus Bitter, Repert. Sp. Nov. 21: 85. 1925.

A densely branched sometimes sprawling spiny shrub with obo-

vate fascicled leaves 1-1.5 (3) cm. long, 4-10 mm. broad; pedicels

8-10 mm. long, clustered; calyx campanulate, about 5 mm. long,

nearly as broad, the broad teeth scarcely 0.5 mm. long; corolla dull

violet, pubescent above, narrowly elongate funnelform (broadly in

some species), 2.5-3 cm. long; filaments adnate below, free for 12

mm., unappendaged; style (stigma small) exserted 7-9 mm., stamens

also or these included. — The style is not included in specimens with

exserted stamens; filaments free nearly to base, glabrous, as corolla

within, this narrower in type than in later specimens which seem

to be espinose but only younger branches present on specimens seen.

Remarkable species intermediate between true Dunalia and the sec-

tion Acnistus (Bitter). F.M. Neg. 2485.

La Libertad: Prov. Santiago de Chuco, Weberbauer 7087; Stork

& Horton 9980 (det. Standley); West 8166 (det. Johnston, D. obo-

vata). Valle de Moche, Nunez 2989.

Dunalia Dombeyana (Dunal) Macbr., comb. nov. Lyciople-

sium Dombeyanum Dunal in DC. Prodr. 13: 492. 1852.

A shrub apparently most similar to D. dolichostyla; type spineless;

leaves (the larger) only 1.5-2 cm. long, 12-14 mm. wide; calyx

obconic-poculiform-campanulate, to 4 mm. long and broad, the broad

unequal lobes whitish scarious marginally; corolla only 4 mm. broad,

13-15 mm. long (red?), glabrous within as the filaments, these in-

serted toward the base; style included but longer than the equal

stamens, anthers cordate. — Ex char, this is not exactly D. obovata

to which Dunal allied it, but diagnostic specific values and their

variation potential in this genus are as yet inadequately determined.

FLORA OF PERU 47

D. Pflanzii Damm., Bot. Jahrb. 50, Beibl. Ill: 57. 1913, from near

La Paz, Bolivia, is D. lycioides Miers, fide Sleumer but may be re-

lated to D. Dombeyana, ex char.; corolla glabrous within, filaments

6 mm. above the base, membranous appendages 1 cm. long, leaves

2.5-8.5 cm. long, 0.5-2.5 cm. wide.

Junin: Near Tarma, (Dombey), type.

Fregirardia Dunaliana Wedd. Chloris And. 2: 100. 1857.

Shrub to 2 meters tall, branches light reddish-brown, apically

pubescent, armed (the type at least) with sharp spines 1.5-2 cm.

long; leaves lanceolate, acute or rarely obtuse, decurrent to petiole,

3-6 cm. long, 5-15 mm. wide, pale green and puberulent both sides;

umbels 2-4-flowered, subaxillary, pedicels 1.5-2 cm. long; calyx about

4 mm. long (after anthesis strongly accrescent), poculiform, pubes-

cent, subtruncate between the remote linear-subulate teeth; corolla

purple, rotate-subfunnelform, very hirsute in throat, scarcely half

longer than calyx; anthers bluish; berry orange, globose, apiculate,

finally 8 mm. in diameter, clasped at base by the enlarged calyx,

the calyx lobes then unequal. — Similar to F. eriolarynx Dunal, 503,

of Bolivia, as to type spineless, the leaves glabrous beneath, corolla

three times longer than calyx; it is probably a small-flowered species

of Dunalia.

Tacna: Palca, (Weddell, type). Bolivia.

Dunalia horrida (HBK.) Macbr. Field Mus. Bot. 8: 109. 1930;

493. Lycium parvifolium Roem. & Schult. Syst. 4: 698. 1819. Ly-

cium horridum HBK. Nov. Gen. & Sp. 3: 52. 1818. Lycioplesium

horridum (HBK.) Miers in Hook. Lond. Journ. Bot. 4: 331. 1845.

Stoutly branched, the approximate uppermost nodes spinose or

producing short divaricate spine-tipped glabrate branchlets (in type

2-4 cm. long) ; leaves 4-7-fasciculate, attenuate into the very short

petioles, obovate or suboblong, obtuse, 8-12 mm. long, about half

as wide, glabrous, lustrous, the veins obscure; flowers violet, pe-

duncles pulverulent, arcuate-pendulous, filiform, 6-8 mm. long; calyx

glabrous, semiglobose-campanulate, about 3 mm. long, obsoletely

denticulate, nearly truncate, glabrous as the tubular corolla, this

violet, slightly curved, to about 1.5 cm. long, equally and acutely

5-dentate, marginally pilosulous, subequaled by the basally inserted

capillary glabrous filaments; style slightly longer than stamens,

anthers oblong; berry black. — Incompletely known but suggests a

desert or upland version of D. Dombeyana or D. dolichostyla; corolla

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

glabrous within in Ferreyra's collection, which matches photo. F.M.

Neg. 2490.

Used for fences (cercos), according to Ferreyra.

Cajamarca: Near Magdalena, Bonpland, type. Cumbre Mayo,

Nunez 1601. — Ancash: Chiquian, Ferreyra 6196. "Churur" (Ferrey-

ra).

Dunalia lilacina Miers, Ann. Mag. Nat. Hist. 2, 4: 32. 1849; 687.

Unarmed shrub with fascicled glabrous spathulate oblong leaves

about 1 cm. wide, 3.5 cm. long, attenuate into petiole 2 cm. long,

and lilac narrowly funnelform flowers about 2.5 cm. long, 8 mm.

broad at throat, borne in axillary fascicles; peduncles 12 mm. long;

calyx campanulate, 5-nerved, the rounded teeth mucronate; corolla

scarcely puberulent without, pubescent below within the nearly en-

tire puberulent limb with 5-6 acute teeth; stamens as style included,

3 shorter, filaments glabrous but with white tomentose fleshy teeth.

— Said to differ from all other species by the bifid very tomentose

filament-appendages. Known only from cultivation at Kew, origin

Peru, fide Index Kewensis.

Peru (perhaps).

Dunalia lycioides Miers in Hook. Lond. Journ. Bot. 4: 334.

1845; 485.

Strongly spinescent shrub 2 meters tall or taller, glabrous unless

the purple more or less nutant solitary or geminate flowers; branches

flexuose, the axillary spines sometimes several cm. long, sometimes

absent; petioles 4-6 mm. long, 1-3-fasciculate; leaves fleshy, lanceo-

late-spathulate, decurrent, rounded apically, about 18 mm. long,

5 mm. wide or larger; pedicels filiform or slender, to about 1 cm.

long; calyx early puberulent, 5-costate, the short subovate mucron-

ulate lobes lanate at tip; corolla deep purple (crimson, Miers), 2 cm.

long, limb 8-10 mm. across, lobes short, marginally floccose, the alter-

nate lobules membranous; filaments purple, adnate one-third, free

8-10 mm., the short acute lateral appendages membranous; anthers

oblong, yellow, scarcely or more or less exserted; berry finally red,

nearly 1 cm. in diameter; seeds compressed-reniform, yellowish, gran-

ules 24-26 (Bitter). — After Dunal (mostly, as usual by Dunal, from

author); perhaps should include similar species, as D. dolichostyla,

etc.; question of the validity of floral characters. Stork & Horton

10708, referred by Standley with query to D. spathulata, may not be

this species; calyx in flower denticulate, corolla 2 cm. long, leaves 4

FLORA OF PERU 49

cm. long; possibly is D. Pflanzii Damm. (see note under D. Dombey-

ana). According to Sleumer, I.e. 141, Bolivian specimens referred

here by him have mature leaves 4-5 cm. long, 1.5 cm. wide. The

stamens, fide Sleumer, are not so far exserted as illustrated by Miers

(pl.2,f.l).

Lima: (Mathews). — Junin: Tarma and Jauja, (Mathews 850, type).

— Ayacucho: Toward Mataral, 3,500 meters, West 3653 (det. John-

ston).— Apurimac: Andahuaylas, West 3745 (det. Johnston, D. obo-

vata) ; Stork & Horton 1 0708? Prov. Avancam, Vargas 2278—Cuzco :

Hacienda Churu, Prov. Paucartambo, Herrera 1023a (vel affine,

Morton). — Arequipa: Above Cotahuasi, 3,700 meters, Weberbauer

6864. Mt. Chiwata, Eyerdam & Beetle 22130 (det. Johnston).—

Puno(?):NearPuno(T),&>tdfc*p#«. Bolivia; Colombia. "Tankar,"

"majra-huayra" (West).

Dunalia obovata (R. & P.) Damm. Bot. Jahrb. 50, Beibl. Ill:

56. 1913; 492. Lycium obovatum R. & P. Fl. Peruv. 2: 46, pi. 183c.

1799. L. peruvianum Hort. ex Dippel, Hand. Laubholzk. 1: 30. 1889,

fide C. L. Hitchcock. Lycioplesium obovatum (R. & P.) Miers in

Hook. Lond. Journ. Bot. 4: 330. 1845.

Glabrous more or less spinose shrub sometimes 2 meters tall or

taller, the erect terete light-colored stems with diffuse or spreading

branches, adult often naked; leaves at first fasciculate, unequal, later

few, obovate, alately decurrent on petioles 4-16 mm. long, obtuse,

3-4 cm. long, 12-15 mm. wide, primary nerves 4-5, faint even be-

neath; peduncles slender, incrassate above, solitary or geminate, 12-

15 mm. long; calyx poculiform, 4 mm. long, 5 mm. across, lobes acute

or subacute, little mucronulate; corolla purple-violet, tubular, 14-

18 mm. long, slightly ampliate to pubescent limb where 8 or 9 mm.

in diameter, the short triangular teeth flocculose marginally; stamens

included or anthers visible, filaments glabrous except for villous base;

style exserted, stigma clavate, subbifid; berry orange, globose, the

small seeds lutescent. — Filaments with obscure to 3 mm. long ap-

pendages (Dammer). Dammer, I.e. 53-58, as Dunal, proposed or

accepted several segregates of D. obovata based largely upon varia-

tions in floral pubescence and development of filament-appendages;

the value of the former character and possibly of the latter is open

to question. Perhaps this should include some if not all of the shrubs

assigned specific names on corolla characters; corolla deep blue, an-

thers white to yellow (Killip & Smith). F.M. Neg. 2521.

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

Junin: Tarma, Ruiz & Pavon, type; Killip & Smith 21976; Velarde

Nunez 637. Chongos Bajo, Soukup 3616. Quebradas de San Mateo,

Isern 2538. "Espino."

Dunalia peruviana (Dunal) Macbr., comb. nov. Chaenesthes

gesneriodes (HBK.) Miers, var. peruviana Dunal in DC. Prodr. 13,

pt. 1 : 488. 1852. lochroma peruvianum (Dunal) Macbr. Field Mus.

Bot. 8: 107. 1930.

Upper branches and fasciculate pedicels densely villous-tomen-

tose; petioles about 1 cm. long; leaves ovate, in type about 5 cm.

long, scarcely half as wide, probably larger in age, apparently gla-

brate above, somewhat pubescent beneath; pedicels slender, incras-

sate above, 3.5-4 cm. long; calyx short, often truncate or unequally

5-dentate, one segment bilobed, the other trilobulate; corolla orange-

red, closely tomentose, 3 cm. long, may be longer; stamens and style

subequal, subexserted, filaments apparently pubescent, stigma capi-

tately bilobulate. — After Dunal (as most descriptions) but ex char,

apparently specifically distinct from the Colombian species of HBK.

with equal rounded mucronulate calyx lobes, style longer, stigma tri-

lobulate; the related /. fuchsioides (HBK.) Miers, 487, of Ecuador

has glabrous corolla; I. cornifolia (HBK.) Miers, 488, seemingly also

/. loxensis (HBK.) Miers, 488, have pubescent filaments, stamens

included, and may be near D. umbellata (R. & P.) Macbr.

Amazonas: Chachapoyas, (Mathews, type).

Dunalia solanoides (Dunal) Macbr. Field Mus. Bot. 8: 109.

1930; 482. Codochonia solanoides Dunal in DC. Prodr. 13, pt. 1:

482. 1852.

A more or less puberulent tomentose shrub; petioles 7-10 mm.

long, stipules geminate in the axils, ovate-oblong, obtuse, 6-8 mm.

long, 4-6 mm. wide; leaves ovate or lanceolate, acuminate, more or

less attenuate to acute or sometimes rounded at base, 4-7 cm. wide,

to 1 dm. long or longer with (11) 12-13 primary nerves, reticulate

veins; flowers in terminal or axillary fascicles or racemiform umbels,

funnelform-campanulate-salverform calyx 4 mm. long, 5-6 mm.

across in anthesis, in fruit 8-10 mm. broad, the broad unequal teeth

obscurely mucronate; corollas reddish toward base, slightly pubes-

cent without, glabrous within, the tube 15 mm. long, 6 mm. wide,

at yellowish apex 10 mm. across, the 5 (6) ovate lobes 5-7 mm. long,

nearly as broad; filaments 8-10 mm. long, glabrous, affixed at base,

anthers 3-4 mm. long; berry black (1 cm. thick) about half enclosed

FLORA OF PERU 51

in the calyx; seeds subreniform, subrostrate and emarginate at base,

ochraceous-rufescent; granules none, fide Bitter. — Type 6-merous

(Codochonia Dunal); approaches the subgenus lochroma but char-

acter probably not specific and if not, there is doubtfully a true dis-

tinction from D. umbellata (R. & P.) Macbr. Corolla 1 cm. long,

greenish-lead color (West). The type, Dombey 335, without locality,

probably came from Lima or Huanuco. F.M. Neg. 6870.

La Libertad: Huamachuco, West 8192. Prov. Otuzco, Lopez

M.0314. Trujillo, Velarde Nunez 1*52. — Lima: Surco, Soukup 4236.

Dunalia spathulata (R. & P.) A. Br. & Aschers. Ind. Sem.

Hort. Berol. App. 6. 1861; 500. Lycium spathulatum R. & P. Fl.

Peruv. 2: 46, pi. 183a. 1799. Acnistus spat(h)ulatus (R. & P.) G. Don,

Gen. Syst. 4: 461. 1838. Huanuca spathulata (R. & P.) Raf. Sylva

Tellur. 54. 1838. D. acnistoides Miers in Hook. Lond. Journ. Bot.

7: 338. 1848?

Glabrous, often rather intricately branched always spineless shrub,

ordinarily a meter or two tall; branchlets divaricate, subangulate or

subarcuate; leaves scattered or fasciculate, usually obovate-spatu-

late, not infrequently oblong-subelliptic, but long-attenuate below

the middle, canaliculately and subalately decurrent on the slender

petiole (this finally 1-2 cm. long), subobtuse, unequal, the larger to

7 cm. long, 3 cm. wide, often smaller, especially when fasciculate;

nerves rather prominent on opaque lower surface, visible on the

slightly lustrous-resinous upper; axillary or subaxillary sessile um-

bels with 1-20 1-flowered slender apically incrassate unequal pedun-

cles often lax in flower, many about 1 cm. long; calyx subtruncate,

obscurely 5-apiculate, cupulate, about 3 mm. long and broad, in fruit

unevenly lobulate, 6 mm. broad; corolla purplish-red or purple, at

full anthesis 10-15 mm. long, slightly ampliate, the 1.5-2 mm. long

acute lobes puberulent, especially marginally; stamens included, fila-

ments about 6 mm. long, inserted 2 mm. above the base, pubescent

below and ordinarily with appendages 3-5 mm. long; anthers oval,

3 mm. long, yellowish-violet; stigma subglobose; berry orange-

yellow, 8 mm. in diameter, the lutescent scrobiculate seeds com-

pressed-orbicular; granules none, fide Bitter, but berry immature. —

The species of Miers may have been based on mixed material; the

chief discrepancy ex char, is the tubular calyx; the smaller (8 mm.

long) corolla and appendages 1 mm. long may be due to error or an

undeveloped example; these characters may have caused Sleumer to

compare the Mathews plant (not seen, at least by Dammer) with

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

D. arborescens, this always with some indument and different leaves;

D. brachystemon A. Br. & Bouche* with shorter stamen parts, the free

filaments and anthers subequal, said to be cultivated and escaped in

hedges as at Tarma, seems to have characters of doubtful taxonomic

validity; Sleumer has suggested that Mathews 849 may be D. arbores-

cens, sens. lat. F.M. Neg. 2523.

San Martin: San Roque, Soukup 3103. — Huanuco: Roadsides,

2312; 3236; 3530; Ruiz & Pawn, type; (Mathews 849, type, D. acnis-

toides). Chulqui, Kanehira 52; Mexia 04103 (det. Johnston). Hua-

cho, Stork & Horton 9402 (det. Standley, D. brachystemon). "Pacha-

pushti" (Mexia).

Dunalia spinosa (Meyen) Damm. Bot. Jahrb. 50, Beibl. Ill:

55. 1913; 494. Atropa spinosa Meyen, Reise 1: 469. 1834. Lycium

(Grabowskia) Meyenianum Nees, Obs. Bot. 390. 1843. Lycioplesium

Meyenianum (Nees) Miers in Hook. Lond. Journ. Bot. 4: 332. 1845.

Grabowskia (Lycium?) Meyenianum Nees ex Walp. Repert. Bot. 3:

113. 1844-1845.

Nodes — those extended as stout spinose branchlets — and young

tips early subfloccose-tomentose; petioles 3-6 mm. long, soon gla-

brate as the attenuate base of the oblong-lanceolate obtuse leaves,

these often 3-4 cm. long, about 1 cm. wide; peduncles puberulent,

1 cm. or longer, 2 cm. in fruit; calyx glabrous, campanulate, rounded

at base, the broad lobes soon unequal, 5 mm. long and broad; ap-

pendages (2) 3-denticulate; corolla violet, 3-4 cm. long, the limb

lightly plicate, 12 mm. long, 5 obscurely puberulent teeth 2 mm.

long, 4 mm. broad, ciliate, alternating with 5 teeth 0.5 mm. long,

scarcely 1 mm. broad; stamens adnate 2 mm. at base of corolla tube,

filaments glabrous, 1-1.5 cm. long, appendages glabrous, bifid, to

7 mm. long; anthers oblong-oval, to 4 mm. long, included (Dammer) ;

fruit orange-red, 8-10 mm. in diameter, calyx 3-4-parted, partly en-

closing the globose berry (Miers). — The Tacna specimen matches

the negative of the type (Berlin) but the appendages seem to be

much smaller. F.M. Neg. 2524.

Arequipa: Near Arequipa, Isern 2014. — Puno: Pisacoma, Meyen,

type.— Tacna: Prov. Tarata, 3,000 meters, Metcalf 30400 (det. Mor-

ton, D. Weberbaueri).

Dunalia tubulosa (Benth.) Macbr., comb. nov. lochroma tubu-

losum Benth. Bot. Reg. 31 (8) : pi. 20. 1845. Habrothamnus cyaneus

Lindley, Bot. Reg. 30 (7): Bot. Misc. 72. 1844, not D. cyanea P.

FLORA OF PERU 53

de Rouvelle ex Dunal, 1852. /. cyaneum (Lindley) M. L. Greene

ex Lawrence & Tucker, Baileya 3: 66. 1955. I. lanceolatum Miers in

Hook. Lond. Journ. Bot. 7: 347. 1848. Chaenesthes lanceolata Miers,

I.e. 4: 239. 1845. /. Weberbaueri Damm. Repert. Sp. Nov. 15: 266.

1918.

Type a 4-meter shrub with more or less densely tomentose branches,

leaves beneath, pedicels (2.5 cm. long) and calyces; petioles 2-2.5 cm.

long; leaves sparsely puberulent above, 9-11 cm. long, 4-6 cm. wide;

flowers blue, clustered; calyx cupulate, irregularly dentate, 6 mm.

long, the obtuse teeth 1-3 mm. long, 2-2.5 mm. wide; corolla puber-

ulent, glabrous within, tubular, little ampliate, plicate limb 10-den-

tate, tube 3.5-4 cm. long, 2-3.5 mm. in diameter, limb 1 cm. across,

5 lobes 3 mm. long, 4 mm. wide, the intermediate 1 mm. long; sta-

mens unequal, included (type), adnate 8 mm. above base, densely

puberulent below, less so above, 3.2, 3 and 2.8 mm. long; stigma

sub-bilobed. — After Dammer; the Peruvian plant according to him

differs from /. tubulosum Benth., 490, of Ecuador in having more

flowers and longer leaves; the latter species has inflated calyx 8 mm.

long, the teeth equal (author) ; also, corolla more deeply lobed, differ-

ences seemingly inconsequential; ex char, nearly D. cornifolia (HBK.)

Macbr. but larger in all parts. His species if distinct will require (in

Dunalid) a new name (not D. Weberbaueri Damm.) . Synonymy after

Lawrence & Tucker, I.e. Illustrated, Bot. Reg. 31 (8) : pi. 20. F.M.

Neg. 2508.

Piura: In shrubs near Rio Quiros, Prov. Ayavaca, Weberbauer

6409. Ecuador.

Dunalia umbellata (R. & P.) Macbr. Field Mus. Bot. 8: 109.

1930. Lycium umbellatum R. & P. Fl. Peruv. 2: 45, pi. 182b. 1799.

Acnistus umbellatus (R. & P.) Miers, Hook. Lond. Journ. Bot. 4: 342.

1845. A. multiflorus Damm. Bot. Jahrb. 37: 336. 1906. Acnistus

oblongifolius Rusby, Mem. Torr. Bot. Club 4: 232. 1895? Kokabus

umbellatus (R. & P.) Raf. Sylva Tellur. 54. 1838. Trozelia and Pli-

cula umbellata (R. & P.) Raf. I.e. and 55.

Becoming a tree similar to D. solanoides but often less pubescent,

the leaves glabrate above, the calyx prominently 5-toothed, the teeth

acuminate and the stamens inserted at or near the middle of the tube;

corolla 1.5 to scarcely 2 cm. long. — Sleumer, I.e. 121, suggested this

might include D. solanoides, which he thought might be referable to

lochroma. Once I distinguished this on sulcate petioles and early

5-dentate calyx but the first character is doubtful and the latter is

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

apparently variable, the extreme in this type. Allied but distinct is

D.ferruginea Sod. & Damm. Bot. Jahrb. 36: 384. 1905 (A. ferrugineus

(Sod. & Damm.) Swan, Misc. Chi. Acad. Sci. 88: 7. 1951), leaves

stellate-pilose above, reddish tomentose beneath, corolla 1 cm. long,

filaments nearly free, anthers 1 mm. long. Miers described Mathews'

specimen: petioles about 2 cm. long, leaves nearly 1 dm. long, 4-

4.5 cm. wide, pedicels 3.5 cm. long, corolla tube 14 mm. long, lobes

8 mm. long (perhaps Dunal's species) ; Weberbauer 2731 has petioles

6-10 mm. long, leaves 6-17 cm. long, 2.5-7 cm. wide, pedicels 2 cm.

long, calyx 4 mm. long and broad, the uneven teeth scarcely 1 mm.

long, corolla lobes 5 mm. long; filaments adnate 7 mm., free 2.5 mm.,

tube 15 mm. long, berry semi-included in accrescent calyx. My col-

lection from a tree 7 meters tall, fruits lead color; Stork & Horton's

from a 15-meter tree, yellow-green corollas with fruit early white-

veined. F.M. Negs. 2499; 2493 (A. multiflorus) .

Cajamarca: Llama, Soukup 4222. Chota, Stork & Horton 10048.

— Amazonas: Prov. of Chachapoyas, (Mathews 3244, det. Miers). —

Ancash: Below Ocros, Prov. Cajatambo, Weberbauer 2731 (type, A.

multiflorus). Near Chiquian, Ferreyra 7386. Macara, Nunez 3239;

3242 (det. Cowan, A. oblongifolius} . — Lima: Obrajillo, Ruiz & Pavdn,

type. Near Matucana, Weberbauer 78. — Huanuco: Cani near Mito,

3452? (fruit only). Bolivia?

lochroma Warscewiczii Regel, Gartenfl. 4: 245, pi. 130. 1855.

Shrub, to 5 meters tall or taller, the terete upper branches, peti-

oles, peduncles (each to about 4 cm. long) and umbellate-fasciculate

pendent pedicels — these at least 5 cm. long, usually longer — more or

less glandular pilose; leaves broadly ovate, attenuate into the petiole,

shortly acuminate, to 14 cm. long, about 1 dm. wide, paler and to-

mentulose beneath, the nerves and veins prominent; calyx finally

subglobose-inflated, 5-costate, 12-18 mm. long and broad, the 5

shortly ovate acute teeth somewhat unequal; corolla lightly curved,

slightly hirtellous, dark violet-blue, the tube about 5 cm. long, 3 mm.

thick, the infundibuliform-campanulate limb 2.5 cm. across or more,

with 5 equal broad apiculate lobes; stamens included, filaments gla-

brous except for a few minute trichomes on the upper free portion;

stigma bilobulate. — The similar Ecuadorian species /. grandiflorum

Benth. has glabrous shorter flowers, clustered apically on angled

branchlets, /. macrocalyx Benth. has hirtellous but subsessile flowers,

longer calyx, and I. tubulosum Benth. has shorter corollas, limb only

1 cm. across, the white-pubescent margin nearly dentate. Revision

FLORA OF PERU 55

of the group, obviously — and naturally, after a hundred years — is

much in order. Type of Warscewicz not seen but the following seems

to be his plant; he collected from the Maranon Valley to Bolivia.

Cajamarca: La Tajona, Prov. Hualgayoc, Weberbauer 1002 (det.

Standley, 7. grandiflorum) . Near San Miguel, Weber bauer 3910; 258.

Rio Maranon, (Warscewicz, type).

Dunalia Weberbaueri Damm. Bot. Jahrb. 50, Beibl. Ill: 53.

1913.

Very near D. obovata (R. & P.) Damm. and D. lycioides Miers

but the filaments hairy at base and their appendages much longer

(6 mm. long); spines (type) 2-4 cm. long; petioles 2-7 mm. long;

leaves 2-4 cm. long, 7-12 mm. wide, obtuse or acute, cuneate at base;

pedicels solitary, 8-18 mm. long; calyx 4x4 mm., acute teeth 1 mm.

long; corolla violet, 2 cm. long, slightly ampliate toward the plicate

limb, the marginally pilosulous teeth acute; filaments 17 mm. long,

adnate one-third, puberulent near base, the appendages one-third

free, the anthers 2.5 mm. long; style 22 mm. long, stigma capitate.

—Perhaps a variant of the shrub of Ruiz and Pavon but appendages

long; granules not solidified, no count (Bitter). F.M. Neg. 2526.

Lima: Tambo de Viso, Weberbauer 117, type.

10. POECILOCHROMA Miers

Saracha Ruiz & Pavon; compare note under that genus.

Shrubs rarely spinescent with coriaceous or sometimes membra-

nous (probably shade-forms) leaves and rather large solitary or fas-

ciculate broadly but deeply campanulate yellowish (rarely white)

more or less purple-spotted flowers; calyx campanulate, very shortly

lobed, sometimes unequally 3-lobed, little if at all accrescent. —

Genus of convenience, the corolla in shape exactly conforming to

the type of Saracha dentata Ruiz & Pavon. An interesting genus to

revise, possibly with many species but probably with fewer than

proposed by Miers, as the apparent differences seem to have little or

no concomitance and therefore they may not indicate genetic values.

The following key is expedient; using the same characters the spe-

cies can be sorted also into other units or varieties on the basis of

pubescence (quality, position, presence of trichomes on filaments

and corolla within), size and shape of corollas, length of stamens.

It is probable that heterostyly exists; the stamens may develop at

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

different times as in Saracha. The genus was referred to Capsicum

by Kuntze; Dunal wrote P. punctatum (R. & P.) Miers. One or more

species attributed to Peru by Miers were collected in Ecuador.

The bizarre beauty of the flowers, sometimes recalling by their

color some Fritillarias, is noteworthy (Weddell).

KEY TO POECILOCHROMA

Corolla glabrous to puberulent or sparsely so within; leaves at most

about twice longer than wide, usually oval or elliptic-oblong.

Plants espinose P. Lobbiana.

Plants spinose P. spinosa.

Corolla (except P. guttata} densely to loosely lanate-floccose within;

leaves mostly or all about 3 times longer than wide.

Corolla subglabrous within; branchlets and leaves glabrous; ovary

tomentose P. guttata.

Corolla closely lanate to lightly floccose within; branchlets and

leaves glabrous (or nearly) as ovary. .P.frondosa, P. punctata.

Poecilochroma frondosa Miers in Hook. Lond. Journ. Bot. 7:

355. 1848; 495. Saracha frondosa Miers, 111. S. Amer. PI. 2: App. 57.

1857.

Branchlets subcompressed, angulate-alate, striate, densely foli-

ose; petioles to 2 cm. long; leaves elliptic, attenuate both ends, to

1 dm. long, about a third as wide, submembranous, glabrous above,

sparsely fulvous tomentose beneath, nerves beneath rubescent, mar-

gin revolute; peduncles fasciculate, glabrous, apically incrassate,

nutant, nearly 2 cm. long; calyx short, red-colored, membranous-

margined, unequally and rounded-lobulate; corolla campanulate,

fulvous puberulent, 5 cm. long, 2 cm. across, limb sinuately 5-angu-

late, ciliate, pubescent within below as tube; filaments lightly pubes-

cent with spreading long-articulate trichomes, unequal, half as long

as corolla, slightly dilated at base; ovary and style glabrous. — Inter-

mediate to P. punctata and P. guttata (author) and points up the

probable unreliability of the characters emphasized, these doubtfully

established genetically.

Amazonas: Chachapoyas, (Mathews, type).

Poecilochroma guttata Miers in Hook. Lond. Journ. Bot. 7:

356. 1848; 495. Saracha guttata Miers, 111. S. Amer. PI. 2: App. 57.

1857.

FLORA OF PERU 57

Similar to P. frondosa; petioles to 3 mm. long; leaves solitary,

rarely geminate, obovate, acute or shortly acuminate, 7-10 cm. long,

about 2 cm. wide, quite glabrous, fleshy coriaceous, veins not appar-

ent, glossy green above, pale beneath, revolute-margined; peduncles

to 2.5 cm. long; calyx lobes rounded, unequal, glabrous except tomen-

tose mucro; corolla 2.5 cm. long, throat 2 cm. across, pulverulent

(type), glabrous within as style and stamens, the latter equaling

three-quarters of the corolla; ovary described as tomentose. — Corolla

white (Kanehira), glabrous as filaments and ovary. F.M. Neg. 8564.

Amazonas: Chachapoyas, (MathewsllSl, type). — Huanuco: Pam-

payacu, Kanehira 187?

Poecilochroma Lobbiana Miers in Hook. Lond. Journ. Bot. 7:

367. 1848; 496. Saracha Lobbiana Miers, 111. S. Amer. PI. 2: App. 57.

1857.

Younger branches floccose (indument copper-colored), the older

blackish, the branchlets divaricate; petioles 6 mm. long; leaves ellip-

tic, acute both ends or obtuse, to about 4 cm. long, half as wide, revo-

lute, lustrous above, pallid beneath, quite glabrous, the primary

pinnate nerves impressed above, reddish and prominent beneath;

peduncles 1-3, incrassate apically, somewhat shorter than leaves;

calyx sometimes 5-angled, 4 mm. long, 3 mm. across, the short

rounded teeth medially mucronulate; corolla finally campanulate,

2.5-3 cm. long, drying orange, pubescent only without or lightly

puberulent toward base, limb sinuately 5-angled, 3 cm. across; sta-

mens and slightly longer style glabrous, included, the stigma clavately

bilobed. — After Dunal as all descriptions here, who (or the composi-

tor) wrote "P. Llobbiana Miers," the author, P. Lobbiana; in litera-

ture regarding the collector (as in Muhlenbergia 7: 100) he is referred

to as Lobb. P, Lindeniana Miers, Ecuadorian, has 3 unequal calyx

lobes; P. maculata Miers, I.e., type by Lobb, without data, differs in

leaves to about 4 cm. long, half as wide, fulvous tomentose beneath,

corolla yellow, spotted, puberulent in and out (F.M. Neg. 33139).

Determinations by Morton. Type (not seen) Lobb 389, "Andes of

Peru."

Ayacucho: Near Huanta, Killip & Smith 22186. — Cuzco: Panti-

calla Pass, Cook & Gilbert 1236; 1859.

Poecilochroma punctata (R. & P.) Miers, in Hook. Lond.

Journ. Bot. 7: 354. 1848; 495. Saracha punctata R. & P. Fl. Peruv.

2: 42, pi. 187a. 1799. Bellinia punctata (R. & P.) Roem. & Schult.

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

Syst. 4: 688. 1819. Atropa punctata (R. & P.) Spreng. Syst. 1: 698.

1825. P. venosa Rusby, Bull. N. Y. Bot. Gard. 4: 424. 1906?

Branchlets early scurfy puberulent as leaves usually slightly be-

neath (Ruiz and Pa von), glabrous in age; petioles several to 8 mm.

long; leaves rarely geminate, ovate-oblong, 3-4 cm. long or longer,

about a third as wide, often larger, veins many, primary nerves 7 or

8 pairs, rather prominent beneath, peduncles fasciculate, few to sev-

eral, more or less nutant, incrassate apically, 2 cm. long or longer;

calyx glabrous, the 5 rounded lobes only 1-3 mm. long; corolla 2 cm.

long or longer, and at least as wide, broadly campanulate, the limb

sinuate with more or less developed somewhat reflexed lobes, gla-

brous or puberulent (rarely lanate), purplish yellow and purple

blotched or spotted within; anthers oblong; filaments and ovary gla-

brous.—F.M. Negs. 2513 (named for Weberbauer, ined.); 23089.

Sometimes becomes a small tree, 7 meters tall, the extremely

bitter leaves used to alleviate pain (Ruiz and Pavon).

Cajamarca: Socota to Tambillo, Stork & Horton 10179 (det.

Standley). — Lima: Obrajillo, (Ruiz & Pavdn). — Huanuco: Northeast

of Huanuco, 2201. Tambo de Vaca, 4412. Muna, Panao, Tambo

Nuevo, Portachuelo, Ruiz & Pavdn, type. Carpish, Swingle 128;

3128; Ferreyra 1726; 8159 (det. Cowan); Stork & Horton 9934 —

Huancavelica: Huari, Nunez 2117; 10335 (det. Standley). — Cuzco:

Santa Ana, Weberbauer 4939 (as ined. sp.). Paso de Tres Cruces,

Pennell 13860; Weberbauer 6974 (filaments glabrous, corolla lanate).

Paucartambo, Vargas 2108; Balls 6774; Soukup 384; 394; 395. Lucu-

mayo Valley, Cook & Gilbert 1285. Cedrobamba, 3,900 meters, Ed-

mund Heller 2191. — Puno: Limbani, Metcalf 30466 (det. Morton).

Puno, Lechler 2080 (P. venosa, fide Rusby) . Bolivia?

Poecilochroma spinosa Damm. Bot. Jahrb. 37: 637. 1906.

Branchlets verruculose, spiny; petioles 1-2 mm. long; leaves in

type obovate, 1-1.5 cm. long, 6-8 mm. wide, coriaceous, glabrous;

pedicels solitary or geminate, 10-12 mm. long; calyx 5 mm. long,

unevenly dentate; corolla yellowish, violet-nerved, puberulent with-

out, pulverulent within at base, 2-2.5 cm. long, teeth scarcely 2 mm.

long; anthers oval; filaments 19 mm. long, glabrous as ovary and

style, this 23 mm. long, stigma 1.5 mm. long. — Weberbauer 6730,

locality unknown, has membranous sparsely hispidulous leaves, 3-

4 cm. long; type described as erect or scandent, 2 meters tall, in river

shrubs. Seems nearly P. Lindeniana Miers, 496, of Ecuador or even

FLORA OF PERU 59

more similar to P. Lobbiana Miers and may be a spinescent state.

F.M. Neg. 2512.

La Libertad: Prov. de Bolivar, Ferreyra 1254- Toward Angas-

marca, West 8162. — Ancash: Prov. Huari, 3,600 meters, Weberbauer

2907, type; 7014- — Ayacucho: Above Quinua, Prov. Huamanga,

Weberbauer 5542.

11. SALPICHROA Miers

Busbeckia Mart. Cat. Hort. Monac. 69. 1829.

Shrubs or small lianas with ovate or ovate-cordate or rarely lan-

ceolate, opposite, geminate, seldom ternate leaves, and (in Peru at

least) usually elongate-tubular or narrowly funnelform solitary (or

loosely corymbed?) greenish- or brownish-yellow or yellow flowers.

Calyx deeply 5-parted, the lobes narrow, little or not accrescent.

Stamens (typically) inserted about the middle of the corolla-tube,

filaments straight, glabrous, anthers narrow; style filiform, erect,

stigma subcapitate; ovary free of fleshy disk, 2-celled as berry; ber-

ries free, typically scarlet or red, in some species greenish-white; seeds

of original species compressed, rugulose, hispidulous. — When a re-

vision is made the shape of corolla lobes, length of filaments and other

characters may be found significant; it is not clear that degree of

pubescence and vegetative differences that have been used as spe-

cific indicators are reliable criteria; indeed, it is my impression that

only about half the names I have had to try to contrast in the key

will be found to represent genetic entities. See Juanulloa microphylla

Dunal, with habit and foliage aspect of this group, also S. breviflora,

S. cuspidata, doubtfully this genus. Miers, in Hook. Lond. Journ.

Bot. 7: 353. 1848, altered the name to Salpichroma.

The fruits of a Paraguayan species are white and sold in the mar-

kets as "cocks' eggs."

KEY TO SALPICHROA

Calyx 2.5-3 cm. long; corolla 7-13 cm. long.

Corolla longer than 1 dm.; calyx 5-parted, segments linear.

S. Didieranum.

Corolla 7-8 cm. long; calyx segments and tube subequal.

S. dependens.

Calyx about 0.5-2 cm. long; corolla 1-5 cm. long.

Calyx lobes subulate-linear, about 1 cm. long or longer; corolla

finally 2.2-5 cm. long.

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

Indument eglandular.

Calyx more or less hirsutulous as usually foliage (somewhat) ;

anthers visible; corolla funnelform S. hirsuta.

Calyx as leaves puberulent; anthers not visible; corolla tu-

bular S. Weberbaueri, S. ramosissima.

Indument glandular at least in part; anthers often visible;

corolla subtubular S. glandulosa.

Calyx lobes 7 mm. long or shorter; corolla at most 2 cm. long.

Corolla 12-15 (20) mm. long; calyx parted nearly to base.

Corolla dilated or evenly ampliate; pubescence obvious.

Indument (ex char.) not glandular.

Calyx lobes more or less hirsutulous, 5-6 mm. long;

lobes oblong-oval.

Plants fragile, lightly pubescent S. dilatata.

Shrub, more or less grayish pubescent S. diffusa.

Calyx, as lanceolate corolla lobes, puberulent, both at

most 4 mm. long S. Gayi.

Indument granular-glandular S. tristis.

Corolla constricted at throat; pubescence essentially lacking.

S. uncu.

Corolla 8-9 mm. long, glabrous as obvious calyx tube.

S. micrantha.

Salpichroa dependens (Hook.) Miers in Hook. Lond. Journ.

Bot. 4: 325. 1845; 472. Atropa dependens Hook. I.e. pi 107. 1837.

Juanulloa microphylla Dunal in DC. Prodr. 13, pt. 1: 531. 1852.

Shrubby stems pendent; leaves geminate, cordate- ovate, shortly

petioled, nearly glabrous but more or less viscid, pale beneath;

calyx tubular, finely pubescent, obscurely viscid, 5-parted, finally

somewhat enlarged and splitting in fruit; corolla long-tubular, sta-

mens and style well-exserted. — This is the original description, not

modified by Miers; corolla truly funnelform and more than 2.2 cm.

long, according to Jaubert; plate shows leaves rounded or weakly

cordate, petioles 7-9 mm. long, blades 2.5 X 1.5 cm., pedicels 8-12

mm. long, corolla tube to lobes (throat little ampliate) 6 cm. long,

1 cm. wide above, lobes ovate-lanceolate, erect or suberect, 12 mm.

long, anthers exserted, exceeded by style, this 8 cm. long; calyx

2.5 cm. long to 3.2 cm. long in fruit which is as long, 1.5 cm. thick,

cleft 1 side in flower, widely so in fruit. — A slender very branching

FLORA OF PERU 61

plant, generally growing from the root of some tall shrub among

whose branches it rambles for support, extending from 3 to 5 meters,

its ultimate pendent extremities covered with large sulphur-colored

flowers (collector). It was collected on the eastern side of the

Cordillera, above potato cultivation. Type, Mathews 829, without

data.

Lima: Pariahuanca, 3,400 meters, Weberbauer 6596 (distr. as S.

glandulosa). — Huanuco: Tambo de Vaca, 4419. — Huancavelica: Sal-

cabamba, Stork & Horton 10382 (det. Standley, S. glandulosa). —

Ayacucho: Near Huanta, 3,200 meters, Weberbauer 7515.

Salpichroa Didieranum Jaubert, Bull. Soc. Bot. Fr. 8: 117.

1861.

A low (2 meters) liana (stem 2 cm. thick near base) with alternate

(uppermost subopposite) lanceolate subscabrous leaves and yellow

long-funnelform nutant flowers, 12-13 cm. long; leaves subrounded

at base, acute, nervose; pedicels short, pubescent; calyx segments

linear, ciliate, 2.5 cm. long; corolla lobes acute, marginally floccose;

anthers hastate; style enlarged to the claviform stigma; capsule

conical, subequaling calyx; seeds reniform. — Related to S. dependens

(Hook.) Miers and S. glandulosa (Hook.) Miers.

Commemorates MM. Ernest and Alfred Grandidier, who, during

1857 and later, toured the American continents, studying natural

science.

Cuzco: Ravine near Mollepata pass, road from Lima to Cuzco,

Prov. of Anta, 4,400 meters, (Grandidier freres) .

Salpichroa diffusa Miers in Hook. Lond. Journ. Bot. 7: 335.

1848; 473. S. foetida Damm. Bot. Jahrb. 37: 639. 1906, as to Peru,

probably.

Flexuose and often divaricately much-branched shrub, the

branches alate; leaves geminate, ovate, obtuse at base, subacute,

both sides hirsutulous, floccose-margined, about 18 mm. long, 10-14

mm. wide; petioles 8 mm. long; calyx as peduncle 6 mm. long, lobes

sublinear, hirsute; corolla yellowish-pubescent, funnelform, tube 12

mm. long, limb 4 mm. long, the oblong lobes reflexed, ciliate; stamens

and style included. — According to Raimondi the very aromatic fruits

are used in helados. S. foetida Damm. of southern Bolivia has

corolla only 1 cm. long; S. Mandoniana Wedd. Chloris And. 2: 98.

1857, from Socota, Bolivia, ex char, has softly pubescent branches,

finely pubescent leaves, exceeded by peduncle, this longer than the

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

2 cm. long flower (corolla 16-18 mm. long, glabrous), calyx less than

half as long. Some or all of the Peruvian specimens of S. diffusa

Miers (type from Bogota by Goudot but the Maclean specimen

(from Peru?) cited by author as the same), may rather be referable

to S. Mandoniana. Illustrated, Miers, 111. S. Am. PI. pi. 28c.

Lima: Near Matucana, Weberbauer 186; 169. — Ayacucho: Huan-

ta, Raimondi. — Arequipa: Above Chuquibamba, Weberbauer 6848.

Cachendo, (Guenther & Buchtien 89, det. Bruns, S. foetida). Mol-

lendo, Weberbauer 1511; 147. Nevada de Chachani, Pennell 13266

(det. Johnston). — Puno: Azangaro, Weberbauer 1+67; 185. Without

locality, (Maclean). Ecuador; Colombia. "Pepinillo."

Salpichroa dilatata Damm. Bot. Jahrb. 37: 639. 1906.

A diffuse subglabrate fragile shrub with angulate branchlets;

petioles 7-9 mm. long; leaves truncate at base, sparsely pilose

both sides, mostly 1.5-2 cm. long, 10-13 mm. wide; pedicels 2-4 mm.

long; linear calyx-lobes slightly hirsute-pilose (to glabrous), 5 mm.

long; greenish corolla elongate-urceolate, the tube 1.5 cm. long, 4

mm. wide, the oval lobes 3 mm. long; filaments dilated, 2 mm.

long, 1 mm. wide, anthers 3 mm. long; style 1 cm. long, stigma

capitate; berry (immature) ovoid, acute, 12 mm. long, 5 mm. thick.—

F.M. Neg. 2940.

La Libertad: Prov. Otusco, Lopez 1025. — Ancash: Near Ocros,

Weberbauer 2679, type. — Ayacucho: Near Ayacucho, Stork & Norton

10804. (det. Standley) ; Soukup 4022. "Cuytulumo" (Lopez).

Salpichroa Gayi Benoist, Bull. Soc. Bot. France 85: 54. 1938.

Scandent, the younger branches pubescent, the older with one

side alate; petioles puberulent, 7-9 mm. long; leaves geminate,

nearly equal, oblong or oblong-triangular, obtuse or subcordate at

base, 14-16 mm. long, 10-12 mm. wide, sparsely and shortly pilose

above, pubescent below, 4-5 secondary nerves each side, the 2

lower originating from base; flowering pedicels 3-4 mm. long; sepals

free nearly to base, linear, acute, 4 mm. long; corolla minutely

puberulent, 12 mm. long, lobes long-lanceolate, 4 mm. long, tube

little inflated in upper half; stamens inserted in upper fourth of tube,

filaments one-quarter as long as anthers, these 2 mm. long, attaining

throat; ovary and style glabrous. — Type, Gay 2308 (not seen),

without data in Herb. Paris.

Cuzco: Prov. Acomayo, near Mayu Huillca, 2,900 meters,

Vargas 255?

FLORA OF PERU 63

Salpichroa glandulosa (Hook.) Miers in Hook. Lond. Journ.

Bot. 4: 325. 1845; 472. Atropa glandulosa Hook. Bot. Misc. 2: 230.

1831.

A low spreading much-branched glandular-pubescent shrub at

most 1 meter tall with geminate strongly cordate to subcordate

(type) leaves and greenish-yellow tubular flowers 3.5-5 cm. long

(not including lobes); leaves in dry seasons about 6 mm. long and

wide but mostly 1-2 (3.5) cm. long, 8-10 (20) mm. wide; petioles

narrow, 6-8 (15) mm. long; pedicels filiform, 6-8 mm. long (type),

apically nutant; calyx lobes sublinear, glandular-hirsute, 8-12 (15)

mm. long; corolla throat 1 (1.5) cm. across, lobes ovate, reflexing,

8 mm. long; stamens finally slightly exserted, exceeded by slender

style; berry greenish- white, glaucous, nearly 3 cm. long and 1.5

cm. thick. — The larger figures are derived from the plate illustrating

Mathews' specimens. The var. longiflora Dunal (type, Pavon, with-

out data, not Mexico as given) has petioles 8-10 mm. long, leaves

2.5-3 cm. long, 8-9 mm. wide, peduncle as calyx 12 mm. long,

corolla 4.5 cm. long, limb 1.5 cm. across, the ovate lobes floccose-

margined. Perhaps a more or less glandular state that should be

drawn to include the similar forms that are obscurely or not at all

glandular. Illustrated, Hooker, Icon. pi. 106.

La Libertad: Huamachuco, West 8127 (det. Johnston, S. Weber-

baueri). — Lima: Banos, Wilkes Exped. Near Casapalca, Ferreyra

6540 (det. Cowan). Rio Blanco, sprawling under great rocks, 3050.

Obrajillo, valley of Canta, (Cruckshanks, type). — Junin: Comas,

Weberbauer 6603. Near Huancayo, Stork 10932; Killip & Smith

22073; Soukup 3688. Vicinity of Cerro de Pasco, (Mathews) —

Cuzco: Paucartambo, Herrera.

Salpichroa hirsuta (Meyen) Miers in Hook. Lond. Journ. Bot. 4:

325. 1845 and 7: 334. 1848; 472. Atropa hirsuta Meyen, Reise 1:

466. 1834.

