V OF
ILLINOIS LIBRARY
AT URBANA-CHAMPAIGN
BIOLOGY
NOV 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 6837Cuzco: 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 2278Cuzco :
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; 4812LoTeto :
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 Tuberarium 1
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,
denser pseudo-axillary cymes, much smaller pale purple (or nearly
white) flowers (about 5 mm. wide) and narrowly ovate calyx lobes
1.5 mm. long to nearly 3 mm. long in fruit. Apparent intermediates
exist and character appears tenuous. The Hitchcock specimen, "after
October rains" has upper leaves about 1 dm. long, corollas nearly
7 mm. long, calyx teeth 2 mm. long. Section Dulcamara.
Piura: Huancabamba, (Bonpland). Cajamarca: Cutervo, Rai-
mondi (det. Werdermann). Amazonas: Chachapoyas, Mathews (det.
Morton). Lima: Valle de Chillon, Velarde Nunez 908 (det. Morton).
Huanuco: Raimondi (det. Werdermann). Junin: Huancayo,
Ledig 35; Soukup 2702 (det. Morton). Tarma, Killip & Smith
21881. Ayacucho: Pomobamba, Raimondi (det. Werdermann).
Cuzco: Sicuani, Cook & Gilbert 88. Urubamba Valley, Herrera 1734
(det. Werdermann); (Herrera, type, S. cymosum). Arequipa: Yura,
Cockerell. Mollendo, Hitchcock 22373. Chala, Raimondi (det. Werder-
mann). Bolivia, Chile. "Kusmaillu" (Herrera); "cushay," "cuspallo,"
"cusmayllu" (Cook & Gilbert).
Solanum Seaforthianum Andrews, Bot. Repos. pi. 504- 1797-
1804; 67.
FLORA OF PERU 189
Glabrous (or nearly), sometimes climbing several meters; petioles
2-3 cm. long, straight or curved; leaves to about 1 dm. long with
mostly 5-9 ovate-oblong or subelliptic leaflets, the 3-5 upper often
only segments by reason of their basal union, all somewhat acumi-
nate, 3-4 (8) cm. long, the terminal little larger; cymes subterminal,
finally lateral, rather few-many-flowered; pedicels slender, incrassate
above, 5 or 6 mm. long; calyx narrowed at base with 5 remote minute
teeth or these nearly obsolete; corolla stellate, violet or lilac, (1) 2-
2.5 cm. across, obscurely or not puberulent; stamens 4 (4.5) mm. long,
exceeded by the glabrous style; berry globose, 6-9 mm. in diameter,
finally reddish. Illustrated, Bot. Mag. 45: pi 1982. Section Poly-
botryon.
Loreto: Yurimaguas, Williams 1+323; 1*877. West Indies; north-
ern South America; Mexico.
Solarium semievectum Bitter, Repert. Sp. Nov. 12: 542. 1913.
A nearly glabrous fleshy or hollow-stemmed herb, sparsely pilose
only on the leaf rachis and midnerves and at the leaf insertions; peti-
oles about 1.5 cm. long; leaflets 3 pairs, the lower lateral petiolate,
the upper subsessile, the former mostly 8.5 X 2.5 cm., the latter
14 X 3.5 cm., the terminal 8.5 X 3.5 to 15 X 5 cm.; half of each
inflorescence axillary, the other half borne from about the middle of
the slender petioles; flowers and fruit unknown in type. Unless by
the inflorescence apparently the same as S. conicum R. & P.; prob-
ably the second inflorescence is also axillary from a young leaf, un-
developed or broken off. Section Polybotryon. F.M. Neg. 2638.
Huanuco: Muna, 4001. Loreto: Yurimaguas, Poeppig, type.
Puerto Arturo, below Yurimaguas, Killip & Smith 27760 (det. Mor-
ton; det. Standley, S. mite).
Solatium semiscandens Bitter, Repert. Sp. Nov. 12: 142. 1913.
Scandent or semiscandent shrub, the younger fistulose stems sub-
floccose with many-celled hairs; petioles 3.5-4.5 cm. long; leaves
ternate, thick, glabrous above, sparsely pubescent beneath; leaflets
broadly elliptic or elliptic-lanceolate, the lateral oblique at base, on
petiolules 6-8 mm. long, the blades 4.5-5 cm. long, 2.2-3 cm. wide,
the terminal 5- nearly 7 cm. long; inflorescence axillary, geminate,
the peduncles 5-10 cm. long, the 4-9 apically congested pedicels
1.5-2 cm. long; calyx lobes 1.5 mm. long; flowers lilac or white with
purple bases, 1-1.5 cm. broad, the fleshy petals pubescent without,
7 mm. long, 4-5 mm. wide at base; anthers sessile, 6x3 mm.; style
190 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
6.5 mm. long, papillose only above. Cf. S. ternatum. Section Poly-
botryon.
Hudnuco: Muna, trail to Tambo de Vaca, 1*279. Mufia, (Pearce,
type).
Solanutn subquinatum Bitter, Repert. Sp. Nov. 12: 144. 1913.
A sparsely pubescent or glabrate herb related to S. ternatum
R. & P. but the leaflets sometimes five, all lanceolate and attenuate
at both ends, the smallest about 1.5 cm., the terminal about 3.5 cm.
long; peduncles 4-5 cm. long, with 4-10 flowers at apex on pedicels
1-1.5 cm. long; calyx 6 mm. across, lobes obtuse, obscure; corolla
12 mm. broad, lobes 6 mm. long, 2.5-3.5 wide, scarcely more than
papillose pubescent; anthers subsessile, 5X2 mm., cordate at base;
style 5.5 mm. long, obscurely or not papillose, stigma obtuse; berry
(young) subconic. Corolla pale yellowish-brown, center yellow;
leaves dark green, lustrous (Woytkowski) . Section Polybotryon.
Amazonas: Chachapoyas, (Mathews, type, Herb. Kew and Brit.
Mus.). Hudnuco: Divisoria, 1,600 meters, Woytkowski 512 (det.
Cuatrecasas, S. semiscandens).
Solatium ternatum R. & P. Fl. Peruv. 2: 38, pi. 172, fig. a.
1799; 67.
A scandent shrub similar to S. semiscandens Bitter but glabrous
and leaflets elliptic to oblong-lanceolate, shortly acuminate to sub-
obtuse, rounded to acute at base, the lateral more shortly and
obliquely, 5-7 cm. long to about 2 cm. wide; peduncles 1-5 cm. long,
the 3-6 subumbellate pedicels often 1 cm. long; calyx 5-crenate;
flowers purplish white to purple-violet, about 1 cm. long; berry
cherry size, ovoid-conic, whitish. Leaves soft, fleshy, or little puber-
ulent on veins beneath (Woytkowski). S. trifolium Dunal, 68, type
by Tafalla from Guayaquil, is distinctive by its subrotund leaflets.
F.M. Neg. 2639. Subgenus Bassovia Bitter; that is, Polybotryon.
Huanuco: Cuchero, Ruiz & Pavdn, type. Junin: Agua Dulce,
Woytkowski 35416 (det. Cuatrecasas). Utcuyacu, Woytkowski 1168
(det. Killip). Colombia.
Solarium Uleanum Bitter, Repert. Sp. Nov. 12: 139, pi. 1. 1913.
S. Uleanum Bitter, var. unipedunculata Bitter, I.e. 140.
High-climbing by scattered and fascicled adventitious rootlets;
densely pubescent, the trichomes mostly simple; petioles 3.5-5 cm.
FLORA OF PERU 191
long; leaflets 6 (-7) pairs, thin, broadly lanceolate or nearly oblong-
lanceolate, subacute, the lowest about 4 cm. long by less than 1.5 cm.
wide, the uppermost and terminal about 6 cm. long, 2 cm. wide;
peduncles geminate or solitary, axillary, 1-2.5 cm. long; flowers sev-
eral to two dozen; pedicels often 8 mm. long (15 in fruit) ; calyx lobes
obtuse; corolla white or yellowish, the lobes about 4.5 mm. long;
filaments 0.7 mm. long, anthers 1.5-2 mm. long, cordate at base;
style 5 mm. long, papillose; berry longer than broad, attenuate both
ends, evanescently pubescent. Var. gracilescens Bitter, I.e. 141, is
more slender; petioles 1.5-2 cm. long; leaflets only 5 pairs, the small-
est about 17 X 8 mm., the terminal about 26 X 8 or 9 mm., all more
sparsely pilose; flowers 7-14. Leaves green both sides (type) or in
variants purple or violet beneath. F.M. Neg. 3178. Subgenus Bas-
sovia according to Bitter, I.e. 17: 331. 1921; that is, Polybotryon.
Rio Acre: Porto Carlos, (Ule). San Francisco, Ule 97561), type;
9756 (type, var. unipedunculata) .
SERIES 4 (mostly Leiodendron, sens, lat.)
Shrubs or small trees, virgate half-shrubs, rarely vines or sub-
scandent, never prickly; leaves entire. Includes for convenience
S. Miquelii, and entire-leaved species of Dulcamara, entire-leaved
species of Polybotryon, the trailing S. oxycoccoides, the lax S. chacha-
poyasense, the 4-merous S. cruciferum (partly at one time Anthoresis,
later Anarrichomenum) ; it is doubtful if these groups, as well as
Anthoresis, Anthopleuris, Indubitaria, for that matter, are natural
associations or at any rate helpful classification or identification aids
maintained separate from Leiodendron. S. caudatum is omitted from
the key as too imperfectly known.
Inflorescence more or less compound, often early terminal, rarely
capitate; species usually somewhat pubescent or defined peti-
oles short or absent (plants allied to S. acuminatum, S. patellare
or forms of these might be sought here; also S. styracioides) .
Leaves not obovate nor alately long-decurrent, sometimes acute at
base; rarely 1.5 dm. long, often much shorter.
Vine, puberulent above; leaves often undulately crisped.
S. styracioides, S. Miquelii.
Shrubs or trees; leaves usually plane.
Leaves rather oblong-lanceolate (sometimes subovate, S. au-
reum); indument beneath often obscure, fulvous, rarely
grayish or absent.
192 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Flowers showy, about 2 cm. long S. patulum.
Flowers much smaller.
Pubescence of leaves distributed, not axillary-tufted.
Leaves not bullate-asperous, sometimes scabrous.
Flowers usually in the acuminate leaves . . S. nitidum.
Flowers usually above the acute or shortly acuminate
leaves S. aureum.
Leaves strongly asperous or (type) bullate-asperous.
S. selachophyllum.
Pubescence tufted in nerve axils S. manicatum.
Leaves rather ovate, canescent-stellate at least beneath.
Calyx subcrenate; leaves bicolor, green but sparsely stellu-
late above S. salviifolium.
Calyx lobed; leaves more or less canescent both sides.
Stiped trichomes dominant on younger parts.
S. umbellatum.
Stiped trichomes not prevalent, at least usually.
S. verbascifolium.
Leaves somewhat obovate (unless aberrant or allied forms), basally
acute or cuneate to alately long-decurrent; often ample, often
sessile or subsessile; never much pubescent, mostly glabrous
or glabrate (many ambiguous specimens exist, species dubious).
Petioles alate-margined even to base by decurrent suboblong-
obovate blades, these often crowded, alternate, (1) 2-3 dm.
long.
Leaves usually 3-4 cm. wide, 1-1.5 dm. long; flowers to 1 cm.
long, congested S. monadelphum.
Leaves finally much larger.
Flowers 1-1.5 cm. long; leaves sessile, even clasping.
S. sessile.
Flowers 5-8 mm. long; leaves subsessile.
Leaves glabrous S. marmellosanum.
Leaves early scurfy tomentose beneath . . S. oblongifolium.
Petioles shortly margined or scarcely, typically defined, often
geminate; corollas 5-7 mm. long.
Pedicels stout, soon incrassate, 5-10 mm. long.
S. dibrachiatum.
Pedicels subfiliform, soon about 1 cm. long S. pulchrum.
FLORA OF PERU 193
Inflorescence simple, axillary, lateral or at branchlet tips, rarely early
terminal (sometimes 1-furcate, S. patellare, S. amotapense, S. lu-
teoalbum, racemes rarely panicled, S. acuminatum) .
Erect or suberect often shortly branched shrubs or half-shrubs,
densely pubescent; corollas 5 mm. long or longer.
Leaves and branchlet tips sericeous pilose or villous; leaves ovate
to elliptic.
Leaves geminate, the smaller stipuliform; trichomes branched.
S. xanthophaeum.
Leaves alternate, stipuliform often on axils; trichomes simple.
S. chachapoyasense, S. sericeum.
Leaves at least beneath and branchlets (except var.) more or less
arachnoid-lanate-tomentose or leaves oblong-lanceolate (see
also third contrast).
Flowers subumbellate, peduncles absent or short, pedicels to
1.5 (-2) cm. long.
Petioles 1-1.5 cm. long or longer.
Calyx lobes low, rounded, firm- margined; indument sub-
persisting at least on branchlets (except var.).
S. ochrophyllum.
Calyx lobes about oblong, subobtuse; indument, except
inflorescence, soon evanescent S. nutans.
Petioles 3-5 mm. long; calyx crenate or lobes rounded.
Leaves all or mostly alternate, about 5-7 cm. wide, ob-
tuse; calyx crenate S. chloranthum.
Leaves geminate, the larger 3^1 cm. wide, acuminate, the
smaller stipuliform; calyx lobes rounded. . .S. solum.
Flowers few, in well-peduncled inflorescences.
Leaves ovate.
Leaves tomentose-stellate both sides; calyx deeply parted;
tomentose S. luteoalbum (var.).
Leaves green above, nerves pubescent.
Calyx gibbous, margin denticulate S. amotapense.
Calyx deeply parted S. Lechleri.
Leaves oblong-lanceolate S. nitidum (var.).
Leaves (beneath) and branchlets rarely hispidulous, early scurfy
or puberulent, usually soon glabrate or glabrous or indu-
194 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
ment localized; corollas 5-10 mm. long (unknown in S. lu-
teoalbum).
Leaves at least mostly alternate, subequal and rarely 1 dm.
long.
Flowers, except S. pseudocapsicum, several, 5-merous; leaves
usually 5-10 cm. long.
Leaves about ovate, obtuse or acute, rarely obtusely acu-
minate.
Calyx lobed or parted.
Leaves rather attenuate to base, obtuse, subobtuse
or subobtusely acuminate, usually pubescent in
nerve axil . . . S. maturecalvans, S. amblophyllum.
Leaves rounded to acute base, acute; indument, if
present, general S. luteoalbum.
Calyx gibbous below, the margin denticulate.
S. amotapense.
Leaves oblong-lanceolate, more or less acuminate.
Pedicels 1-3 in each axil S. pseudocapsicum.
Pedicels racemose S. nitidum.
Flowers 4-merous, 1-2; leaves 2-4 cm. long. .S. cruciferum.
Leaves often geminate and more or less unequal, the larger
mostly 1-1.5 dm. long, even when alternate.
Leaves ovate- or broadly elliptic, rounded to acute or shortly
cuneate at base, little or not narrowed to rounded or
cuspidate tip; smaller leaves (if present) more rounded.
Leaves firm to coriaceous, glabrous to evanescently scurfy,
geminate.
Corollas rather thin, 6 or 7 mm. long. .S. acuminatum.
Corollas fleshy, 8-10 mm. long.
S. psidiifolium, S. oblongum.
Leaves membranous, often puberulent as inflorescence.
Inflorescence racemose, ped uncled; leaves in part gem-
inate S. patellare.
Inflorescence subsessile; leaves often alternate.
S. deflexiflorum.
Leaves oblong-elliptic, subequally attenuate both ends,
often slightly and not or little diverse except in size;
corollas 7 or 8 mm. long S. oppositifolium.
FLORA OF PERU 195
Erect, simple or virgately branched half-shrubs, or sprawling to
scandent, rarely trailing perennial, often rooting at lower
nodes; corollas 2-3, rarely 4 mm. long (longer in oxycoccoides)
(cf. S. deflexiflorum).
Leaves geminate at least some of the upper, all very unequal,
the smaller stipuliform, diverse.
Flowers 2-3 mm. long; leaves not all (rarely none) geminate.
S. loretoanum, S. confine.
Flowers 4 mm. long; leaves (flowering branchlets) all geminate.
S. anisophyllum.
Leaves alternate, subequal, not diverse, sometimes large.
Leaves more or less decurrent; stems glabrous or glabrate.
Corolla lobes about 4 mm. long, broadly ovate-oblong to
narrowly oblong; berry early long-conic, finally 1 cm.
thick S. anceps.
Corolla lobes 2-3.5 (5) mm. long, oblong or suboblong.