A subhirsute diffuse but probably much-branched shrub with

geminate (or fasciculate on shoots) ovate leaves obtuse at base,

unequally acute at tip, to 5 cm. long, nearly 2 cm. wide, petioles

subfiliform, to 3 cm. long; calyx 8 mm. long, the hirsute lobes linear-

subulate; corolla finely pilose, funnelform above, the slender tube

3 cm. long, the ovate lobes 3 mm. long, reflexed; stamen tips and

style exserted; fruit about 1.5 cm. long, 8 mm. thick. — Mostly after

Nees v. Esenbeck; specimen (a single branchlet) named by Meyen,

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

described by Miers, had leaves 18 mm. long, 12 mm. wide, peduncles

16 mm. long.

Apurimac: Andahuaylas, West 8719 (det. Johnston, S. glandulosa) .

— Cuzco: Marcapata, Vargas 3752. — Puno: Near Puno, Vargas 1274,'

Soukup 88. Granja Salcedo, Mexia 7779 (det. Johnston). Near

Pisacoma, about 5,000 meters, Meyen, type. Bolivia; to Colombia?

Salpichroa micrantha Benoist, Bull. Soc. Bot. France 85: 55.

1938.

Ex char, similar to S. Gayi but branches soon glabrous; petioles

11-12 mm. long; leaves ovate, triangular, obtuse or often lightly

cordate at base, slightly inequilateral, obtuse, 15-17 mm. long,

11-14 mm. wide, glabrous both sides; flower pedicels 2-2.5 mm.

long, glabrous (or nearly) as calyx, this 4 mm. long, sepals connate

for lower third, the free part triangular lanceolate, acute; corolla

glabrous, 8-9 mm. long, tube little narrowed below the middle,

lobes 1.5 mm. long; anthers subsessile, 2 mm. long, in apex of throat.

— Type (Paris) Gay 1973, without locality, not seen.

Cuzco: Yucay, Soukup 750.

Salpichroa ramosissima [Mathews] Miers in Hook. Lond.

Journ. Bot. 4: 326. 1845.

A shrub with quadrate nodose branches (the branchlets pu-

bescent) and geminate or ternate nearly glabrous elliptic-oblong

subacuminate leaves, rounded at base but decurrent into slender

petiole 12 mm. long; leaves submembranous, ciliate, in type to about

4 cm. long, 1.5-2 cm. wide; calyx lobes sublinear, slightly pilose;

corolla tubular, contracted at throat, the subacute reflexed lobes

marginally velutinous; berry red, oval, 2 cm. long, 12 mm. thick,

the seeds hirsute. — Length of calyx and corolla in type not given

but latter probably not more than 2-4 cm. long; name in herbaria

by Mathews; very distinct, fide Miers, in form of leaves and corolla

contracted in the throat (where usually widened). It seems doubtful

that the contracted corolla throat and the narrow leaves are sig-

nificant in comparison with S. Weberbaueri.

Lima: Puruchucu, (Mathews 1053, type).

Salpichroa tristis Miers in Hook. Lond. Journ. Bot. 7: 335. 1848.

Type a small depressed shrub with short (to 2.5 cm. long) slender

tortuous nodose branchlets, geminate glandular-pubescent obovate

leaves less than 6 mm. long, at least half as wide, and glabrous

FLORA OF PERU 65

nutant flowers about 1 cm. long; petioles 3 mm. long, compressed;

calyx equally 5-parted, the lobes narrowly subulate, 4 mm. long;

corolla tube 1 cm. long, 3 mm. thick, the short triangular lobes

reflexed; stamens and style included from middle of tube; style

slightly deflexed, incrassate apically, stigma concave; ovary disk

fleshy. — Leaves when dry are black; branches gray, dotted. Type

(Herb. Hooker) from Quito; cited next, Maclean, Peru Andes. In

Peru at least leaves in wet season are larger. S. alata Damm.,

Meded. Rijks. Herb. 29: 26. 1916, of Bolivia might be sought here

because of small flowers but the indument is glandular-pilose and

the branchlets 1-3-alate.

Ancash: In rocks near Huaraz, Weberbauer 2970; 225. — Apurimac:

Saxaihuaman, Ferreyra 2630; 2652 (det. Cowan, with query). —

Cuzco: Calca, Vargas 157. — Puno: Near Puno, Soukup 44-5- Without

locality, (Maclean). Bolivia; Ecuador.

Salpichroa uncu [Dombey] Benoist, Bull. Soc. Bot. France 85:

55. 1938.

Sarmentose, type quite glabrous, the older branches not alate;

petioles 10-14 mm. long; leaves ovate-lanceolate, obtuse or subcor-

date at base, often unequal, subacute, 22-32 mm. long, 14-20 mm.

wide, the lateral nerves often near base, the 2 lower from base; ped-

icels 4 mm. long, calyx 7 mm. long, free nearly to base, sublinear,

attenuate to acute tip; corolla 16 mm. long, upper half distinctly

ampliate, at throat moderately and gradually constricted, lobes lan-

ceolate-triangular from broad base, obtuse, 2 mm. long, style inserted

in upper fifth of tube; filaments nearly as long as anthers, these 2 mm.

long, attaining apex of throat, oblong, clearly attenuate; fruit bac-

cate.— Benoist, I.e. 53-55, described two species from Ecuador, one

from Colombia and one, S. sarmentosa, I.e. 55, from La Paz, Bolivia,

Mandon 436, type. This is the most likely to occur in Peru; petioles

6-8 mm. long, leaves 1.5-2.5 cm. long, 12-24 mm. wide; pedicels 2-4,

calyx 8, lobes 3, anthers 2 mm. long; branchlets alate, pubescent as

pedicels, the leaves both sides and the sepals, these sublinear; corolla

puberulent, distinctly ampliate; anthers oblong, affixed at apex of

throat.

Ancash: Tallenga, Ferreyra 7485 (det. Cowan, S. diffusd). —

Lima: Acotama, Dombey, type. Matucana, 2945. Rio Blanco, 3041 •

— Ayacucho: Above Puquio, Ferreyra 5512; 7209 (det. Cowan, S.

diffusa). Near Chavina, Weberbauer 5785 (det. S. diffusa, Herb.

Dahlem). Ecuador.

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

Salpichroa Weberbaueri Damm. Bot. Jahrb. 37: 640. 1906.

S. diffusa Miers var. longiflora Hicken, Ap. Hist. Nat. Buenos Aires 1 :

175. 1913.

Shrub with scandent minutely puberulent branches and ovate-

cordate rather densely puberulent leaves, 1.5-3 cm. long, nearly

as wide; pedicels 8 mm. long, puberulent; calyx lobes slightly pilose,

1 cm. long; corolla yellow, tubular-funnelform, 3.5 cm. long in-

cluding the 3 mm. long ovate-acuminate lobes; filaments 1 mm.

long, anthers 3 mm. long, style 3 cm. long.

Ancash: Near Ocros, Weberbauer 2693. Huari, Weberbauer 2936.—

Huancavelica: Pana, Tovar 207 (det. Cowan, S. diffusa). — Cuzco:

In fences toward Acomayo, Vargas 4753 (det. Standley). Sicuani,

Cook & Gilbert 92; (Hicken, type, var.). — Arequipa: Chachani, Pen-

nell 3266. "Quitung-quitung," "nuno-nuno" (both, Cook & Gilbert) .

12. JABOROSA Juss.

Dorystigma Miers in Hook. Lond. Journ. Bot. 4: 347. 1845.

Lonchestigma Dunal in DC. Prodr. 13, pt. 1: 476. 1852.

Prostrate and creeping or rosette herbs with subentire or irregu-

larly pinnatifid leaves and pediceled axillary funnelform or long tubu-

lar-funnelform yellow, blue or white flowers. Calyx campanulate,

5-lobed, little or not at all enlarged in fruit. Stamens attached above

the middle of the tube, sessile or filaments free; stigma narrowly 2-

5-lobed (type), clavate or broad and acute. — The segregate genus

apparently was meant as a subgenus on the basis of the stigma

character.

Jaborosa crispa (Miers) Hook, ex Wetts. Natur. Pflanzenf. 4,

pt. 3b: 26. 1895. Dorystigma crispa Miers in Hook. Lond. Journ.

Bot. 7: 333. 1848. Lonchestigma crispum (Miers) Dunal in DC.

Prodr. 13, pt. 1:477.1852.

A glabrous or sparsely crisply pubescent cespitose plant with

short prostrate or ascending stems; leaves subfasciculate, pinnately

laciniate, the divisions shortly mucronate and sinuately dentate,

about 6 cm. long and 2 cm. wide, decurrent into a petiole 2 cm. long

or longer; peduncles 1 cm. long, bracts small, subulate; calyx as

corolla white, 12-15 mm. long, glabrous without except for a few

crisp trichomes, the throat woolly within; style shorter than the

subexserted stamens; berry to 3 cm. in diameter. — J. caulescens

Hook., 476, Bot. Misc. 1: 347, pi. 71. 1830, of Chile has more deeply

FLORA OF PERU 67

toothed leaf -lobes, much longer pedicels; however, there seem to be

several closely allied species or variants as J. leiocalyx Damm., just

south of Lake Titicaca, with bright blue corolla, cordate, bilobed

stigma. Illustrated, Wedd. Chloris And. 2: 101, pi 57. 1857.

Arequipa: Vincocaya, Pennell 13346. — Puno: Pomata, Shepard 33

(det. B. L. Robinson). Arid punas, 3,900-4,000 meters, (Weddell).

— Tacna: Volcan Tacora mountains, Werdermann 1158. Bolivia.

13. WITHERINGIA I/Her.

Ligneous, the leaves alternate or geminate, entire, the 4- or 5-

merous flowers geminate or umbellate, the 1-flowered peduncles

lateral, extra-axillary. Calyx dentate or parted. Corolla rotate-

campanulate, parted, subvillous at filament attachment. Stamens

exserted, anther dehiscence longitudinal. Style filiform, stigma capi-

tate.— Longitudinally dehiscent anthers, lobed calyx not accrescent,

and rotate corolla have been accepted as the diagnostic characters

of the group; cf. Capsicum.

Witheringia microphylla Griseb. in Lechler, Bert. Ann. Austr.

58. 1857.

Glabrous except axillary fasciculate blooms and early an obscure

puberulence on branchlet tips; leaves approximate, subsessile, oblong-

lanceolate, to 2 cm. long, 7 mm. wide, often smaller, lustrous; ped-

icels 8-10 mm. long, subangulate, with a cupulate collar at base;

calyx 1.5 mm. long, 3 mm. across, the lobes rounded, not at all ac-

crescent but irregularly cleft in fruit; corolla 7 mm. long, at least as

broad across the top, the broadly rounded lobes about 4 mm. long,

3 mm. wide; berries black, to 1 cm. in diameter. — A shrub about 1.5

meters tall, with fleshy coriaceous subimbricate leaves, the purplish

corollas fulvous tomentulose. Determinations by Standley. F.M.

Neg. 2542.

Apurimac: In rocks and humus, Marcapata, 4,100 meters, Var-

gas 1361; 9716. — Cuzco: Marcachea, Vargas 11178. — Puno: Macu-

sani, Lechler 2645, type.

14. CAPSICUM [Tourn.] L.

Bassovia Aubl. PI. Guiana 1: 227, pi. 85. 1775, in part; Witheringia

in litt. vix L'Her. Sert. Ang. 1 : 33, pi. 1 . 1788? 402. Brachistus Miers,

Ann. Mag. Nat. Hist. ser. 3, 2: 264. 1849; 682. Fregirardia Dunal,

68 FIELD MUSEUM OP NATURAL HISTORY — BOTANY, VOL. XIII

I.e. 502; 688. Aureliana Sendt. in Mart. Fl. Bras. 6: 138. 1846,

fide Bitter.

Reference: Dunal in DC. Prodr. 13, pt. 1: 402-429, 1852, incl.

synonyms, these at least as to Peru.

Simulates many species of Solatium vegetatively but the anthers

dehiscent longitudinally; leaves often geminate, one smaller; calyx

truncate and entire or with 5 (rarely 10) more or less developed mar-

ginal teeth, little or not accrescent. Flowers always solitary on axil-

lary or subaxillary pedicels, these one to many.

Heiser and Smith, Brittonia 10: 194. 1958, quote Morton who

said (in Standley's "Flora of Costa Rica," 1938) that the genus

should include those species with free glabrous filaments and a shiny

pungent berry, while Standley suggested omitting those with soft

pulp-filled, non-pungent berries; this was concurred in by Heiser

and Smith. The generic names cited above as synonyms may not

apply except as regards Peruvian plants referred to them; and these,

for convenience, are included here, yet it seems probable that one

generic name will be found to express their basic unity. Apparently

a good example of the futility of maintaining Brachistus distinct

(from Capsicum} is found in Witasek, Bot. Exped. Sudbrazil Wien.

9. 1910; her illustrations of the species Bassovia Wettsteiniana, C.

ramosissima, and C. recurvatum point up the confusion resulting from

the generic definition of plants probably all belonging to one group,

that is, Capsicum. Simulation to Solanum, section Lycianthus, is

sometimes nearly exact except for anther dehiscence; C. lycianthoides

Bitter is one example, but the slender calyx teeth are quite marginal.

A. T. Hunziker, Darwiniana 9: 225-247. 1950, gave a careful

account with notable etchings by Alfredo E. Cocucci of the species of

Paraguay and Argentina; he restricted the genus to the interpreta-

tion of Bentham and Hooker and Wettstein, indicating later judg-

ment on the inclusion of Bassovia Aublet, Brachistus and Poecilo-

chroma Miers as by Kuntze, Rev. Gen. 447. 1891, or Bassovia in part

as subgenus Aureliana (Sendt.) Bitter, Repert. Sp. Nov. 17: 331.

1921. Whatever the ultimate definition of these groups calyx and

corolla doubtfully exhibit significant characters: as constituted, Sick-

lera Sendt., Flora 178. 1846, has funnelform corolla, limb deeply

parted, calyx truncate, apparently also setaceous; Brachistus, subro-

tate corolla, limb 4-5-parted, calyx 5-lobed (type) or entire; Aureliana,

limb 5-parted, calyx 5-lobed; Decamerus, corolla rotate-campanulate,

calyx with 5-10 setae; obviously these characters, if salient by them-

FLORA OF PERU 69

selves, must result in the division of Solanum L. into several groups.

Of course, they may have at least sometimes supporting elements.

Charles B. Heiser, Jr., and Paul G. Smith, Amer. Journ. Nat. 38:

362-368. 1951, reviewed the genetics of the cultivated species; basic

chromosome number is 12 (Darlington & Yanaki, Chromosome Atlas

Cultiv. Pis. 1945); diploid number for C. pubescens Ruiz & Pavon

is 24 (Heiser & Smith); for an interesting and informative account

of the cultivated Capsicum peppers by the latter students see Eco-

nomic Botany 7: 214-227. 1953, with an extensive bibliography;

their apparently correct taxonomic conclusions indicate the futility

of proposing new species in other groups, even in Solanum, without

genetic understanding of at least some obviously distinct entity.

Ruiz and Pavon noted both C. frutescens L. and their C. pubescens,

"arnacucho" and "iocotto," as very abundant; prehistoric peppers

have been found in the burial sites of Ancon and Huaca Prieta;

Herrera recorded several cultivated varieties (C. pubescens) in Cuzco.

Pungent (Chile) peppers spread widely in Europe only after Colum-

bus, who, according to Peter Martyns, writing in 1493, brought home

a pepper more pungent than that from the Caucasus. Paprika

(bland), cayenne (pungent) and chili are all the dried ground fruit

of Capsicum; chili powdered is ordinarily a mixture of pungent pep-

per with other flavorings as oregano, garlic and salt. The tasty

appetizer, the hot sauce or "aji" — "rojo" or "amarillo," the latter

milder, sweeter — (any pungent pepper moistened usually with oil

and vinegar), so popular in Peru especially with the flavorful locally

grown "papas," is one of several of these condiments. Herrera re-

corded native names probably referable to more than one cultivated

species: "Marati," "chinchi-ucho" (both Valle de Santa Ana); "qui-

ton-quiton" (Marcapata); "piris" (in all valleys); "ccascca-pupu,"

"sempiri" (Urubamba).

KEY TO CAPSICUM

(after Heiser and Smith as to cultivated species)

Calyx teeth 10, unequal (C. brachypodum) , C. Eggersii.

Calyx teeth 5 (rarely 7) or lacking, rarely unequal.

Flowers many in axillary fascicles, 4-5-merous.

Leaves glabrate C. macrophyllum.

Leaves canescent pilose C. molle.

Flowers solitary, geminate or few.

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

Pedicels filiform, 2-2.5 cm. long; flowers hirtellous C. molle.

Pedicels shorter or at least flowers glabrous unless lobes mar-

ginally.

Leaves rather ovate; pedicels in any case usually short.

Calyx dentate, the teeth 1 mm. long or longer, erect; leaves

beneath, pedicel, calyx somewhat pubescent.

Corolla small, glabrous, on 3-8 pedicels 10-20 mm. long.

C. mendax.

Corolla 1.5-2 cm. long, villous, the 1-2 pedicels 2-3 cm.

long C. pendulum.

Calyx truncate, rarely 5-protuberant, or teeth to only 0.5

mm. long or recurved; plants glabrous or more or less

pubescent.

Corolla lobes purple; seeds black; leaves rather rugose-

nerved C. pubescens.

Corolla lobes as seeds pallid, the former rarely purple;

leaves plane.

Corolla throat as anthers rather yellow or tan; berry

red C. pendulum.

Corolla yellow all over; berry green C. Haughtii.

Corolla uniformly colored; anthers usually bluish or

purple.

Corolla greenish-white or -yellow; pedicels usually

2-5.

Pedicels stout, usually declinate; corolla lobes sub-

erect C. sinensis.

Pedicels slender, erect; corolla lobes spreading.

C. frutescens.

Corolla clear or sordid white, rarely purple; pedicels

solitary (typical) C. annuum.

Leaves lanceolate, attenuate both ends; pedicels to 2 cm. long.

C. lanceaefolium.

Capsicum annuum L. Sp. PI. 1: 270. 1753; 412.

Similar to C. frutescens but pedicels rarely two, ordinarily only

1 fruit developing at a node and corolla not tinted, rarely purple;

fruit nearly always larger. — Perhaps not found in Peru, collections

referred to it apparently rather C. frutescens L., most commonly

FLORA OF PERU 71

cultivated in northern areas. Illustrated, Heiser & Smith, Economic

Bot. 7: 223 (fruits); also Irish, Rep. Mo. Bot. Gard. 9 (many plates

hort. vars.).

Peru (cf. note above). Domesticated species.

Capsicum Eggersii Bitter, Repert. Sp. Nov. 18: 126. 1922.

A very leafy sparsely pubescent shrub a meter or two high with

small brownish-yellow flowers borne on slender peduncles 1-1.5 cm.

long; petioles 3-5 mm. long; leaves obliquely oblong to lanceolate,

cuneate to base, acutely acuminate, 4.5-8 cm. long, 1.5-3.5 cm. wide,

membranous, glabrate, lateral nerves 6-7; calyx 7 mm. long, the 10

unequal setaceous teeth subglabrous, 4 and 5 mm. long; corolla cam-

panulate-rotate, about 6 mm. long, the ovate lobes 3 mm. long; fila-

ments 1.5 mm. long, anthers emarginate, 1-1.3 mm. long, stigma

bilobed. — Leaves thin, tapering into the broadly winged scarcely

distinct petiole. Related to C. brachypodum (Dunal) Ktze., Rev.

Gen. 2: 450. 1891, of Ecuador (type from Guayaquil); it has ped-

icels only 2-6 mm. long, calyx 4-4.5 mm. long, the 10 linear subulate

teeth 3-3.5 and 2 mm. long, corolla 7-7.5 mm. long, filaments 2 mm.

long, anthers cordate, 0.8 mm. long, leaves rather densely pubescent

with many-celled trichomes (Bitter, I.e. 17: 333. 1921). C. lycian-

thoides Bitter, I.e. 332, has leaves oblique at base, petioles to 1 cm.

long, pedicels 8 or 9 mm. long, calyx 2 mm. long, teeth 5, glabrate,

linear-subulate, 4-5 mm. long. S. chacoense Hunziker, Darwiniana

9: 228. 1950, has very unequal calyx teeth, 0.5-1.5 mm. long.

Tumbez : Hacienda La Choza, Prov. of Tumbes, Weberbauer 7684-

Ecuador.

Capsicum frutescens L. Sp. PI. 189. 1753; 413.

Glabrous or slightly pubescent; leaves solitary or geminate, ovate,

acuminate, often a dm. or so long, 3-4 cm. wide; pedicels usually 2-5,

erect in fruit, somewhat incrassate above, 1.5 cm. long or often longer;

calyx costate, cyathiform, in fruit suburceolate, teeth reduced; co-

rolla white, whitish or yellowish, the lanceolate acute lobes widely

spreading; anthers blue- violet, longer than filaments; fruit oblong-

ovoid, reddish, smooth, lustrous, 2-celled, the calyx appressed; seeds

glabrous, smooth, reniform. — More common in the southern hemi-

sphere than C. annuum L. (Heiser & Smith). Calyx 5-pointed or

subdentate, the teeth at most 0.5 mm. long; the common form is var.

baccatum (L.) Irish, Rep. Mo. Bot. Gard. 9: 99. 1898.

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

San Martin: Tarapoto, Woytkowski 85161. Juanjui, Klug 4248

(det. Standley). — Huanuco: Near Huanuco, Stork & Horton 9462;

9463 (det. P. G. Smith). Pozuzo, Ruiz & Pavdn. Tingo Maria,

Ferreyra 922. — Loreto: Palta-cocha, upper Rio Nanay, Williams 1270

(det. P. G. Smith). Domesticated species. "Arnaucho" (Ruiz &

Pavon).

Capsicum Haughtii (Svenson) Macbr., comb. nov. Brachistus

Haughtii Svenson, Amer. Journ. Bot. 33: 480, pi. 19, fig. 2. 1946.

Type a small shrub (5 dm. tall), the terete or subangulate branches

with gray or reddish bark, the strict branchlets yellowish pubescent

as the leaf -nerves on both surfaces; leaves alternate or on short

branchlets, congested, ovate, entire, 1-2.5 cm. long, 4-9 mm. wide,

acute or obtuse, attenuate at base sometimes into petiole 2 mm.

long; peduncles axillary, solitary; calyx pubescent, slightly 5-angled,

3-4 mm. broad, with 5 minute recurved teeth below the margin

angles; corolla rotate, plicate, yellow, 1 cm. across, the limb 5-den-

tate; stamens included; style stout, stigma capitate; berry globose,

1-5 mm. in diameter, green, the 7 seeds brownish-yellow. — After

author, who suggests that relationship is with B. Hookeriana Miers,

Ann. Mag. Nat. Hist. ser. 2, 3: 262. 1849, of Guayaquil with much

larger leaves; the plate, opposite page 485, shows ellipsoid anthers,

presumably dehiscent longitudinally.

Piura: Forming little thickets on grassy summit of Cerro Prieto,

Amotape Hills, (Haught & Svenson 11621, type, Brooklyn Bot. Gard.).

Capsicum lanceaefolium (Miers) Ktze. Rev. Gen. 2: 449. 1891;

689. Brachistus? lanceaefolius Miers, Ann. Mag. Nat. Hist. ser. 3,

2: 267. 1849.

A shrub with rusty-tomentose branches, dichotomous, divaricate

flexuous subherbaceous branchlets, petioles 8-10 mm. long, lanceolate

leaves acuminate at both ends, fulvous puberulent especially beneath,

7-9 cm. long, 2.5-3.5 cm. wide; petioles 8-12 mm. long; pedicels

18 mm. long, solitary or 3-4-fasciculate, axillary, 1-flowered; calyx

urceolate, subentire, 16 mm. across, angled, pilose, with 5 minute

teeth; corolla rotate, nearly 2 cm. wide, the acuminate lobes floccose

on the margins; stamens included, erect, glabrous; style incrassate,

stigma capitate-bilobed. — Locality given as "Vita," probably an

error; may not be a Capsicum but apparently, ex char, and from

drawing, Miers, 111. S. Amer. PI. 2: pi. 36.

Junin: Vitoc, (Maclean). Ecuador.

FLORA OF PERU 73

Capsicum macrophyllum (HBK.) Standley, Journ. Wash.

Acad. Sci. 17: 16. 1927; 402. Witheringia macrophylla HBK. Nov.

Gen. & Sp. 3: 13. 1818. Brachistus macrophyllus (HBK.) Miers,

Ann. Mag. Nat. Hist. ser. 3, 2: 263. 1849. C. solanaceum (L'He"r.)

Ktze. var. macrophyllum (HBK.) Ktze. Rev. Gen. 2: 450. 1897; var.

pubescens Ktze. I.e.

A tall coarse more or less pubescent herb or ligneous below, the

upper geminate leaves with many slender unequal pedicels, 12 (20)

mm. long, in their axils; petioles often very narrow, 2-3 cm. long;

leaves to 2 dm. long, 7-10 cm. wide, the smaller of each pair half as

large, all glabrate or early pilosulous, membranous or firm, sub-

lustrous above, nervose and paler beneath; calyx short, urceolate,

4-dentate; corolla subcampanulate, a few mm. long and wide, the

lanceolate acute lobes spreading; stamens 4, inserted in throat, sub-

exserted, anthers yellowish, ovate, filaments villous; berry red, spher-

ical, about 6 mm. thick, seeds granulose. — "B. tetrandrus (Br. &

Bouch4) B. & H." for Ule 6242 fide Dammer is a name not found

by me. C. solanaceum (L'He*r.) Ktze. has much larger flowers.

San Martin: San Roque, Williams 7808. — Huanuco: North of

Tingo Maria, Stork & Horton 9566. Pampayacu, 5032. Cayumba,

Ferreyra 6740. — Junin: Near La Merced, 5803. — Loreto: Iquitos,

Ule 6242 (det. Dammer, B. tetrandrus}. Rio Mazan, Jose Schunke

69 (det. Standley). La Victoria, Williams 2762. Nanay, Williams

327. Bolivia to Panama and Venezuela. "Toe-mullaca" (Williams).

Capsicum mendax (van Heurck & M. Arg.) Macbr. Candollea

5: 402. 1934. Solanum mendax van Heurck & M. Arg. in van Heurck

Obs. Bot. 61. 1870.

Branchlets, leaves beneath and calyces rather densely short pi-

lose; leaves ovate, cuneate to base, rounded to acute tip, 2-3 cm.

long, about 1.5 cm. wide, lightly crisp-pubescent above; flowers

in fascicles of 3-8; pedicels 10-12 mm. long, slender; calyx campan-

ulate, the tube about 1.5 mm. high, the 5 oblong-subulate teeth a

third as long, not accrescent in fruit; corolla glabrous, 4 mm. long,

about as wide, that is, campanulate, rounded-lobed about one-third ;

style as long; anthers nearly 1.5 mm. long. — F.M. Neg. 22899.

Solanum mendax van Heurck & M. Arg. I.e. was based on Spruce

5050 from Ecuador and not from Peru as listed in Kew Index. Ac-

cording to Bitter, Repert. Sp. Nov. 17: 332. 1921, it is a species of

Capsicum, section Aureliana (Sendt.) Bitter.

Cajamarca(?): Weberbauer 6051.

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

Capsicum molle (HBK.) Ktze. Rev. Gen. 2: 450. 1891; 505.

Witheringia mollis HBK. Nov. Gen. & Sp. 3: 15. 1818. Brachistus

mollis (HBK.) Miers, Ann. Mag. Nat. Hist. ser. 3, 2: 264. 1849.

Fregirardia mollis (HBK.) Dunal in DC. Prodr. 13, pt. 1: 505. 1852.

A slender tortuous-stemmed shrub with geminate ovate acumi-

nate leaves canescent tomentose beneath and 2-4 small flowers in

the upper axils solitary on filiform pedicels 2-2.5 cm. long, these erect

in fruit; petioles 6 mm. long; leaves membranous, reticulate-veined,

sparsely pubescent above, 3.5-5 cm. long, 2-2.5 cm. wide, one of each

pair stipuliform or much smaller; calyx urceolate, villous, 5-dentate;

corolla rotate, hirtellous, limb plicate, angled, teeth acute; filaments

sometimes 6, short, glabrous, anthers oblong, obtuse, somewhat ex-

ceeded by the straight style, stigma subcapitate; berry subglobose,

glabrous. — Williams 7808, imperfect or moldy, apparently belongs

here but flowers many, leaves twice as large. F.M. Neg. 2877.

Cajamarca: Near Cajamarca, Bonpland, type. — San Martin: San

Roque, Williams 7808.

Capsicum pendulum Willd. Hort. Berol. 1: 242. 1809; 425.

More or less ligneous, glabrate or lightly (early more densely)

pubescent; leaves ovate or oblong-ovate, acuminate, 2.5-5 cm. long

or larger; peduncles solitary or geminate, incrassate apically, erect

but calyx (and fruit) more or less recurved or reflexed; calyx trun-

cate but with 5 (6) erect subobtuse teeth; corolla whitish but more

or less seriately yellowish or tan, spotted or blotched within; fruit

suboblong, red, smooth or rugose. — According to Heiser and Smith

this has the floral characters of C. microcarpum Cav., Gen. & Sp. PI.

371. 1803, probably (cf. Hunziker, 239) DC. Cat. Hort. Monsp. 1813,

and C. Schottianum Sendt. in Mart. Fl. Bras. 106: 143-144. 1896,

both natives of southern South America, not Cuba (as Cavanilles

thought; Hunziker, 239-240). Hunziker describes the specimen of

De Candolle: calyx teeth about 1 mm. long, corolla white with a pair

of yellowish spots at base of lobes (these 2.5 X 2 mm.), anthers

yellow, 1.8 mm. long, shorter than filaments, the appendages ob-

scure, not free; fruit suboblong; C. Schottianum, similar, but calyx

teeth obsolete, fruit globose; both illustrated, Hunziker, figs. 4 and 5.

The geneticists concluded: C. pendulum Willd. probably will prove

to be a cultivated form of one of these species, in this case, C. micro-

carpum Cav. Illustrated, Heiser & Smith, Economic Bot. 7: 222

(fruits); 224 (flowers).

FLORA OF PERU 75

In coastal Peru one of the most popular of the cultivated peppers

(Heiser & Smith).

Junin: Puento Yessup, Killip & Smith 26351; 26386 (both aff.

C. Schottianum fide Heiser). Brazil to Chile and Ecuador (Heiser).

Capsicum pubescens R. & P. Fl. Peruv. 2: 30. 1799; 421.

Said to become suffrutescent, in any case always pubescent; stems

angulate, often purplish at nodes, dichotomously branched; leaves

solitary or geminate, ovate, obliquely acute, very veiny, opaque, the

larger mostly 4-7 cm. long, about half as wide or larger; peduncles

incrassate above, early erect, usually 1-flowered, longer than the

somewhat recurved petioles, or to 1.5 (2) cm. long; corolla imbricate,

the plaits between lobes distinct, the latter not joined apically in bud;

filaments subfiliform; fruits yellow, orange (or red?), to about 1.5 cm.

in diameter; seeds purple-black, curved or wrinkled. — West 3739 is

densely yellowish-pilose; Cook & Gilbert and Williams specimens are

glabrate. After, in part, Heiser and Smith, Proc. Amer. Soc. Hort.

Sci. 52: 331-335, 1948, who determined diploid number as 24 and

contrasted characteristics with C. frutescens L., suggesting that be-

sides color of flowers, dull leaves, open buds and conspicuous corolla

folds, the species may have a different response to length of day.

C. guatemalense Bitter, Repert. Sp. Nov. 21: 377. 1924, included in

C. pubescens R. & P. by Heiser, has as to type 5 (rarely 7) subequal

linear-subulate calyx teeth 2.5-3 mm. long, in this character at least

suggesting C. Eggersii Bitter as author noted, with, however, as to

type, 10 calyx teeth. Also similar is the Ecuadorian C. lycianthoides

Bitter, I.e. 17: 332. 1921, with 5 linear-subulate calyx teeth 4-5 mm.

long; all these plants have glands within the calyx and more or less

plicate corollas, and their variations — potential or true character —

await genetic investigation. Charles M. Rich gave a useful general

account of this interesting plant, Bull. Mo. Bot. Card. 38: 36-42.

1950. Illustrated, Heiser & Smith, I.e. fig. 1 (plant, flower bud,

fruit, seeds).

Huanuco: Pozuzo, Ruiz & Pavon, type. — Junin: Cultivated, Pa-

natahuas, Ruiz & Pavon. — Loreto: San Antonio, Rio Itaya, Williams

3405. — Apurimac: Cultivated in garden of Dr. Octano Usandivaras,

West 3739 (det. Hunziker). — Cuzco: San Miguel, Urubamba Valley,

Cook & Gilbert 1017 (det. P. Smith). To Mexico. "Roccoto" (Ruiz

& Pavon); "uchu," "aji" (Williams).

Capsicum sinensis Jacq. Hort. Vind. 3: 38, pi. 67. 1776; 413.

C. toxicarium Poepp. ex Fingerh. Monog. 32. 1832?

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

Stems, leaves essentially glabrous, rarely shortly pubescent; leaves

ovate to ovate-lanceolate, to 1.5 dm. wide, smooth or rugose, light

to dark green; flowers 3-5 per node (except depauperate individuals),

pedicels declinate, rarely erect, at anthesis relatively short, thick,

ratio width to length of pedicels one-eighth to less than one-fifteenth;

calyx teeth none, constriction base of calyx marked (rarely faint);

corolla greenish-yellow, rarely white, 0.5-1 cm. long, lobes not spread-

ing, usually with conspicuous folds at sinuses, sometimes long-atten-

uate, folds lacking; anthers blue to purple (yellow); fruits 1-12 cm.

long, smooth or wrinkled, orange, yellow, red or brown; seeds usually

wavy-margined. — C. pubescens has purple corolla, black seeds, C.

pendulum yellow-spotted corolla, yellow anthers; in C. frutescens,

pedicels long, slender, ratio one-twentieth, erect, corolla lobes spread-

ing or recurved; C. annuum, pedicels solitary, corolla white. Has

been collected in Piura, Lima, Huancayo, Iquitos, Tingo Maria

(Smith & Heiser).

Peru: See list of localities above; known only in cultivation or

adventive.

15. DATURA L.

Brugmansia Pers. Syn. 1: 216. 1805.

Reference: Safford, Journ. Wash. Acad. Sci. 11: 173-189. 1921.

Shrubs, trees or herbs with large solitary erect or pendulous fun-

nelform-campanulate flowers. Calyx long-tubular, sometimes spathe-

like, deciduous with the corolla or circumscissile at base leaving a

disk that enlarges beneath the fruit, this sometimes prickly, 4-valved,

4-celled except at 2-celled top. Filaments and style capillary, the

stamen bilobed. Ovary 2-celled, or pseudo-4-celled (Safford) but

each of the 2 cells is partitioned by placenta walls. — The following

re'sume' was compiled before the appearance of "The Genus Datura"

by A. F. Blakeslee (The Ronald Press Company, New York, 1959),

including a genetic investigation by Avery, Satina and Rietsema and

a "Review of the Taxonomic History of Datura," by Satina and

Avery and a summary of nomenclature by Marie Hele*ne Sachet; the

nomenclatorial notes by Fosberg, Taxon 8: 52-57. 1959, have been

noted. The Fosberg and Barclay papers cited appeared while the

above reference was in press. Arthur S. Barclay was reported to be

preparing a much needed revision of the genus, unfortunately not

available now (1959) ; accordingly this re*sum£ of Safford's work has

been done without inclusion of recent materials. It seems to be

FLORA OF PERU 77

agreed that more studies are needed to clarify the taxonomy, but

without doubt the most confusing and unsettled problem is the tax-

onomy of the Brugmansia group; these students preferred Safford's

order of sections. Giro Napanga Agrero, Bol. Mus. Hist. Nat. 7:

228-235. 1943, gave an interesting account of El Floripondio with

two fine photographs.

For an informative account of the use of the narcotic seeds in

ceremonial rites see Safford, Ann. Report Smithsonian Inst. 1920:

537-567. 1922, including illustrations of the following species, except

D. cornigera.

KEY TO DATURA

Species and characters dubious; perhaps hybrids exist.

Plants herbaceous; ovary (fruits) subaculeate, rarely smooth.

Entire plant glabrous D. stramonium.

Stems, often leaves, somewhat villosulous D. inoxia.

Shrubs or trees; fruits tomentulose.

Corolla red or yellowish-red, subtruncate but with points to 1.5 cm.

long; stamens free; calyx subequally lobed. . . .D. sanguined.

Corolla white, greenish-yellow, rarely reddish suffused, the points

often longer than 1.5 cm.; calyx somewhat spathe-like and

extended.

Calyx obscurely spathaceous, unless in age subequally lobed;

stamens early agglutinate or free.

Fruit ovoid; corolla scarcely 2 dm. long; lobes little defined.

D. arborea.

Fruit oblong-cylindric; corolla 2.5-3.5 dm. long; lobes defined,

sinuses rounded or notched D. Candida.

Calyx clearly spathaceous, one side long extended, finally hamate;

corolla points 2-4 cm. long; stamens free D. cornigera.

Datura arborea L. Sp. PL 1: 179. 1753; 183.

A shrub or small tree; leaves softly pubescent; calyx spathe-like,

deciduous in fruit; corolla white, 15-18 cm. long, with distinct sinuses

or notches between the lobes; peduncle velvety-pubescent; calyx

caducous, often 6.25 cm. long; fruit ovoid. — The type came from

Peru. Often cultivated and highly regarded like D. Candida (Pers.)

Pasq. with which it is confused and to which some of the following

references may apply; this may be the same. — D. arborea of Ruiz and

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

Pavon (Fl. Peruv.) was an incorrect determination for D. arborea L.;

the Ruiz and Pavon plant is D. Candida (Pers.). For illustrations of

this species see Safford, Ann. Report Smithsonian Inst. 1920: pis. 9,

10. 1922.

The leaves are used as poultices to relieve pain and hasten sup-

puration.

Cuzco: Urubamba Valley, H err era 1536 (166). — Junin: Valley of

the Chanchamayo between Huacapistana and Palca, (Weberbauer,

247). "Floripondio," "campanchu," "toe" (Williams).

Datura Candida (Pers.) Pasq. Cat. Ort. Bot. Nap. 36. 1867; 182.

Brugmansia Candida Pers. Syn. 1: 216. 1805. D. arborea L. as to

R. & P. Fl. Peruv. 2: 15, pi. 127. 1799.

Distinguished from D. arborea by the large flowers, 2-3 dm. long,

rounded between the points, and the long-cylindrical pointed fruits;

calyx short, lax, somewhat pointed, more or less persisting. — Like

D. arborea known only in cultivation or at least in the vicinity of

habitations, unless Weberbauer's collection represents a "wild" state.

Tumbez: Deciduous bushwood on a brook, mountains east of

Hacienda Chicama, Prov. of Tumbez, Weberbauer 7656. Also prov-

inces of Cercado, Chancay and Huanuco according to Ruiz and

Pavon. — Cajamarca: Near Socota, Stork & Horton 10177. — La Liber-

tad: Cachicadan, Stork & Horton 9948; 9950. — Junin: Puerto Yessup,

Killip & Smith 26355. — Loreto : Michuyacu, Klug 84-2. Yurimaguas,

Williams 5275. La Victoria, Williams 2646. Caballo-cocha, 2395.

— Cuzco: Prov. Convention, Soukup 811. Alto Urubamba, Diehl

2^98. "Floripondio," "campanilla," "almizclillo."

Datura cornigera Hook. Bot. Mag. pi. 4252. 1846; 183.

A softly pubescent shrub or small tree with white flowers well-

marked by the widely spreading corolla-limb with greatly elongate

"points" and by the tapering horn-like apex of the calyx; corolla at

first greenish-yellow, 15-18 cm. long, the caudate tips 4 cm. long;

filaments free, hirsute below, anthers 2.5 cm. long; fruit nearly 1.5

dm. long, about 4 cm. thick. — Calyx of our specimen not obviously

costate and the calyx-beak not quite as long as illustrated. Stands

out distinctly beside typical D. arborea L. because of smaller green-

streaked corolla, different and stronger odor (Stork and Horton as

to 9948); corolla yellow but appearing red at a distance because

distal part is rose-red (Stork & Horton 9950) ; both collections around

FLORA OF PERU 79

habitations. Isern 2013 from Queguena, Arequipa, appears to be

this species, or all Peruvian specimens may represent a new species

or D. rubella Safford, I.e. 185, of Ecuador, the corolla however, de-

scribed as red, 13-14 cm. long.

Huanuco: Appearing as wild in remote sunny places, Muna, 4061?

Mexico.

Datura inoxia Miller, Card. Diet. ed. 8, no. 5. 1768; 179. D.

guayaquilensis HBK. Nov. Gen. & Sp. 3: 8. 1818, fide Svenson, with

query. D. meteloides DC. ex Dunal, Prodr. 13, pt. 1: 544. 1852, fide

Barclay, Bot. Mus. Leaflets 18: 245-255. 1959.

Cinereous-puberulent or pruinose annual; leaves ovate, acute or

rounded at base, undulate, rarely subdentate; peduncle erect, 1 cm.

long; calyx to 8 cm. long, lobes acuminate, unequal; corolla 1 dm. or

longer, shortly 8-10-cusped. — Name, in silly "correction" has been

written "innoxia." White corolla 1.5 dm. long, leaves, more or less

covered with soft down, variously angled, fruit spines relatively soft

(Svenson). Type from seeds obtained at Vera Cruz, Mexico. Illus-

trated, Barclay, I.e. pis. 50, 51, 52. F.M. Neg. 30800 (D. meteloides).

Piura: Parinas Valley, Haught 271 . East of Capo Blanco, Haught

189. To Colombia and Venezuela; Paraguay.

Datura sanguinea R. & P. Fl. Peruv. 2: 15. 1799; 187. Brug-

mansia bicolor Pers. Syn. PL 1: 216. 1805.

The only known species in Peru with dark red flowers shading

into a yellow tube; leaves entire or repand, puberulent; peduncle

slender; calyx about half the length of the corolla or less, with two

(later more) acuminate teeth; corolla 2 dm. long, lightly pubescent,

the limb about 7 cm. wide; filaments pubescent below; anthers

1.5 cm. long; fruit often in persisting calyx, smooth, seeds verrucu-

lose. — A small tree growing at 3,000 to 4,000 meters. The specimens

from southern Peru may represent a distinct species. Although re-

ferred here by Safford they are described by him as having obtuse

calyx-teeth, glabrous flowers 2.5 dm. long and anthers 2.5 cm. long.

D. Rosei Safford, I.e. 188, of Ecuador has upper leaves angular-

dentate, densely tomentose.

An ointment with reputed healing properties is prepared from the

bruised leaves and a narcotic and intoxicating drink called "Tonga"

is made from the seeds, which, according to Tschudi, may not only

stupefy but cause insanity.

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

Amazonas: Guayabamba and Chachapoyas, Woytkowski 7; Sou-

kup 4122. — Ancash: Near Pichin, Weberbauer 2926 (det. Dammer).

— Huanuco: Chinchapalca, pueblo above Mito, 1601. Huanuco,

Kanehira 224? — Junin: Tarma, Ruiz & Pavon, type. Huancayo, at

3,317 meters, Soukup 3976. — Cuzco: Cultivated, (Hen era, 166}.

Ollantaitambo, (Cook}. Also at Tarma, Xauza, Huarochera, Canta

and Huamalies, (Ruiz & Pavdn}. — Puno: Asiruni, Soukup 927.

Ecuador. "Puca campacho," "floripondio encarnado" (Ruiz &

Pavon) ; "campanulas encarnadas."

Datura stramonium L. Sp. PI. 179. 1753.

A bushy annual glabrous weed with ovate angulate dentate leaves,

tubular, usually white flowers 7-10 cm. long, 5-lobed, and erect, usu-

ally shiny pods. — Var. tatula (L.) Torr. is purplish-tinged, including

the corollas, and the prickles of capsules are often subequal. Ac-

cording to Herrera the root of D. stramonium is smoked with tobacco

as a remedy for asthma.

San Martin: Tarapoto, Williams 5856. Juanjui, Klug 4198.—

Huanuco: Along ditches in Huanuco, 2332; Ruiz. Ollantaitambo,

Herrera 3456. "Chaminco," "chamico."

16. MARKEA Rich.

Dyssochroma Miers, Ann. Mag. Nat. Hist. ser. 2, 4: 250. 1849;

also Ectozoma Miers, I.e. 191, at least as to Peru, fide Cuatrecasas.

Reference: Cuatrecasas, Repert. Sp. Nov. 61: 74-83. 1958.

Shrubby, more or less scandent, the long branches often pendent,

the flowers red, showy, solitary or several in racemes, or panicles.

Calyx tubular or campanulate, often angulate, 4-5-dentate or parted.

Corolla more or less ampliate, the tube short or elongate, the 5 im-

bricate lobes equal or somewhat unequal. Anthers subsessile or on

elongate filaments, perhaps always from a scaly or pubescent base,

dehiscence longitudinal. Fruit baccate-capsular, 2-celled, oblong or

subconical, seeds many, imbricate. — Persoon changed the name (often

Marckea in litt.) to Lamarckia but the reference to the famous nat-

uralist is obvious; Cuatrecasas followed Richard. Peruvian species

are sparsely branched epiphytes with short to greatly elongate inter-

nodes often sheltering ants. Seeds with straight embryo (unknown

for Ectozoma?} ; genus therefore in tribe Cestreae.

Flowers small, pale; leaves glabrous M. Ulei.

FLORA OF PERU 81

Flowers large, dark-colored, at least within; leaves ciliate.

M . formicarum.

Markea formicarum Damm. Bot. Jahrb. 37: 170. 1905; 80.

Type described as tuberiferous; petioles a few mm. to 1 cm. long;

leaves lanceolate, acuminate, 8.5-18 cm. long, 3-7 cm. wide, few-

nerved, glabrous except long-ciliate margins; flowers 1-several or

sometimes solitary at branchlet tips; pedicels 1.5 cm. long, incrassate

above, pulverulent as calyx, this with lanceolate caudate-acuminate

lobes to 23 mm. long, 7 mm. wide (acumen 1 cm. long, ciliate) ; co-

rolla bilabiate, 6.5 cm. long, cylindric tube 1 cm. across, below the

middle narrowed to base where 2 mm. across, the 3 smaller lobes of

limb rounded, the 2 larger obovate, denticulate; stamens at 2.5 cm.

above base, equal, filaments 12 mm. long, villous near dilated base,

anthers linear, subsagittate, 1 cm. long; ovary set in a 5-crenulate

annular disk; style filiform, incrassate toward capitate stigma; cap-

sule oblong, 1.5 cm. long, 1 cm. thick, the seeds 2.5 mm. long, 1 mm.

thick. — Given an herbarium name by Spruce in reference to the con-

spicuous ciliation. Illustrated, Karsten & Schenck, Veg. 3, Reihe,

l:j&f

Loreto: Mishuyacu near Iquitos, Klug 24-3; 808; Killip & Smith

29973. Amazonian Brazil; Colombia.

Markea Ulei (Damm.) Cuatr. Repert. Sp. Nov. 61: 78. 1958.

Ectozoma Ulei Damm. Bot. Jahrb. 37: 170. 1905. E. Pavonii Miers,

Ann. Mag. Nat. Hist. ser. 2, 4: 191. 1849; 111. S. Amer. PI. 2: 43,

pi. 1*8. 1849-57?

Terete branches often rooting; petioles 5-12 mm. long; leaves

membranous or fleshy-coriaceous, lanceolate to spathulate or obo-

vate to rounded, often 0.5-1.5 dm. long, about 2-4.5 cm. wide,

rounded or attenuate at base; inflorescences racemose-cymose in

upper axils, pedicels about 1 cm. long, puberulent as the calyx, this

irregular-margined, 5 mm. long, 2 mm. across; corolla greenish-

yellow or white, hypocrateriform-campanulate, about 7.5 mm. long,

the basal tubular part 2 mm. long, the campanulate upper portion

4-5 mm. long, minutely pilose at throat, not scaly; anthers sub-

sessile at constriction, thick, filling the cavity, dehiscence by longi-

tudinal chinks on upper two-thirds of the cells (Cuatrecasas). —

Bentham and Hooker, Gen. PI. 2: 904. 1883, not finding the perigy-

nous ring described and illustrated by Miers referred the latter's

genus to Juanulloa R. & P.; Dammer observed minute pubescent

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

scales; Cuatrecasas noted that the only very distinctive character of

M. Ulei (probably of E. Pavonii} is the subsessile anthers and it is

scarcely a generic one! The four collections of Klug have yellow-

green flowers; his 126, calyx 5 mm. long, deeply cleft, the lobes re-

flexed, corolla 12 mm. long, may be distinct; E. Pavonii (type from

Guayaquil) ex char, has 5 equal triangular erect calyx teeth, anthers

dehiscing to base, subsessile on inner side of a free ciliate ring; these

apparent characters will probably prove to be manifestations of a

variable androecium. But since Cuatrecasas did not choose to trans-

fer E. Pavonii as the earliest name this may best be done by the

student able to study it from the type region or adjacent Peru where

it may be recollected.

Loreto: Boqueron Padre Abad, Woytkowski 34387 (det. Cuatre-

casas). Mishuyacu from Iquitos, Klug 5; 126; 1127. Balsapuerto,

Klug 2953. Cerro de Ponasa, Ule 32 in part (Herb. Dahlem), but

pedicels 1 dm. long, calyces and bright yellow-green corollas 6 cm.

long and broad. Amazonian Brazil; Ecuador?

17. JUANULLOA R. & P.

Ulloa Pers. Syn. 1: 218. 1805.

Reference: Cuatrecasas, Brittonia 10: 146-150. 1958.

Shrubs, often epiphytic or sarmentose with entire more or less

coriaceous leaves and tubular-funnelform red or yellow flowers little

or much exserted from a more or less deeply divided but inflated,

usually colored calyx. Pedicels stout or incrassate toward apex.

Corolla usually contracted at the dentate apex. Filaments filiform,

elongate, anthers linear, dehiscence longitudinal. Ovary usually

conical, annular disk lobulate, fleshy. Fruit enclosed in calyx. —

Differs chiefly from Markea by the shape of the corolla, this rarely

not contracted but not campanulate apically.

Few of the Andean species have been available; the following

account of those accredited to Peru is therefore entirely from litera-

ture. Named for Jorg Juan and Antonio de Ulloa, companions of

La Condamine, Jussieu and others who were in Peru when on the

expedition to measure a degree of the equator; this for the determi-

nation of the shape of the earth !

Two obscure or imperfectly known species are retained, as de-

scribed, in Salpichroa where they can scarcely belong, but quite

possibly they are equally aberrant here.

FLORA OF PERU 83

KEY TO JUANULLOA

Flowers small, scarcely 1 cm. long (cf. J. microphylla}.

Racemes a few cm. long S. breviflora.

Corymbs open, 5-7-flowered S. cuspidata.

Flowers large (unknown, J. microphylla).

Calyx early inflated and deeply cleft laterally, shortly dentate,

finally 4-5-parted; leaves glabrous or early pulverulent at

least beneath.

Calyx membranous.

Leaves 8-12 mm. wide; corolla probably small, narrow.

J. microphylla.

Leaves much larger; corolla to 6 cm. long, ampliate limb

1.5 cm. wide J. grandiflora.

Calyx fleshy-coriaceous; corolla to 4.5 cm. long, narrow.

J. parasitica.

Calyx equally or subequally parted, not or little inflated in bud,

segments 1.5-2 cm. long.

Calyx submembranous; leaves glabrous J. ochracea.

Calyx coriaceous; leaves tomentose beneath J. ferruginea.

Salpichroa breviflora Dunal in DC. Prodr. 13, pt. 1: 474. 1852.

Glabrous including the flower segments but the rigid branches

more or less lineately glandular, the glands variously developed;

leaves obliquely rhombic-subovate-lanceolate, unequally rounded to

the short petiole, this globosely glandular at apex, acutely acumi-

nate, 2.5-7.5 cm. long, nearly 1-1.5 cm. wide, coriaceous, opaque,

paler beneath, with 4-5 pairs of obscure very slender long-branched

nerves; racemes straight, to 2.5 cm. long, peduncles obsolete, to

7 mm. long, pedicels 5-7, stout, scarcely 0.3 mm. long; calyx crateri-

form, deeply parted, the 5 unequal acute segments to 2 mm. long and

wide; buds cylindric-conic, obtuse, about 5 mm. long, 4 mm. thick;

corolla fleshy, subcampanulate, tube 4.5 mm. long, 3 mm. thick, the

5-parted limb 3-5 mm. across, with ovate-lanceolate recurved acute

unequal lobes nearly 2 mm. long, the subinflexed margins puberulent;

stamens little longer than tube, the filaments 2- nearly 3 mm. long,

anthers yellowish, stout, ovoid, sulcate, 1 mm. long; ovary obconic,

sulcate, annulate at base; style shorter than filaments; stigma dis-

coid, glandular. — This ex char, doubtfully a Salpichroa; more prob-

ably a Juanulloa. F.M. Neg. 6872.

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

Amazonas: Chachapoyas, (type, Mathews?, Herb. DC.).

Salpichroa cuspidata Dunal in DC. Prodr. 13, pt. 1: 474. 1852.

Glabrous or essentially, but branches and branchlets somewhat

marked by oval or oblong glands; leaves shortly petioled, elliptic

cuspidate, about 1.5 dm. long, half as wide, coriaceous, paler be-

neath, the 6-8 slender nerves there prominent, anastomosing; corymb

terminal, 2-3-forked, lax, long-peduncled; pedicels 4-10 mm. long,

glandular, with linear-subulate bracts 4-10 mm. long; calyx coria-

ceous, cyathiform-poculiform, the 5 triangular acute segments 5 mm.

long, 7 mm. wide, reddish in herb.; buds infundibuliform, angled,

limb segments oval-lanceolate, obtuse; stamens included; filaments

in tube, pilosulous, to 0.5 mm. long; anthers oblong-linear, obtuse,

reddish, about 3 mm. long, longitudinally dehiscent; ovary disk

prominent; style filiform, 6 mm. long, stigma cupuliform; calyx little

enlarged about the subglobose berry, this 8-10 mm. in diameter. —

This, as S. breviflora Dunal, doubtfully in this group and possibly,

especially this shrub, not in Peru. F.M. Neg. 34134.

Peru(?): (Atropa? 4-56, Pavon, in Herb. Boissier).

Juanulloa ferruginea Cuatr. in Brittonia 10: 149. 1958.

Unique in the dense reddish-brown tomentum on petioles (1-2 cm.

long), leaves (in some degree beneath), these 13-18 cm. long, 5-7.5

cm. wide, pedicels (1-1.5 cm. long), calyces (1.5-2 cm. long) and short

tubular corollas, these about 2.6 cm. long, not basally constricted,

ampliate, with reflexed lobes 7-8 mm. long; stamens free at middle of

tube, this exceeded by the 4 mm. long anthers, their clefts about

2.5 mm. long. — Type from Puerto Espino, Rio Putumayo, near

Ecuador and Peru. There are several similar plants proposed as dis-

tinct species and the Peruvian one with calyx glabrate (in age),

shorter corolla lobes, grayish tomentum, may be yet another entity;

J. speciosa (Miers) Dunal of Colombia may be represented by Sou-

kup 2920 from the Botanic Garden, Lima; its leaves are rusty tomen-

tose beneath, calyx 3-3.5 cm. long.

San Martin: San Roque, Williams 7081; 7641 (ex char.). Adja-

cent Colombia.

Juanulloa grandiflora Dunal in DC. Prodr. 13, pt. 1: 531. 1852.

Branches straight, subterete, glandular puberulent, tomentulose

toward the foliose tips; petioles canaliculate, rounded dorsally, about

FLORA OF PERU 85

1-2 cm. long; leaves approximately paired, oval-lanceolate, attenu-

ate into petiole, subacute, to about 1.5 dm. long, 5 cm. wide, thin-

membranous, glabrate above, puberulent beneath in the 7-9 primary

nerves and reticulate veins; peduncles subaxillary, scarcely 2 cm.

long, incrassate above, 1 (-few) -flowered, puberulent-tomentulose;

calyx 3 cm. long, ovoid, inflated, pilosulous, 4-5-parted, segments

acute, unequal, one deeply divided; corolla subglabrous, 5-6 cm.

long, limb subinfundibuliform, 1.5 cm. across, the broadly ovate,

acute undulate lobes medially gray tomentulose without; ovary con-

ical, style filiform, as long as corolla tube; stamens nearly attaining

corolla-throat. — After author, the type under name of Pavon in

Herb. Boissier. F.M. Neg. 8570.

Huanuco(?): Ruiz & Pav6n, type.

Juanulloa microphylla Dunal in DC. Prodr. 13, pt. 1: 531.

1852.

Branches glabrous, scarred by the fallen leaves, subangulate,

geniculate-nodose, the short foliose branches puberulent; petioles

6-8 mm. long, pilosulous; leaves geminate, ovate or elliptic or sub-

obovate, obtuse or acute, mucronate, to 2 or 2.5 cm. long, 8-12 mm.

wide, green and obscurely puberulent above, paler and glabrate be-

neath, subrounded or slightly decurrent at base, primary nerves 2-3

pairs; peduncle terminal and axillary, 1-flowered, glabrous, incrassate

above, 10-12 mm. long; calyx 1-2 cm. long, membranous, lustrous,

glabrous, inflated-cylindric, 7 mm. in diameter, the teeth triangular-

oblong, acutely acuminate; ovary ovoid, style straight, terete, enlarged

at base, equaling calyx, stigma thick, 1 mm. across, suborbicular. —

After author, otherwise unknown; in habit and foliage it suggests

Salpichroa; it may, when flowers are discovered, be found to be

lochroma. F.M. Neg. 8569.

Junin: Huasahuasi, Ruiz & Pavdn, type (Herb. Boissier, Geneva).

Juanulloa ochracea Cuatr. in Brittonia 10: 148. 1958.

Shrubby, the pendent branchlets, the petioles (5-14 mm. long)

and the 3-4 subumbellate axillary flowers including the yellow-

ochraceous corollas without more or less pulverulent; leaves ovate-

oblong or oblong-sublanceolate, rounded obtuse or cuneate at base,

acutely acuminate, 8-20 cm. long, 4-7 cm. wide, glabrous, papyra-

ceous but somewhat rigid, secondary nerves (about 6) obscure above,

subprominent beneath, veins faint; bractlets 1.5-2 mm. long; ped-

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

icels 5-15 mm. long; calyx firm-membranous, sub-basally parted,

the 5 oblong-lanceolate acute segments 15-22 mm. long, 2-5 mm.

wide at base; corolla barbate within only at stamen insertion, about

1 cm. above base, 3.5 cm. long, the tube 8 mm. long, 5 mm. thick,

the limb about 8 mm. across, little constricted apically, the oval ob-

tuse lobes reflexed, 2 mm. long; filaments 12-15 mm. long, anthers

8 mm. long, subcordate; disk undulate, 1 mm. high; style lobed, am-

pliate; fruit ovoid, 12 mm. broad. — Resembles Marked Rich. (1792)

but corolla narrowly tubular, slightly broadening medially, con-

stricted below apex; Juanulloa parviflora (Ducke) Cuatr., I.e. 150,

has corolla not constricted nor enlarged at apex; with discovery of

more intermediate species of Juanulloa R. & P. (1794), some botanist

may admit that the generic definitions are arbitrary, a seemingly

obvious probability even now.

Loreto: Florida, Rio Putumayo, Klug 2360. Colombia.

Juanulloa parasitica R. & P. Fl. Peruv. 2: 47, pi. 185. 1799;

528. Ulloa parasitica Pers. Syn. 1: 218. 1805.

Suffrutescent glabrous epiphyte with many often dependent stems,

little or not branched, the younger angulate, purplish; petioles 1.5-

3 cm. long, often curved at base; leaves alternate or subgeminate,

oblong-elliptic or somewhat obovate or lanceolate, entire or sub-

repand, attenuate at base, cuspidate, mostly 1-1.5 dm. long, 5-7 cm.

wide, lustrous (drying opaque), coriaceous, punctulate, paler be-

neath, the 5-8 primary nerves not prominent; racemes sometimes

dichotomous, at least early terminal, more or less pendent, to 1.5 dm.

long, purplish peduncles and pedicels terete, the latter 6-12 mm.

long; calyx orange, fleshy, ovoid, inflated, early 2, finally 4 cm. long,

2 cm. broad, the unequal linear-oblong segments acutely acuminate,

in age divided; corolla below throat inflated-ventricose, 8 mm. in

diameter, a little contracted at throat, 6 mm. wide at base, the limb

1 mm. long, the rounded or subacuminate lobes scarcely 4 mm. long;

filaments densely pilose basally, 8-10 mm. long, anthers equal, 8 mm.

long; ovoid-globose ovary in torulose disk; fruit 2.5 cm. long, nearly

2 cm. thick, puniceous, the violet seeds in a lustrous pulp. — Ovoid-

inflated corolla 3 cm. long as illustrated by Miers, 111. S. Amer. PI.

San Martin: Tarapoto, Ule 6606. Jepelacio, Klug 3529 (det.

Standley, J. grandiflora). — Huanuco: Huamalies, Weberbauer 3^37.

Pozuzo and San Antonio, Ruiz & Pavon, type. — Loreto: Puente Me-

lendez, Tessmann 4876 (det. Werdermann). Bolivia; Mexico?

FLORA OF PERU 87

18. GESTRUM L.

Sessea R. & P. Prodr. 21, pi. 33. 1794.

Reference: P. Francey, Candollea 6: 46-398. 1936 (not 1935;

cf. 7, title page); 7: 1-132. 1936.

Shrubs or trees, often with virgate flowering branches, and usu-

ally greenish-yellow flowers in terminal or axillary cymes, panicles

or racemes, stipuliform leaves often deciduous, probably sometimes

caducous. Calyx campanulate, tubular or poculiform. Corolla sal-

verform or tubular-funnelform, the tube very long and slender, more

or less contorted at throat. Stamens usually equal or subequal (ex-

cept C. conglomeration, C. confertum, C. foliosum), 5, adnate to the

corolla tube, the filiform filaments often pubescent especially below,

sometimes tumid or with a toothlike appendage. Anthers (Peru)

spherical or ellipsoid. Ovary glabrous (or papillose, rarely glabrate),

oblong or globose, or shortly stiped, 2-celled, style filiform, stigma

entire or lobed, rarely exserted. Fruit capsular to baccate, scarcely

or little succulent. — Embryo straight or nearly. Floral parts 4, some-

times 4, 6 or 7. Many species are reputed to have medicinal value

and most emit a very disagreeable or even foetid odor, although the

flowers may be fragrant.

Sessea, as section and genus, recalls Don Martin Sesse", director

of the Botanical Garden of Mexico at the time of Ruiz and Pavon.

Dunal (followed by Francey) divided the genus into two sections:

Habrothamnus (Endl.) Schlecht. flowers somewhat purple to orange

or red, the calyx narrow, often reflexing segments, the Mexican and

Central American species C. elegans (Brongn.) Schlecht. and C. au-

rantiacum Lindl., both cultivated, at least in the botanical garden at

Lima in 1946 (Soukup 2921, 2928), the former with tomentose leaves,

purple flowers 2 cm. long, the latter with glabrous leaves, orange

flower 2.5 cm. long; Eucestrum Dunal, flowers whitish, greenish, yel-

lowish, calyx with broad short erect teeth.

The key includes Peruvian plants described as Sesseas but several

have not been transferred, since their standing may be questioned;

in the description of generic characters not diverse to typical, as gla-

brous ovary, rotund anthers, etc., these plants have not been in-

cluded, and for convenience the stipuliform leaves sometimes developed

are referred to as stipules in the key. This has been transcribed from

Francey's meticulous work but modified in conformity to the floristic

approach of this publication; the well-drawn descriptions are essen-

tially his or his interpretation of Dunal's. In Notizbl. Bot. Gart.

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

Berlin 11: 978, 1934, he gave a key largely copied herewith as per-

taining to Peru based primarily on that of Bitter, I.e., but he noted

that it was not possible for him to place all species in the grouping of

the previous student.

Francey, I.e. 49-51, discussed the character of Sessea in contrast

to that of Oestrum; while he retained the former his remarks indicate

that he did so with considerable doubt; then, too, as a student he

may have been diffident or was counseled to respect tradition. The

conclusion that Sessea constitutes a natural group would be rein-

forced if the family as a whole did not exhibit within well-defined

groups various fruit anomalies that are accepted as merely specific;

even the not closely allied Lycium furnishes examples, as do Datura,

Solatium and Brunfelsia. Accordingly, in conformity with the gen-

eral intent followed in this work, the species of Sessea with 2-valved

capsules, the valves sometimes so deeply bifid as to appear 4 (S. pedi-

cellata is actually 4-valved according to Francey) are treated as spe-

cies of Cestrum. Francey found that the ovary in dissection was not

easily or as easily parted in Cestrum but the distinction is one of

degree and, significantly, not concomitant with characters of pubes-

cence or fruit, the latter, as agreed by Francey, sometimes passing

in design from capsular to baccate, notably, in Peru, in the case of

C. glaucophyllum and probably other species when fruit is known.

Some species, as in true Cestrum, may prove to be individual variants.

KEY TO CESTRUM (including SESSEA)

Indument obvious all over leaves beneath and in inflorescence in-

cluding calyces (or these tardily glabrate; glabrous only, C. flex-

uosum) and (or) corollas; inflorescence or the flowers at tip of

often bracteate or foliose branchlets, these always longer than

the subtending petiole.

Corolla pubescent including the tube; fruit capsular except C. strig-

illatum.

Calyx subtruncate-crenate; ovary apically pilose.

Calyx 7-9 mm. long C. Mancoi.

Calyx 5-5.5 mm. long (species may be one and variable; see

also S. multiflora).

Corolla (type) at least 2 cm. long; pedicels in part 1-1.5 mm.

long C. Franceyi.

Corolla (type) at most 18 mm. long; pedicels none to 0.5 mm.

long S. discolor.

FLORA OF PERU 89

Calyx 3-3.5 mm. long C. osnoense.

Calyx dentate, 7-10 mm. long.

Corolla about 1.5 cm. long; fruit capsular.

Flowers many in broad panicles; ovary pilose at tip.

C. maleolens.

Flowers few, inflorescence simple or panicles narrow; ovary

glabrous C. dependens.

Corolla about 3 cm. long; fruit baccate.

C. cancellatum, C. strigillatum.

Corolla glabrous or the lobes obscurely pulverulent; fruit baccate

or tardily dehiscing at apex.

Indument sparse or fine, stellulate.

Flowers 3.5 cm. long, 1-3; filaments glabrous, 1.5 mm. free.

C. glaucophyllum.

Flowers about 17 mm. long, several to many; filaments pilose,

5 mm. free.

Calyx glabrous C. flexuosum.

Calyx lanate C. Humboldtii.

Indument dense, villous-stellate; filaments glabrous, 5.5-7 mm.

free.

Corolla lobes obtuse; calyx teeth broadly ovate, acute; leaves

subcordate C. lanuginosum.

Corolla lobes acute; calyx teeth 1-1.5 mm. long, 1 mm. wide,

acuminate; leaves subacute at base C. tomentosum.

Indument lacking in age or an obscure puberulence on calyces, co-

rollas and rarely leaves beneath especially on nerves, then often

deciduous.

Inflorescence rarely sessile, always longer than subtending petiole,

sometimes at branchlet tip.

Leaves somewhat pubescent beneath; filaments pilose (except

C. glaucophyllum) , free 5.5-7 mm.

Lateral leaf nerves 20 or more pairs . . (C. venosum) C. petiolare.

Lateral leaf nerves 13 or 14 pairs C. Humboldtii.

Lateral leaf nerves 5-9 pairs.

Calyx 5 mm. long; spikes 5-7-flowered C. flexuosum.

Calyx 10 mm. long or longer; spikes 1-3-flowered.

C. glaucophyllum.

Leaves glabrous, or soon obscurely pulverulent.

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

Calyx 6-10 mm. long; filaments somewhat pilose, denticulate

(C. aberrans), free 4-6 mm. or longer except C. longi-

florum (2 mm.).

Leaves 2-7 cm. long; flowers to 2 cm. long.

Corolla 22 mm. long, glabrous; fruit baccate. . C. Dunalii.

Corolla 1.5 cm. long, calyx puberulent; fruit capsular.

C. aberrans.

Leaves often 1-2 dm. long; flowers 5-6 cm. long.

C. longiflorum.

Calyx 2-4.5 (5) mm. long; filaments glabrous except C. ovato-

lanceolatum.

Filaments denticulate or bent, 3.5 mm. free; calyx nearly

tubular, 3.5-5 mm. long; corolla 2.5-3 cm. long (spe-

cies may be tenuous).

Leaves crisped-undulate; stipules deciduous; filaments

denticulate C. undulatum.

Leaves plane or nearly; stipules none.

Filaments denticulate; leaves (type) abruptly cuspi-

date C. ovalifolium.

Filaments merely geniculate; leaves acuminate.

Corolla about 2.5 cm. long; calyx 4 mm. long.

C. Raimondianum.

Corolla about 3 cm. long; calyx 5 mm. long.

C. pseudopedicellatum.

Filaments smooth, even, free 0.5-3 mm.; calyx obconic-

campanulate, 2-3.5 mm. long.

Stipules none; calyx 2 mm. long, subpulverulent; corolla

about 1 cm. long C. ovatolanceolatum.

Stipules present unless in age; corolla about 2.5 cm. long.

Leaves often 8-15 cm. long, principal nerves 7-8 pairs.

C. reflexum.

Leaves often 5-7 cm. long, principal nerves 10-12 pairs.

C. auriculatum.

Inflorescence rachis more or less developed or (and) the leaves ob-

long-lanceolate or in part about 1 dm. long and often elliptic;

filaments various; see contrast 3.

Leaf nerves many (often 13-21 pairs), subparallel nearly to mar-

gin; filaments uneven, 1-5 mm. free.

FLORA OF PERU 91

Stipules none; leaves rounded at base; adnate part of fila-

ments pilose C. racemosum.

Stipules present; leaves acute to base; filaments glabrous.

C. Mathewsii.

Leaf nerves 7-15, arcuate, uneven, often joined well before

margin.

Stipules conspicuous; leaves rarely 4 cm. wide, oblong-ovate-

lanceolate except C. Weberbaueri; filaments glabrous.

Leaves often 1 dm. long, acute at base; filaments dentate

(known).

Leaves to 2 cm. wide; stipules lunate C. Macbridei.

Leaves to 4 cm. wide; stipules suboblong.

Calyx to 5 mm. long; stipules 3-5 mm. long.

C. peruvianum.

Calyx 6-6.5 mm. long; stipules 1-1.5 cm. long.

C. Weberbaueri.

Leaves 5-10 cm. long, rounded basally or to a subacute

base; filaments smooth C. foliosum.

Stipules deciduous, minute or lacking; leaves all or many

5 cm. wide or wider except C. laevifolium; filaments pilose

to puberulent.

Leaves truly ovate or ovate-lanceolate, mostly 5 cm. wide

below middle; corolla 1 cm. long, filaments free 0.5 mm.

(type) C. ovatolanceolatum.

Leaves oblong-lanceolate, few 5 cm. wide, lustrous; corolla

2 cm. long, filaments free 4 mm C. laevifolium.

Leaves more or less elliptic-lanceolate, many 5-10 cm. wide,

subopaque; corolla 1.5-2 cm. long.

Leaves not decurrent; calyx 4.5 mm. long, clearly costate;

filaments barbate, free 1.5-3.5 mm C. loretense.

Leaves decurrent; calyx 2.5-3 mm. long, subterete or

5-nerved; filaments, except C. obscurum, free to

4 mm., pilose (species, except C. obscurum, seem in-

tangible).

Corolla gradually ampliate, finally 16-20 mm. long;

calyx glabrate.

Calyx subcyathiform; leaves membranous.

C. nemanthum, C. silvaticum.

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

Calyx tubular; leaves firm to coriaceous.

C. Baenitzii.

Corolla cylindric nearly to apex, 15 mm. long; calyx

tubular, glabrous in type C. obscurum.

Inflorescence much reduced, rachis none or obscure, the flowers

(or flower) sessile or subsessile in the axils; leaves to 1 dm.

long or longer, always rather elliptic, but often rigid; filaments

even, except C. Sendtnerianum (cf. C. Mathewsii).

Flowers 1-4; filaments denticulate, pilose. . . .C. Sendtnerianum.

Flowers usually many; filaments even.

Leaves very rigid, to 7 cm. long, nerves 4-8 pairs.

C. confertum.

Leaves more or less flexible, 1-2 dm. long, and (or) many-

nerved.

Stipules none; corolla 15-17 mm. long. . .C. conglomeratum.

Stipules present; corolla (14) 19-21.5 mm. long.

Leaf nerves 18-21, rigid; leaves 2.5-3.5 cm. wide, coria-

ceous C. Mathewsii.

Leaf nerves 12-14, rigid, coarse; leaves 4-7.5 cm. wide,

coriaceous C. ellipticum.

Leaf nerves 10-12, slender; leaves 2.5-4 cm. wide, mem-

branous C. falcatum.

Oestrum aberrans Macbr., nom. nov. Sessea confertiflora Fran-

cey, Notizbl. Bot. Gart. Berlin 11: 880. 1934, not Schlechtendahl.

Branches 2.5-4 mm. thick, glabrous as the shortly fastigiate

branchlets except early puberulent apically; petioles 1-6 mm. long,

1-2 mm. wide, enlarged at base; leaves erect or nearly, oblong-

lanceolate, cuneately long-decurrent at base, obtuse or obtusely

acute, 4.5-7.5 cm. long, 1.5-3 cm. wide, subcoriaceous, revolute,

lustrous above, glabrous, irregularly disposed lateral nerves about 7;

axils fuscous tomentose; flowers capitately congested, pedicels 1 mm.

long, peduncles 1-5 (-8) mm. long, puberulent as the obconic calyx,

this 7.5-10 mm. long, 3 mm. across, broad obtuse unequal teeth

1-1.5 mm. long; corolla glabrous except lobes, whitish-green, 1.5

cm. long, tube 11 mm. long, limb lobes puberulent, orbicular, 2.5

mm. long; filaments adnate, 4 mm. long where pilose, the teeth

bifid; ovary glabrous, 6-ovuled; stigma capitate; capsule obovoid,

included, 6.5 mm. long, 3.5 mm. thick, shortly stiped, dehiscing

FLORA OF PERU 93

by narrowly elliptic valves, the 3-4 mature seeds 3 mm. long, 1 mm.

wide, cotyledons elliptic. — According to Raimondi poisonous to all

domestic animals.

Ancash : Between Huara and Llipe, Prov. Cajatambo, (Raimondi,

type). — Lima: San Juan to Huarochiri, (Raimondi). "Laplacata."

Oestrum auricula turn L'Her. Stirp. 1: 71, pi. 35. 1788; 132.

C. lasianthum Dunal in DC. Prodr. 13, pt. 1: 649. 1852. C. serratum

Dunal, I.e. 665. C. leptanthum Dunal, I.e. 666, all fide Francey.

A slender-branched shrub 1-several meters high, glabrous except

for the often finely puberulent inflorescence branches and flowers;

petioles 1-1.5 cm. long, often biauriculate at base, the auricles

lunate and as long as or much shorter than the petioles; leaves

oblong-ovate or -lanceolate, to 1 dm. long or longer, often shorter,

more or less acuminate, somewhat narrowed at base; flowers in

terminal and axillary corymbose panicles, sessile or scarcely ped-

icellate, to about 2.5 cm. long, yellowish-green with blackish or

reddish-brown overtones; calyx glabrous to pubescent, nearly cyathi-

form, scarcely 2 mm. long, the teeth minute; corolla tube filiform,

enlarged above, often lightly puberulent, filaments free 1.5-3 mm.,

glabrous except adnate part, edentate; fruit fleshy, black. — The

plant described here is that illustrated by L'Heritier, and Dunal

may have erred in referring to it the earlier published C. hediunda

Lam. which more probably is his C. hediundinum, the corolla gla-

brous. It is not clear, however, that more than one variable species

is concerned, as Francey decided, but he did not use Lamarck's

name. Dombey 352 and Gaudichaud 87, both Herb. DeCandolle

and without data except "Peru," are types of Dunal's species. Only

some of the many collections are listed. F.M. Negs. 2970, 2980

(C. hediundinum); 6902 (C. lasianthum); 6985 (C. leptanthum); 8572

(C. serratum).

Cajamarca: Chota, Raimondi. Cascas, Raimondi. — San Martin:

Tarapoto, Woyktowski 35026 (det. Cuatrecasas) ; Williams 6169. —

Ancash: Tambo de Pariocota, 254.5. Chancos, Nunez 324.1. — Lima:

Dombey, type. Lurin, Pennell 12221. Stn. Eulalia, Goodspeed 33102

(det. Killip). Canta, Pennell 11+355; Nunez 950; 954. Prov. Canete,

Vargas 9313 (det. Johnston). Rio Chillon, Pennell 14445. Ato-

congo, Mexia 04037 (det. Johnston). Callao, Gaudichaud. Chosica

to Matucana, 76; 539; Mexia 04092 (det. Johnston) ; Soukup 2055;

2069. — Huanuco: Pampayacu, Sawada 47. — Arequipa: Laspinas,

2,200 meters, Eyerdam & Beetle 22149 (det. Johnston). Tiabaya,

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

Pennell 13089. Near Arequipa, Rose; Isern. Guicacha, Ferreyra

7220. — Puno: Near Puno, Soukup 406 (distr. as C. conglomeratum) .

Ecuador. "Hierba Sancta," "hierba hedionda."

Cestrum Baenitzii Lingelsh. Repert. Nov. Sp. 7: 248. 1909; 309.

Branches erect, 6-8 mm. thick, the erect rigid branchlets longi-

tudinally striate-canaliculate, sometimes pulverulent toward the

apex; stipuliform leaves none; petioles stout, sometimes tortuous,

arcuate, 7-15 (20) mm. long; leaves approximate apically, at ma-

turity oblong-elliptic or subovate, long-attenuate into the petiole,

gradually narrowed to the acute, obtuse or rounded and shortly

cuspidate apex, mostly 1-3 dm. long, 6-10 cm. wide, sometimes

even larger, chartaceous or firmer, often coriaceous, obviously revo-

lute at least toward apex, lustrous especially above, the midnerve

and 7-10 lateral nerves prominent beneath, the reticulation lax;

racemes subspicate, 7-10-flowered, often 3 or 4 in axil, the rachis

3-20 mm. long, puberulent as the 1 mm. or shorter peduncle; floral

leaves none; flowers yellowish or pale green, usually subsessile; calyx

tubular, terete, 2.5-3 mm. long, 1.5 mm. thick, glabrate or minutely

puberulent only without, the 5 teeth strongly unequal, 0.5-1 mm.

long, 1 mm. wide, often connate; corolla infundibuliform, finally

16.5-18 mm. long; lobes 3-4.5 mm. long, glabrous within; filaments

adnate two-thirds the length of tube, free 3.5 mm., smooth, pilose

to above the middle; style 11.5-14 mm. long, pubescent toward

tip; stigma capitate-discoid; fruit obovoid or subglobose, shortly

attenuate at base, 8.5 mm. long, 6 mm. thick, black, violet or green;

seeds 3, 5 mm. long, 3 mm. broad, 2 mm. thick, rounded dorsally,

angled or plane on the inner face. — Perhaps only varietally distinct

from C. megalophyllum Dunal in DC. Prodr. 13, pt. 1: 638. 1852

(Morton, Journ. Arnold Arb. 17: 341. 1936); however, the typical

form of DunaTs plant from Trinidad often has shorter dentate

petioles, shorter flowers, equal calyx teeth, characters that may be

inconstant or inconsequential. Illustrated, Francey, pi. 3, fig. 50

(flower).

San Martin: Tarapoto, Spruce 4355. Jepelacio, Klug 3612 (det.

Standley). San Roque, Williams 6970; 7617. Pongo de Cainarachi,

Klug 2625. — Huanuco: Exito to Balsaplaya, Vargas 5355. Puente

Durand to Exito, Mexia 8155 (det. Morton, C. racemosum). Tingo

Maria, Allard 21944 (det. Lyman Smith). — Loreto: Yurimaguas,

Poeppig 2315; Killip & Smith 28051; 28534; 29034; 29051; Williams

5277; Ferreyra 7839. Rio Mazan, Mexia 6463 (det. Standley).

FLORA OF PERU 95

Pongo de Manseriche, Tessmann 4209. Iquitos, Ule 6240; Killip

& Smith 27453; King 1083. Rio Nanay, Williams 627 (det. Werder-

mann). Bolivia and Brazil to Central America. "Yanagara negra"

(Mexia).

Cestrum cancellatum Dunal in DC. Prodr. 13, pt. 1: 657.

1852; 136.

Branches compressed above, sulcate-canaliculate, 3-4 mm. thick,

apically leprose-tomentose; petioles 5 or 6 mm. long, 1-1.5 mm.

thick, enlarged at base, stellate-tomentose as the inflorescence and

younger leaves beneath, these ovate- or elliptic-lanceolate or oblong,

long-attenuate or cuneate to base, in type acuminate to a very narrow

acute tip, usually 1-2 dm. long, about 4-6 cm. wide, membranous,

scarcely revolute, greenish-olive, glabrous and lustrous above, the

slender lateral nerves 7-8, prominent and reticulate beneath; stipule-

like leaves none; spikes axillary, 4.5-7 cm. long, stellate-tomentose;

flowers sessile, rather fasciculate, the subpetiolate bractlets ovate-

acuminate or cuspidate, 8-12 mm. long, 4 or 5 mm. wide; calyx

lax, cyathiform, angled, 7-9 mm. long, 4 or 5 mm. thick, puberulent

within, 5-nerved, the acute or acuminate teeth 2.5 mm. long, 2 mm.

wide; corolla 3-3.5 cm. long, tube contracted above and below ovary,

cylindric, subfiliform, dilated at throat, infundibuliform-clavate,

about 27 mm. long, not constricted below the limb, yellowish, stellate-

pilose and within pubescent, the ovate acuminate segments 7-8.5

mm. long, 2 mm. wide, marginally introflexed, densely tomentose

without, puberulent within; stamens equal, filaments adnate to tube,

1-2 mm. free, glabrous, smooth, straight; ovary 16-ovuled; style

28 mm. long, slightly tomentulose only above, stigma capitate, bifid;

berry sessile, ellipsoid, at least 6 mm. long with 10 seeds. — The

type by Poeppig fide Dunal from Rio Plata is doubtfully separable

from C. strigillatum R. & P. var. calycinum (Willd.) Ktze. to which

Francey in herb, referred Williams 6570 from Tarapoto; in mono-

graph he referred the specimen to C. cancellatum, at least as to Peru;

C. cancellatum, then, seems at most a variant. F.M. Neg. 2969.

Peru (cf. note above). Brazil?

Cestrum confertum R. & P. Fl. Peruv. 2: 29, pi. 153. 1799; 221.

Glabrous except for some long evanescent lanosity at the growing

tips, this indument reddish and enduring in the leaf axils; branches

many, elongating, virgate, 3-5 mm. thick, becoming sulcate, the

few branchlets 1-1.5 dm. long; leaves spreading, broadly elliptic

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

or obovate, acute or rounded at base, shortly acuminate or rounded

and shortly cuspidate, 3.5-6 cm. long, about 2-4 cm. wide or some-

what larger, coriaceous, rigid, revolute, nitidulous both sides, paler

beneath, the midnerve and 4-6 (-8) lateral nerves there yellowish

and very prominent; stipuliform leaves none; flowers sessile, 6-7

congested in the leaf axils and as many as 20 crowded apically;

bracts linear, acuminate, 2 mm. long; calyx coriaceous, obconic,

cyathiform, 3-4 mm. long, 2 mm. across, glabrous within but merely

glandular without and the 5 broad teeth subacute or obtuse, mar-

ginally tomentulose, 0.5 mm. long; corolla infundibuliform, green

or slightly brown-violet, 18.5 mm. long, the tube contracted below

the ovary, scarcely ampliate to apex or constricted below the limb,

15.5 mm. long, glabrous without, pulverulent within, the lobes

(of the limb) 3-4.5 mm. long, broadly ovate or suborbicular (ex-

planate), glabrous except for the puberulent margins; stamens un-

equal, 14-15 mm. long, the filaments free for 2 mm., smooth, arcuate,

not geniculate nor tumid, the adnate portion puberulent; anthers

subquadrate; ovary attenuate at base, glabrate; style 1.5 cm. long,

filiform, sparsely puberulent to base, stigma capitate; fruit ovoid-

ellipsoid, fulvous, 4-5 mm. long, 2.5-3.5 mm. thick, subtended by

the accrescent calyx; seeds brown, oblong, triangular, 3 mm. long,

1.5 mm. broad, 0.7 mm. thick. — Var. grandifolium Francey, I.e. 22,

seems to be only a particular name given to an individual with leaves

to nearly 1 dm. long, lateral nerves 7-8, the greenish-yellow flowers

noted by the collector as fragrant. The leaves are extremely rigid,

and at each petiole base there is a little cushion of reddish trichomes

(Francey). F.M. Neg. 18392.

Huanuco: Panao, Chaglla and Muna, (Ruiz & Pavdn, type);

3961. Huacachi, 4184. Chaglla to Muna, 2,400 meters, Weber-

bauer 6796. In sunny thicket, 2,100 meters, Muna, 4058 (type,

var.). — Junin: Carpapata, Prov. Tarma, Soukup 3466.

Cestrum conglomeratum R. & P. Fl. Peruv. 2: 29, pi 156.

1799; 227.

In general like C. confertum but the leaves (typically) 1-2 dm.

long, 3.5-nearly 6.5 cm. wide, elliptic-lanceolate, acute at base,

acuminate, subcoriaceous, the lateral nerves 9-11 and, as the reticu-

late venation, prominent beneath; petioles 12-23 mm. long, gibbous

at base; stipuliform leaves none; flowers sessile, (5-) 15-30 glomerate

(typically) in the leaf axils, bracts (typically) elliptic-lanceolate, acute,

glabrous or puberulent, 5 mm. long, bractlets linear, 2-3 mm. long;

FLORA OF PERU 97

calyx tubular-poculiform, 3.5-5 mm. long, 2 mm. across, glabrous

within as without except for the 5-7 acute teeth, these tomentose

marginally and within, 1 mm. long and broad; corolla greenish- white,

infundibuliform, 17 mm. long, tube contracted below the ovary, finally

obconic, little ampliated to apex, not constricted below the limb,

13.5 mm. long, quite glabrous; lobes 5-7, 2-3 mm. long, acute,

lightly puberulent and marginally tomentose only without; stamens

5-7, 10-12.5 mm. long, 1 or 2 shorter, the straight smooth glabrous

filaments free for 2 (-5) mm.; ovary oblong or shortly ellipsoid,

8-ovulate, obconically stiped; style 12.5 mm. long, filiform, obscurely

puberulent toward apex; stigma capitate. — Var. Kunthii Dunal,

DC. Prodr. 13, pt. 1: 620. 1852 (C. reticulatum Willd. ex Roem.

& Schult. Syst. 4: 808. 1819), apparently only an individual variation,

has more oblong-elliptic leaves, to 13 cm. long, to nearly 4 cm. wide,

membranous but rigid; flowers 1.5 cm. long, 6-7-glomerate; fila-

ments free for 5 mm.; bracts 2-3 mm. long; var. simulans Macbr.,

var. nov., ramulis foliisque subtus pileis minusve puberulentis; foliis

ad 1 dm. longis, 6 cm. latis, acutis vel breviter acuminatis; this

suggests C. falcatum with longer corollas, narrower leaves, and C.

bolivianum Francey, I.e. 230, with narrower leaves, flowers 6-7 con-

gested on a peduncle, and C. coriaceum as to Herrera, Synop. Fl.

Cuzco 366. 1941, not Miers. Floral divisions are sometimes 6 or 7.

Illustrated, Francey, 7: pi. 2, fig. 53 (flower); pi 3, fig. 28 (flower).

F.M. Negs. 2971; 2973 (var.).

According to Herrera an infusion of the leaves added to the

popular beverage "chicha" serves as an excellent soporific.

Cajamarca: Huanamarca and Cascas, Bonpland (type, var. Kun-

thii).— Huanuco: Panao, Ruiz & Pavdn, type. Near Muna, 4159.

Yanano, 3714- Vilcabamba, Rio Chinchao, 5151. — Junin: Dos de

Mayo, Pichis Trail, Killip & Smith 25860. — Apurimac: Prov. Aban-

cay, Raimondi 11098. — Cuzco: Calca, Herrera 2084. Urubamba

Valley, Herrera 1124- Quispicanchis near Marcapata, Metcalf 30731

(type, var. simulans). — Puno: Prov. Carabaya, Raimondi. Colombia.

Cestrum dependens (R. & P.) Macbr., comb. nov. Sessea de-

pendens R. & P. Veg. Syst. 44. 1798; Fl. Peruv. 2: 9, pi. 116. 1799;

217. S. Dombeyi Dunal in DC. Prodr. 13, pt. 1: 596. 1852.

A slender tree to 10 meters high with spreading pendent branches,

the younger (and the leaves beneath) very pulverulently floccose-

tomentose; petioles 1-3.5 cm. long; leaves ovate- or oblong-lanceolate,

acute or rounded at base, to 17 cm. long, 5.5 cm. wide, green above

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

and glabrate or somewhat pubescent with few-branched trichomes;

inflorescence in type pendulous, to 3 dm. long, very narrow, simple

or often shortly branched, the lower branches to 6.5 cm. long;

calyx 7-9 (11) mm. long, somewhat unequally lobed, prominently

5-costate; corolla 2.5 cm. long, tube more or less contracted below and

above ovary, broadly obconic, 14.5 mm. long, limb segments 5 mm.

long, when expanded 4.5 mm. wide, margin glandular-tomentose,

obscurely puberulent; stamens subequal, 14 mm. long, adnate 5.5

mm., free part little pilosulous basally; stigma capitate, exserted;

ovary glabrous; capsules 13 mm. long, 4 mm. wide, the 4 valves 6 mm.

long, 2.5 mm. wide. — (Apparently these data, after Francey, I.e. 879,

from Raimondi and Bolivia specimens.) Bitter pointed out that

the Dunal type by Dombey was part of the same Ruiz and Pavon

collection, the apparent differences, as finer leaf nerves, fewer flowers,

unimportant. F.M. Negs. 3003; 6883 (S. Dombeyi).

Huanuco: Distrito Churubamba, Mexia 8210 (det. Morton with

query, S. stipulata). — Junin: San Jose", Andamarca to Pangao, (Rai-

mondi, det. Francey). Huasahuasi, Dombey; Ruiz & Pavon, type. —

Cuzco:Marcapata, Vargas 3709. Bolivia. "Pichi-pichi" (Mexia).

Sessea discolor Francey, Notizbl. Bot. Gart. Berlin 11: 985.

1934.

Branches flexuose, costate, verruculose below, 1.5-3 mm. thick,

puberulent only toward apex; petioles mostly 2-3 cm. long, sub-

quadrate, often contorted basally; stipuliform leaves deciduous, ro-

tund, 2-3.5 mm. long and wide, tomentulose beneath; leaves often

pendent, elliptic-ovate-lanceolate, ordinarily broadest below the mid-

dle, rounded or subacute at base, not decurrent, shortly and acutely

attenuate at apex, 5.5-9 cm. long, 2.5-4.5 cm. wide, membranous,

undulate, plane, opaque and glabrous above except the puberulent

midnerve, beneath whitish-tomentulose, the 10-13 lateral nerves ob-

vious; corymbs terminal, dense, 18 cm. long, 13 cm. broad, the

branches strongly dichotomous, tomentose, the peduncles 7-10 cm.

long; flowers congested, pedicels 0.5 mm. long; calyx attenuate be-

low, tubular, 5.5 mm. long, 2.2 mm. thick, only without minutely

pilose as the 5 minute rounded teeth marginally; corolla yellow,

18 mm. long, tube contracted below ovary, scarcely ampliate apically,

3.2 mm. broad, densely stellulate-pilose, 14.5 mm. long; lobes 2.8

mm. long, acute, puberulent only without, the margins lanate;

stamens unequal, about 14 mm. long, the filaments free for 7 mm.,

slightly geniculate, incrassate and pilose, smooth, the adnate part

FLORA OF PERU 99

obscurely puberulent; ovary densely pilose above, the stipe obconic;

style 14.5 mm. long, glabrous as the capitate stigma. — Leaves more

tomentose beneath than the similar S. muUiflora Bitter; the several

plants designated as species may prove to be variants of one or two;

Balls' collection has flowers 2 cm. long or longer, but sessile or sub-

sessile. This, if distinct from S. muUiflora, requires in Cestrum

a new name (not C. discolor Dunal).

Cuzco: Rocky brushy slopes below Colquipata, 3,200 meters,

Pennell 13782, type. San Miguel, Cook & Gilbert 111*8. Quebrada

de Paucartambo, Vargas 11183 (det. Standley) ; Balls B6673; H err era

2974. Bolivia. "Asnaj-salli" (Herrera).

Cestrum Dunalii Francey, Candollea 6: 353. 1936.

Branches erect, 1-2 mm. thick, pulverulent to puberulent-

tomentulose, notably foliose with oblong-ovate-lanceolate leaves

only 2-3 cm. long or slightly longer, 6-15 mm. wide, decurrent,

acute or acuminate, membranous, reticulate-venose both sides, the

4-6 lateral nerves strongly ascending, conspicuous beneath, where

sparsely puberulent, the upper surface glabrous; petioles 3-5 mm.

long, grooved above; stipuliform leaves none; panicles terminal, sub-

corymbose, often 11 cm. long, nearly as broad, the floral leaves

petiolate, oblong-lanceolate, acuminate, puberulent, 3-9 mm. long;

flowers thyrsoid, 3-5 mm. distant, pedicels 2-3 mm. long, glandular,

verruculose; calyx tubular-cupuliform, lax, angled, stoutly 5-nerved,

9.5 mm. long, glabrous without, papillose within, the unequal teeth

often connate, these 3-5, acute, sinus acuminate; corolla 22.5 mm.

long, tube contracted below and above ovary, obconic, 4 mm. broad

at apex, not constricted below the limb, 17.5 mm. long, glabrous

without, pilose on the adnate filaments, these free for 7.5-8 mm.,

little tumid or geniculate, villous-pilose; lobes of limb 4 mm. long,

acute, glabrous except margins; ovary globose, shortly stiped, 16-

ovulate; style 17.5 mm. long, glabrous, scarcely exceeding stamens;

stigma capitate. — The small leaves are distinctive in the group of

species with pedicellate flowers and long calyces (author). F.M.

Neg. 3251.

Amazonas: Near Chachapoyas, Mathews 3251, type; Raimondi

923.

Cestrum ellipticum Francey, Candollea 6: 216. 1936.

Resembles the allied C. confertum but lax and with cushions

of lanate pubescence in the petiole axils; the much larger leaves

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

(these 1-2 dm. long, about 4-7.5 cm. wide) elliptic-ovate-lanceolate,

long-acute or acuminate, lustrous both sides, with 12 or more lateral

nerves; petioles to 1.5 cm. long; stipuliform leaves ovate-falcate,

at least 1 cm. wide, 1.5-2 cm. long; bracts lanceolate, 3-5 mm.

long; calyx tubular-urceolate, 3.5 mm. long, 1.5 mm. across, gla-

brous except for the marginally puberulent acute teeth; corolla

21.5 mm. long, the long cylindric tube ampliate-clavate toward

apex, 17.5 mm. long, glabrous within as without; stamens 17 mm.

long, often unequal, entirely glabrous. — This species is characterized

by the regularly elliptic leaves, the nerves prominent beneath con-

trasting with the clear green of the leaf blades (author).