Berry even early not conical; pedicels in flower arcuate,
5-10 mm. long S. robustifrons.
Berry early conical; pedicels subascending, to 5 mm. long.
S. theobromophyllum, S. angustialatum.
Leaves not or little decurrent at rounded base, somewhat
pubescent both sides.
Flowers 2-3 mm. long; stems stout. . . .S. hederiradiculum.
Flowers 5-6 mm. long; stems slender, trailing.
S. oxycoccoides.
Solanum acuminatum R. & P. Fl. Peruv. 2: 34, pi. 159, fig. a.
1799; 146. S. hypomicropogon Bitter, Repert. Sp. Nov. 18: 56. 1922.
A glabrous or glabrate shrub or tree with geminate oblong-elliptic
or ovate abruptly cuspidate-caudate leaves, somewhat unequal in
size, the larger to 1.5 dm. long, the smaller of each pair about 8 cm.
long or longer, lustrous above, reticulate-venose beneath, the 7 or 8
lateral nerves there prominent; petioles 10-12 mm. long; racemes
nearly opposite the leaves, simple; peduncle (rachis) cicatrose to
1 cm. above base, becoming 4-5 cm. long; flowers many, secund, in
two alternate series, the slender pedicels 1-1.5 (2) cm. long; calyx
cyathiform, broadly ovate lobes subacute or apiculate; corolla in type
white, about 12 mm. across, the subacute cucullate elliptic-oblong
lobes to 7 mm. long; anthers obovate, the cells gibbously thickened,
196 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
to 4 mm. long; filaments 1 mm. long; style 5- nearly 5.5 mm. long;
berry (Williams) nearly 1 cm. in diameter. Sometimes a few tri-
chomes are in the nerve-axils of leaves beneath; doubtfully distinct
from S. oblongum R. & P. Cuzco and Puno specimens had pale to
dark purple flowers, leaf-nerves in latter more prominent, at least a
variant but probably nearer S. oblongum R. & P. than S. Warmingii
Hiern. Bitter's proposed species was based on tree (10 meters) with
more acuminate leaves, more open inflorescence, larger flowers in
contrast to S. oblongum; these characters, developed in degree, sug-
gest that a single species may in fact be concerned. S. triste Jacq.
Stirp. Amer. 50. pi. 40. 1780, to which a Poeppig specimen was re-
ferred in several herbaria presumably belongs here or to S. oblongum
R. & P., scarcely the same as Jacquin's plant of the Caribbean. An
open shrub or small tree. Illustrated, Mart. Fl. Bras. 10: pi. 2, figs.
20-25. Section Leiodendron. F.M. Negs. 2594; 2610.
Tumbez: Region del Caucho, Velarde Nunez 355 (det. Morton
with query). Cajamarca: Prov. Hualgayoc, Soukup 3882. San
Martin: Lamas, Williams 6352. Tarapoto, Williams 5450. Hud-
nuco: Cueva Grande, Near Pozuzo, 4794. Mufia, 4027; 4155. Chin-
chao, Ruiz & Pav6n, type. Junin: Merced, (Weberbauer 1876, type,
S. hypomicropogon) . Loreto: Yurimaguas, Poeppig 2279 (distr. as
S. triste Jacq,). Near Iquitos, Williams 8242. Masisea, Killip &
Smith 26863 (det. Morton, S. hypomicropogon}. Cuzco: Pillahuata,
Pennell 13951. Puno: Santa Domingo to Chabuca.
Solatium amblophyllum Hook. Bot. Misc. 2: 231. 1831; 157.
S. hypostichopogon Bitter, Repert. Sp. Nov. 16: 95. 1920. S. barbu-
latum Zahlbr. Ann. Naturh. Hofm. Wien 7: 7. 1892.
A stocky short-branched shrub in hills or small open bush or tree
(especially in higher areas), the mature foliage usually glabrous ex-
cept (sometimes absent) loose tufts of pubescence in the axils of the
principal, often (9) 13 leaf -nerves beneath; internodes 2.5-4 cm. long,
sometimes much shorter; leaves yellowish-green, elliptic-lanceolate
to suboblong, obtuse, mostly about 7 (5.5-10) cm. long; inflorescence
opposite, the leaves subterminal, few- (a dozen or so) flowered; pe-
duncles 0.5-2.5 cm. long, pedicels about 1.5 cm. long (-2.5 cm. in
fruit, nutant); calyx lobes subobtuse, (1) 1.5-2 mm. long; corolla
white, (14) 18 mm. across, the suberect or reflexing lobes 6-8 mm.
long, partly papillose-pubescent; filaments minute, anthers (3.5)
5 mm. long, subellipsoid ; style 6-6.5 mm. long, stigma obtuse; berry
about 14 (10) mm. in diameter. Bitter distinguished his plant by
FLORA OF PERU 197
slightly shorter internodes, leaves and anthers; also by the smaller
flowers, remarking that these characters may prove variable; he gave
a description of Weberbauer 209, I.e. 93, from which I have drawn.
F.M. Negs. 2611 (S. hypostichopogon) ; 33048 (S. barbulatum). Sec-
tion Anthoresis.
Piura: Palambla, (Weberbauer). Cajamarca: Cutervo, Jelski 56
(type, S. barbulatum) . Huambos, Prov. Chota, Stork & Horton 10066
(det. Standley) ; Soukup 4.534 (det. Morton, S. oblongum). La Liber-
tad: Prov. Santiago de Chuco, Lopez 1012. Lima: Below Obrajillo,
Wilkes Exped., type. Above Obrajillo, Pennell 144-18. Chicla,
Weberbauer 209. Puente del Infiernillo (Lima-Oroya R. R.), Seler
236 (type, S. hypostichopogori) . Viso, 568. San Mateo and Matu-
cana, Raimondi (det. Werdermann); Isern 2537. Huanuco: Muna
to Tambo de Vaca, 2,500 meters, 4328 (open, spreading shrub). Near
Muna, 4099. At 2,700 meters, Woytkowski 64 (det. Cuatrecasas, tips
tomentose). Junin: Prov. Jauja, Ochoa 531. Carpapata, Soukup
3465 (det. Lyman Smith). Huancavelica: Salcabamba, Stork & Hor-
ton 10273. Apurimac: Trancapata, Vargas 9621. Bolivia. "Shu-
ruco" (Ochoa).
Solatium amotapense Svens. Amer. Journ. Bot. 33: 483. 1946.
Unarmed shrub 1 meter high, the contorted branches (these 5-
6 mm. thick) glabrous, sulcate, the younger branchlets and peduncles
puberulent with simple rarely bifurcate or branched trichomes; peti-
oles 2-4 cm. long; leaves alternate, approximate, cordate-ovate, acu-
minate, to 9 cm. long, 5 cm. wide, sparsely pubescent above on the
nerves, all over beneath, membranous; inflorescence terminal, mostly
racemose, the peduncles 4-5 cm. long; calyx urceolate, 3-4 mm.
broad, gibbous below, the 5 teeth 2 mm. long, strigose; corolla white,
shortly campanulate, 1-1.5 cm. across, the 5 acute lobes pubescent
without; filaments equal, basally adnate, 0.5-1 mm. long, anthers
oblong, 5 or 6 mm. long, narrowed toward the apical pores; style
glabrous, 9 mm. long; fruit globose, glabrous, brown, 1-1.5 cm. in
diameter; seeds deltoid, 5 mm. thick, strongly compressed, yellow.
This has a superficial resemblance to S. luteoalbum Persoon (Sven-
son). The Weberbauer specimen has leaves to twice as large, corolla
at least 2 cm. broad, filaments nearly 2 mm. long. Illustrated, Sven-
son 484, pi. 19, fig. 1.
Tumbez: Amotape Hills, Cerro Prieto, (Haught & Svenson 11634,
type); Haught 195. East of Chicama, Weberbauer 7634?
198 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solanum anceps R. & P. Fl. Peruv. 2: 36, pi. 169. 1799; 155.
Bassovia anceps (R. & P.) Rusby, Bull. Torrey Club 26: 197. 1899.
Glabrous, the cymes obscurely or minutely granular; stems terete,
little branched, about 1 meter high; leaves ovate-oblong-elliptic,
acute or shortly acuminate (type), 1.5 dm. long, 8 cm. wide, or to
2.5 dm. long, nearly 1.5 dm. wide, or much smaller and narrower,
a little decurrent at base into the 1.5-5 cm. long petiole; subumbellate
cymes axillary, binate or ternate, in fruit to 2.5 cm. long, 3-7 (8)
-flowered; pedicels 2-6 mm. long, slightly longer and erect in fruit;
peduncles 12-15 mm. long; corolla greenish, petals 2 mm. long;
stamens about 3 mm. long, the very short filaments capillary;
anthers ovate-elliptic, connivent; style filiform, erect, longer than
stamens, stigma bilobed; berry early conical, a little curved toward
the long beak, whitish, 1.5 cm. long, nearly 1 cm. thick near base,
cuspidate, at maturity about 1 cm. thick. Type imperfect; probably
includes, naturally, S. theobromophyllum, S. angustialatum. Sub-
genus Bassovia (Aublet) Bitter, Repert. Sp. Nov. 17: 329-331. 1921;
Bitter included in this subgenus (as to Peru) S. incurvum (Cy-
phomandra fide Werdermann), S. mite, S. conicum, S. diffusum,
S. ternatum, S. angustialatum, S. hederiradiculum, S. theobromo-
phyllum, S. trizygum, and S. Uleanum as well as S. suaveolens, var.
Endlicheri and S. jasminoides; the subgenus included mostly Poly-
botryon of Dunal, defined by Bitter as perennial herbs or subligneous,
glabrous or nearly, inflorescence axillary, often geminate, flowers
secund, filaments short, glabrous, style nearly, corolla rotate or
stellate, berry globose or conical, sclerotic ("stone") granules lacking.
This is not a subgenus as constituted but may be a not unnatural
subgroup, with modification and expansion. F.M. Neg. (no num-
ber). Bassovioides.
Huanuco: Cuchero, Ruiz & Pavdn, type; Poeppig 1625. Divisoria,
Woytkowski 560. Pampayacu, Poeppig 1469. Below Rio Santo
Domingo, Rio Huallaga Canyon, 4243. Bolivia.
Solanum angustialatum Bitter, Repert. Sp. Nov. 11: 471.
1912.
Stems very or obscurely narrowly alate by the decurrent 8-10
mm. long petioles; leaves broadly lanceolate, attenuate at both ends,
acute, to 2.5 dm. long, about 1 dm. wide, glabrous unless minutely
stipitate-glandular on the 17-18 nerves both sides; peduncles gem-
inate, axillary, to 12 mm. long, 10-14-flowered; pedicels glabrate
but minutely glandular, about 5 mm. long; calyx 2 mm. across,
FLORA OF PERU 199
scarcely lobed; corolla lobes 2-3 mm. long, papillose-pilose; filaments
0.3-0.4 mm. long, glabrous, anthers ellipsoid, 1.7 mm. long; style
to 3.5 mm. long, with a few 1-2-celled trichomes toward the
obtuse stigma, not obviously papillose. Originally placed by author
in section Polybotryon. F.M. Neg. 33045.
San Martin: Tarapoto, Spruce 484-9, type; 6260. Jepelacio,
Klug 3757. San Roque, Williams 6929; 7689. Prov. Mayo, Williams
6260. "Viuda shambu" (Williams).
Solatium anisophyllum Van Heurck & M. Arg. Obs. Bot. 52.
1870.
Similar to S. confine Dunal but as to type all the upper leaves
geminate, the smaller of each pair suborbicular and the flowers
somewhat larger. It seems probable that these are variable char-
acters and unless there are other differences the plant, with S.
loretoanum Bitter, which apparently is intermediate, is actually a
part of one entity. F.M. Neg. 22874. Section Leiodendron.
San Martin: Tarapoto, Spruce 4250, type (once referred by Bitter
to S. pulchrum); also 4830. San Roque, Williams 7616.
Solatium aureum Dunal, Syn. 16. 1816; 102. Vars. latelanceo-
latum Bitter, Bot. Jahrb. 54, Beibl. 119: 13. 1917; angustelanceolatum
Bitter, I.e. S. cutervanum Zahlbr. Ann. Naturh. Hofm. Wien 7: 7.
1892.
A sometimes climbing shrub with alternate narrowly lance-
olate to ovate-elliptic subacute entire leaves, lustrous above in age,
stellate-pilose beneath, and terminal laxly divaricate inflorescences
pulverulent-tomentose with (at least early) rusty-yellow stellate
branched trichomes; petioles to 1.5 cm. long; leaves 3-5 (-7) cm.
long; lateral nerves 5-8, prominent beneath; peduncles dichotomous;
pedicels basally articulate, 4-7 mm. (in fruit -12 mm.) long; calyx
about 4 mm. long, 7-8 mm. across, the triangular lobes acute;
corolla violet, about 17 mm. broad, tomentose without, the lanceolate-
oblong lobes 6x4 mm., revolute margined; filaments minute,
anthers narrow, to 4 mm. long, pubescent only within; style to
6.5 mm. long, curved below the subglobose stigma, papillate toward
base; berry without granules. Perhaps the second of Bitter's vari-
eties merits recognition on the basis of the leaves being narrowed
at each end, the fruit the typical form, somewhat cordate at base;
he gave a detailed description in Repert. Sp. Nov. 16: 83. 1919,
which is in part included here. Perhaps correctly a part of S. nitidum
200 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
R. & P. The var. riobambense Werd., Notizbl. Bot. Gart. Berlin 12:
379. 1936, has leaves glabrous above and beneath except for a few
brownish trichomes on midnerve; inflorescence irregularly dichot-
omous; this is often so in contrast to the usually simple inflorescence
of S. nitidum. Series Endotricha Bitter, anthers pubescent on the
inner side, style papillose at base, and Series Monadelphoidea Bitter
(includes S. monadelphum, S. marmellosanum, S. sessile, S. ptero-
podum, S. amblophyllum, and S. hypostichopogori) are according to
Bitter in 1920 "diverse." F.M. Negs. 2890; 33065 (S. cutervanum).
Section Anthoresis.
Piura: Ayavaca, Soukup 4337 (det. Morton). Cajamarca:
Cutervo, Jelski 30 (type, S. cutervanum). Huanuco: Mito, 1859.
Yanano, 4933. Chaglla, Weberbauer 6700 (type, var. latelanceolatum) .
Ecuador.
Solarium caudatum Dunal in DC. Prodr. 13: 140. 1852.
Shrub with glandular subterete branches and leafless flowering
branchlets, scarred by many large leaf- or flower-bases; leaves gla-
brous, obovate-lanceolate, cuneate, abruptly cuspidate (acumen sub-
falcate, 16 mm. long), about 2 dm. long and nearly one broad,
unevenly verruculose beneath, 8-10 lateral nerves there prominent,
reticulate with the veins; petioles subterete, canaliculate above,
rugulose, 1.5-2 cm. long; peduncle (or perhaps branchlets?) to 3
dm. long, little incrassate toward apex; pedicels (or peduncles) 2-
flowered, solitary or geminate; fruiting calyx 8 mm. across, deeply
parted, the divisions linear, acute; berry globose, 8-10 mm. thick.
Flowers unknown; the long subulate acumen from the rounded
leaf-tip is only partly equaled by Killip & Smith 28087 (sterile
in specimen seen) which Morton in herb, suggested as a new species.
F.M. Neg. 34114. Section Leiodendron.
Peru(?): (Ruiz & Pavdn, type, Herb. Geneva).
Solarium chachapoyasense Bitter, Repert. Sp. Nov. 16: 13.
1919.
A shrubby plant, possibly scandent, the leaves and upper branches
(internodes 2-3 cm. long) densely and softly pubescent with yellowish
simple hairs; petioles 1-1.5 cm. long; alternate leaves elliptic-
lanceolate, 2.5-4.5 (6) cm. long, about 1.5-2 (4) cm. wide, mostly
very oblique at the rounded base; minute (0.5-1.5 mm. long) stipule-
like leaves present in the leaf -axils; inflorescence 5-6-flowered, con-
gested to lax, borne at the tip of small branchlets, these with inter-
FLORA OF PERU 201
nodes 2-7 mm. long; pedicels about 1 cm. long; calyx lobes lance-
olate, often in part connate, 4 mm. long; corolla rotate-stellate,
1.5 cm. across, the lanceolate lobes involute and pilosulous toward
tip, 8 mm. long; filaments 1 mm. long, pubescent below, anthers
narrowly ellipsoid, 4 mm. long, cordate at base, introrse apical
pores oblique; style 7.5-8 mm. long, glabrous or nearly, the stigma
obviously bilobed. A soft, weak herb, corolla white (Stork & Hor-
ton) but stem ligneous. The section, described by Bitter, I.e. 11:
247. 1913, includes in Peru also S. oxycoccoides Bitter and several
Ecuadorian species, all with solitary flowers or few often on short
axillary branchlets, stem usually scandent with nodal roots. F.M.