Ayacucho: Ccarrapa, 1,200 meters, Killip & Smith 23229. Bolivia.

Cestrum falcatum Francey, Candollea 6: 228. 1936.

Similar to C. conglomeratum but more or less puberulent, especially

the younger parts, and the membranous leaves often plicate-falcate;

petioles 6-8 mm. long, sulcate above where densely puberulent;

stipuliform leaves on upper branches ovate-falcate, acute or acu-

minate, to 2 cm. long, half as wide, glabrous in age; leaves spreading

or pendent, oblong-subovate to lanceolate, frequently unequal,

acutely decurrent into petiole, attenuate and shortly acuminate at

apex, 8-10 cm. long, 2.5- nearly 4 cm. wide, rigid, often plicate-

falcate, the slender nerves 10-12, prominent beneath; flowers sessile,

many in the leaf axils, the rachis 5-8 mm. long, densely tomentose;

bracts lanceolate, acuminate, about 5 mm. long, the bractlets 2 mm.

long, densely ferruginous-tomentose; calyx tubular-poculiform, 3.5-4

mm. long, 1.5-2 mm. thick, subterete, completely glabrous except

the densely ciliate margins of the 5-6 triangular acute teeth, these

0.5 mm. long, 0.6 mm. thick; corolla 19 mm. long, tube contracted

below ovary, finally obconic, little ampliate, 15.5 mm. long, glabrous

or within obscurely puberulent; lobes 3.5 mm. long, nearly as wide,

puberulent marginally; stamens 14.5-15 mm. long, free 3.5-5.5 mm.,

straight, smooth; ovary globose, 6-ovulate, the stout stipe glandular;

style filiform, 1.5 cm. long, glabrous, the stigma capitate. — Resembles

strongly C. conglomeratum but the leaves generally plicate (author) ;

may prove to be a geographical variant but flowers longer and

more slender, stipules present; Soukup 3674 (det. Lyman Smith,

C. conglomeratum) has the pubescence and leaves of S. flexuosum but

the inflorescence of C. conglomeratum R. & P. and the ovate-falcate

stipules of C. falcatum; it lacks the many subparallel nerves of

C. Mathewsii, which otherwise it simulates; apparently new at least to

FLORA OF PERU 101

Peru, it may be noted as C. falcatum Francey var. Soukupii Macbr.,

var. nov., stipulis ovato-falcatis, 5-8 mm. longis, 3-5 mm. latis,

foliis subtus sparse stellulatio-puberulis, corollis 1.5 cm. longis.

Used as a soporific (Herrera).

Lima: Conception, Hacienda Ingenio, Soukup 3674 (type, var.). —

Cuzco: Rio Urubamba, 2,800 meters, (Herrera 734, type). Hacienda

de Urcos, Karl Schmidt (det. Standley, C. conglomeratum). "Nucjan"

(Herrera).

Oestrum flexuosum Francey, Candollea 6: 179. 1936.

Branches flexuose, 1.7-4.5 mm. thick, tomentose-pulverulent to-

ward the foliose tips; petioles slender, 9-12 mm. long; leaves oblong-

lanceolate, slightly decurrent at acute base, long-acuminate, often

unequal, about 6-7.5 cm. long, 2-2.5 cm. wide or somewhat wider,

subcoriaceous, minutely revolute, glabrous, smooth, opaque above,

sparsely puberulent all over beneath, reticulate-nerved, the mid-

nerve as the 7-9 lateral prominent; stipuliform leaves none (or

early deciduous); racemes axillary, peduncles flexuose, 2- nearly

7 cm. long, 5-7-flowered; pedicels 0.5 mm. long, glabrous, the sub-

tending bracts oblong, acute, 13 mm. long, 6 mm. wide; calyx poculi-

form, scarcely costate above, 5 mm. long, 3.5 mm. thick, coriaceous,

glandular but glabrous, 7-nerved, papillose within, the 7 unequal

teeth acute, 0.5-0.8 mm. long, 1 mm. wide, marginally pilosulous;

corolla 22 mm. long, the tube contracted below the ovary, cylindric,

gradually ampliate to 4.5 mm. broad, obscurely constricted below

limb, 19 mm. long, glabrous within and without, the 6 lobes 2.2

mm. long, acute or (explanate) obtuse, glabrous except the lanate

margins; stamens 6, equal, 18 mm. long, the filaments free for

5.5 mm., geniculate, little dentate, slightly pilose; ovary globose,

20-ovulate, the filiform style verruculose-pilosulous at apex; stigma

capitate; fruit black, globose, stipitate, 12 mm. thick, with 9 broadly

oblong little compressed seeds 4-^4.5 mm. long, 2-3 mm. broad,

0.7-1 mm. thick; cotyledons ovate.

Huanuco: Muiia to Tambo de Vaca, 2,000 meters, 4310 type.

Cestrum foliosum Francey, Candollea 6: 260. 1936.

Glabrous or essentially, the branches sparsely verruculose, 5 mm.

thick, the virgate elongate branchlets pulverulent only apically but

exceptionally foliose; petioles 2.5-4 mm. long, little enlarged basally,

scarcely canaliculate; stipuliform leaves persisting, geminate, acute,

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

2.5-6 mm. long, 2-3 mm. wide; leaves widely spreading (internodes

only 6-12 mm. long), narrowly ovate-lanceolate, rounded or obtuse

at base where little decurrent, long-acuminate, often 3.5-5 cm. long,

1-1.5 cm. wide, membranous, concolor, nitidulous, reticulate-nerved,

the 7-8 arcuate lateral nerves prominent both sides; racemes axillary,

leafy, 2-3 (-6) -flowered, the rachis 3-16 mm. long, obscurely puber-

ulent, the floral leaves subsessile, ovate-lanceolate, 6-11 cm. long,

1-3.5 cm. wide, the bracts filiform, 1-1.5 mm. long; peduncle 1-1.5

mm. long, apically bracteate; pedicels 1-2 mm. long, glabrous or

obscurely glandular; calyx cyathiform, 2.5 mm. long, 2 mm. across,

glandular pilose only above, glabrous within, the 5 acute teeth

0.3 mm. long, 0.8 mm. wide, the nerves slender; corolla to 27.5 mm.

long, tube below ovary suddenly, above gradually, contracted, long-

cylindric, gradually ampliate three-fourths of its length, clavate at

apex, 3.3 mm. broad, constricted below the limb, 23.5 mm. long,

purplish without, greenish within, quite glabrous; lobes of limb

3 mm. long, glabrous without, glandular within, margins minutely

puberulent; stamens unequal, about 23 mm. long, the filaments

free for 2 mm., glabrous, smooth; ovary ellipsoid, shortly stiped,

6-ovulate; style 24.5 mm. long, very minutely puberulent toward

the capitate stigma. — The flowers resemble those of C. sparsiflorum

Britton, C. papyraceum Rusby and C. rigidum Rusby of Bolivia,

the first with acuminate calyx teeth, stipuliform leaves none (appar-

ently), the second with extremely thin leaves, filaments free for

1 mm., lateral nerves 9-10, and the third with filaments free for

3 mm., lateral nerves 9-13; stipuliform leaves persist in the latter

two; here, as elsewhere in this genus where so many similar plants

have been proposed as species, only much more observation will

determine if the relatively slight variations upon which some have

been based are constant and taxonomically significant. The author

describes the leaves as membranous but also as rigid; in the her-

barium they seem to be chartaceous; a specimen by Stork and

Horton (10153} was referred to C. peruvianum Willd. by Standley,

with query but probably sens. lat. correctly; in some herbaria it

has been referred here but it seemed to me to be nearer C. Raimond-

ianum; possibly more than one species was distributed under this

number.

Huanuco: Villcabamba, Rio Chinchao, 5203. Bolivia.

Cestrum Franceyi Macbr., nom. nov. Sessea pedicellata Fran-

cey, Notizbl. Bot. Gart. Berlin 11: 882. 1933, not Sendt., 986.

FLORA OF PERU 103

Branches slender, early trigonous and densely pulverulent-

puberulent, soon smooth and glabrate; petioles 12-25 mm. long,

narrowly canaliculate, not enlarged at base, puberulent; adult leaves

oblong- or elliptic-lanceolate, unequally acute or subacute at base,

shortly and acutely attenuate at apex, 1.5 dm. long or longer,

about 3.5-5 cm. wide (the narrower younger rounded at base, long-

acuminate), subcoriaceous, minutely revolute, somewhat lustrous

olive-green above, minutely stellate pilose beneath, reticulate both

sides, the 11-13 lateral nerves nearly straight; stipuliform leaves

none; lateral branches of panicles elongate, strongly dichotomous

apically; flowers pale yellow, 21.5 mm. long, pedicels 1-1.5 mm.

long, hirsute-pilose as calyx, this 5 mm. long, glabrous within, with

5 semiorbicular crenations 0.5 mm. long; corolla densely stellate-

pilose, very acute lobes 2 mm. long; stamens unequal, filaments

villous at medial insertion. — Type a 2-meter tall shrub; ovary densely

pilose. Capsule 7 mm. long, 2.5 mm. thick, elongate, parted into

4 valves (Francey); Vargas specimen has shorter (but perhaps not

fully developed) corollas and the leaves are pubescent as those of

S. discolor.

Cuzco: Near Lares, 2,900 meters, Prov. Calca, Weberbauer 7895,

type; Vargas 3604?

Cestrum glaucophyllum Francey, Candollea 6: 149. 1936.

Tree more or less stellulate-tomentulose, the trichomes often fer-

ruginous and extending to the calyces without where more villous in

character, the branches 3-6 mm. thick, the branchlets 1-2 mm., vir-

gate; petioles 3-10 mm. long; leaves erect, narrowly ovate- or oblong-

lanceolate, rounded or subacute at base but not decurrent, narrowly

attenuate-acuminate, mostly 7-16 cm. long, 2-5 cm. wide, membra-

nous, subrevolute, green and lustrous above, the midnerve slightly

puberulent, obscurely reticulate, scarcely paler beneath but the 5-8

lateral nerves and the reticulate veins there prominent; stipuliform

leaves none; spikes axillary, (1) 2-3-flowered, the filiform flexuose

rachis 2.5-6 cm. long; peduncles short; bracts linear, petiolate, about

1 cm. long, marginally pilose; calyx lax, tubular-poculiform, to 14.5

mm. long (!), 4 mm. thick, slightly obconic below, glabrous within,

4-5-dentate, 5-nerved; teeth triangular-lanceolate, acuminate, 4.5

mm. long, often connate; corolla pale green, nearly 3.5 cm. long, the

tube below the ovary manifestly, above scarcely, contracted, long-

cylindric, abruptly ampliate at apex, 4 mm. broad, constricted below

the limb, 2.5 cm. long, glabrous without and within, segments 8.5 mm.

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

long, incurved, narrowly lanceolate, acute or explanate, elliptic-lan-

ceolate, obtuse, papillose-glandular without, glabrous within, or mar-

ginally puberulent; stamens 24.5 mm. long, the filaments free only

1.5 mm., smooth, glabrous; ovary globose, shortly stiped, 12-ovulate;

style 2.5 cm. long, filiform, densely pilosulous at apex; stigma discoid;

fruit oblong, sessile, 1.5 cm. long, 7.5 mm. thick, the 11 mature seeds

oblong-angled acute, 4.5-5.5 mm. long, 0.7-1.2 mm. thick; cotyledons

elliptic. — Suggests in character of calyx, leaves and indument the

species C. calycinum HBK., C. strigillatum R. & P. and C. cancellatum

Dunal; it differs from them essentially by the glabrous corolla tube

(author). The fruit is tardily valvate at apex; thus toward section

Sessea. Illustrated, Francey, I.e. 7: pi. 1, fig. 31+ (flower).

Junin: Pichis Trail, in dense forest, Killip & Smith 257J.O, type.

Cestrum Humboldtii Francey, Candollea 6: 393. 1936.

Related to C. petiolare; stipuliform leaves ovate, 4.5-7 mm. long,

3 or 4 mm. wide; leaves erect to spreading, ovate- or oblong-lanceo-

late, shortly acute or acuminate, 13-17 cm. long, about 7.5-8 cm.

wide, subcoriaceous, not revolute, more or less densely puberulent on

the nerves above, stellate-pilose, especially the nerves, beneath, the

lateral nerves 13 or 14; panicles axillary, rather long-peduncled, ra-

chis branched at base, 1.5-4.5 cm. long, bracts filiform, 4 or 5 mm.

long, pilose; calyx tubular-poculiform, 4 or 5 mm. long, 2.5 mm.

thick, densely pilose without, papillose within, the 5 teeth 0.8 mm.

long and broad; corolla 16.5 mm. long, the tube purple, distinctly

contracted only below the ovary (typically), scarcely ampliate,

3.7 mm. broad, 13 mm. long, glabrous without, the lobes triangular-

lanceolate, 3 mm. long, glabrous or nearly except the margins gland-

ular without; stamens 12.5 mm. long, filaments free for 5 mm.,

slightly geniculate 2 mm. above the insertion, reflexed pilose, not

denticulate; fruit black, 4.5 mm. thick, 3 rugulose seeds 2.5-3 mm.

long, 1.5-2 mm. broad, at least 1 mm. thick. — Var. calycinum

Francey, 394, has oblong leaves, acute at base, petioles to 6.5 cm.

long, calyx 6 mm. long, the often connate teeth 1.5 mm. long, corolla

tube slightly clavate above, constricted below the limb, the lobes

3.5 mm. long, the stamens adnate three-fifths length of tube. With

C. petiolare, this species is allied to Central American species C. calo-

neurum Pittier and C. calycosum Pittier (author). C. Dielsii Werd.,

Notizbl. Bot. Gart. Berlin 12: 373. 1934, from adjacent Ecuador (Rio

Pastaza valley), while of this alliance (author) has a short inflores-

cence.

FLORA OF PERU 105

Huanuco: Pampayacu, Rio Chinchao, 51 29, type. Mufia, trail

to Tambo de Vaca, 2,400 meters, 4332 (type, var.).

Cestrum laevifolium Francey, Candollea 7: 62. 1936.

Type a small slender glabrous (unless floral parts) tree about

4.5 meters tall with elongate branches and narrowly oblong-lanceo-

late attenuately acuminate leaves about 1-1.5 dm. long, 2.5-5 cm.

wide, decurrent on petioles 10-12 mm. long, subopaque above, lus-

trous beneath with 8 or 9 arcuate-ascending lateral nerves; stipules

none; racemes axillary, the 6-8 subsessile whitish-green flowers 2.25

cm. long; bracts many, filiform, 1-1.5 mm. long; calyx tubular-

campanulate, scarcely costate, 3.8 mm. long, papillose within, the 5

unequal teeth 0.6-0.9 mm. long, sparsely pilosulous; corolla pale

green, infundibuliform, barbate within at base and at filament in-

sertion, the lobes 3.5 mm. long, glabrous except the edges; filaments

straight, 4 mm. free; ovary 6-ovulate; stigma capitate, included. —

Extremity of the leaf is characteristic (author).

Junin: San Nicolas, Pichis Trail, Killip & Smith 26074, type.

Cestrum lanuginosum R. & P. Fl. Peruv. 2: 30, pi. 157. 1799;

168.

Erect shrub, known to attain about 2 meters, conspicuously lanu-

ginose or lanate-hirsute except the older leaves above and the corollas,

the latter quite glabrous within and without; branches lax, 1.5-3 mm.

thick, the leafy axillary flowering branchlets 4-9 cm. long, 1-1.5

mm. thick; petioles densely lanate-hirsute, 5-7 mm. long, com-

pressed; stipuliform leaves often obliquely ovate, about 2 cm. long,

half as wide; mature leaves spreading or pendent, cordate-ovate or

incised at base, scarcely decurrent, acute or subobtusely acuminate,

about 6.5-10 cm. long, 4.5-7 cm. wide, membranous, slightly if at all

revolute, early puberulent above, finally glabrate, lustrous, the 8 to 9

lateral nerves and reticulate veins rather prominent beneath; flowers

5-8-fasciculate at branchlet tips and in the axils of foliaceous acumi-

nate bracts, 12-16 mm. long, 4-6 mm. wide, the filiform bractlets

1.5-3 mm. long, densely stellate-lanate; flowers sessile, the obconic-

cyathiform calyx about 6 mm. long, 2.5 mm. thick, glabrous within,

the 4 or 5 broadly ovate acute teeth 1.2 mm. long and wide, the

nerves slender; corolla 24 mm. long, yellowish-green, the tube below

the ovary suddenly, above gradually, contracted, in age obconic,

scarcely constricted below the limb, 21 mm. long; lobes 2.5 mm.

long, ovate, puberulent without, glabrous within, marginally tomen-

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

tose; stamens 4 or 5, 21 mm. long, the filaments free for 7 mm., this

portion basally geniculate, often shortly dentate above, quite gla-

brous; anthers subcordate; ovary globose, glabrous, 10-ovulate, style

capillaceous, 2.5 cm. long, minutely puberulent at tip; stigma capi-

tate; fruit violet-colored, ellipsoid, sessile, 8 mm. long, 5.5 mm.

thick, maturing 6 rugulose somewhat arcuate compressed oblong

often truncate seeds 4.5-5.5 mm. long, 2.5-3 mm. broad, 1-1.5 mm.

thick; cotyledons rotund. — Illustrated, Francey, Candollea 7: pi. 1,

fig. 50; fig. 59 (flower); fig. 61 (calyx). F.M. Neg. 2984.

Huanuco: Near Pozuzo, Ruiz & Pavdn, type. — Arequipa: Atiqui-

pa, (Raimondi 1312}; Ruiz & Pavdn (Tafalla), type.

Cestrum longiflorum R. & P. Fl. Peruv. 2: 28, pi. 154* 1799;

235.

Known to attain 8 meters, the younger parts mostly lightly puber-

ulent, as the leaves glabrous in age, the erect branches to 5 mm.

thick; petioles at least 1 cm. long, arcuate or sometimes uncinate at

base where enlarged; stipuliform leaves none; leaves spreading, ovate-

oblong, subacute or rounded at base, acuminate, often 1- nearly

2 dm. long, nearly 3-7 dm. wide, subcoriaceous but rigid, very lus-

trous above, reticulate-veined both sides, the 18-20 lateral nerves

impressed above, prominent beneath; panicles terminal, lax, 5.5-

12 cm. long, the pulverulent rachis 2.5-5.5 cm. long, dark purple,

peduncles branched, slightly lanuginose, 1.5 to many cm. long, the

numerous flowers sessile or shortly pedicellate; bracts lanceolate-

ciliate, 12 mm. long; calyx poculiform-tubular, basally obconic, 6-

7 mm. long, 4 mm. thick, coriaceous, early obscurely glandular and

very sparsely puberulent without or often essentially glabrous, pu-

bescent within, the 5 nerves obscure, the 5 subobtuse teeth only

0.75 mm. wide; corolla nearly 4-5.6 cm. long, light green, the tube

basally contracted, finally long-cylindric, apically ampliate, arcuate,

3-4.5 cm. long, 2-3 mm. across, quite glabrous; lobes of limb 9 mm.

long, 3-4 mm. wide, acute, glabrous except the lanate margins; sta-

mens equal, the filaments adnate to apex of tube, obscurely pubes-

cent, the free part 2 mm. long, smooth; anthers subelliptic; ovary

multi-ovulate, the disk glandular; style filiform, 4-5.7 cm. long, hir-

sute above; stigma capitate, puberulent; fruit ovoid, 2-celled, pur-

plish, subsessile, 12-16 mm. long, 6-8 mm. thick, the subtending

calyx cupuliform; seeds about 50, 2-3 mm. long, 1.5-2.5 mm. broad,

0.5-1 mm. thick, angled; cotyledons ovate. — F.M. Neg. 18393.

FLORA OF PERU 107

Huanuco: Chinchao, Macora and Cuchero, Ruiz & Pawn, type.

Rio Pozuzo, at the Rio Palcazu, 1,800 meters, Weberbauer 6779. Rio

Hualyaco Canyon, below Rio Santo Domingo, 4257. — Junin: Palca,

Weberbauer 11995. Huacapistana, 5820; Killip & Smith 24101.

"Yerba hedionda."

Cestrum loretense Francey, Candollea 6: 225. 1936.

A small shrub, glabrous except for some pubescence in the ex-

tremely short 5-8-flowered axillary spike; branches 3.5^4.5 mm.

thick, conspicuously verruculose, the erect branchlets broadly sul-

cate, 2-3 mm. thick; petioles stout, 4-6 mm. long, stipuliform leaves

none; leaves ascending, often curved toward apex, elliptic-lanceolate,

attenuate to acute base but not decurrent, long-acuminate, often

1-2 dm. long, 4-7 cm. wide, coriaceous, not revolute, not or slightly

lustrous both sides, the 8-10 lateral nerves impressed above, very

prominent as the reticulate venation beneath; rachis of spikes to

7 mm. long, reddish-tomentose, the lanceolate acuminate bracts 3-

7 mm. long, obscurely puberulent, the flowers sessile; calyx poculi-

form, angled, strongly costate, 4.5 mm. long, 2.5 mm. across, scarcely

pilose without, glabrous within, the 5 triangular acuminate teeth

1 mm. long, puberulent marginally; corolla 2.5 cm. long, tube con-

tracted below ovary, finally cylindric, gradually ampliate to the cla-

vate apex, constricted below limb, 18.5 mm. long, glabrous without,

barbate within on or near adnate portion of filaments, these free for

1.5-3.5 mm., straight, smooth, the acuminate lobes glabrous except

the margins; ovary globose, 7-ovulate, the filiform style densely

puberulent apically, 18 mm. long; stigma capitate; fruit shortly

stiped, oblong, 8 mm. long, 5.5 mm. thick, maturing 4-angled costate

compressed acute seeds 4-5.5 mm. long, 2.5-3 mm. broad, 1.5-2 mm.

thick. — Marked by large leaves and conspicuously nerved calyx.

Loreto: La Victoria, Williams 3129, type.

Cestrum Macbridei Francey, Candollea 6: 262. 1936.

Type a small shrub 1.5 meters high, quite glabrous (but in fruit),

the branches stout, 5 or 6 mm. thick, the elongate branchlets densely

foliose; petioles 4-6 mm. long, canaliculate above, slightly incrassate

at base; stipuliform leaves geminate, persisting, sessile, ovate-falcate,

acute, 2.5-15 mm. long, 1-8 mm. wide; internodes 9-15 mm. long;

leaves erect-spreading or widely spreading, narrowly lanceolate,

acutely decurrent at base, narrowly long-acuminate or cuspidate,

mostly 6-14 cm. long, 1-1.5 cm. wide, coriaceous, nearly concolor,

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

nitidulous, finely reticulate-nervose, the 13-14 arcuate ascending

lateral nerves rather obscure above, obvious and flavescent beneath;

spikes axillary, 4-6-flowered, the stout rachis to 13 mm. long; pedi-

cels 1-2 mm. long, 2-3 mm. thick; fruit in calyx cupuliform, 2 mm.

long, 5 mm. across, glandular without, the 5 nerves thick but not

conspicuous, the 5 teeth triangular, acute, 1 mm. long, 2.5 mm. wide,

margins obscurely puberulent; fruit black, sessile, ellipsoid, 8.5 mm.

long, 6 mm. thick, the 4 immature seeds pale green. — Resembles

C. foliosum Francey in stipules, position of leaves and branches but

differs basically in form and dimension of leaves; also related to

C. rigidum Rusby and C. papyraceum Rusby (author); some fruits

have opened shortly at tip.

Junin: Hacienda Schunke near La Merced, 1,200 meters, 5724-,

type.

Cestrum maleolens Macbr., nom. nov. Sessea stipulata R. & P.

Syst. Veg. 1: 44. 1789 not C. stipulatum Veil.; 200.

Branchlets toward tip, leaves especially beneath and much-

branched subcorymbose panicles more or less floccose-tomentose

with candelabra-branched trichomes, this indument more or less re-

duced, extending to the calyx and ovary at apex; petioles often 2-

2.5 cm. long; leaves membranous, lanceolate or elliptic-lanceolate,

mostly obliquely rounded or more or less cordate, acutely or sub-

obtusely acuminate, the larger often about 1 (-1.5) dm. long (grad-

ually reduced upward), about a third as wide and commonly with

1 (or 2) sessile or petioled eventually deciduous stipuliform leaves

opposite them, the persisting leaves sordid-green or drying brownish-

rufescent above with 11 to 16 curved ascending prominent (some-

what impressed above) lateral nerves; inflorescences 1.5-2.5 dm. long;

calyx narrowly campanulate- tubular, 7.5-8 mm. long, the acute or

acuminate lanceolate teeth 1-1.5 mm. long; corolla 15-17 mm. long,

glabrous only near base, the densely tomentose lobes 2-2.5 mm. long;

filaments 6 mm. long with a few slender trichomes toward base

(Bitter) ; ovary densely pseudostellate-tomentose at the bilobed apex;

stigma obscurely bilobed; capsule exserted, 1 cm. long, glabrous be-

low, 4-valved, the usually 4 alate seeds about 8 mm. long. — Miers

described the filaments as retrorsely pilose. Sometimes a small tree

5 meters tall with a trunk diameter of 1.5 dm. near base; the flowers

as well as the foliage have an extremely disagreeable odor; the cap-

sule, black when mature, dries brown in herbaria. Illustrated, Miers,

111. South Amer. Bot. pi. 15. F.M. Neg. 3006.

FLORA OF PERU 109

Huanuco : Acomayo, Woytkowski 34263 (det. Cuatrecasas) . Ron-

dos, Ruiz & Pavdn. — Junin: Huasahausi, Ruiz & Pav6n, type.

Bolivia.

Cestrum Mancoi Macbr., nom. nov. Sessea Weberbaueri Bitter,

Repert. Sp. Nov. 18: 211. 1922, not C. Weberbaueri Francey, 1936.

Type a shrub 3 meters tall with subvirgate branches, rather sim-

ilar especially in indument to S. stipulate, R. & P. but the olive green

leaves glabrate in age above, little paler beneath and stipule-like

leaves lacking or caducous and calyx teeth obsolete; petioles about

1.5-2 cm. long; lower leaves at most 1 dm. long, 4 cm. wide, all

mostly rounded and broader at base, gradually attenuate to obtuse

or subobtuse tip, firm-membranous, the veins reticulate, impressed

above; calyx tubular, (7) 8-9 mm. long, truncate; corolla sulphur

yellow, about 2 cm. long, tube 5.5 mm. across at top, the narrowly

triangular teeth 3 mm. long; filaments glabrous, scarcely 3 mm. long;

stigma obtusely capitate. — F.M. Neg. 3007.

In Cestrum the name of this shrub may recall Manco Inca — not,

unless by historical association, the legendary Manco Capac (cf.

Prescott's "Conquest of Peru"). Manco Inca was the last of his race

animated by the heroic spirit of the ancient Incas (Prescott) . Now

I have recorded in this work, at least to my own satisfaction, several

of the better-known Inca personalities at about the time of Pizarro's

conquest (cf. Field Mus. Bot. 11: 13, 15, 23. 1931).

Apurimac: Among evergreen shrubs on grass-steppes, 2,900 me-

ters, Andahuaylas to Huancarama, Weberbauer 5913, type; Velarde

Nunez 1388.

Cestrum Mathewsii Dunal in DC. Prodr. 13, pt. 1: 637. 1852;

243.

Branches stout (to 8 mm. thick), broadly sulcate, brown-reddish,

glandular with paler elliptic glands; petioles enlarged at base, scarcely

canaliculate above, glabrous, 13-16 mm. long; stipuliform leaves au-

riculate-ovate or oblong, subobtuse, 5 or 6 mm. long and nearly 3 mm.

wide; leaves widely spreading, oblong-lanceolate, long-attenuate to

the often unequal base, not decurrent, long-acuminate and subcus-

pidate, about 13-16.5 cm. long, 2.5-3.5 cm. wide, subcoriaceous,

rigid, subrevolute, lustrous especially beneath, glabrous both sides,

the 18-21 lateral nerves parallel, little impressed above, prominent

beneath; racemes axillary, subspicate, about 8-flowered, geminate or

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

ternate, sometimes solitary, the 7-11 mm. long rachis tomentulose;

bracts many, linear, acuminate, strongly revolute, 4-6 mm. long;

pedicels 0.5-1 mm. long, glabrous as the tubular-poculiform calyx

without, this pilose within only above, 3-3.5 mm. long, 1.5 mm.

thick, coriaceous, the 5 unequal teeth subacute, 0.7-1 mm. long,

marginally densely pilose, the nerves thickened toward apex; corolla

14-18.5 mm. long, the tube scarcely contracted below ovary, grad-

ually inf undibulif orm toward the middle, clavate to apex, constricted

below the limb, 16 mm. long, completely glabrous; lobes of the limb

ovate, 2.5 mm. long, glabrous without, obscurely puberulent within;

stamens subequal, 14.5-15 mm. long, the filaments adnate (three-

fifths) four-fifths their length, glabrous, the free part not tumid,

smooth; anthers orbicular; ovary 4-ovulate, disk glandular; style

15.5 mm. long, densely but minutely puberulent above; stigma capi-

tate.— The great number of primary nerves is characteristic (Francey).

Simulates C. conglomeration R. & P. but rachis, while short, is some-

what developed and leaves are entirely different; the corolla in type

collection, Washington, as in the Conchan shrub is usually 15-17 mm'

long, the filaments of the latter free for 5 mm. F.M. Neg. 8576.

Cajamarca: South of Conchan, Prov. Chota, 2,500 meters, Stork

6 Horton 10068. — Amazonas: Chachapoyas, Mathews, type, Geneva

Herbarium.

Sessea multiflora Bitter, Repert. Sp. Nov. 18: 212. 1922, not

Oestrum multiflorum R. & S.

Related and similar to C. Mancoi; petioles of larger leaves 2.5-

3 cm. long; leaves very sparsely puberulent above, oblong-lanceolate,

the lower about 1.5 dm. long, 5- nearly 6 cm. wide, often with 1-2

sessile obtuse leaves about 1 cm. long and wide in the axils; lateral

nerves 14-16, the veins scarcely articulate-impressed; panicles am-

ple, to 2.5 dm. long; calyx truncate, tubular-campanulate, 5-5.5 mm.

long; corolla tube 4-5 mm. broad at apex, lobes ovate, 3.5 mm. long;

filaments 7 mm. long, with a few branched and simple trichomes

toward base; stigma obliquely capitate; capsule obovoid-ellipsoid,

7 mm. long, 3 mm. thick, the 4 coriaceous valves glabrous below for

5 mm. — S. Herzogii Damm., Meded. Herb. Linden 29: 1916, of Bo-

livia, has densely flowered corymbs about 5 cm. long, the stellate

pilose leaves obviously reticulate-veined above. Type near a house

where perhaps planted or cultivated (Weberbauer) . My collections

were distributed by me as S. stipulate, R. & P.; they seem to be inter-

mediate to S. multiflora and S. discolor Francey; like the latter, if

FLORA OF PERU 111

valid, it requires in Oestrum a new name. From a tree 3-5 meters tall.

F.M. Neg. 3005.

Huanuco: Mito, 1^86. Yanahuanca, 1235. — Puno: Sandia, 2,400

meters, Weberbauer 599, type.

Oestrum nemanthum Dunal in DC. Prodr. 13, pt. 1: 637. 1852;

306.

Glabrous, or noticeably puberulent only in the very short axillary

spikes, the branches curved, 3-4 mm. thick, the branchlets spreading

or pendent, foliose to apex, floriferous entire length; stipuliform

leaves none; petioles 6-10 mm. long, canaliculate above, incrassate

at base; leaves erect or pendent, deciduous from the branches, ob-

long- or elliptic-lanceolate, narrowed to base and decurrent, subob-

tusely acuminate, 9-15.5 cm. long, 2.5-7 cm. wide, subcoriaceous or

chartaceous, plane, concolor, rather lustrous, the 6 to 7 lateral nerves

only moderately conspicuous either surface; spikes 2-5-flowered, the

pulverulent rachis 1-5 mm. long, the bracts hardly 1 mm. long;

flowers subsessile; calyx tubular above a cyathiform base, annulate,

3 mm. long, 1.7 mm. thick, obscurely puberulent without, glabrous

within, the 5 nerves stout, the 5 acute teeth 0.5 mm. long, 0.7 mm.

wide, apically pilose; corolla 15.5 mm. long, tube contracted above

and below ovary, finally infundibuliform, slightly ampliate to apex,

constricted below limb, 12 mm. long, entirely glabrous; lobes 3.5 mm.

long, tomentulose without, puberulent within, stamens 10 or 11

mm. long, the filaments adnate only to middle of tube, free for 4 or

5 mm., smooth, at middle and base of tube pilose; ovary globose,

obconically stiped; style 11 mm. long, incrassate toward tip, glabrous;

stigma capitate. — This is distinguished from C. Baenitzii Lingels. by

the form of the leaves; these are unique because all the nerves, while

visible, are rather obscure, the leaves thus having a soft smooth

appearance on both surfaces (Francey) ; type, fide Dunal, by Glaus-

sen! So the citation must be a lapsus calami by the author or by

Francey; compare C. silvaticum Francey or perhaps C. loretense.

Huanuco: Chicoplaya, (Ruiz & Pavdn, type, Geneva, fide Francey).

Brazil.

Oestrum obscurum Francey, Candollea 6: 302. 1936.

Glabrous or essentially except the short axillary racemes, these

1.5-2 cm. long, 3-5-flowered, often geminate or ternate; branches

flexuose, 2-3 mm. thick, angulate and somewhat sulcate toward the

dark purple tips; stipuliform leaves none; petioles canaliculate above,

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

7-12 mm. long; leaves spreading, oblong or elliptic, acute or nar-

rowed and decurrent at base, shortly acuminate or very acute, many

9-12 cm. long, about 3-6 cm. wide, membranous but rigid, undulate,

often bullate, subopaque above, pale and opaque beneath, the mid-

nerve there prominent and dark brown as the 9 or 10 arcuate lateral

nerves; rachis about 7 mm. long, filiform, glabrous; peduncles only

0.5-2 mm. long, the bractlets scarcely 1 mm. long, puberulent;

flowers sessile; calyx tubular, 2.5 mm. long, 1.5 mm. thick, quite

glabrous including the 5 ovate acute teeth, these 0.7 mm. long, 1 mm.

wide; corolla green, 15.5 mm. long, the tube not constricted, scarcely

ampliate at apex, 13 mm. long, entirely glabrous; lobes 2-2.5 mm.

long, acuminate or very acute, puberulent only without, the margins

there tomentulose; stamens 12.5 mm. long, free for 2.5 mm., smooth,

long-pilose for three-fifths their length; ovary subquadrate; style

12 mm. long, glabrous, the stigma capitate. — The deep purple of the

branches, the flowering peduncles, the abruptly decurrent leaves on

the petiole and the slender flowers are the unique characters of this

species (author). Illustrated, Francey, I.e. 7: pi. 2, fig, 54 (flower);

pL 3, fig. 52 (calyx).

Junin: La Merced, (Weberbauer 1930, type).

Cestrum osnoense Macbr., nom. nov. Sessea graciliflora Bitter,

Repert. Sp. Nov. 18: 214. 1922, not C. graciflorum Francey.

In indument and many other characters resembles S. multiflora

Bitter; branches suberect, densely yellowish floccose-tomentose, as

the leaves beneath; petioles 2.5-3.7 cm. long; leaves elliptic or ovate-

elliptic, the larger 1-1.5 dm. long to 6 cm. wide, veins impressed

above; calyx tubular, truncate, 3-3.5 mm. long; corolla narrowly

tubular, 22 mm. long, only 3 mm. across at apex, the rhomboid-

triangular lobes 3 mm. long; filaments 5-6 mm. long, rather densely

and simply pubescent; ovary densely pubescent all over; stigma

obliquely capitate. — A shrub, 3 meters high, or probably more or

less, with divaricately much-branched inflorescence. F.M. Neg. 3004.

Ayacucho: Between Tambo above Osno and the Rio Apurimac,

3,200 meters, Weberbauer 5582, type. — Cuzco: Paucartambo, Soukup

396 (det. Standley, S. discolor}.

Cestrum ovalifolium Francey, Candollea 6: 362. 1936.

Generally resembles C. Dunalii; flowering branchlets ordinarily

5 cm. long; leaves widely divaricate, ovate or subelliptic, rounded or

obtuse at base but shortly decurrent and notably shortly acuminate,

FLORA OF PERU 113

3-5.5 cm. long, 1.5-3 cm. wide, membranous but very rigid and frag-

ile, the lateral nerves 5-7, the reticulation lax beneath; racemes axil-

lary and at the apex of terminal branches 5- or 6-flowered, the rachis

1.5 to scarcely 3 cm. long; pedicels 1.5-2.2 mm. long, the lower 1-flow-

ered; peduncles 1 cm. long, the upper 2-3 mm. long; bracts filiform,

acuminate, to 2.5 mm. long; calyx tubular-campanulate, terete,

4 mm. long, 2 mm. thick, the 5 equal teeth marginally puberulent,

the nerves not prominent; corolla 24 mm. long, ampliate three-

fourths the length, above clavate, 3 mm. across, constricted below

the limb, 16.5 mm. long, the lobes 7 mm. long, acutely acuminate;

stamens free for 3.5 mm., denticulate at the insertion, quite glabrous

as the tube all over; style 16 mm. long. — The species is characterized

by the regularly oval or less generally elliptic leaves with a short

apical point (author). Illustrated, Francey, I.e. 7: pi. 2, fig. 32

(flower).

Cajamarca: Cutervo, Jelski 6549, type.

Oestrum ovatolanceolatum Francey, Candollea 6: 279. 1936.

Type a tree 12 meters high, glabrous except the lax many-flowered

axillary panicles, these 2.5-3 cm. long or longer; branches 3.5-4 mm.

thick, the flexuose brownish-purple branchlets broadly sulcate; inter-

nodes mostly 2-3 cm. long; petioles 8-11 mm. long, canaliculate

above; leaves alternate, widely divaricate, ovate-lanceolate, obtuse

or rounded at the scarcely decurrent base, long-attenuate to the

shortly acuminate or very acute apex, broadest at one-fourth their

length, mostly 9-11 cm. long, 3.5-5 cm. wide, membranous, ob-

scurely puberulent above, subopaque and entirely glabrous beneath,

the prominent midnerve there brownish-purple, the 10-13 lateral

nerves obvious as the reticulate venation; stipuliform leaves none;

basal branches of the panicles 3-7 mm. long, 2-4-flowered, the lan-

ceolate puberulent bracts 5 or 6 mm. long, 1-1.5 mm. wide; flowers

sessile, pale, greenish-yellow; peduncles often filiform, the bractlets

acuminate, subfiliform; calyx obconic-poculiform, subterete, 2 mm.

long, 1-1.5 mm. thick, puberulent only without as the 5 often con-

nate acute teeth, these 0.3 mm. long, 0.5 mm. wide, the nerves slen-

der; corolla 10.5 mm. long, tube contracted below ovary, ampliate

to apex, not constricted below limb, 8.5 mm. long, 2.5 mm. broad,

quite glabrous as the acute 2 mm. long lobes except the puberulent

margins; stamens 8.5 mm. long, the filaments free for only 0.5 mm.

long in type, smooth, slightly pilosulous; ovary globose above, cylin-

dric below, 6-ovulate; style 8.5 mm. long, apically pulverulent; stigma

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

capitate. — The form of the leaves, the reduced dimensions of the

flower, and the form of the corolla tube characterize this species

(author). The name was originally written with a hyphen. Klug

specimen with somewhat longer corolla, filaments free for several

mm. is doubtful; it may be rather a variant of C. racemosum R. & P.

Loreto: Balsapuerto, Klug 2942? — Junin: La Merced, at about

600 meters, 5435, type.

Cestrum peruvianum Willd. ex Roem. & Schult. Syst. 4: 807.

1819; 7: 3 (species no. 208). C. affine HBK. Nov. Gen. & Sp. 3: 60.

1818, not Persoon.

Foetid nearly glabrous shrub or small tree, only the triangular

acute calyx and corolla lobes marginally puberulent, the latter all

over without; petioles 5-8 (-12) mm. long, gibbous-enlarged at base;

leaves spreading, elliptic- or oblong-lanceolate, little decurrent at the

acute or subobtuse base, acuminate, 7-17.5 cm. long, 1.5-4.5 cm.

wide, membranous, revolute, lustrous, brownish or somewhat pur-

plish-green, the 14-15 lateral nerves spreading, little prominent

above, the veins reticulate; stipuliform leaves oblong-lanceolate,

3-5 mm. long; spikes axillary, the 6-9 subsessile flowers crowded

apically, 21.5 mm. long; bracts elliptic-lanceolate, 3-5 mm. long;

calyx tubular-poculiform, 5-6-costate-dentate, 4.5-5 mm. long, 2 mm.

thick, the acute teeth 0.5 mm. long; corolla whitish-green or sulphur-

colored, 19 mm. long, tube strongly constricted, pilose within below

filaments, these 6-8 mm. free, more or less dentate; lobes 5 or 6,

glabrous within, 3-3.5 mm. long; stamens 5 or 6, subequal; ovary

long-stiped, 14-1 6-0 vulate; stigma discoid, included. — Known to

attain 4 meters. F.M. Neg. 2964.

Piura: Ayavaca, Bonpland, type. Palambla, Soukup 4287.—

Cajamarca: Chota, Jelski 5477. Nancho, Raimondi. — Amazonas:

East of Chachapoyas, Weberbauer 4442. — Ancash: Llata, 2294.—

Junin: Near Chacapalca, Raimondi. To Colombia. "Yerba Santa"

(Raimondi).

Cestrum petiolare HBK. Nov. Gen. & Sp. 3: 58. 1818; 391.

C. venosum Willd. ex Roem. & Schult. Syst. 4: 807. 1819(7).

More or less densely floccose-tomentose, becoming pulverulent in

age or the leaves above even glabrous; branches stout, angled, 3.5-

5 mm. thick; stipuliform leaves more or less lunulate, 1-1.5 cm. long,

about half as wide, glabrous; petioles 1.5 cm. to nearly 4 cm. long, an-

gled, canaliculate; leaves erect, ovate- or elliptic-lanceolate, acute,

FLORA OF PERU 115

cuneate or rounded at base, shortly acuminate, 1-2 dm. long, 4-9 cm.

wide, or even larger, membranous but rigid, minutely revolute, rather

lustrous both sides but much paler beneath, the midnerve as the

20-24 parallel lateral nerves (Francey) prominent beneath, laxly

reticulate; panicles in axils and terminal, the rachis 1-9 cm. long, the

peduncles 1-5 mm. long; flowers sessile, congested; calyx tubular-

campanulate, 6-6.5 mm. long, 3 mm. thick, glabrous without, pubes-

cent within, the 5 teeth equal, acute, 1.5 mm. long, the nerves stout,

lateral nerves many; corolla white or yellowish-green, 18 mm. long,

the tube not contracted, regularly infundibuliform, the limb contin-

uous, 15.5 mm. long, entirely glabrous as the semi-orbicular lobes

without, these 2.5 mm. long, acute or subobtuse (explanate), puberu-

lent marginally and within; stamens 13.5 mm. long, subequal, the

filaments adnate to the middle, free for 6.5 mm., reflexed pilose, re-

curved, not incrassate, smooth; ovary papillose, clearly stipitate,

8-ovulate, the style 1.5 cm. long, enlarged apically, puberulent, the

stigma capitate, discoid; fruit globose, obtuse, 2 mm. long and thick;

seeds 2, 2.5 mm. long, 1 mm. broad, 0.5 mm. thick; cotyledons cor-

date.— Flowers (and fruit) except calyx unknown in type; description

(as others) after Francey, who uses the name of Willdenow.

Piura: Ayavaca, (Bonpland, type). To Colombia.

Cestrum pseud oped i eel la turn Francey, Candollea 7: 29. 1936.

Glabrous or essentially, except for evanescent puberulence at the

tip of the elongate flexuous branchlets, these only 1-2 mm. thick;

stipuliform leaves none; petioles often uncinate at base, 7-10 mm.

long; leaves spreading or pendent, oblong-lanceolate, acute or cune-

ate at base, shortly acuminate, many 6-8 cm. long, about 3 to nearly

4 cm. wide, membranous, opaque above, slightly lustrous beneath,

the 8 or 9 arcuate nerves and reticulate veins prominent beneath;

racemes axillary in a foliose terminal panicle, 3-6-flowered, the dark

purple branches to 5 cm. long, 1-flowered peduncles 5-12 mm. long,

pedicels 0.7 mm. long, flowers 33 mm. long; bracts oblong-lanceolate,

1-1.5 cm. long, bractlets filiform, 4-9 mm. long, both long-attenuate

to base; calyx (in herb.) coriaceous, subtubular, narrowly obconic-

poculiform, 5 mm. long, 3 mm. thick, obscurely puberulent only

within, the 4 or 5 often coalescent teeth acute, ciliolate; corolla in-

fundibuliform, 32 mm. long, tube contracted below ovary, cylindric

to middle, gradually ampliate to 5.5 mm. across at apex where purple

and glabrous, the often plicate lobes 3.5 mm. long, marginally puber-

ulent, glandular; filaments 8.5 mm. free, gibbous, bent, subdenticu-

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

late, only the adnate part faintly pulverulent; ovary shortly stiped,

14-ovulate; style hirtellous at tip, the exserted capitate stigma puber-

ulent. — Flowers notable in dimensions, the pedicels very short but

the peduncles very long (author).

Peru: (Without data at Geneva, Weberbauer 6675, type).

Cestrum racemosum R. & P. Fl. Peruv. 2: 29, pi. 154. 1799;

273.

Glabrous in age except the racemiform panicles, the younger

branchlets and leaves sparsely pulverulent-puberulent; branches

erect, 3 mm. thick, purplish, lustrous as the spreading branchlets,

these to 2 dm. long, compressed, angled; petioles 7-16 mm. long;

stipuliform leaves none; leaves spreading or pendent, ovate-lanceo-

late, rounded at base, narrowly subobtusely or acutely acuminate,

often 1-1.5 dm. long, 2.5-5 cm. wide, membranous, plane or nearly,

lustrous, especially above, the 13-15 lateral nerves prominent be-

neath; panicles axillary, 1-3, about 10-flowered, the axis 1-2 cm. long,

pulverulent- tomentose; peduncles little branched, 2-5 mm. long, 2-3-

flowered, minutely lanuginose; flowers greenish, sessile; bracts linear-

acuminate, 2-3 mm. long, pilosulous as the tubular poculiform calyx

without, this 1-2 mm. long, 1-1.5 mm. thick; calyx teeth 5, unequal,

0.3-0.7 mm. long, 0.7 mm. wide, pilose at acute apex; corolla infundi-

buliform, about 1.5 cm. long, tube contracted below and above ovary,

inflated at apex, not constricted below limb, 12.5 mm. long, quite

glabrous; lobes of limb 3 mm. long, pubescent only without, the mar-

gins tomentulose; stamens 11 mm. long, the filaments 1-2 mm. free,

smooth, only the adnate part pubescent below; ovary globose; style

11.5 mm. long, glabrous; stigma capitate; fruit ellipsoid, subsessile,

4.5 mm. long, 3 mm. thick, maturing 3 elliptic or ovate-oblong seeds

4 mm. long, 1.5 mm. broad, 0.7 mm. thick; cotyledons oval. — Illus-

trated, Francey, I.e. 7: pi. 3, fig. 20 (flower).

San Martin: Tarapoto, Williams 6620; Spruce 4552. — Huanuco:

Chinchao and Macora, Ruiz & Pavdn, type. Monzon, Weberbauer

3434- — Loreto: Pongo de Manseriche, (Tessmann 4269). Bolivia to

Central America. "Yerba hedionda" (Ruiz & Pavon).

Cestrum Raimondianum Francey, Candollea 6: 382. 1936.