Neg. 23104. Section Anarrichomenum.
Cajamarca: Shrub-land west of Socota, 2,800 meters, Stork &
Horton 10104- Amazonas: Chachapoyas, Mathews, type. Junin:
Carpapata, Soukup 3463.
Solatium chloranthum Spreng. Neue Entd. 28: 1822; 372?
S. arenarium Sendt. in Mart. Fl. Bras. 10: 26, pi 3, figs. 16-18. 1846;
DC. Prodr. 1: 129. 1852, fide Bitter, Repert. Sp. Nov. 16: 400. 1920.
Shrub, the terete branchlets early pulverulent-tomentose with
yellowish, densely branched trichomes, their branches divaricate,
simple, this indument also on leaf venation above (lateral nerves
8-10), leaf-surface beneath and few-flowered simple inflorescences
including the corolla without; petioles 3-5 mm. long, mostly solitary,
the entire coriaceous leaves long-elliptic or lanceolate-elliptic, ob-
liquely cuneate into petiole, obtuse, about 1-1.5 dm. long, nearly
half as wide at or above the middle; racemes subopposite leaves,
peduncles 4-7 mm. long, pedicels 7-9 mm. long; calyx 2.5 mm.
long, 3.5 mm. broad, crenate, the lobes only about IXl mm.;
corolla 8-10 mm. across, the lanceolate lobes 4 mm. long, 1.5 mm.
wide, only the cucullate tip puberulent within; filaments basally
connate, to 0.6 mm. long, anthers ellipsoid, emarginate, 2.2 mm.
long; pores lateral and overhung; style slender, glabrous, 5 mm.
long, stigma capitate. After Bitter, I.e., as to synonymy, who places
plant in Anthoresis-Indubitaria.
Peru (possibly). Brazil.
Solanum confine Dunal in DC. Prodr. 13: 137. 1852.
Glabrous half-shrub or shrub-tree, apparently near to S. oppositi-
folium R. & P. but the lower leaves geminate, the larger ovate-
lanceolate to elliptic, acuminate, unequal at decurrent base, to
202 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
nearly 1.5 dm. long (often apparently smaller) and half as wide,
the smaller of each pair 2.5-5 cm. long, 1-2 cm. wide, sessile or
subsessile, acute or obtuse, reticulate-veined, membranous, lustrous
above, the 10 or 11 lateral nerves rather prominent beneath; cymes
opposite the leaves, in flower to 1.5 cm. long, in fruit about 2 cm.
long; pedicels about 8 mm. long, to 12 mm. long or longer in fruit,
incrassate at tip; calyx urceolate, about 3 mm. broad, the lobes
subovate, mucronate; corolla lobes lanceolate, acute, white, 2.5-3
mm. long; anthers fleshy, 2 mm. long, dehiscing by 2 subapical
chinks; style straight, erect, clavate toward the capitate stigma,
very spheroid, at least 6 mm. long. Sometimes a tree to 7 meters
high (Killip & Smith); sometimes seems to have few, even no gem-
inate leaves, and especially in fruit simulates the larger-flowered
plant of Ruiz and Pavon; an apparent variation is var. curtum
Macbr., var. nov., pedunculis pedicellisque circa 5-7 mm. longis.
Some of the following collections may belong to S. loretoanum
Bitter, scarcely distinguishable, at least in some cases. S. suprani-
tidum Bitter, Repert. Sp. Nov. 18: 69. 1922, of adjacent Brazil
may occur; the younger leaves beneath as the many-flowered in-
florescence are villosulous. F.M. Negs. 6782; 2624 (var.). Section
Leiodendron.
San Martin: Tarapoto, Spruce 4161 (det. Bitter); 4830; Williams
5527; 6179; 6260; 6767 (var.). Juanjui, Williams 6921. San Roque,
Williams 6929; 6965. Huanuco: Puerto Lira, Rio Pachitea, Killip
& Smith 26829 (shade form). Prov. Huamalies, Weberbauer 2624?
Loreto: Yurimaguas, Williams 4377; 3851; Killip & Smith 27610;
28163 (det. Morton); 27861; Poeppig 2279. Iquitos, Killip & Smith
26981; 29878; King 834 (type, var. curtum).
Solanum cruciferum Bitter, Repert. Sp. Nov. 16: 402. 1920.
A low (about 1 meter high) shrub well-marked by its subsessile
(1-) 2-flowered lateral inflorescence and tetramerous calyx and co-
rolla; branches and especially the leaves beneath very lightly tomen-
tulose (trichomes many-celled) and glandular, in age glabrous; leaves
alternate or geminate, subsessile, entire or undulate, oblong-lanceo-
late, often acuminate, 2-4 cm. long, 7-12 mm. wide; pedicels deflexed,
1.5-2 cm. long, little longer in fruit; calyx lobes ovate, acuminate,
minutely glandular, about 2 mm. long; corolla white, stellate, 14 mm.
across, the broadly lanceolate cucullate lobes about 7 mm. long,
partly pilose; filaments and anthers glabrous, the former 0.8 mm.
long, basally connate, the latter narrowly ellipsoid, 4 mm. long; style
FLORA OF PERU 203
straight, 6-6.5 mm. long; berry without granules. F.M. Neg. 2603.
Section Anthopleuris.
Cajamarca: Chugur, northwest of Hualgayoc, Weberbauer 4092,
type.
Solatium deflexiflorum Bitter, Repert. Sp. Nov. 18: 49. 1922.
Shrub, glabrous or glabrate except the pulverulent nodes and
branchlet tips, the leaves beneath, the 11 or so primary nerve-axils,
the short inflorescences including calyces and the corollas, these
white, stellate, especially toward the tip of the acute or subobtuse
lanceolate lobes; petioles 7-12 mm. long; leaves broadly lanceolate,
4-4.5 cm. wide, 9.5-11 cm. long, acuminate into petiole, subacutely
acuminate; inflorescence opposite leaves, simple, subsessile, 5 mm.
long; flowers 12-15, pedicels 12-13 mm. long, deflexed; calyx cam-
panulate, 2.5 mm. long, 4-5 mm. broad, lobes ovate, 1-1.5 mm. long;
corolla 14-17 mm. across, lobes 6 or 7 mm. long, glabrous within;
filaments glabrous, connate at base; style as stamens about 4.5 mm.
long, the stigma capitate. Peruvian collections, det. Morton, "near
S. deflexiflorum," with oval leaves, smaller flowers, so, perhaps var.
diversum Macbr., var. nov., foliis 1-2 dm. longis, 8-10 cm. latis,
corollis circa 5 mm. longis. Section Leiodendron.
Loreto: Creek Carapisa, above Pongo de Manseriche, Mexia 6261
(type, var.). Colombia.
Solanum dibrachiatum Van Heurck & M. Arg. Obs. Bot. 59.
1870. S. campaniforme Roem. & Schult. Syst. 4: 662. 1819, as to
Peru?
Shrub, glabrous or essentially, with geminate or in part alternate
chartaceous leaves, the typically more or less obovate larger often 1.5-
2 dm. long, the smaller (when present) oblong-elliptic, 3-6 cm. long,
nearly as wide, sometimes larger; peduncles forked or dichotomously
branched, 1-4 cm. long, the branches becoming as long or longer;
pedicels 5 mm. long in flower, incrassate but often not much longer
in fruit; anthers to 5 mm. long; otherwise apparently similar to
S. acuminatum R. & P. This plant has the aspect of a Cyphomandra
without anther connective thickening; in Herb. Geneva it was noted
by Bitter as a variety or subspecies of S. pulchrum Dunal. The leaves
apparently vary from ovate-elliptic to oblong-lanceolate; several
Iquitos specimens were referred by Morton to S. campaniforme
Roem. & Schult. perhaps correctly, but they lack the long pedun-
cles and filiform pedicels; on the other hand, while most of the in-
204 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
florescences are at least 1-furcate, the leaves resemble more S. oblon-
gum (and S. acuminatum). Flowers greenish-white (Stork & Hor-
ton); berries 13 mm. in diameter (Williams). F.M. Neg. 23021.
Section Leiodendron.
San Martin : Tarapoto, Spruce 4250, type. San Roque, Williams
7217; 7655. Juan Guerra, Wittiams 6860. Huanuco: Tingo Maria,
Stork & Norton 9486. Loreto: Santa Rosa, Williams 4764; 4932;
Killip & Smith 28971; 28717 (det. Morton). Puerto Arturo, Wil-
liams 5238; Killip & Smith 27920. Yurimaguas, Williams 4097; 4596.
Caballo-cocha, Williams 2095; 2224; 2281; 2310. Iquitos, Killip &
Smith 27054 (det. Morton); Williams 7912; 8066; 8184. Rio Itaya,
Williams 57; 248; 3518; 3385. La Victoria, Williams 2992; 3110.
Pongo de Manseriche, Mexia 6338? (see note under S. pulchrum).
Rancho Indiana, Mexia 6421 (det. Morton, S. campaniforme) .
Solatium hederiradiculum Bitter, Repert. Sp. Nov. 11: 12.
1912.
High-climbing liana, the pubescent branches affixed to tree-trunks
by numerous rootlets; petioles 3-5.5 cm. long; leaves broadly elliptic,
abruptly attenuate at base, subacuminate, to 18 cm. long, 9-10 cm.
wide, at least the veins on both sides most minutely but densely
pubescent; inflorescence axillary, mostly geminate, to 15-flowered;
peduncles about 2.5 cm. long; pedicels 5 mm. (to 9 mm. in fruit)
long; calyx pubescent, scarcely lobed, 2.5 mm. broad; corolla yellow-
ish-white, the spreading lobes about 4 mm. long, densely pubescent
without, especially at the cucullate apex; filaments scarcely 0.5 mm.
long, anthers 2 mm. long; style papillose above, 4 mm. long; berry
conical. Probably related to S. conicum R. & P., notwithstanding
its simple leaves; cf. S. theobromophyllum Bitter, erect, and S. robusti-
frons Bitter, creeping toward or at base. F.M. Neg. 2608. Section
Polybotryon.
Loreto: Yurimaguas, Ule 6276, type. Balsapuerto, King 2864-
La Victoria, Williams 3137. Iquitos, Killip & Smith 27329.
Solanum Lechleri Rusby, Bull. Torrey Club 26: 193. 1899.
Stellate-tomentose, unarmed (type), the mature leaves dark green
and only slightly pubescent above; petioles 1-2 cm. long, narrowly
marginal; leaves 5-10 cm. long, 2-5 cm. wide, ovate, broadly rounded
to subtruncate at base, acutely acuminate, entire, membranous, ve-
nation slender; peduncles in type terminal, 4 cm. long; pedicels sub-
pendent, 7-10 mm. long, weak; calyx 4-5 mm. long, lobed two-thirds,
FLORA OF PERU 205
lobes ovate, subacute as broad sinuses; corolla substellate, at least
1 cm. across, lobes broadly ovate, obtuse; anthers yellow, 3.5 mm.
long, pores subapical, introrse; style much longer, stigma small;
berry dark, 7 mm. in diameter. Branches herbaceous; type (Rusby
790} from Yungas, Bolivia, but according to the author the same as
Mandon 1106 and the Lechler collection, this presumably from south-
ern Peru.
Peru (Lechler 1939, fide author). Bolivia.
Solatium loretoanum Bitter, Repert. Sp. Nov. 18: 70. 1922.
Type an essentially glabrous shrub 1-2 meters high, upper branches
2.5-3 mm. in diameter, internodes 5-6 cm. long; lower leaves solitary,
upper geminate, the larger broadly elliptic-lanceolate, often 1.5-2 dm.
long, 7-9 cm. wide, broadly cuneate to short (5-6 mm. long) petiole,
acutely or subobtusely acuminate, lateral nerves mostly 12, the smaller
leaves obliquely ovate or suborbicular, obtuse or rounded, about
1.5 cm. long, to twice as broad; racemes 6-12-flowered, obscurely
glandular; peduncles 5-17 mm. long; pedicels slender, slightly shorter
than 1 cm.; calyx 1 mm. long, 2 mm. across, acute lobes marginally
puberulent; corollas white, stellate, 8 mm. across, the lanceolate lobes
2.5-3 mm. long, papillose only at short recurved tips; filaments
scarcely 0.5 mm. long, anthers broadly ellipsoid, 1.6 mm. long, apical
introrse pores oblique; style glabrous as little shorter stamens. Cf.
S. confine Dunal; calyx, developed, is probably larger. F.M. Neg. 2619.
Loreto: Yurimaguas, Ule 6277, type. Mouth of Rio Santiago,
Tessmann 3968; 4531 (vel aff., det. Werdermann). Pongo de Man-
seriche, Mexia 62^7; 621 2 A (det. Morton).
Solatium luteoalbum Pers. Syn. 1: 221. 1805; Dunal, Syn. 18.
1816; 121. S. pubescens R. & P. Fl. Peruv. 2: 36, pi. 169, fig. b. 1799,
not Willd. 1794. S. foetidum R. & P. I.e. 39?, not Rottb. 1778.
S. maleolens Macbr. Field Mus. Bot. 8: 111. 1930.
A low shrub, rather sparsely and finely villous (with simple tri-
chomes) ; leaves ovate, subabruptly acute; racemes lateral, the flowers
whitish-yellow; petioles about 1.5 cm. long; leaves mostly 8-10 cm.
long and about half as wide, early more or less puberulent or hispid-
ulous mostly on the 5-7 pairs of nerves; calyx deeply parted, the
acute lobes reflexed in fruit; berry orange, lustrous, globose, size of a
cherry (Ruiz & Pavon), to 2 cm. thick, the fruiting peduncles to
4 cm. long, pedicels 1.5 (-2) cm. long, apically incrassate; seeds
brownish-red or darker, reniform, 6 mm. long, 4 mm. wide, finely
206 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
scrobiculate. The description is drawn in part from my fruiting
material, which apparently is this species. S. foetidum R. & P., at
least as to Dunal, who describes the seeds as smooth, may not belong
here; and if there are two species the glabrous or early puberulent
one with foetid odor, "berries the size of walnuts," may be S. maleo-
lens Macbr. (i.e. S. foetidum R. & P.) . The Cuzco specimens, in fruit,
are canescent with a dense tomentum of stellate trichomes; no flowers
but probably a distinct species, perhaps introduced; for convenience
it may be recorded as var. tunya Macbr., var. nov., foliis utrinque
stellato-tomentosis. F.M. Neg. 29723 (S. pubescens}.
Huanuco: Cuchero, Ruiz & Pavdn, type. Mito, 3273. Yanano,
3795. Tarma, Ruiz & Pavdn (type, S. foetidum}; 1038. Cuzco:
Ollantaitambo, Cook & Gilbert 387a; 804 (type, var.); West 6476
(var.). Brazil? "Ama de casa" (Ruiz & Pavon), "pajarito," "tunya-
tunya" (Cook & Gilbert).
Solatium manicatum Bitter, Bot. Jahrb. 50, Beibl. Ill: 63.
1913.
Type a 3-meter shrub with lustrous purplish branches (decurrent
lines obscure), glabrous except for tufts of branched trichomes in the
axils of the leaf-nerves beneath and about the sleeve-like insertions
of the pedicels on the inflorescence-branches; petioles 1-1.5 cm. long;
leaves oblong-lanceolate, to 2 dm. long, 4-5 cm. wide, narrowed to
each end, the lateral nerves (about 18) and veins impressed above,
prominent beneath; inflorescence (type) nearly 1.5 dm. long and
broad, terminal; pedicels slender, to 2.5 cm. long in flower; calyx
5-7 mm. wide, the rounded lobes 2x2 mm.; corolla white (pale
yellowish, Metcalf), the spreading lobes lanceolate, acute, 1.5 cm.
across, 6 mm. long, subcoriaceous, densely hairy at the cucullate
apex; filaments short, glabrous, anthers deeply cordate, 5 mm. long;
style in part pubescent, 6-8 mm. long; berry yellow-green, about
8 mm. in diameter. Related to S. oligerum Schlechtd. of Mexico
but lacking the winged stems (Bitter); apparently, if correctly in-
terpreted, in pubescence and inflorescence, the latter sometimes re-
duced or perhaps simple. F.M. Neg. 2620. Section Anthoresis.
San Martin: Jepelacio, Klug 3371. Ayacucho: Prov. of Huanta,
between Tambo and the Rio Apurimac, Weberbauer 5643, type.