Allied and similar to C. undulatum; stipuliform leaves none (in

type); petioles 4-7 mm. long, slightly incrassate and uncinate at

base; leaves erect to spreading, oval or oblong, acuminate, shortly

FLORA OF PERU 117

decurrent at acute base, 4.5-6 (9) cm. long, about 2-3.5 cm. wide or

slightly wider, membranous but rigid, obscurely pulverulent on the

midnerve beneath, the lateral nerves 6-8 (type); panicles axillary,

6-8-flowered, the lower subracemose, the upper subspicate; peduncles

obsolete, bractlets filiform, minute; pedicels 1.3 mm. long; calyx

tubular, slightly 5-costate, 4 mm. long, 1.6 mm. thick, papillose with-

in, the 5 acuminate teeth often connate with rounded sinus; corolla

24.5 mm. long, tube below and above ovary contracted, very nar-

rowly obconic, ampliate to apex, 3 mm. broad, constricted below the

limb, 18.5 mm. long; lobes 5.5 mm. long, acute; stamens 17.5 mm.

long, the filaments 4 mm. free, geniculate at insertion, incrassate,

not denticulate; ovary globose, 6-ovulate, style 17.5 mm. long; fruit

sessile, oblong, 6.5 mm. long, 4.5 mm. thick, the subtending calyx

cupulate; seeds (immature) 6. — The species particularly marked by

its relatively large oval or oblong leaves; Stork & Horton collection

has leaves with 12 or more nerves, in this respect suggesting C. undu-

latum R. & P. Illustrated, Francey, I.e. 7: pi. 3, fig. 48 (flower).

Cajamarca: Rio Socota, Prov. Cutervo, Stork & Horton 10153

(distr. as C. affine with query, i.e. C. peruvianum) . — Amazonas:

Lamas, (Raimondi 1055, type). "Yerba Santa."

Cestrum reflexum Sendtn. in Mart. Fl. Bras. 10: 218. 1846; 265.

Branches elongate, flexuose, more or less scandent or at least sup-

ported in other shrubs, glabrous as entire plant except for some pu-

berulence on the younger parts, the leaves beneath near the midnerve

and the panicles; internodes 2.5-3 cm. long, the leaf scars very prom-

inent; petioles slender, often uncinate at base, 6-10 mm. long; stipuli-

form leaves deciduous, ovate, about 5 mm. long, 2-3 mm. wide;

leaves pendent, oblong- or subovate-lanceolate, acute or obtuse, often

unequal at the little decurrent base, acuminate or acute, mostly 8-

10 cm. long, 3.5-5 cm. wide, membranous, plane, lustrous both sides,

the 7 or 8 strongly ascending lateral nerves prominent only beneath,

the veins obviously reticulate on both surfaces; inflorescences axil-

lary, 10-15-flowered, foliose racemes or the upper forming an elon-

gate panicle, the lateral branches 1-3.5 cm. long; floral leaves ovate-

or oblong-lanceolate, 8-14 mm. long, half as wide; terminal panicles

about 1.5 dm. long or longer, the lanceolate or sublinear bracts 2.5-

5 mm. long, the flowers sessile; calyx campanulate, slightly angled

above, 3-3.5 mm. long, 3 mm. across, sparsely puberulent only with-

out, the 5 nerves slender, the 5 spreading teeth 0.5-0.7 mm. long,

1 mm. wide, ciliate; corolla yellowish-tinted, 25.5-27.5 mm. long, the

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

tube constricted below the ovary, abruptly ampliate at apex, cupuli-

form, more or less constricted below the limb, 17.5-21.5 mm. long,

quite glabrous; lobes elliptic- or ovate-lanceolate, 5 or 6 mm. long,

glabrate both sides, marginally subtomentose; stamens 17.5-20.5 mm.

long, the free part of filaments 1.5-2 mm. long, glabrous, smooth;

ovary globose, attenuate into stipe, 16-ovulate, the style 18-21 mm.

long, obscurely puberulent apically, the puberulent stigma capitate;

fruit sessile, ellipsoid, 8.5 mm. long, 6.5 mm. thick, the subtending

calyx cyathiform, 5 mm. long, 6 mm. across; seeds 8-11, oblong,

3.5-5 mm. long, 2-2.5 mm. wide, about 1 mm. thick; cotyledons ellip-

tic.— Illustrated, Francey, I.e. 7: pi. 1, fig. 72 (flower).

Cajamarca: Monte Seco, Prov. Hualgayoc, Soukup 3884- — San

Martin: Tocache, Poeppig. Tarapoto, Williams 5782; 6169; Spruce

4418.— Loreto: Florida, Klug 2061; 2350. Lower Rio Huallaga,

Williams 4286; Killip & Smith 29267.— Cuzco: Quellouno, Prov.

Convencion, Vargas 6331 . Southwestern Brazil to Central America.

Cestrum Sendtnerianum Mart, ex Sendtn. in Mart. Fl. Bras.

10: 215. 1846; 344.

A much-branched shrub sometimes becoming an elegant tree,

quite glabrous except for obscure or sparse indument in the flowers;

branches stout, 3.5 mm. thick, verruculose, the branchlets elongate,

lustrous, brownish-purple toward the tip, the flowers borne at the

apex of the younger, or solitary or 2-4-fasciculate in the leaf axils;

petioles 2-11 (18) mm. long, incrassate at base; stipuliform leaves

lanceolate, subsessile, obtuse, 7-10 mm. long, to 4 mm. wide; leaves

crowded, often plicate-falcate, ovate- or oblong-lanceolate, acutely

or acuminately decurrent, long-acuminate or cuspidate, ordinarily

3.5-10 cm. long, 1.5-5 cm. wide, rarely larger, membranous, not or

obscurely revolute, lustrous especially above, reticulate nervose, the

5-7 lateral nerves prominent only beneath; pedicels to 0.6 mm. long,

bracts lanceolate-acuminate, 4.5 mm. long, pulverulent; calyx ovate-

cylindric, rather strongly 5-nerved, subcoriaceous, 2.5-4 mm. long,

1.5-2.2 mm. thick, glabrous except the ciliate margins of the acute

teeth; corolla purplish or greenish yellow, about 2.5-3 cm. long,

the coriaceous tube contracted below the ovary, slightly tumid

above, finally narrowly obconic and gradually ampliate, constricted

below the limb, 19.5-24 mm. long, glabrous without, only the mar-

gins of the ovate lobes (4-5 mm. long) tomentulose; stamens 19-23

mm. long, the free part 3.5 mm. long, denticulate at or just below

insertion, the teeth and subgibbous adnate portion pilose; ovary

FLORA OF PERU 119

globose, 6-ovulate, with obconic glandular stipe; style 19-23 mm.

long, puberulent at apex, the stigma capitate; fruit dark violet,

sessile, ellipsoid, 13 mm. long, 7 mm. thick; seeds 4 to 6, dorsally

canaliculate, 5.5-6.5 mm. long, 2-3.5 mm. broad, 0.7-1 mm. thick;

cotyledons elliptic.

San Martin: San Roque, Williams 7285; 7638. Tarapoto, Wittiams

6881; Spruce 4277. Tocache, Poeppig 2039. — Loreto: Near Iquitos,

King 436; 877; Williams 1469; Tessmann 3564. Pebas, Williams

1649. Caballo-cocha, Williams 2128. Rio Nanay, Williams 663.

Lower Rio Huallaga, Williams 4522; 6881. Argentina to Costa

Rica. "Yerba Santa."

Cestrum silvaticum Francey, Candollea 6: 316. 1936.

Glabrate or obscurely puberulent on the younger parts and the

leaf midnerve both sides; branches flexuose, 1.5-4 mm. thick, sparsely

glandular-verruculose; stipuliform leaves semi-lunate, glabrous, 1-1.5

mm. long, 0.8 mm. wide, deciduous; petioles 1-1.5 cm. long; leaves

erect to spreading, elliptic-lanceolate, acutely decurrent, acuminate

or shortly cuspidate, about 1-1.5 dm. long, 3-4.5 cm. wide, mem-

branous, obscurely revolute, little if at all lustrous above, paler

and opaque beneath, reticulate both sides, the 8-12 lateral nerves

arcuate ascending; panicles axillary, 5-10-flowered, the basally

branched sparsely puberulent rachis 4-16 mm. long; bracts elliptic-

lanceolate, obtuse, 4 mm. long, the filiform bractlets minute; pe-

duncles at most 2 mm. long, pedicels 0.5 mm. long, glabrous as

the cyathiform calyx, this about 3 mm. long, 1.5 mm. across, ob-

scurely verruculose-glandular; teeth 5, equal, 0.4 mm. long, 0.6 mm.

wide, acute, pilosulous marginally, the sinus acute, the nerves slender;

corolla nearly 2 cm. long, the tube contracted below the ovary,

very narrowly obconic, apically ampliate, 2.8 mm. across, con-

stricted below the limb, 15.5 mm. long, glabrous without, barbate

within for two-thirds the length, adnate portion of filaments free

for 3.5 mm., smooth; lobes of limb 3.3 mm. long, glabrous except

the densely puberulent margins; ovary ellipsoid, 8-ovulate, the disk

glandular; style filiform, 16 mm. long, obscurely pubescent apically,

the stigma capitate; fruit 5.5 mm. long, 3.5 mm. thick but immature.

— Strongly resembles C. tenuissimum Francey, 248, of Bolivia but

differs by the size of calyx with acute sinus, by the short 1-flowered

peduncles and by the somewhat pilose filaments (author) ; however,

see C. Baenitzii; Klug 3612, referred there by Standley, seems to be

intermediate.

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

San Martin: Pongo de Cainarachi, Klug 2690 (det. Morton).—

Junin: Pichis Trail, Killip & Smith 25483, type. La Merced, 5645.—

Loreto: Above Pongo de Manseriche, Mexia 6251. Florida, Rio

Zubineta, Klug 2136. "Mitira-ey" (Klug).

Cestrum strigillatum R. & P. Fl. Peruv. 2: 29. 1799; 142.

C. unibracteatum Dunal in DC. Prodr. 13, pt. 1: 656. 1852, fide

Francey. C. calycinum HBK. Nov. Gen. & Sp. 3: 58. 1815 (as

var. below). Sessea rugosa Rusby, Bull. N. Y. Bot. Gard. 8: 119.

1912, fide Francey.

Typically a virgately stemmed branched shrub, the younger parts

and the leaves beneath, especially the nerves, lightly to lanate-

tomentose; branches to 5 mm. thick, strigose- pulverulent in age;

stipuliform leaves none; petioles 5-10 mm. long, canaliculate above;

leaves spreading, ovate to ovate-lanceolate, basally rounded or cor-

date, acute or acuminate, 0.5-2 dm. long, 4-10 cm. wide, membra-

nous, plane or nearly, glabrous and lustrous above, the 6-8 lateral

nerves prominent beneath; panicles more or less racemose, axillary,

solitary or geminate, several to 10 cm. long, 6-many-flowered, the

rachis to 9 cm., strigose-pulverulent; peduncles 5-16 mm. long, the

1-flowered 1-3 mm. long, filiform, apically much incrassate; flowers

sessile, basal bractlets 3-5 mm. long, linear-acuminate, densely to-

mentose; calyx tubular-poculiform, angled, 10-12 mm. long, 4-4.5

mm. thick, early tomentose without, glabrous within, the 5 ovate

ciliate teeth to 2 mm. long, nearly as wide, slender nerves prominent,

lateral many; corolla yellowish-green, 3-3.5 cm. long; tube con-

tracted below the ovary, cylindric, slender, strongly ampliate at

apex, above calyx strigose-tomentose, glabrous within, not con-

stricted below the limb, the latter with oblong-lanceolate or elliptic-

lanceolate acute lobes 9-12 mm. long, 4-5 mm. wide, tomentose

without, glabrous within; stamens 24 mm. long, the filaments only

free for 1 mm., smooth, glabrate; ovary globose, with obconic stipe;

style 24 mm. long, hirsutulous toward apex, the capitate stigma lob-

ulate; fruit ellipsoid, sessile, 11 mm. long, 6 mm. thick, the calyx ac-

crescent; seeds 7, 5 mm. long, 2 mm. broad, 1 mm. thick; cotyledons

elliptic. — Young fruit has a crossed scar at apex as in Sessea (Francey).

The var. calycinum (Willd.) Kuntze, Rev. Gen. 3: 220. 1893, treated

as specifically distinct by Francey, I.e. 137, and perhaps logically,

is distinguished primarily by more generally distributed, softly stel-

late tomentose indument and especially by the rather crowded flow-

ers, these sometimes subfasciculate or in subspicate 4-7-flowered

FLORA OF PERU 121

racemes 2-4 cm. long. Flowers with fragrance of jasmine; infusion

of leaves used for tonic (Seibert). Illustrated, Francey, I.e. 7: pi. 1,

fig. 32 (flower). F.M. Negs. 18394; 2998 (C. calycinum); 6899 (C.

unibracteatum).

Cajamarca: Gonzanama, (Bonpland, type, C. calycinum). — San

Martin: Bellavista to Banos, Ferreyra 1+71+6 (det. Cowan). San

Roque, Williams 7223; 7353. Tarapoto, Williams 5566; 6570 (var.) ;

Spruce 1+161+; 1+16 1+a (var.). Moyobamba, Raimondi; Woytkowski

35302 (var.). Chazuta, Klug 1+130 (var.). — Huanuco: Cuchero, Dom-

bey (type, C. unibracteatum) ; Poeppig 96. Pozuzo and Rio Chinchao,

1+778; 503 1+; Ruiz & Pavdn, type; Poeppig 1326. — Junin: Palca to

Chanchamayo, Raimondi. Colonia Perene", Killip & Smith 251+20.

La Merced, 5329; 5588; Killip & Smith 23673; 23385.— Loreto:

Yurimaguas, Williams 1+775; 1+857; 5288; Killip & Smith 28223 (var.) ;

2831+7 (var.). Puerto Arturo, Killip & Smith 27780 (var.); 2801+3

(var.). — Ayacucho: Huanta to Rio Apurimac, Killip & Smith 23216

(var.); 23100. — Cuzco: Santa Ana, Raimondi. — Madre de Dios:

Iberia, Seibert 2019 (var., det. A. C. Smith); 2151. Uruguay to

Colombia. "Ucha Panga" (Williams), "Santa Maria" (Seibert).

Cestrum tomentosum L. f. Sp. PI. Suppl. 150. 1781; 171.

A canescently stellate-tomentose shrub rather similar to C.

lanuginosum; petioles 3-12 mm. long, terete; leaves oblong- or ovate-

lanceolate, rounded, obtuse or subacute at base, not decurrent,

attenuately acuminate or acute, 7 to about 12 cm. long, usually

2.5-5 cm. wide, membranous or subcoriaceous, very sparsely stellate-

pilose above, the lateral nerves 7-9, the younger leaves obtusely

acuminate, naturally smaller, the floral oblong, acute, 6-8 mm.

long and half as wide; stipuliform leaves geminate, mostly persisting,

ovate or oblong-lanceolate, about 1-2 cm. long, half as broad; flow-

ers 3-5-fasciculate; bracts narrowly lanceolate, acuminate, 2-10 mm.

long, 0.7-2 mm. wide, puberulent, calyx campanulate-tubular, 5-7

mm. long, 2.5-3 mm. across, truncate at base, villous-tomentose or

subhirsute, the 5 acuminate teeth 1-1.5 mm. long, 1 mm. wide;

corolla 16.5-21.5 mm. long, infundibuliform, often tinged with brown

or purple, the tube a little contracted below the ovary, 14-18 mm.

long, the lobes 2-3.5 mm. long, acute or subacute; stamens 5, 13-17

mm. long, free for 5.5-6 mm., anthers elliptic; ovary subquadrate;

style 13-18 mm. long, filiform; fruit black, 8.5 mm. long, 4.5 mm.

thick with 3 oblong-elliptic seeds 4-5 mm. long, 2-2.5 mm. broad,

about 1 mm. thick. — The flowers fasciculate at the tip of a peduncle

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

or a lateral branch and subtended by bracts or floral leaves; charac-

ters of the flower constant (Francey). Illustrated, Francey, I.e. 7:

pi. 1, fig. 53 (flower); pi. 2, fig. 13 (calyx).

Piura: Ayavaca, Soukup 4295. — Cajamarca: Nancho, Raimondi

6646. Contumaza, Raimondi 6682. Prov. Hualgayoc, Soukup 3885.

— Ancash: Between Samanco and Huaraz, 3,000 meters, Weberbauer

3160. — Junin: Tarma, Raimondi 2461. To Colombia and Venezuela.

Cestrum undulatum R. & P. Fl. Peruv. 2: 28, pi. 155. 1799;

384.

A glabrous shrub developing a granulate erect trunk with terete

spreading branches 1-2 mm. thick, the flowering branchlets ordinarily

1-2 dm. long, flexuose, dark purple, obscurely puberulent apically;

stipuliform leaves lanceolate, long-acuminate, 2-5 mm. long, scarcely

1 mm. wide; petioles 6-10 mm. long; leaves reflexing, ovate-lanceolate,

often unequal at the somewhat rounded decurrent base, acuminate,

about 6-9 cm. long, mostly 2.5-4 cm. wide, membranous, undulate,

slightly lustrous above, minutely glandular beneath, the 7-10 lateral

nerves rather prominent; racemes axillary and terminal, solitary,

lax, 5-10 cm. long, 4-8-flowered, the filiform rachis to 7 cm. long,

peduncles to 6 mm. long, pedicels 0.5-2 mm. long, articulate, the

basal bracts geminate, linear-subulate, deciduous; calyx tubular-

poculiform, 3.5-4.5 mm. long, 2.5-3 mm. thick, glabrous or with-

out scarcely puberulent, the 5 or 6 subequal teeth acute or acuminate,

marginally ciliate, 2 mm. long, half as wide, nerves slender; corolla

lutescent, to 26.5 mm. long; tube contracted below ovary, finally

12 mm. long, apically ampliate, scarcely constricted below the limb,

19 mm. long, entirely glabrous; lobes 7.5 mm. long, acuminate,

only marginally pubescent; stamens 19 mm. long, filaments four-

fifths adnate, subbifidly denticulate; anthers quadrate; ovary

globose-ellipsoid, 8-ovulate, the disk glandular; style 19.5 mm. long,

puberulent below the capitate stigma; fruit violet, ovoid-ellipsoid,

2-celled, 8 mm. long, 6 mm. thick; seeds 5 or 6, angled, ovoid-

truncate, brownish, 5.5 mm. long, 2.5 mm. broad, 1.2 mm. thick;

cotyledons suborbicular. — Sometimes found in gardens, as at Lima

by Raimondi, the flowers fragrant. Illustrated, Francey, I.e. 7:

pi. 3, fig. 7 (calyx). F.M. Neg. 18395.

Huanuco: Near Huanuco, 2045; Ruiz & Pavon, type; Raimondi.

Ambo, 3161. Chaulki, Sawada 74- Tomaiquichua, 2425. — Junin:

Dos de Mayo, Pichis Trail, Killip & Smith 25862. — Loreto: Yuri-

FLORA OF PERU 123

maguas, Poeppig 23. La Victoria, Williams 2589. Rio Mazan,

Schunke 24- "Yerba hedionda," "yerba sancta."

Cestrum Weberbaueri Francey, Candollea 7: 5. 1936.

Rigidly branched shrub, the branchlets and erectly borne leaves

lustrous and glabrous as all parts except the puberulent bracts,

calyces only within (teeth marginally tomentulose) and the corolla

lobes both sides; leaves acute to acuminate at base but not decurrent,

1- nearly 2 dm. long, 3.5-6 cm. wide or wider, scarcely revolute,

the lateral nerves 11-15; stipuliform leaves 1-1.5 cm. long; bracts

obovate, about 4 mm. long; flowers green, 21 mm. long, pedicels

to 1 mm. long; calyx tubular, 6-6.5 mm. long, the 5 acute teeth

0.7 mm. long; corolla not constricted below limb, the lobes 4 mm.

long; stamens equal; stigmas lobate, subexserted. — Otherwise like

the related C. peruvianum Willd.

Cajamarca: Balsas to Celendin, (Weberbauer 4254, type).

19. NICOTIANAL.

Reference: T. H. Goodspeed, The Genus Nicotiana, 1-536. 1954

(Taxonomy, T. H. Goodspeed, H. M. Wheeler and P. C. Hutchison,

328-492). Goodspeed and Mildred C. Thompson, Cytotaxonomy

of Nicotiana, Bot. Rev. 25: 385-415. 1959.

Generally, in Peru, becoming tall subligneous shrubs, sometimes

coarse herbs, rarely diminutive annuals, almost always pubescent,

often viscid-glandular. Leaves alternate, entire. Flowers pedicelled

in terminal panicles with evident central axis, false racemes or these

modified into flat pinnate panicles or variously lax or remotely

glomerate more or less dichotomous systems, exceptionally flowers

foliose bracted. Calyx 5-toothed or -cleft, always persistent, usually

somewhat enlarged in fruit. Corolla slightly if at all irregular, often

with distinct tube and throat, the limb entire, shallowly 5-lobed

or 5-cleft, early contorted-plicate, rarely imbricate. Stamens 5,

the free filaments variously inserted, anther dehiscence longitudinal.

Ovary 2-celled, the base adnate to a thick annular disk, placental

cushions on the central partitions, the many ovules anatropous;

style terminal, stigma slightly grooved. Capsule membranous to

subligneous, the lower part indehiscent, the upper parting septicidally

or very shortly loculicidally, rarely indehiscent. Seeds minute, the

single seed coat variously reticulate. Chromosome number chiefly

12 or 24 pairs. — Genus is named for Jean Nicot, French consul

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

to Portugal in 1560, who sent to France seeds that he obtained from

a Dutchman from Florida. The Spaniards are supposed to have

first become acquainted with tobacco in the West Indies and its

name may have been derived from Tobago or Tabasco, the latter

locality in Mexico. Ruiz and Pavon found tobacco cultivated in

several provinces, observing the long cigars and the incredibly hard

"skeins" or bundles called "tobaco de andullo," which were also

offered me.

The work of Dr. Goodspeed exemplifies, as few other accounts

of the life history of a group of plants, the effectiveness of modern

botanical research when consummated by a student as great in

talent as in erudition. The author's secondary title, "Origins, Re-

lationships and Evolution of its Species in the Light of Their Distribu-

tion, Morphology and Cytogenetics," indicates the scope of his

investigation. In compiling, I hope accurately, the taxonomy as

presented by him and senior botanists of the University of California,

I acknowledge for all students as well as myself their useful contribu-

tion.

The technical key of the monographer and aides is supplemented

by a simpler version, this possible because of the relatively few

species within Peru.

KEY (after Goodspeed)

Corolla clavate-obconic, throat slightly contracted; stamens included

except N. Benavidesii; petiole not alate (obsolete, N. thyrsiflord) ;

corolla limb entire or nearly, yellow or green (subgenus Rustica

(Don) Goodspeed).

Corolla 2-5 cm. long, at least twice as long as broad (section

Paniculatae Goodspeed).

Stems, leaves quite glabrous, glaucous N. glauca.

Stems, leaves more or less puberulent.

Calyx teeth triangular; corolla yellow-green or limb dark.

Vigorous herbs; stamens included.

Corolla glabrous, entirely yellow-green — N. paniculata.

Corolla sericeous, the limb dark green . . . . N. Knightiana.

Shrubby; stamens exserted N. Benavidesii.

Calyx teeth minute; corolla pale greenish-yellow.

N. Raimondii.

Corolla 1-1.5 cm. long, obconic- tubular.

FLORA OF PERU 125

Leaves oblanceolate, petiole not or poorly defined (section Thyr-

siflorae Goodspeed) N. ihyrsiflora.

Leaves ovate-elliptic or cordate, petioled (section Rustica Good-

speed) N. rustica, N. Pavonii.

Corolla salverform to short tubular-salverform, the limb lobed, yellow

or sordid greenish-white above (subgenus Petunioides (Don)

Goodspeed).

Flowers in thyrsoid panicles, not closing in sunlight.

Large calyx lobe soon convex; corolla limb irregular.

N. undulata.

Large calyx lobe plane; corolla limb subregular N. Arentsii.

Flowers in pseudoracemes or panicles, vespertine.

N. plumbaginifolia.

Corolla infundibuliform, throat widely expanded; stamens somewhat

exserted; petiole alate (except N. glutinosa) or obsolete (sub-

genus Tabacum (Don) Goodspeed).

Corolla curved, throat expansion broad as long; subarborescent

(section Tomentosae Goodspeed).

Flowers thyrsoid paniculate; leaves ovate to lanceolate.

Style, stamens well-exserted, ascending N. tomentosa.

Style, stamens declined, little exserted N. Setchellii.

Flowers pseud oracemose; leaves cordate N. glutinosa.

Corolla straight or nearly, throat longer than broad; herbaceous

(section Genuinae Goodspeed) N. tabacum.

SUPPLEMENTAL KEY

Corolla curved, throat expansion broad as long, cupuliform; petioles

alate (except N. glutinosa).

Inflorescence racemiform; leaves cordate N. glutinosa.

Inflorescence paniculate; leaves usually attenuate toward often

auriculate or alate base.

Stamens well-exserted, ascending; petioles, unless uppermost,

defined, narrowly alate N. tomentosa.

Stamens subincluded, declined; leaves sessile or short, petioles

broadly alate N. Setchellii.

Corolla straight or nearly, not long-tubular, throat longer than

broad, or poorly defined (see contrast 3).

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

Petioles obscure to alate or (and) auriculate; corolla 1-3 cm. long

or longer.

Petioles alate, defined; corolla about 3 cm. long or longer.

Flowers paniculate, throat defined N. tabacum.

Flowers often in racemiform group, throat poorly defined.

N. plumbaginifolia.

Petioles obscure; corolla about 1.5-2 cm. long. .N. thyrsiflora.

Petioles well-defined, not alate or auriculate.

Corolla clearly lobed; calyx teeth broad, subobtuse.

Large calyx lobe soon convex; corolla limb uneven.

N. undulata.

Large calyx lobe plane; corolla limb subregular. . . N. Arentsii.

Corolla obscurely lobed; calyx teeth acute.

Calyx about half as thick as long; capsule 7-12 mm. long,

valves widely separating N. Pavonii.

Calyx more than half as thick as long; capsule 13-16 mm. long,

indehiscent or valves little parted N. rustica.

Corolla at least twice longer than thick, easily defined as long-

tubular.

Stems, leaves more or less puberulent.

Corolla 2-3.5 cm. long; stamens included; leaves if cordate

with open sinus.

Corolla sericeous, 2-2.5 cm. long to dark green limb.

N. Knightiana.

Corolla glabrous or nearly, entirely greenish-yellow.

Inflorescence rather open, sub viscid; fruiting pedicels nearly

straight N. paniculata.

Inflorescence often dense, subcolumnar; fruiting pedicels

often arcuate-uncate N. Raimondii.

Corolla 4-5 cm. long; stamens exserted; leaves cordate, sinus

often closed N. Benavidesii.

Stems, leaves glabrous, glaucous N. glauca.

Nicotiana Arentsii Goodsp. in Proc. Calif. Acad. Sci. ser. 4, 25:

297, pis. 26, 27. 1944; 379.

A stout annual or short-lived perennial resembling N. undulata

with erect axis and several rigid ascending branches, the stem early

FLORA OF PERU 127

often reddish-purple with conspicuous weak short silvery trichomes,

glabrate in age, thinly corky, ligneous within; leaves spreading,

pilose to obscurely pubescent, rotund-ovate or subcordate, acute or

acuminate, 1-2.5 dm. long, the petiole less than half as long; panicles

reduced-foliose below, lanceolate bracted above, the well-developed

rachis branching, the pedicels 2- to finally 6 mm. long, calyx pubes-

cent, 10-14 mm. long, broadly cylindric or subcampanulate, lobes

unequal, broadly triangular-ovate, shorter than tube; corolla to limb

16-20 mm. long, slightly asymmetric, tube proper at most 4 or 5 mm.

long, 3 mm. wide, throat about 3 times as long, 2.5 times as wide,

broadly obconic-clavate, pale greenish-yellow, shortly white pilose,

the limb 5-9 mm. wide, dingy straw-colored, pubescent without,

the broad lobes subacute to slightly acuminate; fifth filament sig-

moidly curved, less pubescent than the 4 longer; capsule elliptic-

ovoid, the large calyx lobe plane or nearly; seeds broadly elliptic,

0.6-0.8 mm. long, dark brown; chromosome number 24 pairs. —

Otherwise much like the related species N. wigandioides Koch &

Fintelmann (corolla barely 8-25 mm. wide), 381, of Bolivia; it

gives morphological and cytological evidence of being a natural

occurring amphiploid involving N. undulata and N. wigandioides;

it is distinguishable from N. undulata by the larger rotund ovate

leaves, larger flowers, green nerve, corolla limb and large plane

rather than boat-shaped calyx lobe in flower and fruit, and from

N. wigandioides by smaller leaves, smaller flowers and markedly

uneven calyx teeth; honors George Arents, who made a noteworthy

commentary on the literature and culture dealing with tobacco

(Goodspeed). Illustrated, Goodspeed, I.e. 380, fig. 76.

Puno: Limbani, Vargas 1322; 9666, type; Metcalf 30529. Bolivia.

Nicotiana Benavidesii Goodsp. Univ. Calif. Publ. Bot. 18:

137, pis. 9, 10, 12a. 1938; 345.

Sparsely branched, subligneous, 2-3 meters high, the stout stem

to 3 cm. across, densely puberulent and moderately viscid as the

leaves beneath and sometimes the calyx; leaves 1-3 dm. long, twice

as long as the petiole, apex retuse, cordate, with basal lobes prolonged

inward and often upturned, the upper surface puberulent; panicles

columnar, the pedicels 7-8 mm., in fruit 7-9 mm. long; calyx poculi-

form, densely puberulent, membranes obsolete or obscurely con-

necting the short broad equal teeth; corolla 4-4.5 cm. long to limb,

glabrous or nearly, light greenish-yellow, the tube 5-7 mm. long,

5 or 6 mm. broad, the throat 5-7 times as long, 8-12 mm. wide,

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

clavate-cylindric but with a short slight dilation just below

the somewhat contracted mouth, the limb about 4 mm. wide with

obscure apiculate lobes; stamens commonly of 3 lengths, exserted,

abruptly and densely lanuginose near slight insertion bend at base

of corolla throat; capsule rotund-ovoid, acute, exserted, 9-15 mm.

long, slightly ligneous; seeds broadly suboblong or irregularly angular,

0.6 mm. long, dull brown, reticulate; embryo straight, chromosome

number 12 pairs. — Named for General 0. R. Benavides, a former

president of the Republic of Peru, who gave assistance to the Good-

speed Expedition of 1935-1936. In Berkeley, California, the plant

does not flower until the second year. Illustrated, Goodspeed, 344,

pi. 62.

Apurimac: Near Pachachaca Bridge toward Abancay, West 3979,

type. — Cuzco: Rio Limatambo near Rio Apurimac, West 384-8. Prov.

Anta, Vargas 328. Rio Paucartambo, north of Abancay, Goodspeed

Exped. 105^8.

Nicotiana glauca Graham, Edinb. New Phil. Journ. 5: 175.

1828; 335.

Glabrous or the greenish-yellow flowers minutely pubescent, laxly

often virgately branched ligneous shrub or short-lived tree several

meters high, the younger parts characteristically glaucous, greenish

or blue-purplish, reddish-brown as cork begins to form, finally gray;

leaves about twice as long as petioles, those on younger shoots some-

times as broad as long, 2.5 dm. or more, flexible, cordate-ovate,

elliptic or lanceolate, 0.5-2.5 dm. long; panicles short, glabrous to

puberulent, flat, lower branches elongating, the pedicels 3-10 mm.

long, later thickened-hamate, 7-12 mm. long; calyx 1-1.5 cm. long,

cylindric, sometimes as corolla minutely pubescent, membranes lack-

ing or nearly, teeth triangular, acute, much shorter than tube;

corolla 3-3.5 (2.5-4.5) cm. long to limb, the tube itself 5-8 mm.

long, about 3 mm. wide, the throat 3-6 times as long, cylindric to

clavate, at most 6-8 mm. across, commonly yellow, the limb 3-4 mm.

wide, subcircular-subpentagonal, greenish or yellow; stamens sub-

equal, extending almost to mouth, the glabrous filaments geniculate

just above insertion on base of corolla throat; capsule broadly

elliptic, 7-15 mm. long, included; seeds longer than broad, truncately

angular, laterally compressed, about 0.5 mm. long, brown, foveolate-

reticulate; embryo straight; chromosome number 12 pairs. — Pre-

sumably originated in Argentina, its present wide range now due

in considerable part to introduction; narrow-leafed forms occur in

FLORA OF PERU 129

southern Peru and Bolivia; in California specimens have been known

to live, as trees, for over ten years (Goodspeed). Raimondi thought

it was introduced into Peru about 1865 via Bolivia. According to

Herrera it is a very strong narcotic. Illustrated, Bot. Mag. 55.

pi. 2837; Goodspeed, I.e. 334, pi. 59; Addisonia 23: pi 739.

Cuzco : Urcos, Soukup 1 4- Vilcanota, Pennell 141 87. Huambutio,

West 6495. Prov. Quispicanchis, Herrera 685. — Moquehua: Near

Moquehua, Weberbauer 7453. Argentina, perhaps Bolivia; now

widely distributed in warm temperate climates. "Ccjamata," "supai-

ccarco" (Herrera).

Nicotiana glutinosa L. Sp. PI. 1: 181. 1753; 369.

Coarse annual ordinarily a meter or two high with a rather sturdy

stem, rigid narrowly divergent branches and a viscid pubescence ex-

tending even to the corolla; petioles shorter than the leaves them-

selves, these in general cordate-rounded, the more or less acuminate

tip twisted, the basal lobes often incurved, the larger 1-2 dm. long;

racemiform inflorescence elongating, stout, the pedicels 5-12 mm.

long, little longer in fruit; calyx campanulate, 5-7 mm. long, the

lobes markedly unequal, 1-3 times as long as the tube, triangular,

narrowly acuminate, often recurving; corolla tube about 3-5 mm.

long, 5 mm. wide, the throat light greenish-yellow or red-tinted, the

cylinder 5-10 mm. long, the cup oblique, 1.5-2 cm. long, nearly as

broad, bell-like below the limb, this 8-12 mm. wide, pink, red or

lemon-yellow within, waxen, glabrous but viscid puberulent without,

the lobes usually broadly ovate with shortly acuminate recurved

grooved tips, now and then short, even truncate; stamens all tardily

exserted from throat or one shorter, the filaments sparsely lanate on

lower half, oriented with style above the basal insertion, against

upper side of flower; capsule broadly ovoid, 1-1.5 cm. long, included

or nearly; seeds 0.6 mm. long, dark brown, reticulate, the ridges

unevenly wavy; embryo straight; chromosome number 12 pairs.—

According to Comes, N. glutinosa was first discovered by Feuill^e

in the Lima Valley and seeds were sent to Paris by the younger

Jussieu; cultivated plants from seeds received from Lima had pink

flowers with long, recurved calyx lobes and long, broad ovate corolla

lobes; variations in other races include more strict plants, flat pan-

icles, pendent flowers, shorter calyces with erect segments, shorter

corolla throat and corolla lobes as well as color ranging from yellows

to purplish-reds; plants grown from the Haught collection had lemon-

yellow corollas and more triangular leaves while Metcalf 30782 varied

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

considerably in habit, leaf shape, orientation of pedicels and flowers,

the latter varying also in size, shape and color; the floral variation

tends to parallel that in N. tomentosa R. & P. and related species

(Goodspeed). Characteristically occurs in semi-arid regions of the

lower Andean slopes but has been collected to 2,700 meters, prob-

ably as a weed. Illustrated, Goodspeed, I.e. 370, pi. 73.

Piura: Cerros de Amotape, Haught 85. — Amazonas: Balsas, Met-

calf 30782. — Ancash: Los Zorros, Goodspeed Exped. 9193. — Lima:

Huacho, Weberbauer. Matucana, 258. Rio Chillon, Pennell 14457.

Near Lima, Soukup 3806; Weberbauer. Canta, Soukup 2827. —

Huancavelica: Pampas to Salcabamba, Stork & Horton 10258. Ancco,

Soukup 4028. — Apurimac: Cunyac Bridge near Abancay, Vargas 33 4;

9625. Ecuador.

Nicotiana Knightiana Goodsp. Univ. Calif. Publ. Bot. 18: 139,

pis. ll,12b. 1938; 340.

Similar and related to N. paniculata but perennial in tendency,

becoming subligneous and with shorter broad-throated bicolored co-

rolla; panicles longer than broad, the pedicels 3-4 mm., in fruit 5-

8 mm. long; calyx cylindric, narrowed to base, the membranes often

long, narrow-wedged, the short teeth broadly triangular; corolla pale

yellow-green cinereous with short sericeous trichomes, the limb dark

green, velvety, faintly notched into 5 subtruncate often mucronulate

lobes; filaments slightly curved; capsule broadly ovoid, 6-10 mm.

long. — The name honors the late Lora J. Knight, who gave support

to the first two South American expeditions of the University Bo-

tanical Garden (California).

Arequipa: Nazca to Chala, Metcalf 30336. Atiquipa, Worth &

Morrison 15673. Southeast of Chala, Worth & Morrison 15696.

Near Mollendo, Mexia 04161, type.

Nicotiana paniculata L. Sp. PI. 1: 180. 1753; 339.

A coarse foliose bushy annual (now and then persisting for more

than a year), rarely 3 meters tall, with many vigorous erect or spread-

ing branches equaling or exceeding the original axis; stem toward

base, as petioles and leaves beneath, pulverulent-puberulent, above

viscid pubescent especially in inflorescence; leaves, with petiole, about

1-1.5 (3) dm. long, ovate to cordate; panicles long, the central axis

slender, the pedicels 2-5 mm., in fruit 5-10 mm. long; calyx narrowly

cylindric, 6-10 mm. long, the membranes obscure, narrow, often elon-

gate, the cuneate teeth shorter than the tube; corolla yellowish-green,

FLORA OF PERU 131

glabrous or nearly, 2-3 cm. long to limb, the tube itself 3-5 mm. long,

2 mm. wide, the throat 3-5 times as long, 1.5-3 times as broad, cla-

vate, the limb 2-3 mm. wide, the lobes frequently apiculate; the

stamens subexserted, all or only 4 of filaments slightly geniculate,

pubescent at insertion on base of corolla throat; capsule elliptic-

ovoid, 8-12 mm. long, usually included; seeds oblong, angular-obo-

void or ellipsoid, sometimes compressed, 0.5-0.6 mm. long, blackish-

brown, fluted reticulate; embryo straight; chromosome number 12

pairs. — Almost continuously in flower between 300 to 3,100 meters;

this was the third species to be noted by a botanist, having been de-

scribed and figured from Lima by Louis Feuille'e in 1714 (J. Observ.

Phys. Math, et Bot. 1: 717, pi. 10. Paris). The type was grown from

seed collected in Peru by Bernard de Jussieu. Illustrated, Good-

speed, I.e. 338, pi. 60; Ruiz & Pavon, Fl. Peruv. 2: pi. 129.

Cajamarca (fide Goodspeed). — La Libertad: Otuzco, Gastanadui.

— Lima: (Nation 162}. Chancay, Ruiz & Pavdn; Wawra 2544.; Seler

243. Pachacamac, Mexia 8097; 8329. — Huanuco: Near Huanuco,

Ruiz & Pavdn. — Junin: Palca, Stork 10984- Tarma, 104-4! Killip &

Smith 21927.— Ayacucho: Puquio, Metcalf 30292.— Cuzco: Prov. de

Anta, Hen era 1953. — Arequipa: Tingo, Pennell 13124- Near Mol-

lendo, West 8207. Arequipa, Seler 214; Isern 2012. — Tacna: South

of Tacna, Metcalf 30351. "Tobaco Cimarron," "ccama-sayari."

Nicotiana Pavonii Dunal in DC. Prodr. 13, pt. 1: 561. 1852;

355. N. rustica L. var. Pavonii (Dunal) Goodsp. Nicotiana 355. 1954.

To 12 dm. high, canescent pubescent all over, the prominent main

axis with oblique branches from each axil; leaves broadly ovate to

elliptic-ovate, the uppermost lanceolate, frequently attenuate to

base, obtuse or acute; panicle with many slender many-flowered

branches, in well-developed specimens becoming ample; calyx 8-

12 mm. long, 3-6 (8) mm. wide, oblong-cylindric, the dorsal tooth

(or 2 teeth) elongate, all the teeth acute; corolla to about 17 mm.

long, throat 6 mm. wide, limb 3-4 mm. wide; capsule elliptic-ovoid,

7-12 mm. long, 6-10 mm. across, the valves nearly entire, finally

separating; seeds 0.7 mm. long, angular; embryo straight or nearly.

— Peruvian collections, especially in bud, have calyx, corolla and

ovary somewhat purplish; Metcalf 30800 has calyx 11 mm. long,

corolla 22 mm. long, limb 1.5 cm. wide, appearing crenate but the

obscure broad lobes actually retuse; plants from seed collected at

Huamachuco had corolla 19 mm. long, limb 11 mm. wide, the ob-

scure lobes apiculate (Goodspeed) ; compare remarks under N. rus-

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

tied. Type localities, fide Dunal, are Chavin, Bombon (Junin) and

Chicoplaya (Huanuco). Illustrated, Goodspeed, I.e. 352, fig. 66 (N.

rustica var. Pavonii). F.M. Neg. 3029.

Cajamarca: Near Cajamarca, Metcalf 30780. — Amazonas: Toward

Chachapoyas, Metcalf 30800; Soukup 1*172. — La Libertad: Huama-

chuco, West 8084- — Ancash: Huaraz, (Stadelman 33). — Lima: San

Mateo, (Stadelman 7). — Huanuco: Maria del Valle, 3566. — Junin:

Tarma, Stork 10948; 10983.— Cuzco: Valle del Apurimac, Huara,

Paucartambo, Vargas 331 (det. Standley). — Puno: Cuyocuyo, Met-

calf 30712. Chile, Bolivia, Ecuador.

Nicotiana plumbaginifolia Viviani, Elench. PI. Hort. Dinegro

26, pi. 5. 1802; 403.

An erect slender annual with tuberculate hispid flexuose soon

wiry stem, long basal branches and a few rosulate leaves, these spat-

ulate to obovate or oblanceolate, 1-2 dm. long or longer, hispid;

leaves of lower stem as large or larger, rotund-ovate, broadly alate

at the clasping to auriculate base, abruptly acuminate, usually

twisted, undulate to plicate-erosulate, the upper sessile, auriculate,

lanceolate to linear-lanceolate, strongly undulate-plicate; raceme-

like inflorescences simple or remotely furcate, occasionally tardily

branching, the pedicels 3-7 to finally 5-10 mm. long, flowers vesper-

tine; calyx hispidulous, 8-13 mm. long, the elliptic ovoid tube 10-

ribbed, the membranes long, narrow, the segments subulate filiform,

subequal, about equaling the tube, erect or slightly spreading; corolla

puberulent, the tubular part 2.5-3.5 cm. long, 1.5-2 mm. broad,

greenish-ivory or somewhat purplish, the throat poorly defined but

finally abruptly swollen to 3 or 4 mm. a few mm. below the some-

what contracted mouth, the limb about 1 cm. wide, deeply lobed,

the ovate acute lobes ivory or lavender without, with 5 dark nerves,

white or lavender within; anthers dark purple, 4 subsessile, 2 or 3 mm.

below mouth, sometimes didymous, the fifth about 3 mm. lower, its

filament 1 mm. long; capsule narrowly ovoid, 8-11 mm. long, usually

included; seeds 0.5 mm. long, rotund elliptic, light brown, wavy-

reticulate; embryo straight; chromosome number 10 pairs.

San Martin: Tarapoto, Spruce 4169. Adjacent Ecuador; north-

western Argentina to southeastern Brazil; Mexico and Caribbean,

common, perhaps introduced.

Nicotiana Raimondii Macbr. Field Mus. Bot. 8: 106. 1930; 349.

Semi-ligneous (the thin reddish cork tardily forming), a meter or

two high with several to many erect whitish-green slender stems from

FLORA OF PERU 133

a root-crown, dull velvety-puberulent as the petioles, leaf blades be-

neath and calyces; leaves ovate or slightly cordate, 1-1.5 dm. long;

panicles narrowly columnar, dense, the pedicels 6, finally 7 mm. long;

calyx broadly cylindric, 6-7 mm. long, membranes variable, usually

broader above, the subequal slightly recurved teeth minute; corolla

2.5-3 cm. long or longer to limb, glabrous, the tube itself about 2 mm.

long and wide, the throat 10-14 times as long, to 3 times as broad,

clavate, pale greenish-yellow, the paler limb 2-3 mm. wide, recurved

with obscure shallow apiculate lobes; stamens subexserted, filaments

densely lanate to 8 mm. above insertion at base of throat where

straight; capsule dark brown, ellipsoid, 8-10 mm. long, slightly ex-

serted; seeds angular-obovoid or suboblong, somewhat compressed,

0.6 mm. long, brown, fluted reticulate; chromosome number 12 pairs.

—Inflorescences rather lax to subglomerate; after an thesis many

pedicels become hamate by basal curving; withered and darkened

corolla may persist briefly (Goodspeed). Illustrated, Goodspeed,

I.e., 348, fig. 64.

Cuzco: Urubamba Valley, Hen era 1540, type; (Stadelman 22};

Soukup 16. Huambutio, West 6496. Ollantaitambo, Pennell 13665;

Herrera 233.

Nicotiana rustica L. Sp. PI. 1: 180. 1753; 351.

Robust, the solitary or sometimes several erect stems moist-viscid

pubescent, the indument merely a puberulence on the fleshy leaves,

these ovate to elliptic-lanceolate, sometimes cordate or subrotund,

often unequal at base, 1-1.5 dm. long or longer, the petiole much

shorter than the blade; panicles various but the central axis distinct,

the pedicels 3-4, later 5-7 mm. long; calyx 8-15 mm. long, poculi-

form to cylindric, pubescent, the membranes narrow, one of the

broad acute teeth much longer; corolla greenish^yellow, 12-17 mm.

to limb, puberulent, the tube commonly 3 mm. long, 2 mm. wide,

the throat about 3 times as long, 6-8 mm. wide, broadly obconic,

slightly contracted at mouth, the limb 3-6 mm. wide, obscurely and

obtusely lobed; stamens white pilose for about 2 mm. above basal

insertion, 4 barely included, sigmoidly erect, the fifth shorter; cap-

sule subellipsoid to subglobose, 7-16 mm. long, included or nearly;

seeds 0.7-1.1 mm. long, dusky brown, fluted-reticulate; embryo

straight or bent; chromosome number 24 pairs. — After Goodspeed

(as of course all other descriptions), who notes that his interpretation

represents a composite picture of this highly polymorphic species,

long a cultigen and apparently unknown today in the wild state (with

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

the possible exception of var. Pavonii (Dunal) Goodsp. (N. Pavonii

Dunal), I.e. 355, which his description includes). For the usefulness

of this work it seems preferable to retain N. Pavonii as a distinct

species; otherwise its taxonomic standing is parallel to the formal

variants pumila Schrank and brazilia Schrank; the native N. Pavonii

is more slender, the filaments are slightly curved and especially the

smaller capsules are dehiscent, the valves separating. The descrip-

tion is included here largely to indicate its relationship to N. Pavonii

and because according to Goodspeed genetic evidence has shown that

it is an amphiploid, progenitors of N. paniculata and N. undulata

having entered into its origin, N. Pavonii showing affinity to the for-

mer in narrower flower, to the latter in traces of purple coloration.

Illustrated, Goodspeed, I.e. 354, pi. 67 (var. brazilia Schrank).

N. rustica rather than N. tabacum L. was the first tobacco grown

and exported by the American colonists but today N. tabacum is

cultivated commercially as a source of nicotine products (Goodspeed).

Peru (compare note above).

Nicotiana Setchellii Goodsp. Univ. Calif. Publ. Bot. 18: 195,

pis. 14, 15. 1941; 367.

Soft-woody shrub, 2-5 meters high, the 1-3 stems in age thinly

corky, striated by shallow furrows, marked by leaf scars, much

branched, green, viscid pubescent as the leaves beneath; leaves

shortly decurrent, sessile or with short alate petiole, base somewhat

auricled, obscurely pubescent both sides, oblong, oblanceolate or

lanceolate-elliptic, 4-8 dm. long; panicle congested, primary branches

about 30, mostly simple, the pedicels 10-18, in fruit 15-20 mm. long,

directing the flowers centrifugally, the maturing capsules centripet-

ally; calyx 1.5-2.5 cm. long, broadly cylindric or cylindric-campanu-

late, teeth unequal, narrowly triangular-acuminate, the longest often

equaling the tube; corolla viscid or varnished without, the tube

5 mm. long, slightly wider, the throat pale greenish-yellow, sometimes

red-tinged, the throat cylinder 1.5-2 cm. long, 7-8 mm. wide, the

throat cup twice as broad as the cylinder, 15-18 mm. long, slightly

inclined, widely open; limb 12-15 mm. wide, ascending, red tinged

with bronze, within pink to red, the lobes shallow to obsolescent with

acuminately protracted tips, the entire limb pentagonal; stamens all

or nearly all exserted but included in limb, the filaments inserted

near the base, then oriented with the style against the lower side of

the flower, quite glabrous or below puberulent; capsule elliptic, 1.5-

2 cm. long, included; seeds subspherical to somewhat obovoid, about

FLORA OF PERU 135

0.5 mm. long, dull brown, reticulate, the ridges minutely wavy; em-

bryo straight; chromosome number 12 pairs. — Known only from type,

found along trail on Beauchamp's second day's journey from Cha-

chapoyas; readily distinguished from N. tabacum by its shrubbiness,

simple pattern of inflorescence, stamen orientation and bowl-like co-

rolla limb, which with the throat cup forms a bell. After first year

(at Berkeley) a single plant increased by root sprouting to twenty or

more individuals, some even at a distance of twelve meters. Illus-

trated, Goodspeed, I.e. 368, pi. 72.

The name recognizes the role of the late W. A. Setchell in initi-

ating and contributing to the Nicotiana investigations at the Uni-

versity of California.

Amazonas: Between Chachapoyas and Leimabamba, Beauchamp,

type.

Nicotiana tabacum L. Sp. PI. 1: 180. 1753; 372.

Stout viscid annual or limited perennial 1-3 meters high, the thick

erect stem with few rapidly ascending branches, the decurrent leaves

(the larger at least 5 dm. long) ovate-elliptic or lanceolate, attenuate

to base or alate-subpetiolate, acuminate; panicle with distinct rachis,

the several branches usually compound, commonly shorter than ra-

chis; pedicels 5-10 (15) mm. long, in fruit 10-20 (25) mm.; calyx

cylindric to cylindric-campanulate, 12-20 (25) mm. long, viscid, the

triangular acuminate teeth shorter than or equaling the tube, un-

equal; corolla little if at all curved, puberulent, the tube proper (7)

10-15 mm. long, 2.5-3 mm. wide, the throat (23) 25-40 mm. long, the

lower half cylindric, 3-5 mm. wide, pale greenish-cream as the upper

half or this pink to red, usually abruptly expanded into a deep cup

7-12 mm. wide but occasionally obconic; limb 1-1.5 cm. wide, lobed

or pentagonal, white, pink or red ; stamens inserted on base of corolla

throat, erect, oriented to upper side of flower or evenly spaced, an-

thers or two longer pairs near mouth slightly exserted, the fifth

stamen shorter than either pair; capsule narrowly ellipsoid, ovoid or

orbicular, acute or obtuse, exserted or included, 1.5-2 cm. long; seeds

spherical or broadly elliptical, about 0.5 mm. long, brown, ridges

fluted, embryo straight; chromosome number 24 pairs. — This, of

course, is the species in cultivation that is the commercial source

of the habit-forming drug nicotine, the use of which doubtless some

day will be controlled as in the case of other drugs. Goodspeed, I.e.

373-375, has given an interesting account of the species-origin, vari-

ation and relationship which in part reads: there is today no well-

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

authenticated record of the occurrence of this species in the wild

state; presumably it escaped from pre-Columbian as it certainly has

from present-day agriculture, particularly in Central and South

America; its demonstrable origin in amphiploidy, involving progeni-

tors of a member of section Tomentosae and of N. sylvestris Speg. &

Comes, suggests that its original area of natural distribution was

northwestern Argentina and adjacent Bolivia, where N. otophora

Griseb. is today in contact with the latter or where relatively re-

cently that or other species of section Tomentosae had such a contact.

Still quoting the monographer, the type (from "America") consists

of a lanceolate acuminate leaf 24 cm. long, attenuate into a short

alate petiolar region and a young inflorescence with only flowers;

calyx 16-17 mm. long, cylindric to campanulate, corolla 4.5 cm. long

to limb, throat cylinder 3 mm. wide, abruptly expanded, throat cup

1.5 cm. long, 7 mm. wide; limb lobes elongate, acuminate; major

variations involve shrubby habit, leaves sessile, auricled, or with

non-alate petioles, inflated orbicular calyces, short broad teeth, slen-

der corollas, gradually expanded throat, pentagonal limb. For synon-

ymy consisting mostly of varieties see Goodspeed, I.e. 372. Illustrated,

I.e. 374, fig. 74.

Often persisting as an escape from cultivation as the race "Machu-

picchu" collected near the famous archaeological site above the Rio

Urubamba, Cuzco; tall in habit, it is strongly suggestive of the Bo-

livian N. tomentosiformis Goodspeed, I.e. 363, in leaf, inflorescence

and flower.

Cajamarca: Monte Seco, Soukup 3890. — San Martin: Tarapoto,

Lamas, San Roque, Williams 5529; 6456; 7248. — Huanuco: Llata,

2277. — Junin: Huancayo, Soukup 3978. — Loreto: Rio Nanay, Wil-

liams 428; 1287. — Cuzco: On old terraces, ruins of Machupicchu,

Mexia 8073; 8072.

Nicotiana thyrsiflora Bitter ex Goodsp. Univ. Calif. Publ. Bot.

18: 138, pi. 13. 1938; 349.

Coarse, simple or sparingly branched perennial 1-3 meters high

marked by the densely puberulent thick stem terminating in an erect

stout columnar panicle with slender suberect laterals; pedicels 8-

15 mm. long; calyx 1-1.5 cm. long, deeply campanulate, obscurely

pubescent, the membranes short, narrow, the triangular teeth tipped

with darker green, the longest less than half the length of the tube,

often much less; corolla glabrous, greenish-yellow, 9-15 mm. long

to limb, the tube 2-4 mm. long and wide, the throat 3.5 times as

FLORA OF PERU 137

long, 5-7 mm. wide, broadly obconic or inflated-funnelform, con-

tracted at mouth, the limb 3.5 mm. wide, the lobes apiculate; sta-

mens all subexserted, inserted at base of throat, the filaments there

sparsely pilose; capsule broadly oval, 1-1.5 cm. long, included; seeds

about 1 mm. long, dull sooty brown-black, honeycomb reticulate;

embryo straight or nearly; chromosome number 12 pairs. — Its spicate

habit suggests Verbascum thapsus L.; the corolla limb is a bowl as

broad as the corolla is long; it differs from N. rustica L. in the

crowded narrow sessile leaves, spicate habit, ascendant corolla limb

(or this only in part deflecting) and sometimes purple anthers (Good-

speed). Grass-shrub formation, stony slopes, 2,800 to 3,800 meters.

Illustrated, Goodspeed, I.e. 350, fig. 65.

Cajamarca: Los Tajones Valley, Stork & Horton 10018. — La

Libertad: Between Huamachuco and Angasmarca, Weberbauer 7015,

type; West 8179. North of Cachicadan, Stork & Horton 9990. San-

tiago de Chuco, Stork & Horton 9943.

Nicotiana tomentosa R. & P. Fl. Peruv. 2: 16, pi. 129, fig. a.

1799; 357. Lehmannia tomentosa (R. & P.) Spreng. Anleit. Kennt.

Gew. 2:458. 1817.

Sometimes arborescent and attaining 7 meters, the ascendant

branches forming a crown, the 1-3 stems tardily forming cork, in

age laced by leaf scars, usually more or less viscid pubescent includ-

ing the calyces; leaves decurrent, the short petiole alate, 2-11 dm.

long, the uppermost leaves subsessile, typically viscid-tomentose be-

neath, ovate to elliptic-lanceolate; panicles in type lax, ample, with

10-15 remote rigid branches, the bracts deciduous, the pedicels 10-

12 mm., in fruit 15-20 mm. long; calyx about 1.5-2 cm. long, poculi-

form-campanulate, typically loosely enclosing the corolla throat, the

longest of the rather broad acute teeth not exceeding the calyx tube;

corolla sparsely and minutely glandular only without, the tube obso-

lete or nearly, the throat 2-3.5 cm. long, light greenish-yellow, 6 or

7 mm. broad below, narrowed above, one side shorter, the throat cup

as long as the cylindrical part, about twice as broad, declinate, widely

open, 5-10 mm. wide, usually pink or red and the darker and lighter

areas forming a pattern, the lobes in type poorly defined, abruptly

acuminate; stamens far-exserted, filaments at base then oriented

against upper side of flower, curved, glabrous; style somewhat curved

toward apex; capsule broadly ovoid-oblong, 10-18 mm. long in type,

included or nearly, subligneous; seeds oval to nearly spherical, about

0.4 mm. long, dark brown, reticulate, the ridges irregularly wavy;

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

embryo straight; chromosome number 12 pairs. — Limb margin withers

with cup. The seed is the smallest in the genus; approaches through

var. Leguiana (Macbr.) Goodsp. I.e. 361 (N. Leguiana in Field Mus.

Bot. 8: 105. 1930), N. tomentosiformis Goodsp. I.e. 363, of Bolivia,

the throat cup of the smaller flower shorter than the cylinder, and

N. otophora Griseb., of Bolivia and Argentina, unique in withering

of limb margin in anthesis (Goodspeed). The variety (named in

honor of a former president of Peru) which replaces the typical form

in southern Peru and adjacent Bolivia has leaves chalky- or downy-

tomentose beneath to obscurely pubescent or nearly glabrous in age;

panicles usually compact, branches 12-60, rarely all remote; calyx

1-1.5 cm. long, ordinarily closely investing and shorter than the

cylindrical part of the corolla throat, the teeth variable; corolla limb

white, pink or red, deeply lobed, the lobes obtuse, acute or acuminate;

capsules at least half exserted. It seems to be a consistently and rea-

sonably well-established species occupying its own geographic area

in spite of its obviously close relationship to the three plants already

mentioned.

"N. colossea Andre" appears to be the most widely distributed

garden representative of the polymorphic variety. It is spontaneous

in gardens in Arequipa and cultivated in warmer botanical gardens

everywhere; Goodspeed, I.e. 363, has given a detailed description of

this horticultural race. All collections are the variety except those

from Huanuco. Illustrated, Goodspeed, I.e. 358, fig. 68; 360, fig. 69

(var.); Bot. Mag. 118, pi. 7252 ("N. colossea" as N. tomentosa).

F.M. Neg. 18237.

Huanuco: Near Chinchao, Mexia 04144- Maria del Valle, 3548.

Muiia, 3990; Ruiz & Pawn, type. Acomayo, Mexia 04110. Hua-

mincha, Woytkowski 34221. Puente Durand, Stork & Horton 9879. —

Junin: La Merced, Killip & Smith 24070 (distr. as N. Rusbyi). Tarma

to Carpapata, Stork 10964; Soukup 3964- — Huancavelica: Salca-

bamba, Stork & Horton 10281. — Ayacucho: Aina, Killip & Smith

23118. — Apurimac: Talavera, West 3736. Prov. Abancay, Good-

speed Exped. 10579. — Cuzco: Machupicchu, Herrera 3213; Goodspeed

4527. Valle de Santa Ana, Herrera 3616. Ollantaitambo, Pennell

13638. Paucartambo, Vargas 332; 333. Chuyani, Vargas 9744-

Towards Pillahuata, West 7104. Yacay, Soukup 769. Medias

Mayo, Vargas 11160. Cedrobamba to Maquina, Valle de San Mi-

guel, Herrera 1991 (type, N. Leguiana). — Puno: Near Sandia, Met-

calf 30716. Bolivia.

FLORA OF PERU 139

Nicotiana undulata R. & P. Fl. Peruv. 2: 16, pi. ISO, fig. b.

1799; 377. N. breviflora Poir. Suppl. Lam. Encycl. Meth. 4: 94. 1816.

N. undulata var. angustifolia Comes, Monogr. Nicot. 30. 1899.

Somewhat fleshy annual, to 2 meters high, the rather stout more

or less ridged velvety canescent stem often viscid above, the many

branches more slender; radical leaves rosulate, elliptic to lanceolate,

2-8 (18) cm. long, puberulent, the lowest petioles one-half as long,

progressively shorter to obsolete above, the upper leaves broadly

ovate or ovate-elliptic to lanceolate, sometimes cordate, usually ob-

tuse, ribbed and commonly corrugated, 8-20 (30) cm. long; panicle

axis and branches racemosely extended, the pedicels 1-3 mm., finally

2-6 mm. long; calyx nearly as long as corolla, frequently viscid, the

tube cupulate, membranes scant, teeth broad, obtuse or subacute,

one long and convex, the rest very short; corolla sordidly greenish-

yellow, usually with a blue-purple tinge, the tube proper 2-3 mm.

long and broad, the throat 3 times as long, 1.5-2 times as broad,

obconic-cylindric, clavate, pubescent, the limb 2-6 mm. wide, re-

flexed, irregular, the short broad lobes acute; 4 stamens barely in-

cluded, the filaments subgeniculate near insertion at base of throat

where spreading pilose, the shorter fifth filament glabrous or nearly,

arcuate; capsule broadly oval, 5-10 mm. long, included, the large

calyx segment galeate; seeds oblong or suboblong, about 0.8 mm.

long, nearly black, fluted reticulate, embryo straight; chromosome

number 12 pairs. — N. undulata is a polymorphic species; apparently,

a tall (averaging 1 meter) pyramidal form with larger corolla, limb

4-6 mm. wide, is common from the Cuzco region to near Cajamarca,

while southward from Lake Titicaca the average height approaches

5 dm. Habit is commonly a clump of stems, corolla tube is 2-3 mm.

wide and inflorescence is simpler; the species bridges subgenus Pe-

tunioides (as evidenced by the acute lobe, irregular greenish-white

corolla limb) and subgenus Rustica as shown in habit, in leaf-type

and in the velvety canescent stem (Goodspeed). Occurs in cold dry

lower puna, 2,700 to 4,200 meters. Illustrated, Goodspeed, I.e.

fig. 75. F.M. Negs. 18236; 18391.

Cajamarca: Toward Celendin, Metcalf 30781. — Lima: Chicla,

(Ball). Viso, 584- — Junin: Pampa, Bombon, Ruiz & Pavdn, type.

La Oroya, Stork 10445. — Huancavelica: Near Huancavelica, Weber-

bauer; Soukup 2773. — Ayacucho: Puquio to Lucanas, Metcalf 30328.

— Cuzco: Pisac, West 7173. Near Cuzco, Soukup 17; 18. San An-

dres, Vargas 9843. — Puno: Granja Saleedo, Vargas 9627; Soukup 902.

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

Near Puno, Soukup 632. — Tacna: Candarave, Weberbauer 7375. To

northwestern Argentina.

20. FABIANA R. & P.

In Peru a viscid-resinous compactly and fasciculately branched

shrub with almost minute somewhat imbricate leaves. Peduncles

solitary, the single tubular-funnelform flower 5-merous. Corolla

tube more or less ampliate, the plicate limb shortly lobed. Stamens

equal, included, the anthers longitudinally dehiscent. Stigma bi-

lamellate. Disk if present fleshy. Capsule included, 2-valved, api-

cally bifid.

Fabiana densa Remy, Ann. Sci. Nat. ser. 3, 8: 227. 1847; 589.

Bark rugose, the erect branches rather slender, somewhat flexu-

ose, puberulent; leaves many, alternate, sessile, narrowly linear,

slightly incrassate apically, 2-3 mm. long, scarcely 0.3 mm. wide;

pedicels axillary, shorter than leaves, cylindric, glandular; calyx

tubular-campanulate, segments lanceolate-linear, obtuse, granulose-

viscid-puberulent as the violet-green corolla, this three times longer,

narrowed medially, throat inflated, oval lobes subacutely narrowed ;

disk biparted, orange; style longer than stamens, inflated stigma

simple; capsule ovoid. — The Peruvian shrub is var. ramulosa Wedd.

Chloris And. 2: 95, pL 57. 1857, type a meter tall, the yellowish-

brown flowers about 1 cm. long.

Tacna: Tacora, (Weddell, type, var.). Candarave, Weberbauer

7363; 7363a. Tijaca, Raimondi. Bolivia.

21. NIEREMBERGIA R. & P.

Reference: R. Millan, Darwiniana 5: 487-547. 1941.

Herbs or ligneous below, the Peruvian species with filiform repent

stems, short ascending-erect branches. Flowers subsessile, solitary,

extra-axillary or opposite the entire leaves, the corolla tube slender,

elongate. Capsule valves deeply bifid. — Otherwise like Fabiana

R. & P., the ovary stiped. Juan Eusebio Nieremberg was a Spanish

naturalist (1595-1658).

Nierembergia repens R. & P. Fl. Peruv. 2: 13, pi. 123, fig. c.

1799; 498. N. spathulata HBK. Nov. Gen. & Sp. 3: 8. 1818, fide

Millan; cf. below.

FLORA OF PERU 141

Branches and younger solitary or subgeminate leaves slightly

puberulent; leaves elliptic-subspathulate, rounded or obtuse at apex,

narrowed to petiole, 1-nerved, glabrous in age; calyx tubular, pilose

below, the lanceolate segments acute; corolla white or lilac, glabrous,

the slender tube 4 times longer; limb abruptly dilated, subcampan-

ulate, plicately 5-lobed, the rounded lobes equal; capsule rounded-

ovoid. — The presumably Peruvian plant is var. pilosella Dunal in

DC. Prodr. 13, pt. 1: 583, 1852, petioles margined, 6-30 mm. long,

leaves smaller, subapiculate, 8-18 mm. long, 4-5 mm. wide, pilosu-

lous; corolla tube 2.5-3 cm. long, abruptly dilated into the campan-

ulate limb; seeds oblong, subangled, brownish. Corolla tube 3-6 cm.

long, limb 2.5 cm. wide (Millan). Millan has reasoned that it is an

error to accept Nicotiana minina Molina, Sag. Nat. Chile 153. 1782,

as the earliest name for this plant as proposed by Johnston, Contr.

Gray Herb. 70: 92. 1924; the plant of Molina is included without ques-

tion by Goodspeed in his list of insufficiently known species of Nicoti-

ana. However this may be, the occurrence of the plant in Peru is

to be verified, as apparently there is only the one doubtful record.

Likewise, the viscid N. petunioides Dunal, I.e. 506, with subfiliform

tube gradually dilated to the limb, is even more doubtfully Peruvian,

but in all probability it was collected in Chile and may indeed be a

species of Petunia. Possibly occurring is N. boliviana Millan, I.e.

537, pi. ISA, from adjacent Bolivia, an ascending (scandent?) plant

with viscid sublinear leaves and calyx lobes, glabrous corolla tube

8 mm. long, limb small; it may be Ule 9762 from Rio Acre, Brazil,

given an herbarium name by Bitter in reference to its habit. Francey,

in Diels' Contr. Flora Ecuador, Biblioteca Botanica 288. 1938, ob-

served that the leaves and flowers of Ecuadorean specimens repre-

senting species of HBK. are only a third to a half as large as those of

N. repens, ex char. Illustrated, Millan, I.e. pi. 1. F.M. Neg. 540.

Peru(?): Without locality, Dombey (Ruiz & Pavori). Ecuador?

Colombia? Chile.

22. SALPIGLOSSIS R. & P.

Leptoglossis Benth. Voy. Sulph. 143. 1844. Leptofeddea Diels,

Repert. Sp. Nov. 16: 193. 1919.

Slender-stemmed viscid-pubescent or nearly glabrous annuals or

half-shrubs with more or less evanescently subrosulate basal leaves,

the upper peduncled or sessile, narrow, entire (Peru) . Flowers tubu-

lar, with 5-lobed somewhat bilabiate spreading limb. Corolla often

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

net-veined. Fertile stamens 4 or 2 (Leptofeddea) or two reduced in

size (Leptoglossis') ; no botanical name is needed to honor the owner of

the useful serial in which Diels published his segregate "marked by

the contrasting basal and stem leaves."

The popular garden flower S. sinuata R. & P. of Chile may per-

sist; it is known by the generic name; its almost infinite variations

in color and corolla lobing have been studied cytologically as by Dale,

Amer. Journ. Bot. 24: 651-656. 1937, and by Dale and Rees-Leonard,

Genetics 24 : 356-367. 1939, who, curiously, did not investigate the

species.

Fertile stamens 4 (2 sometimes smaller) ; calyx glabrous.

Corolla colored; calyx straight S. acutiloba.

Corolla white with yellow center; calyx curved S. albiflora.

Fertile stamens 2; calyx puberulent or viscid.

Evanescent annual; calyx puberulent S. lomana.

Persisting but flowering as an annual; calyx viscid.

S. schwenkioides.

Salpiglossis acutiloba Johnst. Contr. Gray Herb. 85: 179. 1929.

S. linearis Johnst. I.e. 81: 96. 1928, not Hook.

Annual to 3 dm. high, inconspicuously glandular- villous at base

and on corollas without; rosulate basal leaves oblanceolate, 2-3 cm.

long, the cauline linear, 2-6 cm. long, 1-1.5 mm. wide, all entire;

flowers at branchlet tips in an irregular cyme; calyx 4-7 mm. long,

to 10 mm. long in fruit, the upper lobes longer; corolla tube orange-

yellow or greenish, more or less brown- or purple-veined, especially

the lobes, 17-20 mm. long, the tube 11-15 mm. long, to 0.5 mm. wide

at base, 2 mm. wide above, the lower lip lobes oblong-lanceolate,

bilobulate, to 2.5 mm. long, the upper 3 lobulate, the lobes narrowly

ovate, 2 mm. reflexed, 2 mm. long; 2 upper anthers perfect, filaments

villous toward base, lowest stamen (in type) also fertile but reduced,

stigma entire; capsule ellipsoid, to 4 mm. long, seeds angulate. —

Section Leptoglossis Wettstein, Pflanzenfam. 4, Abt. 3b: 36. 1891,

but corolla tube abruptly contracted along the base; the fifth stamen

may be present or absent in most species, as it probably is here

(author).

Arequipa: Tiabaya, Pennell 13081; 13063, type. Tingo, Pennell

13119. Yura, Vargas 7984. — Moquehua: Hills southeast of Moque-

hua, Weberbauer 71*56 (det. Johnston).

FLORA OF PERU 143

Salpiglossis albiflora Johnst. Contr. Gray Herb. 85: 178. 1929.

Similar to the preceding species but differs in lobes of white corolla

semiorbicular or broadly ovate, 2-3 mm. broad, ascending, calyx

somewhat curved, costate, shorter, and stigma emarginate, bilobed.

— The corolla is white with yellow eye; the stigma resembles that of

S. schwenkioides (author).

Ancash: Pampa Romas, Weberbauer 3178. — Lima: Matucana, 375

(det. Johnston). Near Chosica, Weberbauer 5325. San Bartolome',

Weberbauer 5297 (det. Johnston). — Moquehua: Mt. Estuquina,

Weberbauer 7424a, type. Between Moquehua and Torata, Weber-

bauer 7424- — Ayacucho: Near Nazca, Ferreyra 5465 (det. Cowan),

—lea: Huaytara, Weberbauer 5400.

Salpiglossis lomana (Diels) Macbr., comb. nov. Leptofeddea

lomana Diels, Repert. Sp. Nov. 16: 193. 1919.

Type to about 1 dm. tall, simple; basal leaves (petioles 2-5 mm.

long) more or less suborbicular, 3-8 mm. across, veinless, the stem

leaves near or with the cyme, linear-oblanceolate, to 12 mm. long,

1.5-2.5 mm. wide; cymes pubescent; calyx tube 4 mm. long, 3.5 mm.

wide, linear lobes 3 mm. long, 1 mm. wide; corolla lurid yellow or

cream-colored (Mexia), the tube about 1.5 cm. long, 4-5 mm. broad,

the lobes 2.5-3 mm. long; stamens 7-8 mm. long, staminodia 4 mm.

long; ovary 3 mm. long; style much dilated at apex, emarginate. —

The type a slender annual, the leaves mostly subrosulate near the

base. F.M. Neg. 3064.

Arequipa: Mollendo lomas, Weberbauer I486, type; Gunther 12246

(det. Morton). Mexia 7776 (det. Johnston); 04176.

Salpiglossis schwenkioides (Benth.) Wettst. Pflanzenfam. 4,

Abt. 3b: 36. 1891. Leptoglossis schwenkioides Benth. Voy. Sulph.

143. 1844.

Persisting as a perennial, ligneous below, herbaceous and, espe-

cially above, viscid-pubescent, branching; leaves sessile, lanceolate

or linear-lanceolate, acuminate, 1-1.5 cm. long or longer; flowers

crowded at branchlet tips, variously cymose; calyx 3 to (in fruit)

about 5 mm. long; corolla tube 15-18 mm. long, limb 6-8 mm. across,

fertile stamens 4, included; style apically dilated, lobes orbicular;

capsules membranous, valves finally bifid ; seeds many, subangulate.

La Libertad : Above Trujillo, 1,000 meters, Ldpez M. 0739. — An-

cash: (Weberbauer, 167). — Lima: Toward Huaral, south base of Hua-

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

choc Pass, 1,000 meters, Hutchison 1032. Huaman tango, (Barclay,

type). Sta. Eulalia, 1,200 meters, Goodspeed 33098. San Bartolome",

Weberbauer 5290. Without locality, (Mathews 1011; Cumings 1010).

Near Canta, Ferreyra 7255 (det. Cowan).

23. SCHWENKIA L.

Erect or scandent herbs or somewhat ligneous with alternate en-

tire leaves and racemose-paniculate long-tubular yellowish-green

flowers in the upper axils. Corolla limb erect, plicate or 5-denticu-

late, the minute or setaceous teeth often expanded into laciniform

entire or bifid appendages. Fertile stamens 2-4, anthers 2-celled,

coherent. Style subexserted, stigma entire. Disk cupulate or some-

times obsolete. — The almost filiform corolla base of the only species

in Peru distinguishes it handily from Salpiglossis and Browallia;

otherwise, the induplicate scarcely imbricate aestivation has been

accepted as its chief character.

Schwenkia grandiflora Benth. in DC. Prodr. 10: 193. 1846.

Becoming scandent and suffruticose toward base, branches terete,

finely striate and pilosulous as the petioles (6-10 mm. long) and leaf-

nerves beneath; leaves suboblong, membranous, to 6 cm. long or

longer, 3.5 cm. wide, acuminate, the uppermost much reduced; pan-

icles ample, lax, the bracts minute; pedicels filiform, scarcely as long

as calyx, this campanulate, about 3-4 mm. long, teeth linear, acu-

minate, 1.5 mm. long; corolla 2.5-3 cm. long, tube little ampliate to

limb, setosely denticulate and with much longer acuminate append-

ages; anthers 2, visible, reaching to tip of corolla lobes, style slightly

exserted; filaments pubescent; capsule papillose-punctate, valves mem-

branous; seeds few, striate, rugulose. — Flowers clear green (Schunke) ;

yellow (Klug).

Loreto: Rio Mazan, Jose Schunke 24. Near Iquitos, Klug 751.

To British Guiana.

24. BROWALLIA L.

Reference: Bentham in DC. Prodr. 10: 197-198. 1846.

Slender glabrous or viscid-pubescent annuals (or rarely peren-

nials) with thin entire alternate or subopposite leaves and salverform

solitary or racemose flowers. Calyx (in flower) narrowly to broadly

campanulate. Corolla limb oblique, slightly bilabiate, aestivation

FLORA OF PERU 145

imbricate. Perfect stamens 4, filaments ciliate to lanate, unequal.

Style sublobulate. — Frequently grows in cultivated lands and along

trails, the yellow-eyed blue, violet or white flowers attractive. The

Chilean Schizanthus pinnatus R. & P., with pinnate or bipinnate

leaves and purple-blotched white or lavender flowers with cleft or

laciniate lobes, may persist from cultivation, which probably was

the origin of the collection by Woitschach from Tacna.

John Browall was Bishop of Abo (Sweden) in 1743; originally an

intimate friend of Linnaeus, the latter, according to Hooker, Bot.

Mag. 58: 3069. 1831, immortalized their subsequent disagreement

by naming the three known species B. elata L., this to express the

degree of their union, B. demissa L., its cessation, and B. alienata

L. (Ruellia species) ; while these names intimate the uncertain char-

acters of the plant, they also imply the friends' differences; botanists

have often made known their disappointment in each other, as Ruiz

and Pavon and any number of our time, but rarely if ever so subtly!

It seems to me there is no clear concomitance of morphological

characters used to define species and therefore their genetic status is

questionable, in turn their taxonomic significance; the expediency of

my key will be apparent in the sorting of any considerable number

of collections; it may be useful to remark that as in so many annuals

with weedy tendencies, early and later vegetative developments may

differ greatly (one or more species surely persist as semi-ligneous

plants) and apparently size of flowers as well as luxuriance of entire

plant may depend at least partly on environment; sometimes corollas

considerably smaller may be among the last produced and there

seems to be a glandular and eglandular phase for at least some spe-

cies, as the character itself is variable. Obviously my scepticism of

the species is great — it may be too great.

[Mr. Macbride's scepticism is perhaps amply justified. We note

that Mr. Langhammer, who has a revision of Browallia in prep-

aration, has annotated most of our Peruvian specimens of the genus

as Browallia americana L. (with two subspecific designations). If

Mr. Langhammer is justified in making the reductions which his

annotations on our specimens would indicate, the Browallias of

Peru should be reduced to two species : B. americana (with two sub-

species) and B. speciosa.

Mr. Macbride could not be consulted when this publication was

ready to go on the press. In light of information not available to

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

Mr. Macbride when he saw the page proofs — and information sup-

plied by telephone by Conrad V. Morton, astute student of the

Solanaceae — it was decided to take out of the manuscript two un-

published specific entities which were credited to Mr. Morton.

The manuscript, other than the change mentioned and necessary

revision of the key to reflect the modification, has been left as

Mr. Macbride wrote it.

The specimens which Mr. Macbride cited under the two specific

entities which we have deleted from the Flora have been annotated

by Langhammer as Browallia americana L. (broad sense).

These specimens, as Mr. Macbride cited them, are:

Huancavelica: Surcubamba, Weberbauer 6496. — Ayacucho: Es-

trella, Kittip & Smith 23083, type. Puquio, Ferreyra 5467 (flowers

large); 22683 (flowers large, det. Standley, B. grandiflora). — Cuzco:

Tunguimayo, Balls. Potrero, Vargas 1750.

Piura: Amotape Hills, Haught 56. East of Talara, (Haught &

Svenson 11555). Piura to Nomala, Weberbauer 5942. La Brea,

Horton 11585. Negritos, Haught 8. Near Sullana, Haught 152.

East of Cape Parina, Haught 120. South of Paita, Haught 236.—

Lima: Obrajillo, (Cruckshanks, det. Hooker). Ecuador.

For those who may wish to treat the Peruvian species of Browallia

more conservatively the following key will distinguish the two specific

concepts maintained according to Langhammer's annotations on

our specimens. The concept of B. americana L. is, of course, in the

broad sense.

Corolla tube about 4 cm. long, the limb 4-5 cm. broad; calyx in

flower about 2 cm. long, distinctly asymmetrical, the lobes

linear-lanceolate and about half as long as the calyx tube.

B. speciosa.

Corolla tube mostly less than 2 cm. long, the limb about 2 cm.

broad; calyx in flower about 1 cm. long, the lobes narrowly

oblong to narrowly triangular or sometimes ovate, usually less

than half as long as the tube B. americana.

Editor's note.]

KEY TO BROWALLIA

Calyx lobes ovate-subulate, acuminate, subequaling tube; solitary

flowers to 4 cm. long, 5 cm. across B. speciosa.

FLORA OF PERU 147

Calyx lobes oblong-elliptic or suboval, obtuse to subacute, much

shorter than tube; pedicels more or less elongating in fruit, then

at least twice as long as calyx.

Floriferous branchlets or solitary flowers from or above often re-

duced sessile or subsessile leaves.

Pedicels, especially the lower, much longer in fruit.

B. peduncular is.

Pedicels, even the lower, rarely more than twice longer than

fruiting calyx B. Dombeyana, B. abbreviata.

Floriferous branchlets or solitary flowers from axils of little re-

duced shortly petioled leaves . . . .B. speciosa, B. grandiflora.

Calyx lobes ovate-lanceolate (or suboblong), acute to acuminate;

pedicels and calyx subequal even in fruit, the former unless the

lowest not or little elongating; leaves usually only slightly re-

duced above, sometimes remote (cf. B. speciosa).

Calyx lobes somewhat divergent or pedicels approximate, the

latter sometimes a little longer in fruit.

Pedicels at least as long as calyx; leaves glabrate.-B. grandiflora.

Pedicels shorter than calyx; leaves hirtellous B. viscosa.

Calyx lobes erect or suberect; solitary leafy-bracted pedicels little

longer in fruit B. americana.

Browallia abbreviata Benth. Voy. Sulph. 144. 1844.

Similar to B. grandiflora Graham but typically distinguishable

by the short broadly campanulate calyx, the oval or ovate-oblong

divisions early subequaling the tube. — In the Dombey specimen (in

flower, Herb. DC.) the calyx lobes are about half as long as the tube!

It is B. Dombeyana in part but the leaves are petiolate; Metcalf

30320 has calyx 6 mm. long, broad teeth scarcely 2 mm. long, leaves

sessile to petioled; calyx as in B. peduncularis often purple-lined or

-spotted; these specimens may be phases of B. peduncularis Benth.

or (and) B. viscosa HBK. depending on the significance of the genetic

characters. The calyx is about half the size of B. viscosa. F.M.

Neg. 7787 (Dombey).

La Libertad: Hacienda Campoden, Nunez 437 (toward B. vis-

cosa in calyx). — Lima: Huaman tango, (Barclay, type). Near Lima,

(Cuming 1078}. Puruchuco, (Mathews 712). Chosica, Weberbauer

5348 (det. Johnston, B. viscosa). Prov. Canete, Vargas 9317. Viso,

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

Goodspeed Exped. 11527. Lomas of Asia, Grant 7494. Matucana,

194 (distr. as B. viscosa); Raimondi (det. Werdermann, B. demissa).

Toward Canta, Ferreyra 7267 (det. Cowan). Valle de Chillon, Nunez

909. Surco, Ferreyra 3456; 6104 (det. Cowan, B. viscosa). Without

locality, Dombey 321 (det. Bentham). — Huanuco: Edge of cultivated

field, 4217; 1459 (toward B. viscosa). — Ayacucho: Puquio to Luca-

nas, Metcalf 30320 (det. Morton). — lea: Above Pisco, Weberbauer

5398.

Browallia americana L. Sp. PI. 631. 1753. B. demissa L. Syst.

ed. 10. 1118. 1758-59.

Slender hollow-stemmed annual, lightly pilose or essentially gla-

brous, rarely slightly glandular even to the calyx; pedicels about

5 mm. long in fruit, the calyx teeth then ovate, acute, less than half

as long as the tube or about 2 mm. long; corolla to about 2 cm. long;

seeds brown, densely foveolate. — Stems and leaves soft, the flowers

solitary (Svenson). Description from Raimondi 1871; all his speci-

mens det. Werdermann.

Tumbez: Toward Matapato, Ferreyra, Cerrate & Tovar 10682.—

Cajamarca: Cascas, Raimondi. — San Martin: Moyobamba, Rai-

mondi 1871; King 3376 (det. Morton); Woytkowski 35305 (det. Cua-

trecasas); San Roque, Williams 7126; 7167; 7407; 7622.— Lima:

Hills of Lurin, Ruiz & Pavdn. — Huanuco: Cuchero and Pillao, Ruiz

6 Pavdn. — Pasco: Quillasu, Soukup 3302 (det. L. B. Smith). — Junin:

Biscaton, Raimondi. — Loreto: Iquitos, Williams 8105. Widely dis-

tributed in subtropical South America.

Browallia Dombeyana Damm. Bot. Jahrb. 37: 642. 1906.

Branches scarcely 1 mm. thick; leaves sessile or nearly (petioles

to 1 mm. long), ovate-lanceolate, obtuse, 8-15 mm. long, 3-8 mm.

wide, glabrous; pedicels 1-2 mm. long, pulverulent; calyx campanu-

late, minutely glandular pilose as corolla, the oval lobes scarcely

1 mm. long; corolla hypercrateriform, tube cylindric, 1.5 cm. long,

1 mm. in diameter, throat 3 mm. long, 2 mm. in diameter, lobes

unequal, obtuse; stamens 4, filaments 2 mm. long, pilose; ovary api-

cally pilose, style 1 cm. long, stigma broadly lobed. — The 2 upper

filaments connate in type but doubtfully always (author); almost

certainly a phase of B. abbreviata. F.M. Neg. 3050.

Lima: Cerro Amancaes, Raimondi (det. Werdermann); Ferreyra

4048 (det. Cowan). Matucana, Raimondi. Lurin, Pachacama,

Ruiz & Pavdn; (Dombey, type).

FLORA OF PERU 149

Browallia grandifiora Graham, Edinb. Phil. Journ. 10: 168.

1830. B. cordata G. Don, Gard. Diet. 4: 478. 1838.

Diffuse (at least in cultivation) branching annual, the soft stems

and leaves usually glabrous or the upper as the calyces early viscid-

or glandular- villosulous or -puberulent; leaves ovate, acute or acumi-

nate, more or less attenuate or perhaps sometimes subcordate at base,

the nerves prominent beneath; flowers solitary on straight peduncles

in the axils of the reduced subsessile upper leaves, thus also laxly

foliose-racemose at the ends of branches; calyx 5-lobed, nervose,

about 1 cm. long, the segments sometimes spreading, typically ob-

tuse or retuse, usually shorter than tube; corolla tube at least 2 cm.

long, slightly ampliate, in type glandular puberulent, greenish-yellow,

dark striate, limb white or pale lilac with yellow throat, bilabiate,

3 cm. across or larger; upper filaments dilated, rather recurved, pu-

berulent, exceeding the glabrous apically curved style, the green

peltate stigma obscurely lobed. — The black seeds of specimens de-

termined as this species are striately ridged-tuberculate. Illustrated,

Bot. Mag. 58: pi 3069; Bot. Reg. 16: pi 1384. (B. cordata?).

Piura: Canchaque, Ferreyra 3153 (det. Cowan with query). —

Lima: Yazo near Canta, (Cruckshanks, type, from seeds); Weber-

bauer 5295. Puruchuco, (Mathews 511). Chosica to Matucana,

Goodspeed 113321 (det. Johnston); 209; 2896. Rio Santa Eulalia,

Goodspeed & Stork 11485; Horton 10999. Rio Rimac, Goodspeed &

Metcalf 30235 (det. Morton). Rio Chillon, Pennell 1U87 (toward

B. peduncularis) . Surco, Soukup 3733; Weberbauer 5210; Ferreyra

6939. — Huanuco: Huamincha, Woytkowski 34204- (det. Cuatrecasas).

— Junin: Utcuyacu, Woytkowski 35361 (det. Cuatrecasas, B. viscosa).

San Ramon, Constance & Tovar 2277. Satipo, Soukup 2860.

Browallia peduncularis Benth. Bot. Voy. Sulph. 143. 1844.

Similar to B. grandiflora but leaves (typically) scabrous-hirtellous

both sides, the upper sessile; flowers few in reduced leaves, axillary

or loosely racemose; calyx-teeth oblong typically, about equaling the

tube, usually shorter; peduncles mostly elongate, 2.5-8 cm. in fruit.

— Apparently always viscid-pubescent. The very sessile or clasping

leaves are noteworthy in type but character apparently variable;

probably a phase of B. abbreviata or B. grandiflora Graham. F.M.

Neg. 23042 (Mathews).

La Libertad : Usquil, Prov. Atuzco, Ferreyra 7361 (det. Cowan) .

—Lima: Huamantanga, (Barclay, type). Puruchuco, (Mathews 512,

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

det. Bentham). San Buenaventura, Pennell 14514; 14517; 14569.

Obrajillo, Wilkes Exped. Canta, Velarde Nunez 920. Trapiche, Prov.

Canta, sandy wash, Hutchison 1021. Lomas Patirilca, Goodspeed

Exped. 9229. Ambar, Goodspeed Exped. 9229. Quive, Pennell

14311.

Browallia speciosa Hook. Bot. Mag. 73: pi. 4339. 1847.

Somewhat suffrutescent, erect, branched, glabrous or essentially;

petioles stout, 5-10 mm. long; leaves ovate-elliptic, acute at base,

acuminate, 5 or 6 cm. long, 2-2.5 cm. wide, nerves prominent be-

neath; peduncles solitary, about 2 cm. long in flower; flowers to 4 cm.

long, 5 cm. across; calyx lobes subalate or ovate, long-acuminate,

10-12 mm. long, nearly as long or longer than the tube; corolla tube

at least 2.5 cm. long, lobes ovate, acute or some broadly acuminate,

1-1.5 cm. long, striate, dark purple above, pale beneath, the throat

white; filaments ciliate; style nearly as long as tube, stigma 2-lipped,

4-lobed within the lip (Hooker). — Distinguished by author from

B. grandiflora Graham by the larger flowers, the segments not re-

tuse nor bifid.

Huanuco: Pozuzo, Raimondi (det. Werdermann). Colombia.

Browallia viscosa HBK. Nov. Gen. & Sp. 2: 373. 1818.

Diffuse herb several dm. tall, the attenuate branches terete, the

younger as the peduncles and calyces viscid-hirtellous; petioles about

1 cm. long; leaves subrotund-ovate, obtuse, 2.5- nearly 4 cm. long,

2-3 cm. wide, membranous, reticulate- veined, hirtellous both sides;

peduncles solitary in the axils, approximate toward apex of branch-

lets, in fruit 6 mm. long; flowers violet, 18 mm. long, calyx violet-

tinged, lobes lanceolate, erect; tube of glabrous corolla twice as long

as calyx, lobes unequal, obovate, emarginate, the larger white-

spotted at base; filaments pilose as ovary, the capsule pubescent

apically; seeds brown, angled, scabrous. — After authors; Bentham

noted calyx teeth as subdivergent, oblong, shorter than tube, ped-

icels and calyx subequal, upper racemose; this seems to describe the

Peruvian specimens and may suggest that they merge with B. grandi-

flora Graham; the following specimens are less pubescent than the

Ecuadorian (type).

Cajamarca: Cutervo, Jelski 162; Raimondi (det. Werdermann).

Huambos, Soukup 4507 (det. Cowan). Monte Seco, Soukup 3854-

— Amazonas: Chachapoyas, (Mathews 3174, fide Bentham). — Lima:

FLORA OF PERU 151

Canta Valley, (Cruckshanks) . Pillao, Lima & Cheuchin, Ruiz &

Pawn (in Herb. Madrid). Between Matucana and Surco, Weber-

bauer 5210. — Huanuco: Valley of Rio Chinchao, Stork & Horton 9430;

9867 (both det. Standley, B. americana). Mito, 3426. — Junin: Hua-

capistana, Killip & Smith 21*125. Tarma, Raimondi (det. Werder-

mann). Ecuador.

25. STREPTOSOLEN Miers

A finely scabrous pubescent or hirsutulous shrub related to Brow-

allia and Brunfelsia but with yellow and burnt orange flowers, the

corolla tube spiralled, especially toward the greenish base.

Streptosolen Jameson! (Benth.) Miers, Ann. Mag. Nat. Hist,

ser. 2, 5: 209. 1850. Browallia Jamesoni Benth. in DC. Prodr. 10:

197. 1846.

Usually leafy, 1-2 meters high; leaves ovate-elliptic, rugose, about

(1) 1.5- about 3 cm. long, 9-15 mm. wide; flowers 2 cm. long, in sub-

corymbose cymes. — Inflorescence strigillose; in the var. hirtella Vatke,

hirsutulous (Weberbauer 4152, Cajamarca). Corolla is orange ex-

cept for greenish tube, and the fine white trichomes are glandular

(collectors).

Piura: Abra Porculla Pass, Prov. Huancabamba, 2,100 meters,

Hutchison 1384- Valley of the Quiros, Prov. of Ayavaca, Weberbauer

6344; Soukup 4339. Canchaque, Ferreyra 3137. — Cajamarca: Cu-

tervo, Raimondi (det. Werdermann). Callocate, Jelski (det. Zahlbr.).

Chugur, Weberbauer 4152. Socota, Stork & Horton 10117 (det. Stand-

ley). Huambos, Soukup 4499 (det. Cowan). Ecuador; Colombia.

26. BRUNFELSIA [Plum.] L

Smooth shrubs or small trees with entire leaves and showy ter-

minal or clustered flowers. Corolla similar to Browallia except in

size. Perfect anthers 4. Fruit fleshy or coriaceous, tardily or not

dehiscent. — Otto Brunfels was a German physician and botanist of

the sixteenth century. Monachino, Phytologia 4: 342-347. 1953,

gave a clear and succinctly organized list of the species names, noting

probable validities, in connection with his new species B. mire of

adjacent Bolivia — a helpful account, closing with the pertinent re-

mark: For a true understanding of the species of Brunfelsia a critical

revision of the whole genus is required.

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

Leaves oblanceolate.

Corolla tube 2.7-3.5 cm. long; calyx glabrescent, to 14 mm. long.

B. mire.

Corolla tube about 2.5 cm. long; calyx pubescent, longer.

B. hydrangaeformis.

Leaves elliptic or oboval-oblong.

Pedicels to 2 cm. long; corolla lobes narrow B. maritima.

Pedicels typically short; corolla lobes broad.

Corolla tube 2-2.5 cm. long B. bonodora.

Corolla tube 3-4 cm. long B. grandiflora.

Brunfelsia bonodora (Veil.) Macbr. Field Mus. Bot. 8: 112.

1930. Besleria bonodora Veil. Fl. Flum. 261. 1825. Icon. 5. pi. 80.

1827. B. latifolia (Pohl) Benth. in DC. Prodr. 10: 199. 1846. Fran-

ciscea latifolia Pohl, PL Bras. Icon. 1: 3, pi. 1. 1827.

Branchlets glabrous, sub terete, longitudinally rugose; petioles to

4 mm. long; leaves ovate-lanceolate or oblong, often somewhat obo-

vate, mostly acuminate, more or less abruptly contracted at base,

apically crowded, subundulate, firm-membranous, glabrous or ob-

scurely pubescent on nerves beneath, minutely punctate, often 1-

2 dm. long, at least half as wide, nerves rather prominent, veins

obviously reticulate; peduncles usually developed; pedicels 4 mm.

long or longer; calyx tubular-campanulate, membranous, drying cori-

aceous, reticulate-veined, the slender corolla tube at least twice

longer, this curved above and with spreading rounded violet or white

lobes; capsule fleshy (Bentham). — Mostly after J. A. Schmidt in

Mart. Fl. Bras. 8, pt. 1: 258. 1862. As implied by Monachino, the

name of Vellozo may not apply to the Peruvian tree, the name, as

he remarks, transferred by me, "following the leader" (in this case

Bentham) without judgment as to the extent of its application; ap-

parently it is the earliest name for a species, perhaps only of Brazil,

with a lax inflorescence; perhaps then the following collections are

wrongly determined or require a new name, but the species may be

more variable than as yet realized; most of the following specimens

have pedicels 5 (-10) mm. long, calyces 5-10 mm. long, corollas with

tube 2-3 cm. long; mostly referred by Standley (often with query)

to B. maritima Benth., questioned also by Morton.