Apurimac: Pachachaca Valley, West 3789 (det. Johnston, S. con-
fine}. Cuzco: Marcapata, Metcalf 30729 (det. Killip).
Solanum marmellosanum Bitter, Repert. Sp. Nov. 16: 89.
1920.
FLORA OP PERU 207
Shrub, glabrous except for some obscure punctate glandulosity
and a dense papillosity on the cucullate corolla lobes; leaves approx-
imate, entire, sessile, broadly oblong-elliptic, becoming 3 dm. long,
1.5 dm. wide or larger, narrowed abruptly to base, gradually to acute
apex, the 14-16 subparallel nerves prominent beneath; inflorescence
terminal, 3-forked, many-flowered, peduncles 3 cm. long, pedicels at
least 1 cm. long; calyx 6-7 mm. long, 7-8 mm. across, the lobes about
3 mm. long; corolla campanulate-stelliform, to 18 mm. across, the
lanceolate lobes 8 or 9 mm. long; filaments shorter than 1 mm., an-
nulate at base; anthers ellipsoid, emarginate basally, about 5 mm.
long; style 3.5 mm. long, enlarged toward subclavate stigma. Near
S. sessile R. & P. and probably merely a variant with slightly smaller
flowers and apparently merges also with S. dibrachiatum Van Heurck
& M. Arg. The Peruvian specimens have a small rounded leaf at the
base of the larger but one is present on a similar specimen (Klug 2883}
referred by Standley to S. sessile R. & P. and with the larger flowers
of that species. Section (Anthoresis) Leiodendron.
Loreto: Soledad, Killip & Smith 29564; 29712; 29779 (det. Mor-
ton). Amazonian Brazil.
Solarium maturecalvans Bitter, Bot. Jahrb. 11, Beibl. Ill: 64.
1913.
Similar to S. amblophyllum Hook, but the tufts of pubescence
felt-like and occurring irregularly on the leaf veins beneath and on
the young branchlets; a low shrub, otherwise glabrous (minutely
glandular on nerves above), with elliptic-lanceolate subcoriaceous
leaves 5-9.5 cm. long, 2.5-3.5 cm. wide, and white flowers (about
6 mm. long) borne in short at least early terminal inflorescences of
3-6 flowers each; pedicels to 17 mm. long, in fruit 2 cm. long or longer;
calyx lobes 2-2.5 mm. X 2 mm., rounded; corolla lobes 5-5.5 mm.
long, densely pulverulent toward tip, papillose without; filaments
minute, glabrous; anthers about 3.5 mm. long, the little longer style
glabrous; stigmas subclavate; berry 12 mm. in diameter, granules
absent. Probably will prove to be a part of S. amblophyllum Hook.
F.M. Neg. 2622. Section Anthoresis.
Ayacucho: Prov. of Huamanga, above Quinua, Weberbauer 5543,
type.
Solarium Miquelii Morton, Contr. U. S. Nat. Herb. 29: 43.
1944. S. laetum Miq. Stirp. Surinam 135. 1850, not Kunze, 1842.
S. sempervirens Dunal in DC. Prodr. 13: 88. 1852, not Miller, 1768.
208 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
S. crispum R. & P. Fl. Peruv. 2: 31, pi. 158, fig. a. 1799, at least as
to Peru.
Vine, the upper slightly angled stems and terminal panicles pilos-
ulous-puberulent, in Peru stellulately, typically also punctiform-
glandular; petioles more or less curved, 6-10 mm. (in Peru to 3 cm.)
long; leaves glabrous, subrepand to plicate-undulate, subcoriaceous
(in Peru membranous), few-nerved and reticulate beneath, broadly
ovate to ovate-elliptic-lanceolate, unequally cordate or rounded at
base, obtuse to acuminate, many very unequal, often 5-7 cm. long,
3-5 cm. wide; panicles to 1 dm. long, foliose at base, branches few,
short, divaricate; pedicels 4-6 mm. long; calyx 3 mm. broad, the
broad subquadrate lobes mucronulate; corolla bluish or lilac, 1 cm.
long, segments suboblong, puberulent; anthers oblong-linear, to
6 mm. long, obliquely dehiscent. S. crispum R. & P. as to type is
Chilean and has anthers only 3 mm. long; it may be a variable species.
Hudnuco: Forest edge, Divisoria, Woytkowski 550 (det. Cuatre-
casas, S. crispum) . Loreto : Rio Nanay, Williams 51 1 ; 54-9. Florida,
Rio Zubineta, Klug 2320; 2071 (det. Morton). Mouth of Rio Tigre,
Kittip & Smith 27589 (det. Morton). Chile(?); Guianas.
Solatium monadelphutn Van Heurck & M. Arg. Obs. Bot. 42.
1870.
Shrub, glabrous except the white flowers within; closely related
to S. sessile R. & P. but leaves mostly much smaller, sublanceolate,
8-17 cm. long, 1.5-4.5 (6) cm. wide, lateral nerves yellowish, 9-10;
inflorescence 2-3-forked, many-flowered; peduncles to 4 cm. long,
pedicels to 7 mm. long, probably larger in fruit; calyx about 5 mm.
long, 6 or 7 mm. wide, unequal, 3 lobes 2-2.5 mm. long; corolla
about 12 mm. long, 14 mm. across, two-thirds connate, the free por-
tion of the lobes broadly triangular, acute, 5-7 mm. long, apically
only papillose-pilose; filaments 5, connate at base for nearly 2 mm.,
to 2.7 mm. long; anthers ellipsoid, emarginate at base, 5 mm. long,
1.5-1.8 mm. wide, glabrous as the straight style, this 4-6 mm.
long, the stigma obtuse. After Bitter, Repert. Sp. Nov. 16: 87.
1920; Allard determinations by L. Smith. F.M. Neg. 22902. Sec-
tion Anthoresis.
San Martin: Tarapoto, Spruce 4051, type; Ule 6632; Williams
6734. Boqueron Pass, Allard 21719; 22111. Huanuco: Tingo Ma-
ria, Allard 21153; 21728 A. Chaglla, 3656. Loreto: Puerto Yessup,
Kittip & Smith 26400 (det. Morton). Yuroc, Allard 22148.
FLORA OF PERU 209
Solatium nitidum R. & P. Fl. Peruv. 2: 33, pi. 163. 1799; 93.
S. angustifolium R. & P. I.e., not Lam. 1797. S. pulverulentum Pers.
Syn. 1: 223. 1805, not L.; 100. S. gnaphalioides Pers. I.e.; 127.
S. calygnaphalum R. & P. I.e. 31; 127. S. stenophyllum HBK. ex
Dunal, Syn. 15. 1816; 101. S. havanense Jacq. Enum. Carib. 15.
1760, as to Peru.
Typically glabrous except the inflorescence (this also glabrous in
a var.), the branches above subherbaceous, lustrous as the leaves,
these ordinarily oblong-lanceolate, subrepand, acute or shortly de-
current at base, more or less acuminate, mostly 1-1.5 dm. long, 2.5-
4 cm. wide, the many slender subparallel lateral nerves reticulate
with the veins and rather prominent beneath; petioles 5-15 mm.
long; racemes early terminal, the lower opposite the often crowded
leaves pulverulent to stellate-tomentulose (except var.), often in-
cluding the calyces; pedicels subumbellate, articulate at base, api-
cally incrassate, arcuate in fruit when at least 1 cm. long, calyx
cyathiform, the 5 ovate-subulate segments acute, scarcely 2 mm.
long; corolla lilac-purple, deeply 5-parted, 1.5-1.8 cm. across, the
lanceolate-oblong segments subobtuse; stamens equal, anthers linear,
subemarginate, dehiscing by short finally lateral chinks; style straight,
longer than stamens, the stigma subglobose-capitate; berry 8-10 mm.
in diameter, dark red when mature, the small dark seeds angular.
Variable in size, especially of leaves, and in extent of pubescence
(S. calygnaphalum, S. pulverulentum) ; this, apparently correctly, was
treated as a variety of S. calygnaphalum by Bitter in Herb. Madrid,
and for convenience ought to be recorded as a variant of S. nitidum.
Bitter once wrote S. calygnaphalum var. nitidum (R. & P.) Bitter.
S. gonocladum Dunal, 93, type from La Paz, is apparently similar
and may be Hutchison's 1243, but extreme in being densely canes-
cently pubescent. S. stenophyllum HBK., 101, is doubtfully more
than a narrower-leaved individual (leaves to 8 cm. long, 1.5 cm. wide)
of the puberulent form. S. havanense Jacq. has broadly elliptic co-
rolla lobes, blue berries. Most extreme is P. C. Hutchison's 1243,
which may be distinct, here recorded as var. Hutchisonii Macbr.,
var. nov., glaberrimum; petiolis 2 cm. longis, foliis 5-6 cm. longis,
ad 1.5 cm. latis, subovatolanceolatis, veniis obscuris; pedunculis 1-
2 cm. longis, pedicellis 1.5 cm. longis; floribus glabris calyce irregu-
lariter 2-3-fido, laciniis plus minusve connatis, subulato-apiculatis;
petalis glabris vel margine breviter ciliolatis, circa 8 mm. longis.
F.M. Neg. 29726.
210 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
The berries are used as a yellow dye (Raimondi) and as a cos-
metic and are saponaceous; leaves when toasted expel thorns (Ruiz
and Pavon).
Piura: Olleros to Yauta, (Bonpland, type, S. stenophyllum) .
Ancash: Chiquidn, Ferreyra 5836. Recuay, 2515. Lima: Oyon,
3,000 meters, Ferreyra 3536. Chicla, Safford. Rio Blanco, Kittip &
Smith 21714. Banos, Wilkes Exped. Huanuco: Acomayo, Ruiz &
Pavdn (type, S. angustifolium). Mito, 1670. Near Tambo de Vaca,
Woytkowski 117; 144. Junin: La Quinua, 2011; Ochoa 328. Tarma
and Acobamba, Ruiz & Pavdn (type, S. angustifolium, S. calygna-
phalum). Tarma, 1075; Soukup 2539; Ruiz & Pavon, type. Car-
papata, Killip & Smith 24365. Ocopa, Kittip & Smith 22002. Huan-
cayo, Ledig 3; Isern 2062; Killip & Smith 23846. Huancavelica:
Pampas, Smith & Horton 10241; 10833; Soukup 2772. Ayacucho:
Tortorabama, Weberbauer 5483. Lucanas, Hutchison 1243 (var.).
Ccarrapa, Killip & Smith 22468. Apurimac: Cunyac, Prov. Aban-
cay, Vargas 461; Ferreyra 2792. Cuzco: Panticalla Pass, Cook &
Gilbert 1889. Puquiura, Cook & Gilbert 1937. Near Cuzco, Hen era
684; Pennell 13709. Paucartambo, Pennett 14148; Hen era 1070.
Valle de Vilcanota, Velarde Nunez 1325. Puno: Near Puno, Soukup
360; Mexia 04200 (det. Johnston, S. pulverulentum) . Chuquito,
Metcalf 30678. Limbani, Metcalf 30470 (det. Killip). Tacna: Azapa,
Eyer dam 24645. Candarave, Metcalf 30378. "Nununya," "rapace"
(both Ruiz & Pavon), "campucassa," "huiscacassa," "tacachilla,"
"catruincho," "cahuincho," "illauru."
Solatium nutans R. & P. Fl. Peruv. 2: 34, pi. 166, fig. a. 1799;
128.
A shrub, rusty tomentose-pulverulent, especially the young branch-
lets, leaves beneath, calyx and corolla, the flexuose branches glabrate
in age as leaves above; pubescence branched on younger branchlets,
sublanuginous; petioles to 1.5 cm. long; leaves elliptic-ovate, mostly
1-1.5 dm. long, 4.5-6 cm. wide or larger, shortly acuminate, rather
prominently reticulate-veined especially beneath; lateral nerves many;
inflorescence opposite the leaves, to 2.5 cm. long, in fruit to 3 cm.,
closely few-flowered, the peduncle none or recurved, pedicels apically
incrassate, 2-5 mm. long; calyx subcampanulate, 5-7 mm. across,
glabrous within, the oblong segments subobtuse, corolla about 8 mm.
long, 1 cm. broad, pulverulent without, glabrous within, 5-parted
nearly to base, the ovate-lanceolate segments acutely acuminate,
cream-colored or white; stamens equal, subsessile, half as long as
FLORA OF PERU 211
corolla, the stout dark purple anthers dehiscing by 2 anterior chinks;
ovary subangled, glabrous, the straight erect subtrisulcate style
longer than the stamens, the stigma obscurely trilobed; fruit yellow,
globose, 2 cm. in diameter. F.M. Neg. 2627. Section Anthopleuris.
Cajamarca: Cutervo, Raimondi. Huanuco: Pillao, Ruiz & Pa-
v6n, type. Southwest of Huanuco, 2103; Dombey. Yanahuanca,
1214- Northwest of Mito, 1925. Mito, 1473. Mufia, 4278; 4325.
Tambo de Vaca, 4416? (fruit). Gauhuncho, Sawada 52. Junin:
Pichis Trail, Kittip & Smith 25835? "Chuculate" (Ruiz & Pavon),
"campucassa," "huiscacassa."
Solanum oblongifolium HBK. Nov. Gen. & Sp. 3: 27. 1818; 96.
S. hypomalacophyllum Bitter, in Pittier, Man. PI. Us. Venez. 137.
1926?
Branches smooth, glabrous, the branchlets (type) as peduncles,
pedicels (4-6 mm. long) villous pubescent; petioles 12-16 mm. long,
canaliculate, auriculate, with small sessile suborbicular leaves in
the axils; leaves geminate, elliptic-oblong, acute, narrowed to base,
subentire, 1.5 dm. long, nearly half as wide, the smaller about half as
large, firm-membranous, glabrous and green above, paler and (typ-
ically) pubescent beneath on the midnerve and veins, or fulvous-
tomentose (Bitter) ; flowers many, in subterminal bifid pilose recurv-
ing racemes; pedicels incrassate apically; calyx subglabrous, 6-7 mm.
across, the 5 equal spreading segments subrotund-ovate, obtuse;
corolla white, glabrous, about 1.5 cm. across, the ovate-oblong obtuse
lobes spreading; stamens equal, half as long as corolla, subsessile,
anthers connivent, oblong-linear, obtuse, apically bipored, equaling
erect style, stigma obtuse. Bitter's description reads: Tall robust
fistulose herb, large lanceolate leaves glabrous above, fulvous tomen-
tose beneath, flowers small in terminal corymbs; thus essentially a
nomen nudum; type from Merida, Venezuela; HBK. type from
Colombia. The Peruvian variant has leaves to 3 dm. long, nearly
1 dm. wide, alately long-decurrent to a broad short petiole, small
leaves at axils, larger leaves (in each pair) not very unequal, scurfy
tomentose beneath, perhaps varying in development but worthy
of note as var. Soukupii Macbr., var. nov., petiolis subalatis,
f oliis subequalibus, subtus furfuraceo-tomentosis.
Piura: Ayavaca, Soukup 4336 (type, var.). Cajamarca: Huam-
bos, 2,000 meters, Soukup 4493 (det. Cowan). Colombia; Venezuela?
Solanum oblongum R. & P. Fl. Peruv. 2: 34, pi. 165, fig. b.
1799; 148.
212 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Allied to S. acuminatum R. & P.; glabrous except for a glandular
exudate and often a sparse puberulence on inflorescence, on the
young branches and beneath near the midrib; petioles 1-3 cm.
long; leaves strongly geminate, usually coriaceous, veins reticulate-
impressed above, more or less prominent beneath, ordinarily with
10-12 primary nerves; larger leaves oblong-elliptic, subobtuse to
acuminate, 1-2 dm. long, half as wide, the smaller similar or broader
but shorter, often 5-8 cm. long scarcely half as wide; flowers
crowded, the racemes 1-4 cm. long, peduncles 1-1.5 cm. long (rarely
none?); pedicels to 1.5 cm. long, rather stout, somewhat longer
and incrassate above in fruit; calyx campanulate, lobes short, rounded
but minutely mucronulate; corolla white or faintly purplish, stellate,
1-2 cm. across, the oblong-lanceolate lobes puberulent at cucullate
tip, 8-10 mm. long; anthers oblong-ellipsoid, 4.5-5.5 mm. long,
the filaments 0.3-1 mm. long; style 6 mm. long, glabrous as ovary,
the stigma bilobed. My material is glabrous, the anthers shorter,
the filaments longer than in the specimens described by Bitter, Bot.