San Martin: Moyobamba, Klug 3326; Raimondi (det. Werder-

mann); Mathews (det. Bentham, B. latifolia). Tarapoto, Williams

5482; Ferreyra 7778. Near Bellavista, Ferreyra 4760; 4812—LoTeto :

FLORA OF PERU 153

Yurimaguas, Killip & Smith 28056; 28178. Masisea, Hobroyd 19.

Rio Mazan, Jose Schunke 8. Rancho Indiana, Mexia 6444- Balsa-

puerto, Klug 2880. Bolivia; Brazil. "Chiric-sanango," "z(s)anango."

Brunfelsia grandiflora D. Don, Edinb. Phil. Journ. 86. 1829.

Small glabrous tree, the leaves apically crowded or scattered on

the flowering branchlets; petioles stout, 5-10 mm. long; leaves sub-

coriaceous, oblong-elliptic, slightly narrowed to acute base, shortly

and subacutely cuspidate, 1.5-2 dm. long, 5-8 cm. wide, the 5-7

slender lateral nerves and laxly reticulate veins somewhat conspicu-

ous only beneath; bracts caducous; pedicels 1 cm. long; calyx 1.5-

2 cm. long, the ovate teeth obtuse or mucronulate; corolla tube

3.5-4 cm. long, the limb 3-5 cm. across, the lobes subrotund. —

Bailey & Raffill, Standard Cycl. Hort. 581. 1914, referred this to

B. calycina Benth., var. macrantha (Lem.) B. & R., but from a study

of cultivated plants. The genus needs revision. The type locality

is near Dept. Huanuco. Cuzco and Puno specimens have smaller

flowers but calyces about 1.5-2 cm. long.

San Martin: Uchiza, (Ruiz & Pavdn, type). — Huanuco: Divisoria,

Woytkowski 34536 (det. Cuatrecasas) ; Ferreyra 985. — Cuzco: Echa-

rate, Goodspeed Exped. 10461. — Puno: Near Puno, Soukup 446 >

Brunfelsia hydrangaeformis (Pohl) Benth. in DC. Prodr. 10:

195. 1846. Franciscea hydrangaeformis Pohl, PI. Bras. Icon. 1: 717.

1827.

Similar to B. mire but with somewhat glandular hirsute calyx,

2.5 cm. long or longer, longer corolla, also slightly glandular and,

especially, prominently nerved and finely reticulate-veined. — The

Peruvian collection, not seen since Monachino's study, may rather

be referable to his species. Illustrated, Bot. Mag. pis. 72, 4209.

Loreto: Tierra Blanca, middle Maranon, Tessmann 4921 (det.

Werdermann). Bolivia? Brazil. "Chirisanangu" (Tessmann).

Brunfelsia maritima Benth. in DC. Prodr. 10: 200. 1846.

Type glabrous; leaves subsessile, rounded (rarely narrowed) at

base, scarcely acuminate, less than 5 cm. long; cymes few-flowered;

calyx lax, turbinate-campanulate, shorter than 2.5 cm., slender co-

rolla tube half again as long. — Type from Taipu (Rio Janeiro) by

Lund. J. A. Schmidt, in Mart. Fl. Bras. 8, pt. 1: 258. 1862, referred

here Spruce 1495 and other collections from the upper Amazon, some

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

with much longer pedicels and with pubescent leaf -nerves; species is

included only because several Peruvian specimens have been referred

here, as Mexia 6444-, Schunke 8, from Loreto, with query by Standley.

Cuzco: Echarate, Goodspeed Exped. 10461? (cf. B. bonodora).

Brazil.

Brunfelsia mire Monachino, Phytologia 4: 342. 1953.

Glabrous (unless the bracts and flowers), the tube of the violet

hypocrateriform corolla about twice the length of the campanulate

calyx, this at most 14 mm. long; petioles 3-8 mm. long; leaves oblan-

ceolate, narrowed to base, acuminate, 18-26 cm. long, 5-8.5 cm. wide,

subcoriaceous, pale beneath with 12 or 13 pairs of lateral nerves, the

vein reticulation prominent, areolae broad; flowers many, crowded,

bracts sparsely and minutely pubescent, pedicels glabrous, to 8 mm.

long; calyx reticulate venulose, lobes about 3 mm. long, corolla tube

to 3.5 cm. long, slightly curved above, glabrous or subglabrous, limb

3.2-5 cm. across. — Has been confused with B. hydrangaeformis (Pohl)

Benth. which it resembles superficially in leaves; it differs in the vena-

tion, the smaller glabrescent calyx, the longer extended corolla; the

affinity is rather with B. maritima Benth. and relatives (author);

possibly the specimen referred by Werdermann to B. hydrangaeformis

(Pohl) Benth. is rather referable here.

Plant used in Bolivia as a paralysant of the voluntary muscles

and against cutaneous parasites; see Rusby, Journ. Am. Pharm.

Assoc. 13: 101-102. 1924; Youngken, I.e. 14: 195-200. 1925 (Mona-

chino) .

Peru (possibly). Bolivia. "Mire."

27. SOLANUM L.

Lycianthes (Dunal) Hassl. Ann. Cons. Jard. Bot. Geneve 20:

180. 1917, also Bitter, Abh. Nat. Verein Bremen 24: 292-520. 1919,

fide Morton, Contr. U. S. Nat. Herb. 29: 54. 1944. Lycopersicum

Miller, Gard. Diet. Abr. ed. 4: 3. 1754, fide Wettstein, Pflanzenfam.

VI, Abt. 3b: 24. 1891.

Reference: Dunal, in DC. Prodr. 13, pt. 1: 27-387. 1852.

Plants highly varying in duration, habit and vegetative char-

acters, but ordinarily calyx entire to 5-10-apiculate, dentate or

bilobate, corolla rotate to campanulate, stamens equal or unequal,

more or less connate or connivent, anthers dehiscing by pore, chink

or slit, berry most often globose.

FLORA OF PERU 155

For practical purposes of identification — the basic purpose of

this work as stated in the preface — the group divisions, except

Tuberarium (to be contributed later) and Lycianthes (after Morton)

have not been defined here as proposed or adapted by Dunal because

they are vague in greater or less degree, due no doubt to lack of

information and/or understanding. Morton, U. S. Nat. Herb. 29:

41. 1944, in reference to Dunal wrote: The same species is some-

times described under different names in different sections. Bitter

in his incomplete studies made some progress in more natural de-

marcation of certain groups but he rarely clarified the classification

of many related Peruvian species; it is of interest that he himself,

even as I, in preparing an account of the species of one country

presented a partly expedient key. Morton, I.e. 54-55, remarked

that the two subgenera are the natural cleavage, yet also in these

principal divisions (here three with inclusion of Lycopersicum) there

are a few exceptions of complete concomitance of characters con-

sidered to be diagnostic. Wettstein, I.e., designated five sections,

including the three subgenera; these are not always in usable con-

trast to each other but the following subdivisions approximate them.

Apart from anther dehiscence he regarded equal or unequal filaments

as indicating relationship, also presence of prickles; neither in them-

selves are of value, occurring, as simple or divided leaves, in other-

wise probably similarly derived species. Probably no single char-

acteristic is anywhere reliable, always constant, in this large genus;

certainly, for example, position and nature of inflorescence is obscure

or secondary in merit. Pubescence absent to richly presented in

great variety, often more than two or more sorts intermixed, has

of course been the basis or an associated character for species defini-

tion; apropos is the paper by Reed C. Rollins, Rhodora 60: 145-152.

1958, illustrating the taxonomic insignificance of presence or absence

of trichomes — at least often; he observes that characteristics under

control of the simplest gene systems are expected to be least reliable

as a basis for classification; obviously proposal of new species in

this group would have contributed nothing to their ultimate correct

classification; reluctantly here and of course in all of this work

I have had to retain species proposed by my contemporaries in

circles into which angels would surely fear to tread. But perhaps

the following groups as devised here, while only general in their

connotation, will facilitate determination of the Peruvian species

proposed to date (1958), which, so graphically observed by Morton,

was impossible from the (nevertheless meticulous and detailed)

pioneer work of Dunal.

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

In the berries of some species there occur "stone granules"

possibly roughly similar in origin to the stone cells in fruits of

Pyrus (Morton). Bitter thought that their presence or absence,

number, position and size were constant in a given species; Morton

doubted the taxonomic significance of this character, in any case,

still to be proved, but Stebbins and Paddock, Modrono 10: 70-81.

1949, in an incompleted study of the Solanum nigrum complex

used it as a supplementary aid to specific identification; their work

is the most important contribution that has come to my attention;

they note the need of extensive field work and the difficulty of prac-

tical application (as regards the granules) in the study of herbarium

materials, an opinion entirely in accord with mine. Bitter's prin-

cipal accounts of stone cells in various genera and species are in

Abh. Nat. Verein Bremen 23: 114-163. 1914, and Bot. Jahrb. 45:

483-507. 1911. Only genetic investigations similar to that of

Stebbins will substantiate or disprove the validity of many of the

characters now used for group and species demarcation; so, ex-

pediency has compelled me to key out with doubt many plants

cited as species.

An informative illustrated account of the ornamental species

grown in Argentina has been prepared by Raul Martinez Crovetto,

Technical Publication 26 from the Revista de Invest. Agric. 2:

179-196. 1948; it includes eleven species, among them S. auriculatum

Aiton, S. melongena L., S. Rantonnetii Carr., S. dulcamara L., S.

Seaforthianum Andrews, S. capsicastrum Link, and S. pseudocap-

sicum L., all of which are probably to be found in Peruvian gardens.

In Gardener's Chronicle for 1940 E. K. Balls has an interesting paper,

"Potatoes and Other Plants in the Andes."

Species probably not Peruvian or not noted in the following

text included, cymosum Herrera, Contr. Fl. Dept. Cuzco 2: 168. 1921,

described as an annual, leaves deeply pinnatifid, flowers blue, used

as a sudorific and laxative under the name "Kusmaillu." S. havanense

Jacq. Enum. Carib. 15: 159. 1760, of Cuba, recorded by Ruiz and

Pavon, Fl. Peruv. 2: 39. 1799, from Acobamba and Picoy near

Tarma must be an error in determination. S. macrophyllum Dunal,

Hist. 199. 1813, is S. lancifolium Miller of Mexico, fide Schulz

in Urban, Symb. Ant. 6: 243-244. 1909. S. micranthum Willd. ex

Roem. & Schult. Syst. 4: 663. 1819, as to Peru is some other species

according to Schulz, I.e. 166, but to it Sendtner in Mart. Fl. Bras. 10:

19. 1846, referred a collection by Poeppig from Manaos, this some

species of section Leiodendron.

FLORA OF PERU 157

1. Subgenus LYCOPERSICUM

Anthers fusiform, connate, the oblong cells sterile-appendaged to

base; leaves pinnate or bipinnate.

2. Subgenus PACHYSTEMONUM

Anthers stoutly cylindric or ovoid-ellipsoid, not or little elongate

nor attenuate, at least only obscurely; pores terminal, often

oblique and subequaling the cell-diameter, usually during an-

thesis becoming lateral chinks, rarely extending to base. Species

all unarmed except one species of Lycianthes, with branchlets

sometimes apically indurate.

KEY TO SECTIONS OR EXPEDIENT GROUPS (SERIES)

OF SUBGENUS Pachystemonum

Section Tuberarium1

Leaves mostly unevenly pinnatifid, pedicels clearly articulate at

or above base, subterranean stems often tuberous. — An excep-

tion in Peru is S. muricatum Aiton, the leaves entire; several

species apparently are not tuberiferous; S. montanum L. with

merely repand leaves has a tuber-like root but is allied to

species listed in the next group.

Section Morella (Dulcamara, part)

Leaves entire or somewhat lobed but never alately decurrent to

stem (often more or less decurrent on petioles) and lobes never

alately joined as in some species of next group with more or

less divided leaves; flowers minute to about 1 cm. long, few

to many in pedunculate lateral inflorescences; anthers early

apically bipored; annuals or perennials, rarely partly ligneous,

often diffuse (Dulcamara, mostly Morella, key, p. 164).

Series 3

(Artificial group of various species, mostly compound leaves)

Leaves usually at least in part compound (ternate, pinnate, multi-

pinnate) sometimes only more or less divided, the divisions

1 Text for this section is being contributed. When available, it will be published

separately in continuation of the present number. (Ed.)

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

alately joined, or if merely repand-lobed always alately decurrent

on stems (S. phyllanthum) or root tuberiform (S. montanum) ;

calyx sometimes truncate, teeth minute; anther pores small or

medium, terminal or shortly lateral; plants various, sometimes

elongate, diffuse or subscandent. — See Leiodendron for related

species with entire leaves and Cyphomandra in fruit. In part

Polybotryon (Bassovia), Regmandra, Dulcamara (key, p. 177).

Series 4

(Mostly Leiodendron, sens, lat.)

Shrubs or small trees, virgate half-shrubs, rarely diffuse, or sub-

scandent, then simple or nearly, a few vines, never prickly,

leaves entire; inflorescence various, calyx usually well-lobed or

dentate, not truncate (Peru) ; anther pores ordinarily large, ob-

liquely introrse. Entire-leaved Dukamara, Polybotryon, trailing

or 4-merous Annarichomenum, Anthoresis, Anthopleuris, Leioden-

dron (key, p. 191).

Section Lycianthes

Calyx truncate but usually with 5 or 10 more or less developed teeth,

these sometimes minute or obsolete; erect or scandent shrubs,

rarely repent herbs; leaves entire, often geminate; flowers axil-

lary or subaxillary, 1-several, infrequently on a short peduncle;

anther pores various (Lycianthes, key, p. 225).

3. Subgenus LEPTOSTEMONUM

Anthers cylindric, usually long- or clearly attenuate, the pores

minute or small, terminal or sometimes widening or rarely

elongating; plants mostly aculeate (key, p. 242).

1. Subgenus LYCOPERSICUM

Annual or perennial herbs with pinnate or bipinnate leaves,

yellow flowers in simple extra-axillary racemes or furcate cymes,

pedicels articulated near or above the middle, but, all species con-

sidered, marked only by the development of an apical sterile tip

to the anther cells, these dehiscent their entire length as some-

times in otherwise similar species (L. lycopersicoides, S. Pennellii,

section Tuberarium); the former has been hybridized with S. Lyco-

persicum L. cf. Rich. Proc. Nat. Acad. Sci. 37: 741-744. 1951;

Correll, Modrofio, 14: 232-236. 1958.

FLORA OF PERU 159

KEY TO SUBGENUS Lycopersicum

Reference: Cornelius Muller, U. S. Dept. Agric. Misc. Publ. 382:

1-28. 1940.

Leaflets all or mostly crenate, lobulate, or incised, pubescent or

puberulent.

Plants canescently puberulent; corolla at least 1 cm. long, orange-

yellow; berry (as seeds or these only at tip) pubescent.

S. peruvianum.

Plants greenish, more or less divaricately hirsute; corolla yellow;

berry or seeds or both pubescent.

Stems slender, sprawling or lax; corolla lobed.

Leaves bicolor; berry glabrous; seeds usually lanate.

S. lycopersicum.

Leaves concolor; berry pubescent; seeds subglabrous.

S. Corneliomulleri.

Stems stout, erect or trailing; corolla subentire or acuminate-

lobulate S. agrimoniaefolium.

Leaflets all entire or subentire, glabrate or puberulent; corolla small,

usually yellow, lobed; berry usually glabrous as seeds.

S. pimpinellifolium.

Solarium agrimoniaefolium R. & P. ex Dun. in DC. Prodr. 13,

pt. 1: 24. 1852. Lycopersicum hirsutum Humb. & Bonpl. in Dun.

Sol. Syn. 4. 1816; 21, not S. hirsutum Dunal, I.e. L. agrimoniae-

folium R. & P. ex Dun. I.e. 24.

Well-marked typically by the abundant hirsute pilose indument,

most trichomes 2.5-3.5 mm. long, shorter, glandular and a few

appressed stellate ones intermixed; pseudo-stipules present; larger

leaflets petiolulate, medium and smaller sessile, all subentire and

usually repand-dentate; cymes dichotomous, all parts bracted unless

the 10-12 pedicels, these early filiform, 1.5-2.5 cm. long, often

abruptly deflexed below the flower after anthesis; calyx lobes 8-9

mm. long, acuminate or finally obtuse and 1.5 cm. long, the lobes

3-4 mm. wide; corolla yellow, rotate, about 3 cm. across, the broad

acuminate tipped lobes 5-6 mm. long, 6-8 mm. broad; anther sacs

7 mm. long, the appendages 4 mm. long, rather firmly joined,

staminal column 3-4 mm. thick; style more or less exserted; fruit

1.5-2.5 cm. in diameter, coarsely long-hirsute, greenish-white,

purplish-striped; seeds dark brown, glabrous except minutely at

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

apex. — L. hirsutum forma glabratum Muller, 22, from Ecuador,

besides being less pubescent, has very slender calyx lobes, corolla

2 cm. across, lobes 3-4 mm. broad. Illustrated, Muller, I.e., pis. 3

and 9. F.M. Neg. 2595.

Cajamarca: Inca baths, (Blood & Tremelling 136}. Hualgayoc,

Soukup 388. Cascas & Nancho, Raimondi (det. Werdermann). —

Ancash: Chacchan, 2553. — Lima: Matucana, Weberbauer 79. Huara,

Goodspeed 11342. Ambar, Stork 111+63 (det. Johnston). Canta,

Soukup 2826. Surco, Soukup 3707. — Junin: Incahuasi, Soukup 3196.

Ecuador.

Solanum Corneliomulleri Macbr., nom. nov. Lycopersicum

glandulosum Muller, U. S. Dept. Agric. Misc. Publ. 382: 23. 1940,

not S. glandulosum R. & P.

Character in general that of S. peruvianum but not at all canes-

cent; stems densely pubescent, the trichomes of three types, the

first short, 1-celled, mostly simple, the second capitately resinous-

glandular, the third elongate attenuate, rarely capitately stellate;

pseudo-stipules at base of leaves similar to the leaflets, these (larger)

5 or 6 pairs erosely dentate or round-lobate, the minor ovate 1^

mm. long, entire or minutely erose, all closely puberulent, the tri-

chomes 1- or 2-celled, very sparsely stipitate glandular, the rachis

glandular pubescent like the stems; inflorescence rarely simple, 6-12-

flowered, pedicels 10-18 mm. long, to 2.5 cm. long in fruit, the

commonly distributed bracts 5-12 mm. long; calyx 5-parted nearly

to base, narrowly lanceolate lobes 5-6 mm. long, in fruit 12-15 mm.

long, slightly glandular; corolla about 2 cm. across, medially parted

into broadly triangular reflexed lobes, apically subcaudate; staminal

columns 7-10 mm. long, the appendage tube recurved, 2-3 mm.

long; ovary puberulent, especially at apex; berry globose, 10-12

mm. in diameter, pale with lavender stripes about the top where

early densely pilose, puberulent below. — A very stable species ex-

hibiting little variability even under different cultural conditions,

most closely related to L. hirsutum Humb. & Bonpl. but distinguished

by vining stems, smaller and more dissected leaflets. Like the

related species it rarely occurs at an altitude as low as about 1,200

meters (author). According to Muller, collected at several local-

ities in the Department of Lima by Blood and Tremelling. Illus-

trated, Muller, pis. 3 and 10.

Lima: Prov. Canta, 7 km. north of Trapiche, gravelly plain,

dry wash, flowers yellow, Hutchison 1013. Rio Rimac, at km. 81,

FLORA OF PERU 161

Goodspeed & Metcalf 30234.. Yangos to Cantas, near Lima, Blood

& Tremelling 14., 15 (type sheets U. S. N. A. Herb.). Above Santa

Eulalia, above Chosica, West 8600. Chosica to Matucana, Mexia

4.077. — Arequipa: Laspinas, Eyerdam & Beetle 22150 (det. Johnston).

Solatium lycopersicum L. Sp. PI. 185. 1753; 10. Lycopersicum

esculentum Mill. Gard. Diet. ed. 8. 1768.

Becoming a vigorous herb, flowering as an annual but often

persisting, the stout more or less succulent or below subligneous

stems puberulent to pilose-hirsute, not infrequently viscid-glandular

especially above, including the often ample odd-pinnate to bipinnate

leaves; stipules none, principal leaflets 2-4 pairs, ovate to lanceolate,

petiolate, entire-lobed or pinnate, puberulent, or the rachis glandular-

pilose, 3-10 cm. long, about half as wide, the smaller entire, a few

mm. to 2 cm. long, sometimes lacking; racemes simple, now and then

furcate, 4-5 (-10) -flowered, pedicels 1-1.5 cm. long, articulate at or

above the middle; calyx 5-8- or 10-parted nearly to base, the sub-

ulate lobes about 1 cm. long, 1 mm. wide; corolla lemon-yellow,

medially to three-fourths 5-10-parted, the lanceolate acuminate re-

flexed lobes about 1 cm. long or longer, glabrous or more or less

pilose above and below especially toward tip; stamens 5, 8 or 10,

usually soon parted into groups of 2 or 3 at anthesis, the subsessile

anthers 5 mm. long, appendages 2 mm. long; style slightly exserted,

ovary glabrous; berry 3-10-celled, commonly depressed globose;

seeds flat, oval, 3 mm. long, 2-2.5 mm. wide, densely silvery lanate. —

Illustrated, Muller, I.e. pis. 1-4 and 8.

This is the basic or cultivated form of the species; two recognized

forms or varieties have been described as Lycopersicum esculentum

Mill. var. pyriforme or f. pyriforme (Dun.) C. H. Mull., the flowers

rarely more than 5-merous, berry 2-celled, pyriform; much more

distinctive and native to Peru and probably elsewhere in the Andes

is the var. esculentum (Mill.) Voss, in Vilmorin, Blumengartn. (ed. 3)

1: 721. 1894, or S. lycopersicum Mill. var. cerasiforme (Dunal)

Fosberg, Phytologia 5 : 290. 1955, the raceme more or less elongate,

corolla about 1 cm. long, lobes 6 mm. long, staminal column narrow,

berry 2-celled, globose, 1.5-2.5 cm. in diameter, the less compressed

seeds only marginally pubescent. Muller decided that this plant,

known in English as the cherry tomato, is the form from which the

cultivated plant originated; it is widely cultivated in undeveloped

regions. Fosberg, I.e., wrote: there seems to be no more reason for

maintaining a separate genus for the tomato than for many other

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

groups commonly retained within the genus Solanum; note also the

recent publication of S. Pennellii Correll with the author's remark:

this and S. lycopersicoides Dunal are living evidence in support

of the union of the two genera. Type locality, the coast of Peru.

Widely distributed at least as an adventive.

Piura: ftormala, Prov. Paita, Haught 2. — San Martin: Tarapoto,

Spruce 41 4^- San Roque, Williams 7264 (det. Standley). — Huanuco:

Tingo Maria, (Blood & Tremelling 62; 68; 70). — Junin: La Merced,

(Blood & Tremelling 49; 50; 52; 53}. — Loreto: Puerto Arturo,

Williams 5170 (det. Standley). — Apurimac: Prov. Abancay, (Vargas,

det. Muller). — Cuzco: Echarate, Goodspeed Exped. 10482. Santa

Ana, Cook & Gilbert 1603.

Solanum peruvianum L. Sp. PL 186. 1753; 16. Lycopersicum

peruvianum (L.) Mill. Card. Diet. ed. 8. 1768.

Various in habit but usually more or less decumbent, always

densely, and more or less canescently, pilose-villous, usually eglan-

dular except the inflorescence; stipules present or lacking; leaves

oblong-lanceolate or narrowly ovate, simply or odd pinnate, major

leaflets 3-5 pairs, typically unequally rounded at base, subentire

to erosely dentate, the minor if present ovate, subsessile, 1-5 mm.

long; inflorescence commonly a cyme of two 5-12-flowered racemes

or these again dichotomous (rarely simply), racemes 5-9 cm. long,

pedicels 5-12 mm. long, the peduncles (3-10 cm. long), rachis and

pedicels ordinarily bracted (pedicel bracts sometimes reduced or

wanting); calyx parted nearly to base, the 5 linear-lanceolate lobes

5-6 mm. long; corolla bright orange-yellow, 10-13 mm. long, parted

medially into triangular acuminate lobes so strongly reflexed that

the limb is turned inside out (Muller), dorsally pubescent; staminal

column 6-9 mm. long clearly bent to one side, the entire anther

sacs 4-6 mm. long; style exserted, stigma capitate; ovary sparsely

pubescent, especially near apex, fruit 1-2 dm. in diameter, puber-

ulent, pale with lavender stripe, seeds oblanceolate, glabrous ex-

cept at tip, pale brown, type "Peru." — L. peruvianum var. dentatum

Dunal (L. chilense Dunal; L. atacamense Phil. Fl. Atac. 42. 1860,

both fide Muller) usually has stipules, larger leaflets deeply lobed

or pinnatifid or leaflets subentire, petiolate and cordate; var. humi-

fusum Muller, 19, distinguished by the author as densely and very

shortly pubescent, minor leaflets usually lacking, the larger entire,

ovate-lanceolate. Muller notes that var. dentatum varies greatly

even to seeds in size and shape; the trichomes are more frequently

glandular. Illustrated, Muller, I.e. pis. 3, 6 and 7.

FLORA OF PERU 163

Piura: Talara, Haught 93. Truxillo, (Blood & Tremelling, var.

dentatum). — Ancash: Chacchan, 255 4. — Lima: Chosica, West 3600.

Matucana, 159. Amancaes, Balls 7073. Chancay, (Blood & Tremell-

ing}. San Juan, Magdalena, Piscocucho, (Blood & Tremelling, all

var. humifusum, type, 142). — Huanuco: Cuchero, Poeppig 1449. —

Junin: San Rafael, 31 41- — Arequipa: Tingo, Pennell 1311*7, var.

dentatum. Yura, Carl Schmidt. Mollendo, Johnston 3557. — Cuzco:

Via de Santa Ana, Cook & Gilbert 1405. — Moquehua: Estaquina,

Weberbauer 7445; 7W. Chile, and northward to Bolivia and Ecua-

dor.

Solatium pimpinellifolium Jusl. in L. Cent. PI. 1: 8. 1755;

15. Lycopersicum pimpinellifolium (Jusl.) Mill. Gard. Diet. ed. 8.

1768.

Puberulent, the prostrate ascending stems rarely somewhat pilose

or glandular, leaves narrowly ovate with usually a pair of small divi-

sions alternating with larger ones, the latter 3-paired, unequally cor-

date or rounded, subentire, dark green above, paler beneath, the

smaller ovate to rotund sometimes wanting; racemes rarely fur-

cate, 12-many-flowered, to a dm. long, glandular-puberulent, the

two-ranked filiform pedicels very regularly spaced, 3-15 mm. long,

articulate about 3 mm. below the flower; calyx lobes 5, lanceolate-

acuminate, 2.5-4 mm. long, glandular-pubescent both sides; corolla

bright lemon-yellow or slightly orange, 12-16 mm. across, deeply

5-parted, the narrowly lanceolate lobes long-attenuate, strongly re-

flected at anthesis, dorsally puberulent anthers subsessile, column

5-7 mm. long, the connate portion slightly shorter than the anther

sacs; style slightly exserted, ovary glandular or puberulent, usually

becoming glabrous; berry 2-celled, 1-1.5 cm. in diameter, ordinarily

glabrous, lustrous red or orange-red, the raceme elongating some-

times 2 dm., the accrescent calyx finally 6 or 7 mm. long, seeds thick,

obovate, 3 mm. long, 1.5-2 mm. broad, glabrous except at tip. —

Original locality Peru. Related to S. lycopersicum var. cerasiforme

but it is readily distinguished by its more slender habit, more

elongate racemes and smaller fruit (Muller, who illustrated it, pis.

3 and 5).

Piura: La Brea, Horton 11582. Truxillo, (Blood & Tremelling'). —

Lima: Pachacama, Mexia 8328 (det. Morton). St. Eulalia, Good-

speed 33147. East of Lima, Blood & Tremelling 76. Chacallo, Balls

858. Quillabamba, (Blood & Tremelling}. Santa Ana, Cook & Gil-

bert 1405; 1702.

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

Section MORELLA

Leaves entire or somewhat lobed but never alately decurrent to

stem (often more or less on petioles), and lobes never alately joined

as in species with somewhat divided leaves; flowers few to many in

peduncled inflorescences, minute to about 1 cm. long; anthers early

bipored apically. Annuals or perennials frequently finally ligneous

below, rarely toward apex, usually only firm above where ordinarily

hollow-stemmed and the leaves often much shorter than 1 dm. The

following species center around the ubiquitous S. nigrum L. and prob-

ably there are only a few of them that are not dubious. The key

attempts by the use of characters scarcely significant to lead to some

of the many names proposed.

Corolla, unless in S. nigrum, scarcely 3.5 mm. long, usually shorter.

Plants soon glabrate or glabrous; berries (ripe) black, free.

Flowers few, subumbellate or subracemose, the peduncles usu-

ally simple; anthers soon 2-chinked, about 2.5 mm. long or

shorter.

Calyx lobes distinct, early somewhat reflexing; flowers sub-

umbellate S. inconspicuum, S. nodiflorum.

Calyx lobes not equally distinct, suberect to appressed; flowers

subracemose S. nigrum.

Flowers small, many, in forked racemes; leaves firm, oblong-

elliptic; anthers 2-pored S. corymbosum.

Plants villous-hirsute; berries not black, partly enclosed.

S. sarachoides.

Corolla in full anthesis at least 4 mm. long, the peduncles normally

at least bifid; anthers often about 3 mm. long or longer (see also

glabrate S. nigrum, S. nodiflorum) ; peduncles simple and villous

(S. sarachoides).

Leaves often less than twice as long as wide, broadly ovate to sub-

rhombic, more or less strongly or unevenly repand-dentate to

sublobulate, frequently somewhat caudate, tip entire; tri-

chomes in part glandular.

Calyx lobes ovate, 2-3 mm. long S. excisirhombeum.

Calyx lobes sublinear, 4-5 and 5-7 mm. long.

S. sinuatiexcisum.

Leaves about as above; trichomes eglandular or sub viscid; pedun-

cles usually furcate S. fragile.

FLORA OF PERU 165

Leaves often subentire or remotely few repand-dentate (cf . S. frag-

ile) never, in Peru, very irregularly dentate, only sometimes

not much longer than wide; peduncles (normally) at least

1-furcate, unless S. sinuatiexcisum.

Indument (younger parts) somewhat viscid-glandular villous,

rarely only viscid-puberulent.

Corollas rotate-stellate; peduncles (types) furcate.

Peduncles much-forked (type) S. probolospermum.

Peduncles 1-2-furcate S. juninense.

Corollas campanulate-rotate; peduncles (type) simple.

S. sinuatiexcisum.

Indument eglandular, rarely obscurely viscid in inflorescence.

Trichomes simple or early 1-few-furcate, rarely lacking or

nearly.

Leaves usually much shorter than 1 dm. (expedient char-

acter, species).

Leaves rather oblong-lanceolate or ovate-lanceolate; flow-

ers (type) finally about 1 cm. long.

S. Pentlandi, S. arequipense.

Leaves ovate or broadly ovate-elliptic, often subrotund

at base, mostly 5-9 cm. long, about half as wide;

flowers often smaller. .S. Zahlbruckneri, S. furcatum.

Leaves mostly 1-1.5 dm. long, a third to more than half as

wide S. insolaesolis.

Trichomes at least early mostly or many often pseudo-stellate.

S. sandianum, S. pallidum.

Solarium arequipense Bitter, Repert. Sp. Nov. 11: 204. 1912.

S. atricoeruleum Bitter, I.e. 10: 563. 1912, at least as to Peru.

A meter or more high, often clambering, with long divaricate strict

or tortuous branches; leaves lanceolate-rhomboid, mostly 5-8 cm.

long by about 3 cm. wide, attenuate into a petiole 1-2 cm. or so long,

acute, with 2-3 small teeth below the middle, sparsely pubescent;

peduncles lateral, 10-12-flowered, 1.5-3 cm. long, forked once; ped-

icels 6 mm. long; corolla white (or purple), full grown about 1 cm.

long and broad, the lobes about 4 mm. long; filaments 1.5 mm. long,

anthers nearly twice as long, pubescent as also the elongate style;

anthers ellipsoid, nearly 3 mm. long, with oblique subapical introrse

pores; berry 7 or 8 mm. thick, stone cells 2. — Type has a rotate-

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

stellate corolla about 1 cm. long; a more nearly entire narrower-

leaved variant of S. furcatum Dunal (to which some Junin specimens

have been referred by Lyman Smith and by Killip) or S. coerulescens

Bitter; it probably has an earlier name in S. dianthum Rusby, Bull.

N. Y. Bot. Card. 4: 420. 1907; the Rusby plant as to type had only

2-flowered cymes, slender weak pedicels but was probably a reduced

state from a shady place as the habitat was "wet mould." The prob-

able variant S. atricoeruleum Bitter, I.e. 10: 563. 1912 (Bolivia), was

found (by author) to have 5 granules; fruit with 2 subapical stone

cells (Bitter). F.M. Neg. 2597. Section Morella.

Cajamarca: Huaraz, Sandeman 4655 (distr. as S. nigrum). — La

Libertad: Near Cerro Uruchalda, West 8170. — Lima: El Ingenio,

Soukup 3661. Rio Blanco, 665; 2966. — Huanuco: Southeast of

Huanuco, 2079. — Pasco: Quillasu, Soukup 3321. — Junin: Huancayo,

Soukup 2703; 3233. Carpapata, Killip & Smith 24430; Ochoa 292.

Huacapistana, Killip & Smith 24-181 (det. Morton, S. sandianum).

— Apurimac: Chincheros, West 3704 (flowers purple). — Cuzco: San

Sebastian, Pennell 3631. San Miguel, Urubamba Valley, Hen era

1539. Pillahuata, Pennell 13949. Huasco, Herrera 1022. Achirani,

2,600 meters, Vargas 11135 (det. Asplund, S. dianthum, flowers pur-

ple). San Sebastian, PenneU 13631. Paso de Tres Cruces, 3,800

meters, Pennell 13824. Cerro Machupicchu, Herrera 3236; Mexia

8079 (distr. as S. nigrum). — Arequipa: River-cliffs near Arequipa,

Seler 204, type. Outskirts of Arequipa, Sandeman 3781 (distr. as

S. nigrum). — Puno: Araranca, Pennell 13464- Near Puno, Mexia

04251; Metcalf 30690; Soukup 94; 95; 975. Bolivia? "Ccaya-ccaya"

(Mexia) ; "ccjaya-ccajaya" (Herrera) ; "mancato jiechi."

Solanum corymbosum Jacq. Coll. 1 : 78. 1790. Ic. Rar. pi. 40.

1786; 73. S. corymbiferum Gmel. Syst. Nat. 2: 384. 1791. S. parvi-

florum Usteri, Ann. Bot. 6: 61. 1793. S. cymosum R. & P. Fl. Peruv.

2: 31, pi. 160. 1799. S. leptanthum Moc. & Sesse", var. parvifolium

HBK. Nov. Gen. 3: 24. 1818.

Stem woolly below, flexuose, angled, smooth, sprawling to erect,

often subligneous below, 3-10 dm. high; leaves ovate-lanceolate, long-

decurrent into the petiole, scarcely acute, entire, repand, or rarely

1-2-lobed, glabrous, 4-7 cm. long; racemes subsessile, cymose, half

shorter than the leaves, slightly pubescent; pedicels about 3 mm.

long; calyx lobes ovate, subacute; corolla scarcely 2 mm. broad, blue

or bluish; berry about 6 mm. in diameter, orange-red. — Often in cul-

tivated ground. Section Dulcamara.

FLORA OF PERU 167

Cajamarca: Magdalena, (Bonpland, type, S. leptanthum, var.).—

Amazonas: Chachapoyas, Mathews (det. Morton). — Lima: Provinces

of Lima and Chancay, Ruiz & Pawn (type, S. cymosum) ; Dombey.

Matucana, 189; 315. Valle de Chillon, Velarde Nunez 874 (det. Mor-

ton).— Huanuco: At Huanuco, Kanehira 207; Ledig 11 (det. Morton).

Solatium excisirhombeum Bitter, Repert. Sp. Nov. 11: 1. 1912.

S. atriplicifolium Gillies ex Nees, Nov. Act. Caes. Carol. 19, Suppl. 1:

366. 1843, as to Peru.

Viscid-pubescent, especially on the stems and leaf -veins beneath,

with spreading many-celled, mostly gland-tipped trichomes; petioles

alate, about 1.5 cm. long; leaves nearly rhombic, about 6 or 7 cm.

long, 5-6 cm. wide, deeply dentate or sublobulate, the 3-4 lobes

sometimes 1 cm. long; inflorescence simple (type), 4-6 (8) -flowered,

the peduncle about 1.5 cm. long, the pedicels straight or nearly,

finally nodding, about 6 mm. long (-10 mm. in fruit) ; calyx 4 mm.

wide in flower, 9 mm. in fruit, the lobes early 2, finally 3 mm. long;

corolla lilac, 7-9 mm. long and broad; filaments 1.5 mm. long, an-

thers 2.3-3 mm. long; style 5 mm. long, stigma capitate. — Perhaps

S. coerulescens Bitter is an eglandular state and, like this, a straggling

herb, sometimes ligneous at base, flower white or usually white and

pink-purple. S. atriplicifolium Gillies may be the correct name but

according to Bitter, Repert. Sp. Nov. 11: 213. 1913, that Chilean

plant has subregularly 8-10-lobulate leaves, inflorescence mostly

furcate, calyx in fruit with lobes 7 mm. long, corolla white, 14 mm.

across, anthers 4.5-5 mm. long, shorter gland-tipped trichomes on

stems and leaves, early dense on calyx without; the important

distinction seems to be in the smaller anthers of the Peruvian plant.

F.M. Neg. 2604. Section Morella.

Used as a remedy for toothache (Weberbauer).

Ancash: Near Tallenga, Weberbauer 2868, type. — Lima: Rio

Blanco, Kittip & Smith 21540; 21630 (both det. Morton). Canta,

Soukup 2829 (det. Morton; cf. note under S. coerulescens'). — Junin:

Tarma, Killip & Smith 21871. — Cuzco: Ollantaitambo, Cook & Gil-

bert 370 (det. Morton). Near Cuzco Experimental Estac., Vargas

673. — Arequipa: Nevado de Chachani, Pennell 13260. "Japichina,"

"muyuceaya" (Cook & Gilbert).

Solatium fragile Wedd. Chloris And. 2: 105. 1857. S. atriplici-

folium Gillies, var. minus Dunal in DC. Prodr. 13 : 55. 1852. S. coeru-

lescens Bitter, Repert. Sp. Nov. 10: 554. 1912, at least as to Peru.

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

Stems diffuse, slender, herbaceous or ligneous at base, pubescent;

petioles 1-2.5 cm. long, alate about medially; leaves membranous,

rhomboid-elliptic, decurrent into petiole, obtuse, 2-5 (8) cm. long,

coarsely 1-3- mostly irregularly or sometimes sinuate -dentate, pu-

berulent and ciliate; peduncles 1.5-2.5 cm. long, mostly terminal,

umbellately few- to many-flowered; pedicels 5-10 mm. long; calyx

subvillous, lobes ovate-oblong, subobtuse, in fruit 1-1.5 mm. long;

flowers pale to violet, striate, 12-15 mm. wide, lobes triangular-ovate,

obtuse, subvillous, 4 mm. long; anthers yellow (pores lateral), shorter

than the somewhat pubescent style. — Has been referred to S. atripli-

cifolium of Chile, a shortly glandular plant with long (4-5 mm.) an-

thers and calyx enlarged in fruit, characters probably not shared by

Weddell's plant, but it is not obviously distinct from S. Pentlandi

Dunal, or S. coerulescens Bitter. Plants 1-3 dm. across, with many

ascending stems, a few often divaricate branches; leaves remarkable

by the large teeth, these subobtuse, recalling Chenopodium hybridum

L.; pubescence a little viscid on leaves or rest of plant (Weddell).

Perhaps should include S. excisirhombeum Bitter as a glandular vari-

ant; it is much like it but typically eglandular; pedicels often arcu-

ate-tortuous; of course, may be a part of S. furcatum Dunal, range of

variation not determined. F.M. Neg. 2598 (S. atriplicifolium var.

minus). Section Morella.

Ayacucho: Prov. Lucanas, Metcalf 30239 (det. Killip, S. coerules-

cens).— Cuzco: Trailing in brush below Machupicchu, West 8028.

Near Sicuani, Cook & Gilbert 128. Ollantaitambo, Cook & Gilbert 297

(det. Morton, S. coerulescens}; Hen era, 3^27. — Arequipa: Ditch bank,

Arequipa, Pennell 130^8. Rock slides near Chala, Worth & Morrison

15704 (leaves repand). Laspinas, Eyerdam & Beetle 22152 (leaves

subentire). Above Atiquipa, Worth & Morrison 15668. — Puno: Lake

Titicaca, Meyen (type, var. minus). — Tacna: Candarave, Metcalf

30373. Mountains of Tacora, 4,000 meters, Weddell, type.

Solatium furcatum Dunal ex Poir. Encycl. Suppl. 13: 750.

1814; 51. S. chenopodioides Lam. Illustr. 2: 18. 1793 in part, fide

Stebbins & Paddock. S. violeceistriatum Bitter, Repert. Sp. Nov. 10:

550. 1912, at least as to Peru.

Glabrous or nearly, the herbaceous upper stems subdichotomous,

angulate; leaves ovate, dentate, glabrous; racemes furcate (Dunal).

As to Peru: becoming a tall or sprawling (sometimes 2 meters) half-

shrub, the flowering branches or stems, usually once-forked inflores-

cences and leaves beneath more or less densely puberulent with sim-

ple or slightly furcate mostly appressed trichomes, often absent, or

FLORA OF PERU 169

mostly, in age; leaves usually ovate, acute but often rather rounded

at base, abruptly and shortly or scarcely decurrent, varying from a

few cm. to rarely a dm. or so long, about half as wide, entire, undu-

late-repand or weakly repand-dentate; peduncles lateral, ordinarily

2-several cm. long and forked, frequently fewer than 10-flowered;

pedicels crowded or subumbellate, recurved in fruit, usually a cm.

long, nearly that in flower; calyx lobes obtuse or subacute, minute or

small, in fruit broadly ovate, somewhat accrescent; corolla rotate-

stellate, white or purplish, exserted, slightly puberulent below as

unequal filaments; anthers 2.5- about 3 mm. long; berry finally

5-7 cm. in diameter, not persisting after maturity, 7-34-seeded and

with many large stone cells (Stebbins and Paddock). — The Dombey

type may be Chilean and according to Dunal is glabrous except the

racemes; it probably is the earlier name for S. arequipense. Section

Morella.

Lima: El Ingenio, Soukup 3653 (det. Killip, S. minutibaccatum) .

Canta, Soukup 2832. Matucana, 380 (toward S. coerulescens} . Rio

Blanco, Killip & Smith 21752. Huaros, Pennell 14708. — Huanuco:

Mito, 1630; 1865. Cani, 3385. Piedras Grandes, Woytkowski 153.

Near Muna, 3879. Tingo Maria, Allard 21855 (sens, lat.) . Boqueron

Pass, Allard 22112 (sens. lat.). — Junin: Tarma, Killip & Smith 21791.

La Oroya, Kalenborn 26 (det. Morton, S. Pentlandi) . — Huancavelica :

Salcabamba, Stork & Horton 10264. Chile? "Gapichinia."

Solatium inconspicuum Bitter, Repert. Sp. Nov. 11: 204. 1912.

Pubescence at first dense, short, subappressed; branches in type

many, 3-4 dm. long; leaves nearly ovate, abruptly contracted to a

petiole 5-8 mm. long, attenuate at apex, with 1-2 short obtuse lobes

below the middle, 1.5-2.5 cm. long, 10-12 mm. wide, peduncles lat-

eral, 7-8 mm. (-12 in fruit) long, 2-3-flowered; pedicels 3-4 mm.

(-7 in fruit) long; calyx lobes broadly lanceolate, suboblong, 2 mm.

long; corolla white, 3.5-5 mm. wide, lobes lanceolate, 1.5-2 mm. long,

finally spreading; filaments less than 0.5 mm. long, anthers 1 mm.

long; style less than 2 mm. long, pubescent at the middle; berry

globose, black, 4-5 mm. thick, seeds 45. — Cf. S. nigrum L., from

which it is not clearly distinct, the small-leaved tiny-flowered type

perhaps an ecological state more or less duplicated in shady or wet

places in close competition with other plants. F.M. Neg. 2612.

Section Morella.

La Libertad: Chicama Valley, Smyth 19 (det. Killip). Trujillo

to Salaverry, Worth 8893. — Lima: Moron, above the city of Lima,

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

Seler 222, type. — Junin: Edge of road, San Ramon, Constance &

Tovar 2278. — Loreto: Lower Rio Huallaga, Williams 4592a.

Solatium insolaesolis Bitter, Repert. Sp. Nov. 10: 563. 1912.

S. medianiviolaceum Bitter, I.e. 562. 1912. S. violaceistriatum Bitter,

I.e. 550. 1912. S. brevipedunculatum Rusby, Bull. N. Y. Bot. Gard.

4: 421. 1907?

Type a half shrub, the upper stems and leaves both sides some-

what pulverulent with multi-celled trichomes; petioles alate, to 1 cm.

long; leaves ovate-lanceolate, to 5 cm. long, 1.5- about 2 cm. wide

(or larger), entire, attenuate both ends, tip subobtuse, peduncles to

2.5 cm. long, the branches, if present, 3-7 mm. long; pedicels 6 mm.

long, nearly twice as long in fruit, violet-tinged as calyces, these cam-

panulate, about 4 mm. across, subobtuse lobes 1.5 mm. long; corolla

violet, rotate, to 12 mm. across, the lobes 4 mm. long; filaments 1 mm.

long, anthers 2.5 mm. long, pores obliquely subapical; style about

5 mm. long, puberulent above base, stigma capitate; berry sub-

globose, clearly sulcate medially, 4-5 mm. long, 5-6 mm. broad;

seeds about 40, stone cells 8. — After Bitter; probably varies as to

size of leaves, anthers, number of stone cells and may be part of

S. furcatum Dunal; the similar S. violaceistriatum Bitter, as to fruits

examined, had only 2 stone cells. The Rusby plant, to which Killip

allied Soukup 3618, may be the same; type had, ex char., petioles

3-4 cm. long, leaves 1-3 dm. long, nearly half as wide, rounded at

base, acute both ends, calyx truncate, corolla stellate, lobes 1 cm.

long, anthers 6 mm. long. It may be a race of S. fragile Wedd., or

the collections from central Peru may be rather luxuriant examples

of S. furcatum Dunal but the leaves are well-decurrent; several speci-

mens, perhaps significantly, are from stream thickets. Section Morella.

Lima: Canta, Soukup 2829. San Buenaventura, Pennell 14538. —

Huanuco: Carpish, Stork & Horton 9898 (det. Standley, S. coerules-

cens). — Junin: Tarma, Killip & Smith 31912. Chongos Bajo, Soukup

3618 (aff. S. brevipedunculatum Rusby, fide Killip). — Cuzco: Pacbar,

Pennell 13689. Valle del Apurimac, Herrera (Feb., 1929). Uru-

bamba Valley, Herrera 1539. San Miguel, Cook & Gilbert 872. Isla

del Sol, Lake Titicaca, Buchtien, type. Bolivia. "Suito-mullucaya"

(Cook & Gilbert).

Solatium juninense Bitter, Bot. Jahrb. 54, Beibl. 119: 11. 1916.

A sprawling shrub or the angled branches scandent; younger

branches puberulent villous-glandular toward apex as petioles, these

FLORA OF PERU 171

7-10 mm. long, and the ovate leaves especially on the prominent

veins beneath, these in type mostly 3.5-4 cm. long, 2-3 cm. wide, on

most specimens longer, acute or subacuminate, entire or with 2-4

teeth on each side toward the subtruncate base; inflorescence at first

compact, terminal, in type about 15-flowered, glandular-villous, the

1.5-3 cm. long peduncles forked; pedicels about 1 cm. long; calyx

5 mm. long, 6 mm. broad, the lobes lanceolate, acute; corolla blue-

violet, rotate-stellate, about 1.5 cm. wide; filaments about 1 mm.

long, style in part densely pubescent (to glabrate), 7 mm. long;

stigma globose-capitate; berry 8 or 9 mm. in diameter, seeds (imma-

ture) with basal appendage at least 1 mm. long. — Vicinity of S. sandi-

anum (author) ; varies (I think) especially in size of leaves, abundance

and length of trichomes, degree of viscidity, size of flowers, these well-

developed, showy; cf. the probable extreme in leaf size, S. probolo-

spermum Bitter; simulates also S. glandulosipilosum Bitter, Repert.

Sp. Nov. 11: 213. 1913, of northern Argentina. F.M. Neg. 2613.

Section Dulcamara.

Huanuco: Yanahuanca, 1213. — Junin: Near Huancayo, Weber-

bauer 6598, type; Killip & Smith 23360 (det. Morton). Huariaca,

3089. La Quinua, 3,400 meters, Ochoa 336.

Solatium nigrum L. Sp. PI. 186. 1753; 50.

Glabrous or somewhat pubescent annual with lax stems and

branches sometimes subligneous below; leaves membranous, ovate,

2-several cm. long, entire or more or less repandly serrate; peduncles

lateral, usually subracemosely few-flowered, the pedicels 5-10 mm.

long; corolla 5.5-7.5 mm. long, white or bluish; anthers 2.1-2.5

mm. long; calyx lobes unevenly joined, little increased in fruit; berry

dull black, the many seeds 1.9-2.3 mm. in diameter, stone cells always

lacking (Bitter!).— After Stebbins & Paddock, Modrono 10: 79. 1949,

who confirmed chromosome count as 2N=72. Apparently the Peru-

vian plant so named is at least mostly the similar S. nodiflorum Jacq.

Charles B. Heiser, Jr., Ceiba 4: 293-299. 1955, defined four entities

in Costa Rica as pertaining to "the S. nigrum L. complex." Should

not some geneticist study the plants already represented by many

names (74 according to Heiser) before trying even tentatively to in-

terpret the characters? Besides the following there are several col-

lections from the lomas of Arequipa by Giinther and Buchtien,

according to Bruns. Section Morella.

The young shoots contain gluten which is chewed and known as

"ckausillo" and is said to be a nerve soporific (Herrera).

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

Loreto: Lower Rio Nanay, Williams 396. Caballo-cocha, Wil-

liams 2283. Pongo de Manseriche, Mexia 6298 (det. Morton). —

Cuzco: Machupicchu, Mexia 8079 (det. Morton). Santa Ana, Cook

& Gilbert, 515; (Herrera, 162}. Cosmopolitan. "Yerba-moro" (Ruiz

& Pavon), "ccaya-ccaya" (Mexia).

Solatium nodiflorum Jacq. Icones PL Rar. 2: 288. 1786; 46.

S. chenopodioides Lam. Illustr. 2: 18. 1793, in part, fide Stebbins &

Paddock. S. minutibaccatum Bitter, Repert. Sp. Nov. 10: 549. 1912,

at least as to Peru.

A diffuse glabrous or glabrate perennial often with somewhat

angulate stems and repand dentate leaves; flowers subumbellate,

small but variable in length as filaments and anthers, the latter 1.5-

2.4 mm. long; calyx lobes all distinct, reflexing after anthesis and

below the lustrous black berry, this usually without stone granules

(Bitter) or only 1-4; seeds 1.2-1.8 mm. long. — After Stebbins &

Paddock, Modrono 10: 77. 1949, who noted the chromosome num-

ber as 2N=24, the species, as S. nigrum L., highly variable; type

from the island of Mauritius. Most of the specimens cited were dis-

tributed as S. nigrum L. including one in Madrid without locality

by Ruiz and Pavon. F.M. Neg. 33098 (Jelski £9, sp. ined., Bitter).

Section Morella.

Piura: Parinas Valley, Haught 138. — Cajamarca: Tambillo, Jel-

ski 49. — San Martin : Tarapoto, Williams 6050. San Roque, Williams

7221; 7744. Juan Guerra, Williams 6861.— Lima: Rio Chillon, Pen-

nell 14460. Santa Clara, Rose 18741. Miraflores, Vargas 1231.

Callao, Wilkes Exped. — Huanuco: Santa Maria, Allard 22459; 22042

(distr. as S. minutibaccatum). — Junin: Chanchamayo, Ochoa 639.

La Merced, 5207; Killip & Smith 23424; 24058. Tarma, Killip &

Smith 21862. Rio Perene", Killip & Smith 25218. — Loreto: Rio

Nanay, Williams 338. Pongo de Manseriche, Mexia 6298. Iquitos,

Williams 7928. Santa Rosa, Killip & Smith 28827. Yurimaguas,

Williams 4153. — Ayacucho: Kimpitiriki, Killip & Smith 22970. —

Apurimac: Prov. Abancay, Goodspeed Exped. 10576. Tropics gen-

erally and widely distributed as a weed elsewhere. "Yerba-moro,"

"ccyaya-ccyaya" ( Herrera), "ayac-mullaca," "aji."

Solatium pallidum Rusby, Mem. Torrey Bot. Club 4: 228.

1895. S. planifurcum Bitter, Repert. Sp. Nov. 11: 2. 1912.

Type a loose shrub (1 meter), usually a half shrub, rather densely

pubescent, with forked pseudo-stellate trichomes; petioles about 1 (-2)

FLORA OF PERU 173

cm. long; leaves alternate or paired, scarcely undulate, ovate-oblong,

5-10 cm. long, 3-7 cm. wide, but often much smaller, sometimes

larger, at the base abruptly, toward the apex gradually attenuate;

peduncles lateral, 1.5-2.5 cm. long or longer, 1-3-forked, few- to

many-flowered; pedicels 5-10 mm. long; calyx lobes ovate-lanceolate,

0.6-2 mm. long; corolla pale lilac, the lanceolate lobes 5-7 mm. long;

anthers 4.5 mm. long, dehiscence soon lateral, filaments pubescent

within as style below; berry 8 mm. in diameter. — Varies in leaf size

and degree of pubescence, this slightly viscid especially in the inflo-

rescence; some forms suggest S. juninense Bitter with different indu-

ment; apparently S. planifurcum is only a small-leaved more pubescent

state. Bitter found 8 stone cells in S. planifurcum, 18 in S. pallidum.

Section Morella.

Ayacucho: Pampalca, Killip & Smith 22243. — Apurimac: Am-

pay, Vargas 777 (det. Standley). — Cuzco: Paucartambo, Balls 6782;

6783 (det. Morton, S. planifurcum); Pennell 13795 (det. Standley).

Cerro de Cusilluyoc, Pennell 13950; 14147. Near Marcapata, Met-

calf 30725 (det. Killip, S. planifurcum). — Puno: Sandia, Weberbauer

685 (type, S. planifurcum). Yunguyo, Soukup 589. Near Limbani,

Metcalf 30431 (det. Killip). Bolivia. "Achihuay" (Vargas).

Solatium Pentlandi Dunal in DC. Prodr. 13, pt. 1: 51. 1852.

Stems dichotomous, slightly angulate-winged, pubescent above;

leaves cuneate at base, ovate-lanceolate, acuminate, somewhat an-

gulate-repand, about 5 cm. long, pilose-scabrous above, subhirsute

on the nerves beneath; petioles 4-6 mm. long; peduncles and pedicels

filiform, like the calyces hirsute-pubescent; racemes furcate, branches

subumbellate, about 2.5 cm. long; pedicels 4-6 mm. long, umbellate;

calyx lobes ovate, acute; corolla stellate, 2-3 times longer than the

calyx, puberulent without, lobes ovate-lanceolate, acute; style elon-

gate, in type at least twice as long as anthers, deflexed at apex.—

A variant by Dunal in his synonymy of S. furcatum.

Lima: Chicla, Ball? (also in Herb. Jussieu without data). Bolivia.

Solanum probolospermum Bitter, Bot. Jahrb. 54, Beibl. 119:

10. 1916.

A somewhat sordid- or yellowish-villous hollow-stemmed (simple

trichomes very unequal, multi-celled) half shrub with divaricate scan-

dent angled branches; petioles 1.5-3 cm. long, winged above by the

decurrent leaf -blades; leaves ovate, acute or acuminate, about 1-

1.5 dm. long and 6.5-8.5 cm. broad, mostly membranous, mostly sub-

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

repand, densely pubescent on the prominent veins beneath; corymbs

(type) at first terminal, many (60-70) (or few?) -flowered; peduncles

about 6 cm. long, much forked; pedicels incrassate apically, about

1 cm. long, in fruit 1.5 cm. long; calyx campanulate, 5 mm. long,

6-7 mm. across, the lobes lanceolate, acute (ovate, acuminate, West),

shortly stipitate glandular within, 4 mm. long; corolla violet, 2-2.3 cm.

wide, rotate-stellate, the subtriangular lobes about 7 mm. long, pu-

bescent with simple trichomes without, shortly so within only on

midnerve and toward apex; stamens affixed about 1.5 mm. above

base, filaments pubescent, 2 mm. long; anthers 3.7 mm. long, cor-

date-ellipsoid; ovary glabrous; style 8 mm. long, pubescent toward

base, stigma subglobose; berry about 1 cm. thick; seeds 60-65, retic-

ulate, tailed at base with a process 2-2.5 mm. long. — Belongs equally

to the Morellae and Dulcamarae of Dunal (author) ; may prove to be

a luxuriant state (due to montana habitat) of S. juninense; a Pincos

specimen, bifid peduncle, few larger flowers but young, may not be-

long here at all; West 8031 has lanceolate acuminate calyx lobes.

Section Morella.

A tea prepared from the leaves is taken for flatulency (Weber-

bauer).

Huanuco: Valley of the Rio Pozuzo, 3,400 meters, Weberbauer

6789, type. Tambo de Vaca, ^9 — Ayacucho: Near Huanta, 3,200

meters, Weberbauer 751 6a; 7517. — Cuzco: In dense brush-forest, Rio

Urubamba, La Maquina, West 8031. "Shopta."

Solatium sandianum Bitter, Bot. Jahrb. 50, Beibl. Ill: 62.

1913.

A sparsely pubescent shrub, the type nearly a meter high; tri-

chomes branched, mostly on the leaves beneath and these often con-

fined to the veins; petioles 8-17 mm. long; leaves ovate or elliptic,

5-6.5 cm. long, 2-2.5 cm. wide, scarcely undulate, the lower apically

rounded and obtuse, the upper gradually narrowed and subacute;

peduncles at first terminal, 1.5-4.5 cm. long, forked, 6-11-flowered;

pedicels articulate a little above the base, about 12 (after anthesis 15)

mm. long; calyx 5-6 mm. across, the 5 or 6 lobes lanceolate, about

2 mm. long; corolla violet, rotate, about 2 cm. broad, the broad (7 mm.

at base) lobes to 8 mm. long; filaments 2 mm. long, pubescent on the

inner side, anthers early 2-pored, elliptic, 4 mm. long; style 7 mm.

long, pubescent below, the stigma capitate; pedicels and calyces more

or less suffused (in type) with purple; fruit unknown. — May be an

eglandular state of S. juninense Bitter; the type is soon glabrate

FLORA OF PERU 175

but other specimens referred to it suggest S. pallidum or S. plani-

furcum except for form of leaves. F.M. Neg. 2636. Section Dul-

camara.

Cuzco: Limatambo, Vargas 742. Near Cuzco, Hen era, 819 (det.

Morton). Saxaihuaman, Hen era 2178; 3092 (det. Morton). — Puno:

Above Cuyocuyo, Weberbauer 930, type. "Chinchi-chinchi" (Herrera).

Solatium sarachoides Sendt. in Mart. Fl. Bras. 10: 18, pi. 1.

1846; 62.

Villous angulate-stemmed annual, the unequal many-celled tri-

chomes typically glandular at least above or in the shortly (3-6 mm.

long) pedunculate 3-8-flowered inflorescence; leaves ovate, subcor-

date to abruptly contracted at base, attenuate to obtuse (type) apex,

4-5-repand, subentire, often to 5 or 6 cm. long, nearly as wide or

larger; flowers 2-8, subumbellate; pedicels nutant or deflexed, finally

6-10 mm. long, strongly incrassate apically; calyx about 4 mm., in

fruit 7 mm. broad, the oblong lobes then 4 mm. long; corolla rotate,

5-7 mm. across, lobes 4-5 mm. long; anthers about 2 mm. long; fila-

ments (anther pores large, introrse) and style pubescent, stigma cla-

vate; berry greenish, globose, opaque (type), about 6 mm. in diameter,

about 45-seeded, 4-6 granules. — In part after Bitter, Repert. Sp.

Nov. 11: 208. 1913, who limited his detailed description to the type

and one other specimen. Similar, and also to be expected (as an

introduction from Europe), is S. villosum Mill., 58, with often fewer

flowers, pedicels and calyx slightly enlarged in fruit, this ripening

yellow or reddish. Marked by viscosity and enlarged calyx; the

Peruvian plant, immature, may be the plant of Miller; it was col-

lected in weedy growth near river.

Apurimac: Casinchihua, Prov. Abancay, Goodspeed Exped. 105^8

(10584.}. Brazil and widely distributed or established.

Solarium sinuatiexcisum Bitter, Repert. Sp. Nov. 10: 558.

1912. S. hyoscymoides Bitter, I.e. 11: 236. 1913.

Somewhat glandular pilose (trichomes simple, spreading) half-

shrub often a meter or more high; petioles often 4-6 cm. long or

longer, the upper usually geminate; leaves membranous, more or

less repand-lobate, rather abruptly contracted at base, attenuately

acuminate, ordinarily 8-15 cm. long, somewhat more than half as

broad, in age the indument confined to veins; inflorescence lateral,

extra-axillary or subopposite leaves, mostly 5-7-flowered; peduncles

2-4 cm. long; pedicels subumbellate in flower, about 1 cm. long,

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

nutant; calyx campanulate, the lanceolate acute lobes unequal, about

4-7 mm. long; corolla rotate-campanulate, lilac or violet, 10-13 mm.

long, pilosulous, the lobes scarcely evident; filaments 1.5-2 mm. long,

anthers 4-5 mm. long, dehiscent by subapical chinks; stigma capi-

tate.— Corolla pale violet; in clearing border of the camino (Vargas) ;

identity not certain to me as the corollas seen seem to have longer

lobes than described. The author himself suggested that his names

may represent one entity, the leaves of the later (1913) one more

nearly entire. Very possibly this will prove to be S. probolospermum

Bitter but fruit unknown; the author separated this from Morella

as Section Campulisolanum Bitter, I.e. 11: 234, characterized (if at

all effectively) by the campanulate corolla; in view of the variable

corolla shapes from rotate-campanulate to deeply stellate, this at

most is a specific identification.

Cuzco: Vilcabamba, 2,550 meters, Vargas 4016 (det. Killip).

Bolivia.

Solatium Zahlbruckneri Bitter, Repert. Sp. Nov. 11: 203. 1912.

Suffrutescent (type), the subobvious decurrent lines of the younger

branches, leaves especially beneath and inflorescences rather densely

subcanescent with a crispate indument; internodes 1.5-2.5 (-4) cm.

long; petioles about 1 cm. long; leaves ovate-lanceolate, entire or ob-

scurely repand angulate, subobtusely acuminate, abruptly or shortly

decurrent at base, 4-5 cm. long, 1.5-2.5 cm. wide; peduncles about

1.5 cm. long, in fruit 2-3 cm. long, forked, pedicels finally 7 or 8 mm.

long; calyx lobes lanceolate, equal, early 1.5 mm. long, in fruit 2-

2.5 mm. long; corolla 12-16 mm. across, the lanceolate lobes violet

without, 4 mm. long, 3.5 mm. wide at base, only the interpetiolar

tissue glabrous; filaments 1 mm. long, puberulent as style below with

long multi-celled trichomes, the style exceeding the elliptic-lanceolate

cordate-based anthers by 5-7 mm.; stigma globose, capitate; berry

globose, 5 mm. in diameter, perhaps greenish, seeds about 22, 1 mm.

long, 1.4 mm. thick, granules none. — After author, who named it for

an eminent Austrian botanist; probably genetically a part of S. fur-

catum Dunal and segregates, as S. arequipense Bitter, S. atricoeruleum

Bitter, I.e. 10: 563. 1912, and others of northern Bolivia; some speci-

mens cited under Dunal's name, as my 665, 2966; Soukup 3661, all

from Department Lima as well as several from Puno, as West 04,251,

Metcalf 30698, may, if this is distinct, belong here. F.M. Neg. 33122.

Section Morella.

FLORA OF PERU 177

Cajamarca: Cutervo, Jelski 46, type. Huambos, Prov. Chota,

Soukup 4553. — Ancash: Recuay, 2514-

SERIES 3 (Expedient Group)

This vegetative key pertains to species from various groups but

brought together conveniently because of the common character of

somewhat divided leaves, except S. phyllanthum and the tuberiform-

rooted S. montanum. This arbitrary separation here simplifies con-

siderably the keys in which they would otherwise be distributed or

similarly grouped artificially.

Leaves usually at least in part compound (ternate, pinnate, multi-

pinnate, sometimes only more or less divided), the divisions alately

joined or if merely repand-lobed always alately decurrent on stems

(S. phyllanthum) or root tuberous (S. montanum); plants various,

sometimes elongating, diffuse or semiscandent. Includes in part

Polybotryon, Regmandra, Dulcamara (cf. Cyphomandra, in fruit).

Leaves usually subentire; herb with a fibrous tuberous root; petioles

more or less alate but not alately decurrent on stem.

S. montanum.

Leaves repand-lobulate, rarely subentire, always alate-decurrent on

stems; half shrubs in age S. phyllanthum.

Leaves mostly or all bipinnate or simply pinnate-lobed or -dentate,

succulent; flowering as annuals, the base in age subligneous.

Corolla 1-1.5 cm. across, simply 5-lobulate (always?).

S. multifidum.

Corolla 1.5-2 cm. across, sinuses of lobes minutely lobed.

S. Murphyi.

Leaves ternate, pinnate or pinnately 3-7-lobed and segments alately

joined, often only the terminal; plants subdecumbent, diffuse or

scandent.

Leaves pinnately 3-7-lobed, lobes subequal, glabrous; corollas 5-

10 mm. long S. radicans, S. quercifolium.

Leaves compound with 3-many leaflets, rarely 3 terminal joined.

Leaflets 3, rarely 5 on a few leaves; corollas 7-12 mm. long

(dubious segregates of S. ternatum).

Leaflets oblong-lanceolate, equally narrowed both ends, less

than 2 cm. wide S. subquinatum.

Leaflets ovate-lanceolate to elliptic, oblique at base, mostly

at least 2 cm. wide.

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

Plants glabrous or essentially S. ternatum.

Plants clearly but often lightly pubescent.

Peduncles solitary; leaves 3 times longer than wide.

S. dendrophilum.

Peduncles geminate; leaves twice as long as wide.

S. semiscandens.

Leaflets regularly 5-many, rarely all, or the terminal 3, alately

joined.

Corollas shorter than 1 cm.; leaflets never all joined (the

names probably refer to about half as many entities).

Leaflets 5-9, even the terminal at most 2.5 cm. wide, mostly

narrower.

Corolla in anthesis 6-8 mm. long; berry subglobose, apic-

ulate; plants scandent S. diffusum.

Corolla about 3 mm. long; berry conic-acuminate; type

decumbent-ascending S. chamaepolybotryon.

Leaflets 7-many, if narrower than 3 cm., 11 or more; flowers

3-6 mm. long.

Leaflets suboblong, 9 or more.

Leaflets acuminate, often wider than 2 cm., glabrous

or midnerve puberulent.

Flowers white, 3 mm. long; berry subglobose.

S. mite.

Flowers lilac, 6 mm. long; berry ellipsoid-conic.

Leaflets glabrous; berry not alate. . . .S. conicum.

Leaflets glabrate; berry alate-angled.

S. alatibaccatum.

Leaflets subacute, about 1.5-2 cm. wide, pilose; berry

acute both ends S. Uleanum.

Leaflets rather obovate, subcaudate, more or less pubes-

cent, usually 7.

Flowers greenish-yellow-white; berry globose.

S. huallagense.

Flowers lilac; berry conical S. semievectum.

Corollas usually at least 1 cm. long; leaves with 3-several

often joined segments (cultivated vines, leaves also en-

tire at least in part).

FLORA OF PERU 179

Leaves with a terminal segment much larger than the two

lateral S. Dulcamara.

Leaf-divisions or leaflets about alike.

Calyx repandly lobed S. jasminoides.

Calyx subtruncate, teeth minute S. Seaforthianum.

Solatium alatibaccatum Bitter, Repert. Sp. Nov. 12: 68. 1913.

Glabrous or nearly, at least the firm-herbaceous fruiting upper

part, this with alternate 4-5-pinnate leaves 2-3 dm. long, the peti-

oles to 4.5 cm. long or longer, shortly petiolulate leaflets oblong-

lanceolate or -elliptic, oblique at base, shortly acuminate, lowest to

6 cm. long, a third as wide, upper somewhat larger (in Peru medial

to 5 cm. wide, 1.5 dm. long), membranous, pulverulent on midnerve

and veins above, glabrous or nearly beneath where clearly puncticu-

late; inflorescence binate (-3) in axils, 4-7-flowered (type), in fruit

1-1.5 cm. long; pedicels 1 cm. long, incrassate apically; calyx lobes

obtuse, 1 mm. long and broad; berry conical, to 22 mm. long, half as

thick, prominently alate laterally, the wings subcrenulate, continu-

ous into a compressed beak about 6 mm. long. — Corolla in Peru

greenish-white, scarcely 3 mm. long, on pedicels 3.5-4 mm. long.

Placed by author in species group Polybotryon but allied by him to

his S. theobromyphyllum, probably correctly; as he suggests, relation-

ship is certainly not indicated necessarily by simple or compound

leaves.

Ayacucho: Estrella, 5,000 meters, Killip & Smith 23055 (det.

Killip). Ecuador.

Solatium chamaepolybotryon Bitter, Repert. Sp. Nov. 11:

471. 1912.

Low decumbent-ascending, woody and with roots toward the

base, sparsely pubescent or glabrate except petioles, leaf-rachis and

midnerve of leaves slightly puberulent; internodes about 3 cm. long;

petioles 1.5-3.5 cm. long; leaflets 2 (-3) pairs, lanceolate, attenuate

at both ends, the lateral about 3.5-6.5 cm. long, 12-17 mm. wide,

the terminal 4-7.5 cm. long, to 1.5 cm. wide, tapering into petioles

3-4 (-10) mm. long; pedicels few-6 mm. long, in fruit 1 cm. long, in-

crassate apically; calyx 2 mm. broad, very short; corolla lobes about

2.5-3 mm. long; anthers 1.8 mm. long, emarginate both ends; berry

(immature) conical, acuminate, 8 mm. long, half as thick. — The leaves

are mostly composed of 3 terminal leaflets and only one pair besides;

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

the small flowers are apparently not fully developed, in which case

this may be a young plant of S. diffusum R. & P. F.M. Neg. 33057.

Section Polybotryon.

San Martin: Tarapoto, Spruce 4432, type.

Solanum conicum R. & P. Fl. Peruv. 2: 38, pi 172. 1799; 66.

An herbaceous very sparsely pilose-stemmed plant about 8 dm.

high; leaflets about 11 pairs, oblong-lanceolate, acute, unequal at

base, the larger 8-12 cm. long or little spinulose-pilose above, soon

glabrous; peduncles glabrous, axillary, the racemes often bifid or tri-

fid; flowers bluish, 3-4 mm. long, secund, with ovate, acute lobes;

berry olive-like, white. — Similar to S. mite; stems in type pilose

above, angulate; petioles to 7 cm. long; leaves to 2.5 dm. long or

longer, with 11 suboblong narrowly acuminate leaflets, these scarcely

or not narrowed at the unequal base, the petiolules 2-4 mm. long,

the larger blades about 7-10 cm. long, 2.5- nearly 3 cm. wide, the

lower 7 cm. long, 2 cm. wide, all glabrous, green above, paler beneath,

the 6 or 7 primary veins purplish, little prominent; peduncles 1-3,

in fruit stout (1.5-2 mm. thick, 4-6 cm. long, pedicels at least 1 cm.

long, apically incrassate), the racemes in flower often 3-5 cm. long,

flowers secund; calyx 5-parted; corolla pale blue or lavender, 12 mm.

across (Dunal), narrowly ovate segments acute; berry white, conical,

the size of an olive (Ruiz & Pavon). — My specimen was single-

stemmed, slender, nearly 3 dm. tall, essentially glabrous. Section

Polybotryon.

Huanuco: Chinchao and Cuchero, Ruiz, type. Huacachi near

Muna, 4134.

Solanum dendrophilum Bitter, Repert. Sp. Nov. 12: 143. 1913.

Herb, creeping on trunks, pubescent, with many-celled trichomes;

petioles 7-10 cm. long; leaflets 3 (rarely 5), broadly lanceolate, the

terminal attenuate at both ends, the lateral very oblique at base,

6-7 cm. long, 2.3-2.5 cm. wide, the petiolule 3-4 mm. long, the ter-

minal larger, all membranous; calyx 5 mm. broad, the lobes very

short; flowers purple, to 23 mm. across, the lobes 11-12 mm. long;

pedicels 1.5-2 cm. long, crowded, subsessile at the top of solitary

axillary peduncles, these about 3.5 cm. long; anthers 6 mm. long,

1.5 mm. thick, only subemarginate both ends; ovary conical, gla-

brous as the 6-7 mm. long style, the stigma clavate. Section Poly-

botryon.

San Martin: Campana, (Spruce 4385, type, Herb. Kew).

FLORA OF PERU 181

Solatium diffusum R. & P. Fl. Peruv. 2: 37, pi. 171. 1799; 67.

S. Feddei Bitter, Repert. Sp. Nov. 12: 67. 1913, fide author.

Suffrutescent with virgate and diffuse or scandent branches; leaf-

lets (2) 3-4 pairs, thick, glabrous or nearly so, oblong-elliptic, acumi-

nate, the larger lateral 3-4 cm. long and 12-15 mm. wide; leaves to

1 dm. long, petioles (2-4 cm.) and rachis more or less pilose as also

peduncles, these 2 cm. long with several (-10) violet (sometimes

white) flowers 6-8 mm. long on pedicels 5 mm. long, to 1.5 cm. in

fruit; calyx crenately lobed, 4.5 mm. across, in var. subtruncate and

teeth joined by a membrane; filaments obscure, glabrous as ovary

and style, this 6.5 mm. long (var.), the stigma little enlarged; berry

subglobose, apiculate (my collection). — The var. miozygum (Bitter)

Macbr. Field Mus. Bot. 8: 111. 1930 (subsp. miozygum Bitter, Bot.

Jahrb. 54, Beibl. 119: 14. 1917) is less pubescent, leaflets 2-3 pairs,

calyx nearly truncate. Subgenus Bassovia, according to Bitter, I.e.

17: 331. 1921, as also S. conicum R. & P. Section Polybotryon.

F.M. Neg. 12996.

San Martin: Jepelacio, King 3665 (det. Standley). — Huanuco:

Huacachi, near Muna, 4698. Prov. of Huanuco, Ruiz & Pav6n, type.

Mufia, (Pearce 135, type, S. Feddei, fide Bitter, I.e. 143). Valley of

Rio Pozuzo, Weberbauer 6783, type, var.). — Junin: Pichis Trail, Dos

de Mayo, Killip & Smith 25811 (det. Morton). — Cuzco: Pillahuata,

Pennell 14012.

Solanum Dulcamara L. Sp. PI. 1: 264. 1753; 78.

Flexuose-stemmed vine, the leaves cordate-ovate or the upper

often 3-divided, the middle segment broadly ovate, much longer

than the unevenly oblong lateral segments; flowers 12 mm. across,

the petals 10-spotted; calyx with obtuse lobes; berry ovoid or ellip-

soid, more than 1 cm. in diameter when red. — Variable; probably

grown for ornament. Section Dulcamara.

Loreto: Iquitos, Williams 8241 (perhaps; in fruit). Old World.

"Asnapanga."

Solanum huallagense Bitter, Repert. Sp. Nov. 12: 138. 1913.

Stem 1 meter or so high, subligneous below; rather densely but

finely pilose all over except in type, pubescence on the leaves beneath

confined to veins; leaflets 3-4 pairs (rarely 2), broadly obovate-

lanceolate, about 11 or 12 cm. long and 4-6 cm. wide, or the terminal

sometimes 14 X 7 cm.; flowers small, yellowish-green, often 20-25 in

a binate or ternate axillary inflorescence; peduncle about 1 (-4) cm.

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

long; pedicels 2-4 mm. long; calyx 1.5-2.5 mm. wide; corolla 4-5 mm.

across, the lobes broadly lanceolate, 2.5 mm. long; anthers ellipsoid,

1-2 mm. long; style 3 mm. long, clearly incrassate to capitate stigma;

berry globose, 7-8 mm. long and thick. — Near S. trizygum Bitter,

I.e. 11 : 470. 1913, of Venezuela, a less pubescent plant with smaller

leaves, fewer flowers, cordate anthers and conical fruit (author).

Some of the San Martin specimens are rather densely pilose, sug-

gesting S. dendrophilum or S. Uleanum, the latter typically with

much smaller and more numerous leaflets, the former with fewer,

and, ex char., not obovate. F.M. Neg. 33079. Section Polybotryon.

San Martin: San Roque, Williams 6956. Tarapoto, Williams

6045. Jepelacio, Klug 3686 (det. Standley, S. Uleanum). — Loreto:

Yurimaguas, Spruce 3882, type; Williams 426 %; 5351; Killip & Smith

27614. Santa Rosa, Williams 4905. Balsapuerto, Klug 2872 (det.

Standley, S. Uleanum}. La Victoria, Williams 2829; 2923. "Sapote

yacu" (Williams).

Solarium jasminoides Paxton in Paxt. Mag. 8: pi. 5. 1841; 82.

Many-stemmed, twining, glabrous (except young branches and

pedicels), the leaves subcordate, ovate, acute or acuminate, mostly

entire or 2-5 divided or pinnately parted, the suboblong divisions

subobtuse, ordinarily several cm. long, about a third as wide;

petioles sometimes strongly curved at base, even clasping adjacent

stems, 6-14 mm. long; racemes in terminal or lateral dichotomous

panicles; peduncles and pedicels pilosulous, the latter about 10 mm.

long, often arcuate; calyx green, to 4 mm. across, the broadly ovate

subciliate teeth acute, spreading; corolla pale blue or white, to 2

cm. broad, spreading, deeply divided, the 5 ovate segments acute,

6 or 7 mm. long, nearly half as wide; stamens 4 mm. long; anthers

connivent in a tube, pores obovate; ovary and style whitish, the

latter often somewhat villous, 6 mm. long. — Crovetto, Rev. Invest.

Argent. Agric. 2: 188, has illustrated the similar (or probably merely

a form) S. boerhaaviaefolium Sendt. treated as a variety by Kuntze,

Rev. Gen. PI. 3, 2: 226. 1898. As Dr. Crovetto remarks, the relation-

ship of the plants must await critical studies. Similar species per-

haps grown for ornament include S. Dulcamara L. and S. Seaforthia-

num Andrews; the first has terminal leaflet much the largest, corolla

12 mm. across, calyx lobed, fruit oblong-ovoid, the second the

leaflets subequal, corolla 2-2.5 cm. across, calyx teeth minute,

berry subglobose. Illustrated, Bot. Reg. pi. 33; Crovetto, I.e. 189;

Journ. Linn. Soc. Bot. 9: 43 (petiole around stick).

FLORA OF PERU 183

Loreto: Rio Itaya, Williams 3382. Southern South America.

"Jasmin."

Solatium mite R. & P. Fl. Peruv. 2: 38, pi 175. 1799; 66.

A tall essentially erect or semi-erect glabrous herb; leaflets 4-5

pairs, oblong-lanceolate-acuminate, very oblique at base, petiolate

(petiolules 6-15 mm. long), mostly 7-10 cm. long, 2.5-3 cm. wide;

racemes short, clustered in the axils, about 1-3 cm. long; pedi-

cels filiform, incrassate at apex, recurving, secund, calyx minute,

5-parted, the teeth round-ovate, subacute; corolla white or nearly,

almost 2 mm. long, 4-6 mm. across, the lanceolate segments acute;

stamens half as long as the corolla, subsessile, the stout anthers

connivent, 1-1.5 or scarcely 2 mm. long, dehiscing by two apical

chinks; style subulate, 1-1.5 mm. long, papillose, deflexed above,

obscurely punctulate, the small obconic stigma subemarginate; berry

white, subglobose. — Belongs to the subgenus Bassovia according to

Bitter, Repert. Sp. Nov. 17: 331. 1921; the Killip and Smith material

from erect herbs 1-3 meters high. Perhaps should not be restricted

so narrowly; Bitter proposed a subspecies hexazygum, Repert. Sp.

Nov. 11: 10, 19. 1912, glabrous, 4 pairs of leaflets, anthers nearly

2 mm. long, style 3.5 mm. long; there are doubtless fewer genetic

species; several names probably will be found to represent individuals

only. Section Polybotryon. F.M. Neg. 2625.

San Martin: Tarapoto, Williams 6045. Juanjui, Klug 4252 (det.

Standley). San Roque, Williams 7035. — Huanuco: Tingo Maria,

Allard 22193 (det. Lyman Smith); Stork & Horton 9536 (det. Stand-

ley); Ferreyra 880; 1575 (det. Morton); Soukup 2210 (det. Killip).

Cayumba, Mexia 8326 A. Pozuzo, 4676. Tulumayo, Ferreyra 2138

(det. Morton). — Junin: Chanchamayo, Isern 2241. Pichis Trail,

Killip & Smith 26140 (det. Morton). Near Peren4 Bridge, Killip

& Smith 25331. La Merced, 5267; Killip & Smith 23561; 24066 —

Loreto: Yurimaguas, Killip & Smith 27614; 28108 (all det. Morton,

S. semievectum) ; Williams 4264; 5351.

Solatium montanum L. Sp. PL ed. 2, 1: 336. 1763; 61. S.

tuberiferum Dunal in DC. Prodr. 13: 63. 1852. S. tuberiferum Dunal,

var. avenarium Dunal, I.e.

Low ascending-erect herb from a shallow set subglobose tuber

bearing many fibrous roots; indument sparse, evanescent, spiculose,

mostly persisting only on the bracted or ebracteolate sometimes

furcate few-flowered inflorescences; petioles usually or all well-

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

defined, often elongating to several cm., narrowly alate toward

apex by the more or less decurrent (basally long-attenuate) leaf-

base, commonly more or less amplexicaul but not decurrent on the

weak stems, entire to repand-dentate or lobulate; leaves ovate-

subrotund, ordinarily 5 or 6 cm. long, at least half as wide (-3),

peduncles to 1 dm. long, pedicels about 1.5 cm. long, nearly filiform;

calyx lobes suboblong-obovate, unequal or equal, subobtuse, partly

enclosing the pale berry, scarcely 5 mm. long; corolla white or

violet, rotate-substellate, about 1 cm. long, nearly as broad at

top; anthers oblong-ellipsoid, 4 mm. long, more or less exceeded

by the style, the stigma conical; berry about 5 mm. in diameter.—

DunaPs plant was characterized by a shallow corolla finally opening

stellate, the variety particularly by the presence of bracts, apparently

intangible characters in this instance; his diagnosis of the Linnaean

plant seems to be after the plate of Feuille'e in the latter's work,

pi. 46. 1766, and his variety on plate 160 of Ruiz and Pavon (2: 32);

Bitter proposed in herb, several under-species names. Variable in

foliage, the key character even may not be constant and indeed

some young specimens, as my 5860 and 5864, could be this species;

cf. in this connection S. phyllanthum Cav. S. montanum L. however

is at least characteristically a loma species. F.M. Negs. 2645;

8592; 33119. Section Regmandra.

Ancash: Lomas de Monzon, Goodspeed Exped. 9165; 9166 (det.

Johnston). — Lima: Chorillos, 5860; Weberbauer 5687. Near Lima,

Wawra; Feuillee, type. Prov. Lima and Chancay, Ruiz & Pavdn

(type, S. tuberiferum et var.); Weberbauer 7486b; Goodspeed Exped.

9098; 9100; 9238; 04013; 809 9a (all det. Johnston). Mount St.

Augustin, Weberbauer 5224- Atocongo, Pennell 14785. — Arequipa:

Mollendo, Weberbauer 1454> 1577. Near Chala, Goodspeed Exped.

15609 (det. Johnston). "Papas de lomas," "papas de montana."

Solatium multifidum Lam. Illus. 2: 17. 1793; 66. S. multifidum

R. & P. Fl. Peruv. 2: 37, pi. 17, fig. a. 1799. Witheringia multifida

(R. & P.) Dunal, Syn. Sol. 3. 1816. S. pinnatifidum R. & P. I.e.

pi. 170, fig. b. W. pinnatifida (R. & P.) Dunal, I.e. 2. S. senecioides

Dombey ex Dunal in DC. Prodr. 13: 65. 1852. S. Tafallae Macbr.

Field Mus. Bot. 8: 111. 1930 (S. multifidum R. & P. I.e. not Lam.

Illus. I.e.). S. callianthemum Bitter, ined.

Succulent herb, glabrous or obscurely evanescently pilosulous,

flowering as a slender annual with fibrous root, soon densely leafy,

the root and base becoming stout and stems more or less broadly

FLORA OF PERU 185

alate by the mostly bipinnatifid leaves, these to 2 dm. long, about

half as wide, the ultimate divisions obtuse; in age stems more or

less tufted, cicatricose and subligneous; racemes terminal, 2-6 cm.

long, simple, 2-3-branched or corymbose, the peduncles exceeding

the foliage and sometimes naked, sometimes with an entire or a

pinnately lobed bract, even on the same plant; pedicels capillary,

1-2 cm. long; calyx lobes suboblong, obtuse or subacute, 2 mm. long;

corolla blue, somewhat angulate, 1-scarcely 1.5 cm. broad; anthers

exceeded by style, yellowish, to 4 mm. long, dehiscence lateral;

fruit pale yellow on arcuate or recurved pedicels, 5 mm. in diameter;

stone cells not found by Bitter in this and related species, except

perhaps S. phyllanthum Cav. — Dunal remarked that in spite of

confusion in application of names he thought three distinct species

were concerned (he saw only four specimens); he stressed presence

or absence of bract on peduncle, this sometimes remotely glandular-

verruculose, development of inflorescence and other mostly vegetative

characters; the Arequipa form may be varietal, the corolla larger

(not always), the peduncles obscurely lineately glandular (?).

S. pinnatifidum Lam. Illus. pi. 115, f. 4- 1793 (S. rundnatum

L'He*r. ex Dunal, 68) with fibrous roots was S. pinnatum Cav.

according to Bitter in Herb. Dahlem; it is said also to include

S. Feuillei Dunal, I.e. 70, also of Chile, the root developed as stout,

fleshy; S. pinnatum Cav. with simply pinnate leaves is unknown

in Peru unless in hort. Bruns referred Gunther & Buchtien 107

and 107 a (not seen) from Mejia and Cachendo, Arequipa, to the

Chilean S. Remeyanum Phil., Cat. PI. Itin. Tarapaca 66. 1891,

with puberulent corollas 8-9 mm. long; the determination is doubtful.

A beautiful plant, characteristic of the dryer seaward slopes of

sandy lomas. F.M. Negs. 29724; 6747 (type, S. senecioides) ; 2599

(Weberbauer 1458}. Section Regmandra.

Ancash: Lomas de Monzon, Goodspeed 9182. — Lima: Chancay,

Dombey, type. Lima and Chancay, Ruiz & Pavdn (type, S. pinnati-

fidum). Pasomayo, Stork & Vargas 9343 (det. Johnston). Mt.

San Augustin, Weberbauer 5239; Asplund 13790. Chorillos, Weber-

bauer 5691 . Lurin, 5928. Lomas de Pasomayo, Vargas 1239. Lomas

de Lachay, Soukup 1851. Near Lima, Rose 18561; 18583; Gaudi-

chaud. — Arequipa: Mollendo, Worth & Morrison 15731; 15753 (det.

Johnston); Guenther; Weberbauer 1458 (type, S. callianthemum) ;

1572. Lomas de Checa, Raimondi. Pongo, Prov. Camand, Tafalla

(type, S. multifidum R. & P.). — Tacna: Near Tacna, Werdermann 724.

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

Solatium Murphyi Johnst. Contr. Gray Herb. 95: 34. 1931.

Glabrous (except corolla), succulent, erect, 1.5-3 dm. tall, sub-

dichotomously branched, the usually simple branches to 12 cm.

long; ovate or ovate-oblong leaves 1.5-2.5 cm. long, 5-8 mm. wide,

abruptly contracted to alate petiole, 5-7-lobed, the lobes obtusely

2-3-pinnate, the margins evidently revolute; inflorescence 1-2-

flowered, often terminal on the branches, pedicels slender, 5-9 mm.

long, reflexed in fruit; calyx 4-4.5 mm. long, the 5 oblong obtuse

lobes erect, 3-3.5 mm. long, 1-1.5 mm. wide, stipitate-glandular

within; corolla purple-blue, cupuliform or rotate-infundibuliform,

1.5-2 cm. across, about 12 mm. long, villosulous only without,

the rounded apiculate lobes about 8 mm. broad, 1.5 mm. long

with rounded lobules about 2 mm. wide, 0.9 mm. long in the sinuses;

filaments 4 mm. above base, anthers equal, oblong, 2-2.5 mm. long,

bipored; style 5.5 mm. long, stigmas compressed; fruit globose, di-

ameter 5-7 mm. — Seems to be related to the Chilean S. Feuillei

Dunal (S. pinnatum Cav.). According to Bitter, species apparently

without intermediate lobules in corolla sinuses. A distinct species

or a variant may be represented by Weberbauer 7960, the leaves

simply pinnate, the small lobules entire. Illustrated, Johnst. I.e.

pi. 7 (photo).

This pretty plant, specifically distinct or not, recalls to botanists

the ornithologist Robert Cushman Murphy, whose consciousness

of all nature is so pleasantly apparent in his "The Bird Islands of

Peru" (Putnam, 1925).

lea: Among rocks, foggy summit, Viejas Island, (Murphy 3219,

type). Near crest of San Gallen, (Murphy 3479). Bahia de la

Independencia, 500 meters, Weberbauer 7960?

Solatium phyllanthum Cav. Icones 4: 35, pi. 359. 1797; 63.

Witheringia phyllantha (Cav.) Dunal, Syn. Sol. 2. 1816. S. rhopalo-

stigma Bitter, Abhandl. Nat. Verh. Bremen 23: 148. 1914, form.

A low (vigorous plants sometimes about 1 meter long) at first

often somewhat ashy pubescent herb, persisting and becoming a

half-shrub, the ovate-rhombic to suboblong entire to sinuately lobed

leaves decurrent at base into the winged petioles and stems; leaves,

including the ordinarily scarcely defined petiole, usually 5-8 (11.5)

cm. long, about a third as wide; racemes few-many-flowered, dichot-

omous, seemingly terminal, the wing-based subtending leaf often

entire; flowers light purple or mauve, rarely white, 1-2 cm. wide;

pedicels flexuose or arcuate, 1-2 cm. long; calyx cyathiform, seg-

FLORA OF PERU 187

ments oblong, subobtuse, 5 mm. long, with somewhat rounded

membranous sinus in fruit; corolla plicate, spreading, angulate,

1-1.5 (2) cm. across; anthers yellow, early bipored, promptly laterally

dehiscent, well-exceeded by style, the stigma more conical; berry

whitish or reddish, finally about 1 cm. in diameter, subapical stone

cells 2 (Weberbauer 1554, fide Bitter). — Variable, especially in foliage,

degree and permanence of the simple indument, size of flowers.

An attractive flower in March and April, mostly on rocky slopes;

applied to wounds (Hinkley). Bitter has proposed, in herb., two

segregate species. Glabrate forms and young plants readily con-

fused with S. montanum (S. tuberiferum) with which it sometimes

grows and perhaps some undeveloped specimens referred here on

key character without roots may belong actually to S. montanum.

The southern state is often more puberulent but not always, my

331 and Munz 15517 being similar. F.M. Negs. 2634; 5317 (both

Weberbauer with ined. names). Section Regmandra.

Lima: Lomas, Pasomayo, Vargas 1227. Provinces of Lima,

Chancay and Lurin, Ruiz & Pavdn, type. Matucana, 331; Weber-

bauer 5252. Chosica, Weberbauer 5317 (form). Pachacamac, Mexia

04054 (det. Johnston, S. tuberiferum). Huarochiri, Goodspeed 33111

(det. Killip, S. tuberiferum). Atocongo, Ferreyra 3891 (det. L. B.

Smith, S. tuberiferum). — Arequipa: Quequena, Eyerdam & Beetle

22174- Mollendo, Hitchcock 22374. Chachani Mt., Hinkley 9.

Arequipa, Pennell 13049; 12343; Rose 19004; Sandeman 3777; Munz

15517; Seler 213; Balls 5869; 5871. Yura, T. D. A. Cockerell. Along

railroad to Puno, Shepard 249. Mollendo, Weberbauer 1554 (type,

S. rhopalostigma) . Near Mejia and Posco, (Gunther & Buchtien 90;

91 ; 92) . — Tacna : Near Tarma, Meyen. Candarave, Weberbauer 7368;

Metcalf 30372. Chile. "Papa cimarrona" (Balls), "nuccho bianco"

(Hinkley).

Solanum quercifolium L. Sp. PI. 264. 1753; 71. S. reclinatum

L'Her. ex Pers. Syn. 1: 225. 1805?; 68. S. infundibuliforme Phil.

Anal. Mus. Nat. Chile 65. 1891, as to Peru?

A clambering glabrous or nearly glabrous herb with thickish pin-

nately mostly (3) 5-7-lobed leaves; lobes ovate-oblong to linear-

oblong, entire or the terminal toothed, the sinuses rounded; leaves

5-8 cm. long, decurrent into the petiole; calyx teeth broadly ovate,

acute, early 1 mm. long, to 1.5 mm. in fruit; corolla lavender, about

1 cm. long and broader; fruiting pedicels even 1 cm. long. — Has

been characterized as having more open pseudo-terminal cymes and

188 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL XIII

mostly 7 leaf segments in contrast to S. radicans L. f., seemingly and

probably a fanciful sorting of specimens; here may belong Gunther &

Buchtein 102, 102a and 1401 from Pasco and Mollendo, Arequipa,

determination Brims, S. infundibuliforme Phil. I.e. with corolla 2 cm.

long (ex. char.) . Original by Jussieu from Peru, probably near Lima;

in Herb. Willd. ex Thunberg as from Peru; type of S. reclinatum, by

Dombey, possibly not from Peru, scarcely distinct, the leaf segments

linear-oblong; there are similar Bolivian plants, possibly an exag-

gerated development in shade, terminal leaf lobe to 1 dm. long!

F.M. Negs. 2908; 6746 (S. reclinatum). Section Dukamara.

Lima: Near Lima, Soukup 1923 (det. Killip) ; Matucana, 138; 381

(both det. Johnston). — Huanuco: Near Huanuco, Stork & Horton

9372 (det. Standley); Sawada 62. — Junin: Tarma, Killip & Smith

21 881 . — Loreto : Rio Morona, Dennis 291 64 (det. Morton) . Paraiso,

Rio Itaya, Williams 3382. — Cuzco: San Geronimo, Pennell 14205.

Ollantaitambo, Pennell 13687; Cook & Gilbert 371 (both det. John-

ston).— Arequipa: Tingo, Pennell 13100.

Solatium radicans L. f. Dec. pi. 10. 1762; 71. S. cymosum

Herrera, Contr. Fl. Dept. Cuzco 2: 168. 1921.

Perhaps differs from S. quercifolium L. in having 5 leaf segments,