Jahrb. 54, Beibl. 119: 13. 1916, from Weberbauer and Ruiz and
Pavon collections. My 3732 has obtusely acuminate leaves, cuneate
at base, while 5205 has puberulent branches, submembranous acu-
minate leaves, peduncle obsolete; it can be recorded as var. abrup-
tum Macbr., var. nov., ramulis pulverulentis, pedunculis obsoletis
vel abrupte pedicellatis, pedicellis 1-3; foliis subabrupte acuminatis.
May be new but the large fleshy corollas seem to be typical; ob-
viously there are either more specific entities in this complex (that
includes S. triste Jacq. of the West Indies with capitate stigmas)
or fewer. F.M. Neg. 2628. Section Leiodendron.
San Martin: San Roque, Williams 7330. Huanuco: Pillao, Ruiz
& Pavdn, type. Yanano, 3732. Rio Chinchao, 5205? Chaglla to
Muna, Weberbauer 6709 (det. Bitter). Junin: Vitoc, Raimondi (det.
Werdermann).
Solanum ochrophyllum Van Heurck & M. Arg. Obs. Bot. 50.
1870.
Branchlets toward apex, solitary or geminate leaves beneath
(above quite glabrous) and the several-flowered umbelliform cymes
opposite them in type densely lanate- or arachnoid-tomentose with
more or less fulvous trichomes; petioles 1.5-5 cm. long or longer;
geminate leaves moderately unequal, all approximate, lanceolate,
narrowed to base, acute, often 8-10 cm. long, half as wide; peduncles
to 1 cm. long, usually shorter, pedicels about 5, to 1.5 cm. long;
FLORA OF PERU 213
calyx cupulate, coriaceous, nearly 3 mm. long, the 5 short rounded
lobes broader than high, the margin cartilaginous; corolla 7 mm.
long, parted nearly to base, the lobes long-oval; stamens subsessile,
anthers 4 mm. long, the 2 large terminal pores promptly elongating
into lateral chinks; berry 1 cm. in diameter. Not closely allied
to any species but approaches S. nutans R. & P. (authors); type,
Mandon ^16 from Sorata, Bolivia. Apparently S. hypomalacothrix
Bitter, Repert. Sp. Nov. 18: 70. 1922, is allied or a variant with
calyx subpatelliform, the strongly apiculate lobes membranously
joined below; especially interesting because the author himself as-
signed it to section Anthopleuris but remarked: gehort wohl noch
zu Leiodendron. The Peruvian specimens (in flower) listed are less
densely pubescent especially on the leaves beneath, the veins not
concealed, and approach specimens of S. nutans R. & P. from the
upper edge of its range; this southern shrub may prove to be a
variant here or of S. nutans in which case the Bolivian plant is
also perhaps another variation; the collection of the late eminent
zoologist and humanist Karl P. Schmidt may become the type of
S. ochrophyllum Van Heurck & M. Arg. var. Schmidtii Macbr., var.
nov., foliis pedicellisque glabratis, veniis conspicue reticulatis. The
specimens in early fruit already have glabrate submembranous leaves
but perhaps the scant indument was due to habitat, a wet quebrada.
Ayacucho: Choimacota, Weberbauer 7551 (var. Schmidtii}. Above
Ocros, West 3671 (toward S. nutans}. Cuzco: Hacienda de Urcos,
Schmidt (type, var. Schmidtii). Urubamba, Soukup 36. Cerro de
Cusilluyoc, 3,200 meters, Pennell 14-152. Marcapata, 2,500 meters,
Vargas 9662 (det. Standley) Calca, Prov. Anta, Vargas 4,39. Bolivia.
Solarium oppositifolium R. & P. Fl. Peruv. 2: 35, pi. 168, fig. 8.
1799; 137. S. urceolatum Pers. Syn. 1: 223. 1805?
A glabrous shrub with elliptic-oblong, acuminate leaves, acu-
minate to acute base, all alternate and 2-several flowered racemes
borne opposite them; leaves a little undulate-repand, lustrous above,
pale green, chartaceous, 1.5 dm. long, often 5-6 cm. wide, reticulate-
veined, primary nerves 9 or 10, slender, prominent beneath; petioles
4-12 mm. long; peduncles 5-10 mm. long; racemes about 2 cm.
long to twice as long in fruit, the pedicels 4-6 mm. long, after
anthesis to 2 cm., and incrassate, especially apically; calyx 3 mm.
broad, enlarging in fruit, urceolate, broadly and irregularly 5-dentate;
corolla white, 8 mm. long, 1.5 cm. broad, the lanceolate-oblong lobes
somewhat uncinate at tip; anthers oblong, 5 mm. long, apically
214 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
dehiscent from elongate chink, filaments very short. Werdermann
in Herb. Madrid uses the Persoon name, but it seems unnecessary
to discard the cognomen of Ruiz and Pavon; simulates narrow-
leaved forms of S. dibrachiatum Van Heurck & M. Arg. but in-
florescence simple. F.M. Neg. 29729. Section Leiodendron.
Junin: Prov. Tarma, Woytkowski 353 (distr. as S. nitidum).
Chanchamayo Valley, Schunke 258; 259; 272; 534. Near La Merced,
5768. San Ramon, Williams 45^5. Vitoc, Tafalla, type.
Solatium oxycoccoides Bitter, Repert. Sp. Nov. 16: 14. 1919.
A small trailing slender-stemmed perennial, woody below, rooting
at the nodes; pubescence of branches and leaves rather sparse, simple,
the ovate leaves beneath between the veins also minutely glandular;
petioles 1-3 mm. long; leaves rounded, truncate or subcordate at
base, narrowed to obtuse apex, 0.5-1.5 cm. long, 3-9 mm. wide;
flowers solitary, terminal, on glabrate pedicels about 6 mm. long;
calyx to 8 mm. broad, the acute lobes spreading, 2-2.5 mm. long;
corolla at first violet, later white, stellate, 1.5 cm. across, the lan-
ceolate lobes about 5 mm. long, pilosulous at the little cucullate
tip; filaments nearly glabrous, 1-1.3 mm. long; anthers ellipsoid,
emarginate both ends, 4 mm. long, the oblique introrse pores finally
somewhat laterally dehiscent; style 7-8 mm. long, obscurely papillose,
incrassate to the globose-capitate stigma. My specimen has longer
petioles and larger flowers. See S. chachapoyasense Bitter regarding
the section. F.M. Neg. 2629. Section Anarrichomenum.
Ancash: Near Yungay, Weberbauer 3279, type. Huanuco: Chas-
qui, 3293.
Solatium patellare Van Heurck & M. Arg. Obs. Bot. 53. 1870.
Shrub, sparsely to rather densely puberulent-hispidulous on the
leaf-veins beneath, on the flowering branchlets and on rachis and
pedicels of the short (about 1.5 cm. long) inflorescences; leaves
notably thin, oblong-elliptic, all the smaller ovate (6-8 X 4.5 cm.),
the larger a dm. long or longer by about 6 cm. wide, all moderately
acuminate, acute at base, the petioles 8 mm. (small leaves) to 2 cm.;
racemes crowded or pseudocymose, sometimes furcate; flowering
pedicels 8 mm. long; calyx shallowly lobed, the broadly triangular
acute teeth about 1 mm. high; petals narrowly ovate-oblong, 8 mm.
long; anthers ellipsoid, 2.5 mm. long. The Klug collections have
branchlets and leaves beneath on nerves hispidulous, longer calyx
FLORA OF PERU 215
teeth and anthers and may be distinct. F.M. Neg. 22904 (Spruce
4914}' Section Leiodendron.
San Martin: Tarapoto, Spruce 4845, type; 4914 (det. Bitter).
Jepelacio near Moyobamba, Klug 3262? Loreto: Florida, Rio Putu-
mayo, Klug 2214? (det. Standley with query, S. pukhrum). "Manao-
ey" (Klug).
Solatium patulum Pers. Syn. 1: 223. 1805; 95. S. lanceolalum
R. & P. Fl. Peruv. 2: 33, pi. 164- 1799, not Cav., 1797.
A shrub or small tree to 5 meters high, young branchlets, leaves
beneath and inflorescence yellowish-puberulent or subtomentose;
branches flexuose, yellowish, lustrous in age; petioles about 1-2 cm.
long; leaves lanceolate, glabrate and somewhat lustrous above, atten-
uate at base and apex, acutely acuminate, or obtuse or rounded on
the lateral branchlets, mostly 1-1.5 dm. long, 2-3.5 (4.5) cm. wide;
inflorescence at first terminal, subpaniculate; peduncles 1-3 cm. long;
pedicels 1-2 cm. long; calyx campanulate, deeply lobed (lobes lan-
ceolate, acute or obtuse), 6-9 mm. long; corolla violet (or pure white,
forma album Macbr.), rotate-stellate, 1.5-2 cm. long, 4-5 cm. broad,
the broad lobes abruptly acuminate; stamens nearly 2 mm. inserted
above base; ovary minutely papillose (as style or this also pubescent
toward base); stigma capitate (in type). A beautiful species with
an abundance of flowers, a rather light wistaria color; berry black,
1 cm. in diameter. Var. pilosistylum Bitter, Bot. Jahrb. 54, Beibl.
119: 9. 1916, has style pubescent and papillose toward the base, the
stigma manifestly bilobed; sclerotic granules lacking. In part after
Bitter, I.e. 8. Section Dulcamara.
Huanuco: At Muiia and Tambo Nuevo, Ruiz, type. Tambo de
Vaca, 4351; 4441 (type, forma album). Huancavelica: South of Sur-
cubamba, Weberbauer 6477 (type, var.). Tayacaja, Velarde Nunez
2008. "Urahuacta."
Solatium pseudocapsicum L. Sp. PL 184. 1753; 152.
A small glabrous shrub the wood of the branches rather soft
with repand oblong-lanceolate acutely acuminate leaves about 6 cm.
long and 1-1.5 cm. wide; the petioles apically alate, 6-8 mm. long;
pedicels 1-3 in each axil, 6 mm. long in flower and more or less cernu-
ose, twice as long and ascending in fruit; corolla white, 6 mm. long,
8-10 mm. broad, the calyx lobes ovate-lanceolate; anthers yellow,
connivent, the oval linear cells apically dehiscent; berry red or yel-
low. Spontaneous or cultivated in tropical and subtropical regions.
216 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Illustrated, Rev. Invest. Agric. Argentina 2: 193. 1948. Section
(Pseudocapsicum) Leiodendron.
Huanuco: Waste land in the pueblo Tomaiquichua, below Ambo,
2439.
Solatium psidiifolium Rusby, Bull. Torrey Club 26: 194. 1899.
S. divernicascens Bitter, Repert. Sp. Nov. 11: 483. 1913, fide Bitter,
I.e. 16: 408. 1916. S. Lindenii Rusby, Mem. Torr. Bot. Club 6:
88. 1896?
Except as to type (S. Lindenii) branchlets (these stout) and leaves
beneath more or less stellulate puberulent; leaves oval-ovate; abruptly
contracted at rounded base into the broad marginal petioles (1-1.5 cm.
long), subobtuse or subacute, heavy coriaceous, the slender nerves
(about 12 pairs) somewhat grooved above, prominent beneath; pe-
duncles erect, thick, to 2 cm. long; cymes more or less open, 2-3 cm.
across, or larger, sometimes 25-flowered, pedicels flaccid, to 1.5 cm.
long; calyx as whitish corolla fleshy, 4 mm. long, 5 mm. broad, black-
ish tube dark, lobes semicircular, about one- third as long; corolla
stellate, 1 cm. across, the lobes ovate; style stout, angulate, exceed-
ing stamens 1 mm.; anthers oblong, straight, 4-6 mm. long, the large
elongated pores subterminal as in S. brevipedunculatum Rusby.
After author as to type of S. psidiifolium. S. Lindenii as to type is
glabrous, leaves narrower; the later described plant may be distinct
but the leaves vary in width, sometimes are inequilateral, obtuse or
acute, sparingly to rather densely stellate-pubescent, cymes in short
dense or longer-peduncled loose cymes (Rusby); it seems probable
that these plants are part of S. oblongum R. & P. Stone cells of
berries 2 (S. psidiifolium). Section Oppositifolia, fide Bitter, i.e. An-
thoresis, part.
Junin: Carpapata, Killip & Smith 24364 (det. Morton). Bolivia.
Solatium pulchrum Dunal, Syn. 15. 1816; 96. S. pulchrum
Dunal, var. peruvianum Dunal in DC. Prodr. 13, pt. 1: 96. 1852.
Type with rugose terete lustrous branches 6 mm. thick; leaves
geminate, or (in Peru) often alternate, lanceolate-elliptic or oblong,
cuneate to petioles about 1 cm. long, reticulate-veined (primary
nerves 7-9, prominent and typically puberulent beneath), the larger
about 2 dm. long, 7-10 cm. wide, the smaller (sometimes absent)
acute to rounded, about a fourth as large (leaves not so large in
known Peruvian collections); panicles becoming lateral, to 2 dm.
long and broad, often much smaller, puberulent at apex, peduncles
FLORA OF PERU 217
several cm. long, pedicels crowded, early filiform, 10-12 mm. long,
in fruit 1.5-2 cm. long, a little incrassate below the globose berry,
this 1 cm. in diameter; calyx subhemispheric, about 6 mm. across,
semi-parted into 5 short unequal subcoriaceous lobes, sublucid at
apex; corolla (1) 1.5 cm. across, the lanceolate segments acute,
scarcely 5 mm. long (longer in type), twice as long as the equal sta-
mens, anthers 5 mm. long, subcylindric-conical; style filiform, 7 mm.
long, stigma subclavate. After Dunal, the Peruvian plant not, as
indicated, quite in agreement, especially in the considerably smaller
flowers; here or hereabouts, may be undescribed as suggested by
Morton, in herbaria is Mexia 6338, Pongo de Manseriche, with mem-
branous unequal geminate leaves alately marginal to base, shortly
pedunculate 1-2-forked inflorescences in axils, 3-6 cm. long, pedicels
10 mm. long. Illustrated, Mart. Fl. Bras. 6: pi. 4, figs. 24-30.
F.M. Neg. 2843 (var.). Section Anthoresis.
San Martin : Pongo de Cainarachi, Klug 2659 (distr. as S. mona-
delphum). Loreto: Florida, mouth of Rio Zubineta, Klug 2063 (det.
Morton) . Puerto Arturo, Yurimaguas, Williams 524-6; Poeppig 2093
(type, var.). Pro on the Rio Amazon, Williams 1993. Brazil.
Solanum robustifrons Bitter, Repert. Sp. Nov. 11: 473. 1912.
Stems stout, creeping and rooting below, the erect portion about
2 dm. high; petioles 5.5-7 cm. long; leaves glabrous, broadly rhombic-
lanceolate, attenuate at both ends, about 2.5 dm. long, 1.5 dm. wide
to at least twice larger; peduncles opposite leaves, solitary, branched,
1-1.5 cm. long, in type to 50-flowered; pedicels early 5 mm. long
(slightly incrassate and 1.5 cm. in fruit) ; calyx lobes lanceolate, early
1.5 mm. long, in fruit 2-2.5 mm. long; corolla lobes lanceolate, acute,
about 3 mm. long (to 6 mm., Klug 3662), a third as wide, papillose;
filaments (0.6 mm. long) and style glabrous (in one example 1.2 mm.
long, perhaps longer); berry globose, at least 1 cm. in diameter.
The large-flowered material may not belong here or the species is
variable at least in this respect. Cf. Bitter, I.e. 17: 330. 1921; orig-
inally referred to Polybotryon. F.M. Neg. 33105. Section Leiodendron.
San Martin : Jepelacio, Klug 3632; Klug 3491 ? (peduncle long) .
San Roque, Williams 6952? Huanuco: Tingo Maria, Allard 21667
(det. with query, L. B. Smith). Loreto: Yurimaguas, Poeppig 2483,
type; Killip & Smith 28032.
Solanum salviifolium Lam. Illus. 2: 14. 1793 or 1778?; 107 (as
S. radula Vahl., 1798) . S. citrifolium Willd. ex Roem. & Schult. Syst.
218 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
4: 662. 1819, and HBK. Nov. Gen. & Sp. 3: 452. 1819; 109. S. radula
Vahl. Eclog. 2: 16. 1798. S. clathratum Sendt. in Mart. Fl. Bras. 6:
35. 1846; 120. S. oblitum Dunal in DC. Prodr. 13, pt. 1: 126. 1852.
Branches slender, terete, flexuose, pulverulent-tomentose and
subleprose, as the partly geminate leaves beneath and corymbs in-
cluding the flowers; petioles of the larger leaves 5-8 mm. long;
blades ovate-lanceolate, more or less obliquely rounded at base,
acute or acuminate, green but minutely and sparsely stellulate above,
the (5) 6-8 lateral nerves prominent beneath with laxly reticulate
veins, mostly 8-12 cm. long, half as wide, the smaller resembling
stipules 1-3 cm. long, often nearly as wide, rounded or obtuse;
corymbs opposite the leaves, 1-10 cm. long, finally dichotomously
branched, the cymose pedicels articulate at base, nodding before an-
thesis, erect in flower, 6-10 mm. long, slightly longer and somewhat
arcuate in fruit; calyx cyathiform, 5-sulcate, accrescent and lacerate
in fruit, the segments early ovate-crenate; corolla glabrous within,
spreading or reflexed, 10-12 mm. across, the lanceolate-oblong acutely
acuminate segments 6-8 mm. long; stamens equal, half as long as
corolla, filaments 0.5 mm. long or minute, papillaceous, anthers con-
nivent, 2.5 mm. long, subsagittate, dehiscing by two chinks; ovary
ovoid, tomentose-villous-stellate as the filiform style at base, this
somewhat curved or apically deflexed, less than twice as long as sta-
mens, stigma depressed-capitate or punctiform; fruit about 6 mm.
in diameter. Mostly after Dunal's description of his species, which
seems to represent in the Andes the plant of Lamarck and probably
includes that of Willdenow. Type from Guayaquil, the specimen at
Geneva indicated as S. salviifolium Lamarck by Bitter, who found
2-3 stone cells in S. citrifolium, apparently the same. Killip and
Smith noted their collection as a tree 5 meters tall with white flowers,
deep purple fruit. S. distichophyllum Sendt., I.e., of Amazonian Bra-
zil has smaller leaves and flowers, as to type. F.M. Negs. 33059
(S. clathratum); 34125 (S. oblitum). Section Anthoresis.
San Martin: Tocache, Poeppig 1891 (type, S. clathratum). Jepe-
lacio, Klug 3306 (det. Standley). Near Tarapoto, Williams 6822.
Huanuco: Boqueron Padre Abad, Woytkowski 34.370 (det. Cuatre-
casas, S. citrifolium). Junin: Pichis Trail, Killip & Smith 26069.
To the Guianas; Cuba; Mexico.
Solatium selachophyllum Bitter, Repert. Sp. Nov. 16: 82.
1919.
A shrub or small tree to 5 meters high, well-marked by the closely
bullate-stellate upper leaf-surfaces and the dense long-stipitate stel-
FLORA OF PERU 219
late reddish pubescence (2.5-3 mm. long) of the branches; petioles
1-1.5 cm. long; leaves mostly alternate, oblong-lanceolate, narrowed
at each end, acute, usually about 1.5 (2) dm. long, 4-5.5 cm. wide,
reticulate- veined and scabrous-stellate beneath; corymbs early ter-
minal, shortly rusty stipitate-stellate pubescent, many-flowered; pe-
duncles 6 dm. long, pedicels 5-8 mm. long; calyx cupulate, 4-5 mm.
long, the lobes broadly triangular, acute, about 2.5 mm. long; corolla
pale lilac to dark blue, stellate, nearly 2 cm. across, the lanceolate or
oblong lobes to 1 cm. long, except tube (0.5 mm. long), stellate-
pubescent without, below within sparsely glandular; filaments gla-
brous, 1.5-2 mm. long, style 4.5 mm. long, more or less pubescent
nearly to stigma, this subglobose; anthers emarginate, 4x2 mm.;
berry sparsely stellate-pubescent. Apparently not far from some
individuals of the S. verbascifolium L. complex, especially if this
Williams plant may be correctly placed as var. seorsum Macbr.,
var. nov., foliis asperatis vix vel haud bullatis, ellipticis, 6-8 cm. latis,
floribus 1.5 cm. longis (Williams 2606, type). Section Anthoresis.
San Martin: San Roque, Williams 7470. Huanuco: Southwest
of Monzon, Weberbauer 3399, type. Villcabamba, on Rio Chinchao,
5196. Toward Tingo Maria, Swingle 74. (det. Morton). Carpish,
2,800 meters, Ferreyra 2101; 2338; Ochoa 418 (det. Killip) . Junin :
Oxapampa, Soukup 2547 (det. Morton). Loreto: La Victoria, Wil-
liams 2606 (type, var.); 2956; 2981. Pumayacu, Klug 3203.
Solanum sericeum R. & P. Fl. Peruv. 2: 33, pi. 161. 1799; 122.
Tall shrub with diffuse slender or filiform subterete branches,
these whitish sericeous as the ovate-lanceolate leaves both sides;
petioles 1 cm. long; leaves subacute, 2-2.5 cm. long, about 1 cm. wide
with 5 primary nerves prominent beneath; branchlets foliose, the
younger axillary; racemes laxly 2-6-flowered, 2.5-3 cm. long, the
peduncles, pedicels (6-10 mm. long) and cyathiform calyx densely
pubescent, the ovate- triangular lobes subobtuse; corolla violet-blue
within, canescent sericeous without, 3 times longer than calyx, the
ovate lobes subacutely acuminate; stamens about half as long as
corolla, exceeded by filiform erect style, the stigma bifid; berry glo-
bose, red, size of a pea. After Dunal, who saw a specimen in Paris.
Type said to be imperfect (not found by me) ; it may be the earlier
name for S. chachapoyasense Bitter and is one of many types that
should be recollected at original locality. Section Dulcamara.
Junin: Huasahuasi, Ruiz & Pavdn, type.
220 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solanum sessile R. & P. Fl. Peruv. 2: 35, pi. 167, fig. a. 1799; 98.
A glabrous shrub or small tree with sessile cuneate-oblong-spath-
ulate leaves, mostly 2-several dm. long, and terminal corymbose ra-
cemes sometimes congested, sometimes dichotomously branched, of
white flowers; leaves solitary and geminate, undulate (fleshy in my
collection 4815) and especially toward the base crisp or plicate, more
or less subcordate at base, chartaceous, particularly in age noticeably
reticulate-veined beneath; pedicels articulate at base, slightly incras-
sate at apex, 2-8 (12) mm. long; calyx firm to membranous, sub-
hemispheric, the subrotund somewhat scarious-margined lobes un-
equal, 4-7 mm. long; corolla deeply parted, at least 1 cm. long, the
segments ovate- or oblong-acuminate; stamens unequal, filaments all
filiform, anthers linear, obtuse, subequal, dark purple (Dunal), de-
hiscing by 2 apical chinks; ovary ovate-connate; style capillaceous,
recurving at tip, scarcely longer than stamens, the stigma puncti-
form; berry white. Sometimes there are one or more stipuliform
leaves; an herbarium name by me applying to specimens with capi-
tately corymbose inflorescences is probably S. monadelphum, which
may be only an individual variation; actually the type was only
slightly open; an extreme form is the Constance and Tovar plant with
much-branched inflorescence a dm. across; all the material from
Junin and Soukup 3318 have somewhat smaller flowers, leaves and
more open inflorescence; the character is variable, suggesting the
probable invalidity of S. marmellosanum Bitter. F.M. Neg. (without
visible number). Section Anthoresis.
Huanuco: Mufia, Ruiz & Pavdn, type; Weberbauer 6719. Cushi,
4,815. Chinchao, Poeppig 1554. Pillao, Woytkowski 34168. Pasco:
Quillasu, Soukup 3318 (det. Lyman Smith). Junin: La Merced,
5443; Schunke 321; Soukup 2488. San Ramon, Constance & Tovar
2316. Huacapistana, Killip & Smith 24111; 24327 (det. Morton).
Loreto: Santa Rosa, Killip & Smith 28992. Yurimaguas, Williams
4983. Balsa Puerto, Klug 2883 (det. Standley). San Antonio, Kil-
lip & Smith 29492 (pedicels 1.5 cm. long). Puerto Arturo, Williams
4983Cuzco: San Miguel, Urubamba Valley, Cook & Gilbert 1170.
Rio Acre: Vie 9760 (det. Bitter). Adjacent Brazil.
Solanum solum Macbr., sp. nov.
Ramis (6 mm. crassis), ramulis lignosis (circa 2 mm. crassis, dm.
longis), foliis praecipue subtus et floribus ubique plus minusve brevi-
ter stipitato-stellulato-hispidulis cum pilis subrubescentibus; petiolis
3-7 mm. longis; foliis geminis, oblongo-lanceolatis, majoribus ad
FLORA OF PERU 221
basin oblique acutis, acute acuminatis, 6-11 cm. longis, 2-3.5 cm.
latis, minoribus suboblongis, subobtusis, 1-1.5 cm. longis, 4-7 mm.
latis; floribus racemose 1-7-umbellatis, extra-axillaribus, pedicellis
5-7 mm. longis, rufo-stellato-tomentosis, calycis dentibus rotundatis;
corollae albae, laciniis oblongo-ovatis, circa 3 mm. longis; antherae
crassae, apice biporosae. The small, deeply parted flowers, few in
short racemes or several in pseudo-umbels, suggest a relationship
with S. Karstenii Dunal, 151, of Colombia.
A strict rusty pubescent shortly branched shrub about 2 meters
tall that stood alone in sunny bush.
Junin: Hacienda Schunke, La Merced, 5764, type.
Solatium styracioides Rusby, Mem. Torr. Bot. Club 4: 230.
1895.
Glabrous except for the puberulent inflorescence; woody stems
flexuose, apparently reclining or supported, becoming hollow, deep
purple; petioles 2-5 cm. long; leaves ovate, blunt, thin, 1-1.5 dm.
long, 6-10 cm. wide; panicle terminal, 2-3 dm. long; pedicels 5-
10 mm. long; calyx truncate with 5 obscure teeth; corolla velvety
without, thick, divided nearly to the base into linear-lanceolate lobes;
filaments very unequal, one elongating to 6 mm.; anthers densely
granular, 7 mm. long; style nearly 1.5 cm. long. Possibly a Lycian-
thes but in appearance of inflorescence and downward broadened
filaments suggesting Cyphomandra according to the author, who pro-
poses for it a new section, Andropedas; Poeppig specimen was a high-
climbing shrub with a terminal panicle of large flowers, the leaves
smaller than the Bolivian plant (Bitter).
San Martin: Tarapoto, (Spnwe 4327). Huanuco: Cuchero, Poep-
pig 1533 (det. Bitter). Bolivia.
Solanum theobromophyllum Bitter, Repert. Sp. Nov. 11: 472.
1913; var. procerius Bitter, I.e. 12: 145. 1913.
Entirely glabrous erect half-shrub 1-2 meters high; petioles 1-
3.5 cm. long; leaves broadly elliptic-lanceolate, attenuate at each end,
13-15 cm. X 5-6 cm. (or 2.5 dm. X 9-13 cm. in the typical form) ; in-
florescence geminate at the leaf-axils, 1-1.5 cm. long; pedicels 2.5 mm.
long (to 5 or 6 mm. in fruit); calyx lobes minute; flowers greenish-
yellow (or white in the variety), the corolla lobes 1.3 mm. broad,
about 2 mm. long, subcucullate; filaments to 0.5 mm. long, glabrous,
anthers broadly ellipsoid, 1.6 mm. long, obscurely emarginate, style
222 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
2.5 mm. long, papillose below middle; berry rounded or narrowed
above, tipped with a conical process. Originally referred by author
to Polybotryon, apparently correctly as including Bassovioides; deter-
minations mostly by Morton; perhaps large-leaved specimens of S.
anceps R. & P. F.M. Neg. 2640.
Huanuco: Tingo Maria, Allard 22522 (det. L. S. Smith). Junin:
Pichis Trail, Killip & Smith 26203. Puerto Yessup, Killip & Smith
26203; 26239; 26261. Near La Merced, Killip & Smith 23839
Loreto: San Antonio, Killip & Smith 29420; 29493. Soledad, Kil-
lip & Smith 29584- Yurimaguas, Killip & Smith 29076. Boqueron
Pass, Allard 22077. Rio Mazdn, Jose Schunke 280 (det. Standley,
S. anceps). Brazil. "Muyacasacha" (Schunke).
Solatium umbellatum Miller, Card. Diet. ed. 8, no. 27. 1768;
372. S. dichotomum R. & P. Fl. Peruv. 2: 34, pi. 166b. 1799, not
Lam., 1790. S. riparium Pers. Enchir. 1: 221. 1805; 144. S. rugosum
Dunal, in DC. Prodr. 13, pt. 1: 108. 1852, of Brazil, i.e. S. asperum
Rich, at least as to Peru, both with trichomes all or mostly sessile,
the latter with petioles 2-4 mm. long, but constancy of these char-
acters is dubious over wide areas. S. cortex-virens Dunal, I.e. 372.
Similar to S. verbascifolium but greener, leaves often firmer, usu-
ally more distinctly reticulate-veined, even somewhat rugosely, and,
especially, branches apically strongly stellate-tomentose, the tri-
chomes stiped, flowers often sessile or subsessile (pedicels to 3 mm.
long), indument often fulvous, denser on petals; ovary glabrous
(ex char.). It is doubtful at least as to the Peruvian specimens
that this is a species distinct from S. verbascifolium L., probably as
variable as is usually the case for widely distributed entities. The
corolla is white to violet, the berry yellow, glabrate or glabrous.
Plate of Ruiz and Pavon plant is poor; seemingly it is about the
same as type of Miller from Mexico, at least plant of Dunal; some
material determined as S. verbascifolium L. may rather represent this
or some simulating form and many specimens distributed as S. bicolor
Willd. seem to belong here or to S. verbascifolium. If pubescence
characters presence of simple trichomes, glandular, eglandular stel-
late trichomes, these stiped, sessile, subsessile are consistently in-
herited there are many species within this circle of plants, but
probably names are already available on extra-Peruvian types.
Nevertheless, an unusual collection with glistening pubescence in
the inflorescence (Killip & Smith 24969), perhaps distinct, may, for
present convenience, be noted : var. f ulgens Macbr., var. nov., foliis
FLORA OF PERU 223
fere ad basin alato-attenuatis ad apicem conspicue caudato-acumi-
natis, ad 6.5 dm. longis, 1 dm. latis, supra subglabris, nitidis. Sec-
tion Leiodendron (Anthoresis) .
San Martin: Zepelacio, Klug 3576. Huanuco: Prov. Huanuco,
Ruiz & Pavon (type, S. dichotomum). Villcabamba, 4981. Pozuzo,
4657. Cueva Grande, 4783. Rio Chinchao, 4981. Tingo Maria,
Attard 22023; 20347; 20569 (det. Morton); Soukup 2209; Stork &
Horton 9600. Pasco: Quillasu, Soukup 3320 (det. L. B. Smith).
Junin: Oxapampa, Soukup 2542. Chanchamayo, Isern 2418. Vitoc,
Soukup 2546. Loreto: Mishuyacu, Klug 268; 2564 (det. Morton,
S. rugosum); Klug 568 (det. Morton); Kittip & Smith 29911. Iqui-
tos, Kittip & Smith 27052; 277339 (both det. Morton, S. rugosum,
det. Standley, S. tricolor); Williams 3712; 8182. Rio Nanay, Wil-
liams 290; 744; 773; 1271; 1273. Rio Itaya, Wittiams 3229. Pro,
Williams 1967. Caballo-cocha, Wittiams 2053. La Victoria, Wil-
liams 2598. Cuzco: Valle de Cosnipata, Nunez 1417; 1418 (both
det. Morton). Urubamba Valley, Herrera 1558.
Solanum verbascifolium L. Sp. PI. 1: 184. 1753; 114. S. bicolor
Willd. ex Roem. & Schult. Syst. 4: 661. 1819.
Shrub or tree sometimes 10 meters tall or taller, the upper branches
as leaves stellate tomentose; petioles 1-7.5 cm. long; leaves entire,
oval to elliptic, mostly rounded at base, acute, often 1-1.5 dm. long,
6-9 cm. wide, sometimes smaller, not infrequently larger, gray- white
tomentose above to merely scabrous or glabrate, often straw-colored
beneath, usually with 8 or 9 lateral nerves; terminal inflorescence
ordinarily 5-10 cm. across, 1-2.5 dm. long, and 40-75-flowered, also
few-flowered, becoming lateral or pseudolateral in age; pedicels finally
4-6 mm. long; calyx to 4 (-5.5) mm. long, 6-7 mm. across, the equal
or unequal (S. bicolor) acute lobes 2.5 mm. long; corolla white or
nearly, ordinarily 13-15 mm. across, the lobes 5-6 mm. long, 4-5 mm.
wide; filaments nearly free, smooth, 1-1.5 mm. long, anthers 3 mm.
long, 1 mm. thick, the apical pores early small; ovary typically stel-
late, sometimes pilose only at apex or glabrous, style subglabrous,
5-5.5 mm. long, stigma capitate; pedicels enlarged in fruit, calyx
lobes 5-6 mm. long; berry 8-12 mm. thick, stellate, the many yellow-
ish reticulate seeds accompanied by 2 stone cells. Mostly after
Bitter, Bot. Jahrb. 55: 66. 1919; the Peruvian specimens (for the
most part determined by Morton) are greener above than beneath;
many, at least, belong to the variant S. auriculatum Alton (var. by
Kuntze, subsp. by Hassler) with one or several small axillary leaves
224 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
simulating stipules, as in the similar (if distinct) S. abutiloides
Bitt. & Lillo of Argentina; cf. Lillo, 1924. S. bicolor Willd. is doubt-
fully a true species on the basis of somewhat smaller floral parts,
glabrous ovary; in view of nonconcomitance and variation of these
characters probably, as S. riparium Pers. (S. dichotomum R. &
P., not Lam.), belongs here if not to the doubtfully distinct S. um-
bellatum Miller. Said to be cultivated in all warmer regions as an
ornamental. Illustrated, Rev. Invest. Agric. (Argentina) 2: 183 (S.
auriculatum) . Section Anthoresis.
Tumbez: Pampa Hospital, Velarde Nunez 327. San Martin:
Tarapoto, Williams 6193; 5671. Juanjui, King 4238. San Roque,
Williams 7470. Huanuco: Near Pampayacu, Kanehira 97. Junin:
Tarma, Herrera 786. La Merced, Killip & Smith 23396; 23729.
Perene" Bridge, Killip & Smith 25270. Pichis Trail, Killip & Smith
25486. To Argentina; West Indies; tropical Africa. "Tai mollaca,"
"funga" (Herrera).
Solatium xanthophaeum Bitter, Repert. Sp. Nov. 16: 401.
1920.
A shrub, type about 3 meters high, shortly yellowish-villous even
to the corolla lobes without, the trichomes mostly branched; leaves
(upper) geminate, the larger oblong-lanceolate, acuminate, 1-3 dm.
long, 3-9 cm. wide, petioles 2-5 mm. long, the smaller sessile, broadly
elliptic or suborbicular, 1-3.5 cm. long, lateral nerves 9-12; inflores-
cence opposite the leaves, simple, few to 17-flowered; peduncles 3-
4 mm. long; pedicels 6-7 mm. long; calyx broadly campanulate,
3.5-4 mm. long, 4 mm. broad, the elliptic-lanceolate lobes about
2 mm. long; corolla white, stellate, to 12 mm. across, the broadly
lanceolate lobes about 4 mm. long, nearly half as wide, cucullate;
filaments 0.8 mm. long, little connate, the ellipsoid emarginate an-
thers 2-2.5 mm. long, style nearly straight, 5 mm. long, the clearly
enlarged stigma capitate. F.M. Neg. 2643. Section Anthopleuris.
Huanuco : Between Monzon and the Huallaga, Weberbauer 3690,
type.
Section LYCIANTHES
Bitter classified the following species, Abhandl. Nat. Verh. Bre-
men, 24: 292-520. 1919, as comprising a genus (Lycianthes) ; however,
the group can be defined with difficulty as a section, as observed by
Morton, all species of Solatium even in Peru considered. Bitter's
FLORA OF PERU 225
account, detailed and informative, is without a key to species; his
divisional key may be correct but is sometimes vague; for instance,
the filaments show a range of development from subequal to more or
less unequal, one or three being longer, this character sometimes in-
transigent; furthermore, subgenus Eulycianthes (Dunal) Bitter, 319,
rests on the division of the drupe-like fruit into sclerotic cells, sub-
genus Polymeris (Dunal) Bitter, 331, with or without granules. Prob-
ably, as Morton observed, these are characters not taxonomically
significant.
Morton, Contr. U. S. Nat. Herb. 29. 1944, presented a series of
keys referring to species then known within several countries; obvi-
ously they were carefully and accurately worked out and I have
largely copied from the one pertaining to Peruvian entities and de-
vised from it a second more artificial key as a supplementary aid to
identification. Many species proposed are not understood, and they
may actually be variants of others or have been incorrectly limited;
perhaps the precise position and characteristics of calyx lobes and
pubescence as well as the obviously and variously unequal filaments
should be investigated for a large series of specimens in the field as
well as in the herbarium.
Trichomes at least in part stellate or furcate, often obscure, simple
in L. acutangulum, S. cyathocalyx, S. stenolobum; filaments of
last ex char, subequal or unequal.
Leaves often narrower than 2 cm. (rarely 2.5 cm.); berry 8-celled;
branches often spine-tipped S. lycioides.
Leaves usually much wider than 2.5 cm.; berry 2-celled; sometimes
with several sclerotic grains; no spines.
Calyx teeth soon 4-8 mm. long.
Trichomes simple, of ten few or none; branchlets as leaves be-
neath typically soon glabrescent or glabrous; corolla to
1.5 cm. wide.
Leaves subequal, acute, to about 3 cm. wide; corolla often
white S. acutangulum.
Leaves mostly clearly unequal, subabruptly cuspidate, the
larger 4-8 cm. wide; corolla violet S. stenolobum.
Trichomes (often lacking, S. Rantonnetii, corolla 2-2.5 cm.
wide) at least in part stellate or furcate.
Indument obscure or absent; flowers 2-2.5 cm. wide.
S. Rantonnetii.
226 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Indument obvious; flowers smaller.
Calyx glabrate or puberulent, teeth somewhat unequal.
S. heterochondrum.
Calyx at least in flower densely pubescent, teeth sub-
equal.
Leaves densely pubescent; trichomes minutely gland-
ular S. Sprucei.
Leaves soon glabrate, eglandular S. luxurians.
Calyx teeth even in fruit to 2.5 (3) mm. long or obsolete.
Calyx teeth developed, subterete.
Trichomes, sometimes few, stellate, more or less stiped.
Leaves cuspidate, 3-6 cm. wide; trichomes stiped-stellate.
S. Poeppigii, S. glandulosum.
Leaves subacute, 2-3 cm. wide; trichomes in part simple.
S. Jelskii.
Trichomes simple (and furcate?); leaves acute.
S. cutacense.
Calyx teeth obsolete S. cyathocalyx.
Trichomes simple, often conspicuous (S. coffeifolium, subglabrous) ;
filaments equal.
Erect or subscandent shrubs.
Leaves strigose or villous at least midnerve beneath, sometimes
sparsely; anthers free.
Calyx teeth exceeding calyx margin.
Leaves to about 5 cm. wide, often narrower; primary nerves
at most 10.
Leaves, unless veins, glabrous above.
Flowers yellow (purple, Ruiz & Pa von) ; leaves strongly
dissimilar S. biformifolium.
Flowers white; leaves subsimilar, strigose beneath.
S. lineatum.
Leaves at least sparsely all over pubescent above.
Indument mostly appressed; leaves to 3 cm. wide.
Corolla to 2 cm. wide; pedicels usually solitary,
elongate S. acutifolium.
Corolla to 1 cm. wide; pedicels 2-4, only 2-6 mm.
long S. brachylobum.
FLORA OF PERU 227
Indument mostly somewhat spreading; leaves to 5 cm.
wide.
Calyx teeth 5-7 mm. long S. chrysothrix.
Calyx teeth 2 mm. long S. densestrigosum.
Leaves much larger, with 12-18 nerves.
Indument appressed strigose S. conspicuum.
Indument spreading.
Calyx teeth almost capillary, 7-10 mm. long; leaves
subsessile S. medusocalyx.
Calyx teeth broader, shorter; leaves petioled . . S. Ulei.
Calyx glabrate, the teeth obsolete or obscure.
Leaves ample, many cm. long and wide S. holocalyx.
Leaves linear-lanceolate, about 1 cm. wide. S. brachylobum.
Leaves essentially glabrous; anthers more or less connate; calyx
teeth minute S. coffeifolium.
Repent herb rooting at nodes S. asarifolium.
LYCIANTHES, VEGETATIVE KEY
Leaves strongly pubescent, with simple (rarely 1-2-forked) trichomes
at least beneath even in age; leaves usually wider than 2 cm.
Calyx teeth obvious, sometimes minute.
Calyx teeth soon 4-10 mm. long, at least the longer if unequal.
Leaves often less than 5 cm. wide; nerves at most 10 pairs.
Leaves glabrous above S. lineatum.
Leaves pubescent both sides.
Trichomes simple.
Pedicels elongate; corolla 1.5-2 cm. wide.
S. chrysothrix, S. acutifolium.
Pedicels 2-6 mm. long; corolla to 1 cm. wide.
S. brachylobum.
Trichomes in part furcate S. Sprucei.
Leaves much larger, with 12-18 nerves S. medusocalyx.
Calyx teeth even in fruit only 1-3 mm. long (types).
Leaves to about 5 cm. wide; nerves about 10.
Pubescence mostly somewhat spreading . . S. densestrigosum.
Pubescence mostly appressed.
228 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Pedicels usually solitary, often 2 cm. long.
S. acutifolium.
Pedicels 2-4, often 1-1.5 cm. long S. cutacense.
Leaves much wider; nerves 12-18.
Pubescence appressed, strigose S. conspicuum.
Pubescence spreading, mostly on nerves S. Ulei.
Calyx teeth obsolete; leaves ample S. holocalyx.
Leaves soon glabrate or if pubescent in age or densely the trichomes
mostly furcate-stellate; leaves sometimes less than 2 cm. wide.
Leaves often much less than 2 cm. long, seldom as wide, usually
oblanceolate, rarely broadly obovate, often fasciculate; tri-
chomes if present stellate S. lycioides.
Leaves usually wider or alternate, geminate and (or) diverse,
rarely long-linear-lanceolate.
Shrubs.
Calyx teeth obsolete or subspathulate or rarely 3 mm. long
even in fruit.
Leaves elongate, linear-lanceolate S. brachylobum.
Leaves rather ovate.
Trichomes at least in part stellate, often minutely.
Indument stiped-stellate; leaves 3-6 cm. wide.
S. Poeppigii, S. glandulosum.
Indument in part simple; leaves 2-3 cm. wide.
S. Jelskii.
Trichomes if present simple.
Calyx teeth (0.5) 1-2 (3) mm. long.
Leaves subequal, glabrous or nearly . S. coffeifolium.
Leaves very unequal, nerves pubescent S. Ulei.
Calyx teeth obsolete.
Leaves subequal, the larger to about 1 dm. long.
S. cyathocalyx.
Leaves unequal, the smaller stipuliform.
S. holocalyx.
Calyx teeth soon 4 mm. long (unless S. heterochondrum) , usu-
ally longer, narrower.
Indument obscure or lacking.
Leaves to 3 or 4 cm. wide, acute; corolla often white.
S. acutangulum.
FLORA OF PERU 229
Leaves in part wider, cuspidate; corolla violet.
S. Rantonnetii.
Indument obvious.
Leaves subequal, little diverse.
Indument evenly distributed on leaves beneath.
S. heterochondrum.
Indument mostly on leaf -nerves beneath.
Indument usually sparse but clearly stiped-stellate.
S. luxurians.
Indument usually dense, mostly simple or trichomes
furcate S. Sprucei.
Leaves very unequal and strongly diverse.
Leaves to about 1 dm. long, a third as wide; stems not
strigose S. biformifolium.
Leaves often much larger; stems strigose.
S. conspicuum.
Herb, repent, rooting at nodes S. asarifolium.
Solanum acutangulum Griseb. in Lechl. Berb. Am. Aust. 58.
1857. Lycianthes acutangula (Griseb.) Bitter, Abhandl. Nat. Verh.
Bremen 24: 357. 1920. S. compressibaccatum Bitter, Repert. Sp.
Nov. 12: 456. 1913, fide Morton. L. acutangula (Griseb.) Bitter,
subsp. compressibaccata Bitter, Abhandl. Nat. Verh. Bremen 24: 358.
1920. L. longidentata Bitter, I.e. 356. S. longidentatum (Bitter)
Morton, Contr. U. S. Nat. Herb. 29: 63. 1944.
Flowering branches more or less divaricate, perhaps always scan-
dent or supported in other shrubs, somewhat quadrate, the decurrent
lines rather prominent; internodes at least 5 cm. long; petioles to
1 cm. long; leaves ordinarily geminate, glabrous or nearly except
often barbellate in axils beneath, lanceolate, attenuate both ends,
larger sometimes 7 or 9 cm. long, usually 2-3 cm. wide, smaller about
half as large; flowers mostly 4-6, pedicels to 3 cm. long, longer in
fruit; calyx tube linear-subulate, 10 teeth, the longer in fruit about
6 mm. long or longer, the shorter 4 mm. long or longer; corolla sub-
rotate, lobes lanceolate, 3 mm. wide, glabrous except toward tip;
filaments unequal, 1.5 mm. long, or longest twice as long, all gla-
brous; anthers 3.5-4 mm. long; berry at least in some forms (as
Bitter's subspecies) compressed, 9 mm. long, 7 mm. thick, seeds as
many as 39, apical granules 2. After Bitter, mostly, for his S. com-
pressibaccatum; flowers white (Ule). F.M. Neg. 2582 (L. longidentata).
230 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Loreto: Cerro de Ponasa, Ule 6800 (type, S. compressibaccatum) .
Puno: San Govan, Lechler 2354, type. Rio Acre: Ule 9733 (type
of L. longidentata) . Brazil.
Solatium acutifolium R. & P. Fl. Peruv. 2: 33, pi. 162, fig. b.
1799. Lycianthes acutifolia (R. & P.) Bitter, Abhandl. Nat. Verh.
Bremen 24: 453. 1920; 182. L. Weberbaueri Bitter, I.e. 446. L. tar-
mensis Bitter, I.e. 451. L. alopecoclada Bitter, I.e. 456. L. chryso-
thrix [Poeppig] Bitter, subsp. dolichopoda Bitter, Repert. Sp. Nov.
18: 317. 1922.
A woody plant sometimes 2 meters tall with flexuose hispid-
strigulose branches, very unequal often dissimilar, somewhat deflex-
ing (in age) leaves more or less pubescent both sides with simple
grayish-yellowish trichomes and usually solitary nodding white flow-
ers; leaves mostly geminate, rarely ternate (the petioles 2-10 mm.
long), oblong- or ovate-lanceolate, acute, to about 8 cm. long, 0.5-
2.5 cm. wide, the smaller (in the upper branches) 1-2 cm. or smaller,
subobtuse, primary nerves 6-8, prominent beneath; peduncles lateral
and terminal, filiform, slightly incrassate above, to 2 cm. long or
longer; calyx campanuliform, hispidulous, the subfiliform teeth sub-
equal, soon 3.5-5 mm. long (about 2 mm. in flower in type) ; corolla
white, stellate, to about 2 cm. wide, 1.5 cm. long, the 7-8 mm. long
lanceolate acute lobes hispidulous; filaments 1-2 mm. long, anthers
3 (-5, Weberbauer) mm. long; berry orange. Varieties or possibly
distinct entities may be included, but more material is necessary to
determine the matter; Bitter observed some 5-celled trichomes on
Poeppig 1075, only 3-celled on at least some otherwise similar plants,
and only 3- or 4-celled on Weberbauer 7041; the former may be part
of the scarcely distinct S. chrysothrix, to which the first three speci-
mens cited below are more or less aberrant intermediates. F.M.
Negs. 2591 (L. tarmensis); 2593 (L. Weberbaueri).
Amazonas: Bagazan to Almirante, Weberbauer 4452 (type, L.
Weberbaueri). La Libertad: Prov. Pataz, Weberbauer 7041 (type,
L. chrysothrix, subsp. dolichopoda}. Huanuco: Cuchero, Poeppig
1075 (type, L. alopecoclada}. Muna, Ruiz & Pavdn, type. Junin:
Huacapistana, Weberbauer 2141 (type, L. tarmensis); Killip & Smith
24322. Carpapata, Killip & Smith 24394. Cuzco: Machupicchu,
Vargas 3348 (det. Killip, L. reflexa Rusby).
Solatium asarifolium Kth. & Bche\ Ind. Sem. Hort. Berol. 10.
1845; Linnaea 19: 386. 1847. Lycianthes asarifolia (Kth. & Bche\)
FLORA OF PERU 231
Bitter, Abhandl. Nat. Verh. Bremen 24: 423. 1920. S. chodatianum
Huber, Bol. Mus. Goeldi 4: 602, fig. 7. 1906, fide Morton.
A creeping stoloniferous herb, the branches, petioles, these some-
times elongate, and the subaxillary solitary 1-flowered peduncles
pilose; leaves geminate, very unequal, 1 petioled, broadly or openly
ovate-cordate or reniform, rounded at apex, the upper sessile, much
smaller, orbicular, both glabrous except the slightly ciliate margins;
flowers single, nodding, the pilosulous calyx turbinate-urceolate, trun-
cate, 5-denticulate; corolla whitish, rotate, the 5 lobes rounded; an-
thers yellow, distinct; ovary and style glabrous, the latter exceeding
the stamens. This herbaceous species is readily distinguished by its
creeping habit and asarum-like foliage. The similar S. violiifolium
Schott ex Spreng., Syst. Veg. 4: 403. 1827, of Brazil has leaves deeply
cordate or reniform and must be known as S. repens (Spreng) Dunal
(Boldon repens Spreng, I.e. 1: 179. 1825). F.M. Neg. (sheet 602865,
S. chodatianum).
Loreto: Contamana, Huber 1336 (type, S. chodatianum). Rio
Acre: Ule9761 (det. Bitter). Paraguay; Bolivia; Venezuela. "Tahu-
ampa" (Huber).
Solanum biformifolium R. & P. Fl. Peruv. 2: 32, pi. 161, fig. a.
1799; 182. Lycianthes biformifolia (R. & P.) Bitter, Abhandl. Nat.
Verh. Bremen 24: 443. 1920.
Shrub with erect or sprawling stems, angled branches, flexuose
branchlets, these hispid with simple subappressed trichomes as the
petioles (2-3 mm. long, canaliculate above, margined); peduncles
solitary (Dunal), filiform, with a single flower (3-5-flowered, fide
Ruiz & Pavon), 4-16 mm. long, pilose; leaves geminate, the larger
ovate-oblong-lanceolate, rounded or subacute at base, acutely acu-
minate, about 5-10 cm. long, 2-3 cm. wide, glabrous except the
obscurely hispidulous midnerve and 5-7 primary nerves beneath;
the smaller leaves sessile, subcordate, suborbicular, glabrous and
green above, pale and hispid on the veins and nerves beneath, 3-
4 mm. long, 4-6 mm. wide, to twice as large; calyx suburceolate, more
or less hispidulous, 4-5 mm. across, the subfiliform segments (5-7)
6 mm. long; corolla violet-purple (Ruiz & Pavon), the lanceolate
oblong acute segments 8 mm. long; filaments 1 mm. long, anthers
2.5-2.8 mm. long; berry reddish, enclosed in the accrescent calyx.
The flowers noted as yellow by other careful collectors at Chinchao,
who found it erect in undergrowth of forest or sprawling.
Huanuco: Chinchao, Ruiz & Pavon, type. Cushi, 4831. Puente
Durand, Rio Chinchao, Stork & Horton 9852; Mexia 8246.
232 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solan um brachylobum Van Heurck & M. Arg. in Van Heurck
Obs. Bot. 71. 1870. Lycianthes brachyloba (Van Heurck & M. Arg.)
Bitter, Abhandl. Nat. Verb. Bremen 24: 457. 1920.
A shrub with shortly pubescent branches and linear-lanceolate
often unequal leaves, the larger 4.5-8 cm. long, 1 cm. wide (usually
narrower), narrowed to a petiole 7-8 mm. long; leaves above scarcely
pubescent (trichomes simple) except on the veins; flowers 4- or 5-
merous, 2-4 on pedicels 11-13 mm. long; corolla whitish, 12 mm.
wide, the lobes about 6 mm. long, densely short-pilose toward the
cucullate tips; filaments 1 mm. long; anthers lanceolate, 4 mm. long;
calyx teeth 8 or 10, very minute, scarcely exceeding the edge of the
2-2.5 mm. long calyx cup; berry 5-6 mm. in diameter, seeds many,
strongly applanate. Distinguished readily from L. chrysothrix [Poep-
pig] Bitter by the minute calyx teeth and small leaves. F.M. Neg.
22876.
San Martin : Tarapoto, Spruce 459, type.
Solarium chrysothrix (Poeppig ex Bitter) Morton, Contr. U. S.
Nat. Herb. 29: 67. 1944. Lycianthes chrysothrix [Poeppig] Bitter,
Abhandl. Nat. Verh. Bremen 24: 448. 1920.
Resembles S. acutifolium but at least typically with more spread-
ing and denser villous-hirsute indument, drying brownish-yellow
(violet in youth), and often larger leaves but apparently these vary-
ing from 5-10 cm. long, half as wide, the smaller more or less conform-
ing or suborbicular in the flowering branchlets as in the related
species; pedicels 1-3, often 2-3 cm. long; corolla more or less pubes-
cent with furcate (or simple) trichomes, about 1 cm. long, 1.5 cm.
wide; calyx teeth capillary, densely villous, soon 5-7 mm. long,
longer in fruit; filaments to 1 mm. long, anthers 3 mm. long. In
S. acutifolium and its segregates the smaller leaves are often sub-
orbicular on the younger branchlets; my 8712 with leaves 5 cm. long,
2 cm. wide, suggests closely S. acutifolium R. & P., sens, lat.; it is
doubtful if this species is distinct from the plant of Ruiz and Pavon.
Huanuco: Casapi, Poeppig, type. Muna, 4021. Yanano, 3712.
Pampayacu, Kanehira 278. Cuzco: Achirani, 2,750 meters, Vargas
11152 (det. Standley).
Solanum coffeifolium (Bitter) Morton, U. S. Nat. Herb. 29:
64. 1944. Lycianthes coffeifolia Bitter, Abhandl. Nat. Verh. Bremen
24:432. 1920.
FLORA OF PERU 233
A practically glabrous shrub; petioles 3-7 mm. long; larger leaves
elliptic-oblong, acuminate, 1-1.5 dm. long, about a third as wide, the
smaller diverse only in size; flowers 2-5, on pedicels 8-10 mm. long,
in fruit 1-1.5 cm.; corolla white, to 1.5 cm. wide, the lobes 6 mm.
long; filaments of yellow anthers equal; calyx teeth 10, unequal,
(0.5) 1 and 1.5 mm. long to 2 mm. long in fruit; berry orange; seeds 8.
Distinguished from L. medusocalyx Bitter and relatives by the
short calyx teeth and few seeds. In the Peruvian specimens the
minute, somewhat reflexing calyx teeth are below the narrow (0.5 mm.
wide) membranous or chartaceous truncate margin. The type is
Ule 9737, Rio Acre.
Huanuco: Tubumayo River, Tingo Maria, Stork & Horton 9565.
Junin: Puerto Bermudez, Killip & Smith 26635 (det. Morton).
Loreto: Puerto Arturo, Williams 5270. Brazil.
Solatium conspicuum Morton, Contr. U. S. Nat. Herb. 29:
67. 1944. Lycianthes tarapotensis Bitter, Abhandl. Nat. Verh. Bre-
men 24: 440. 1920, not S. tarapotense Van Heurck & M. Arg., 1870.
Suffrutescent; leaves 1.5-3 dm. long, 5-15 cm. wide, very unequal,
the smaller often stipuliform, 1-2 cm. long and wide or sometimes
larger, pubescent beneath, as the upper branches, with long simple
appressed yellowish hairs; flowers white, 2-5, on pedicels 7-10 mm.
long; corolla lobes 4-4.5 mm. long; filaments about 1 mm. long, an-
thers 3 mm. long; calyx teeth 10, densely pubescent, 1-1.2 (3) mm.
long, scarcely longer in fruit; berry dark red. Distinguished from
L. Ulei by the appressed pubescence. F.M. Neg. 2590 (Ule 6483).
San Martin: Tarapoto, Spruce 4182, type; 4836 (in some herb.
4830); Ule 6483. Loreto: Pongo de Manseriche, Mexia 6347 (det.
Morton) .
Solatium cutacense HBK. Nov. Gen. & Sp. 3: 38. 1818; 173.
Lycianthes cutacensis (HBK.) Macbr. Field Mus. Bot. 8: 110. 1930.
A shrub, probably scandent, with tomentose-pubescent branches
and ovate acute or subacuminate leaves more or less sericeous-pilose
above, canescent beneath, the trichomes typically simple; petioles
6 mm. long; leaves geminate, the larger to 5.5 cm. long, 1.5-2 cm.
wide; flowers 2-3 (4), pedicels 8-15 mm. long; corolla about 1.5 cm.
long, 2-3 cm. wide; filaments little unequal; calyx teeth 10, the longer
1-2 mm. long, in fruit 3 mm. long, the 5 shorter sometimes obscure.
The species is very near S. lentum Cav. of Mexico and northern
234 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
South America (Brazil); compare the similar S. cyathocalyx Van
Heurck & M. Arg. and S. Jelskii Zahlbr.
Piura: Rio Cutaco near Ayavaca, (Bonpland).
Solatium cyathocalyx Van Heurck & M. Arg. Obs. Bot. 65.
1870. Lycianthes cyathocalyx (Van Heurck & M. Arg.) Bitter, Ab-
handl. Nat. Verh. Bremen 24: 400. 1920.
Soon quite glabrous, only the young parts sometimes ashy pubes-
cent with short simple trichomes; leaves geminate, more or less sub-
equal, petioled, lanceolate-elliptic or -ovate, 5-11 cm. long and about
half as broad, shortly cuspidate, the base acute; flowers in type 2-5
in sessile axillary umbels, in Weberbauer specimens also in open
panicles, the filiform pedicels to 3.5 cm. long in fruit; calyx cyathi-
form, 2.5 mm. long, entire, rarely 1 or 2 gibbous teeth; corolla 10-
12 mm. long; 2 filaments 1 mm. long, 3 about 2 mm. long, anthers
3-4 mm. long; berry 8 mm. in diameter. An abundant white-
flowered subscandent shrub, the flowering parts of the branches con-
stantly elongating so that fruits are normally left behind to mature
(usually singly) in the angles of the branches; flowers slightly irreg-
ular, as shown in the filaments (two are only 1 mm. long) and the
upcurved style; cuneiform seeds wedged together to form a hard core,
the seeds themselves stony in texture due to a surface layer of verti-
cally elongated sclerotic cells (Svenson, Amer. Journ. Bot. 33: 485.
1946). Type from near Chanduy, Spruce 6501. The Loreto collec-
tions suggest this species as to calyx and pubescence. Illustrated,
Svenson, pi. 19, fig. 4-.
Tumbez: Granitic rocks, Quebrada Mogollon, Amotape Hills,
(Haught & Svenson 11528). East of Hacienda Chicama, Weberbauer
7636; 7678. Cajamarca: Socota to Tambillo, Stork & Horton 10164-
Loreto: La Victoria, Williams 2667; 2884? Florida, Klug 2249?
Ecuador.
Solanum densestrigosum Bitter, Bot. Jahrb. 50, Beibl. Ill:
65. 1913. Lycianthes densestrigosa Bitter, Abhandl. Nat. Verh. Bre-
men 24: 443. 1920.
A shrub with densely long strigose younger branches, rather softly
yellow-strigose subequally alternate or geminate elliptic-lanceolate
leaves narrowed to both ends, 11-13 cm. long, 4.5-5 cm. wide; peti-
oles 1.5-2 (2.5) cm. long, trichomes simple, 3-4-celled; inflorescences
sessile, opposite the leaves, with about 5 deeply 4- or 5-parted flow-
ers; pedicels 3 cm. long; calyx teeth 8 or 10, subfiliform, 2 mm. long,
FLORA OF PERU 235
pubescent; corolla to nearly 2.5 cm. wide, the membranous lanceo-
late lobes 10-11 mm. long; stamens glabrous, equal, filaments 1.5 mm.
long, anthers 3.6 mm., cordate; style 7 mm. long, stigma obtuse.
The type (Berlin-Dahlem) was without locality indication; it is doubt-
ful if the calyx teeth are short or that the smaller leaves are con-
stantly less dissimilar than in S. chrysothrix (Poeppig) Morton or
S. acutifolium R. & P., sens. lat. F.M. Neg. 2569. Section Polymeris.
Peru: (Weberbauer, type).
Solatium glandulosum R. & P. Fl. Peruv. 2: 35, pi. 167, fig. b.
1799. Lycianthes glandulosa (R. & P.) Bitter, Abhandl. Nat. Verh.
Bremen 24: 348. 1920. S. vitocense Dunal in DC. Prodr. 13: 176.
1852.
Shrub, often supported in other plants, the younger parts some-
what stellate-pubescent to essentially glabrous; leaves ovate-elliptic-
lanceolate, cuspidately acuminate, membranous, typically pale rusty
stellate-pubescent, especially beneath, the larger 7-10 cm. long or
longer, about half as wide; flowers 3-6, axillary; pedicels 8-10 mm.,
in fruit to 2 cm. long; calyx 4.5-5 mm. long, the 10 (-12?) unequal
teeth linear-oblong-spathulate, 1.5-2.5 (3) mm. long; corolla white,
rotate, about 1.5 cm. long, scarcely 2 cm. wide, the lanceolate lobes
glabrous or slightly stellate-puberulent; filaments 2 mm. long except
one 4.5-5 mm. long; anthers 5-5.5 mm. long; berries with 2 stone cells
(Bitter). Sometimes glabrous (type) and Klug 3991 (given an her-
barium name by Standley), this possibly S. Poeppigii, if that is dis-
tinct; more pubescent plants simulate S. luxurians, with longer more
slender calyx teeth; S. hypomalacum (Bitter) Morton of adjacent
Brazil has more gradually acuminate leaves, nearly glabrous above
and may be Williams 5222 (only in bud). Lycianthes hylophila
Bitter, I.e. 336, of Bolivia might be sought here; its leaves are mostly
3.5-5.5 cm. long, 1.5-2 cm. wide; fruit unknown. F.M. Negs. 2574;
34129 (S. vitocense).
San Martin: Boqueron, Allard 21723 (det. Lyman Smith). Cha-
zuta, Klug 3991. San Roque, Williams 7330? (calyx tube obscure).
Huanuco: Divisoria, Woytkowski 530 (det. Cuatrecasas, S. sub-
inerme); Allard 21778 (det. Lyman Smith, S. luxurians). Cuchero,
Poeppig 1432. Junin: Vitoc, near Tarma, Ruiz & Pavdn, type.
Loreto: Santa Rosa, Williams 4956. Rio Mazan, Jose Schunke 178.
Pucallpa, Ferreyra 1135 (det. Morton); 4153. Puerto Arturo, Wil-
liams 5222? Brazil.
236 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solatium heterochondrum Bitter, Bot. Jahrb. 54, Beibl. 119:
15. 1916. Lycianthes heterochondra Bitter, Abhandl. Nat. Verb. Bre-
men 24: 395. 1920.
A shrub with mostly branched pubescence especially on the
young branchlets and on the young leaves beneath, the older gla-
brescent; petioles 7-18 mm. long; leaves ovate, subacuminate, 4.5-
9.5 cm. long, 3-5.3 cm. wide, herbaceous; flowers often 2-4; pedicels
about 2-2.5 cm. long, to 3 cm. in fruit; calyx rather densely pubes-
cent with usually branched trichomes and with some minute shortly
stipitate glands within; calyx teeth 10, linear, unequal, 3.5 (?) and
6 mm. long; corolla white, rotate, 17-20 mm. wide, the pubescence
(if present) simple; stamens nearly glabrous, about 5 mm. long, the
longer filaments 1.75 mm., the shorter 0.5 mm. long; style glabrous,
scarcely longer than stamens, incurved above; fruiting calyx some-
what ampliate at base; berry 11-13 mm. thick with 4-5 stone cells
(2 subapical larger), about 15-22 seeds. After author, who allied it
to S. Rantonnetii Carr.; calyx lobes appear from scrap of type and
from following material, apparently the same, to be less unequal
than described. F.M. Neg. 2576.
Huanuco: Ambo, 3163? (fruit only). Apurimac: Lambrama,
Vargas 9802. Cuzco: Quillabamba, Goodspeed Exped. 101+1+6. Are-
quipa: Lomas de Pongo, Velarde Nunez 1515? (calyx lobes short).
Puno: Sandia, (Weberbauer 582, type).
Solanum holocalyx (Bitter) Morton, Contr. U. S. Nat. Herb.
29: 60. 1944. Lycianthes holocalyx Bitter, Abhandl. Nat. Verh. Bre-
men 24: 459. 1920.
Subligneous, 3-6 meters tall, the upper stem, petioles (these about
1 cm. long) and leaves beneath, especially on the 13 or more promi-
nent nerves and midnerve, more or less appressed strigose; leaves
often somewhat falcate-oblong, usually about 1.5-2 dm. long, 5-
10 cm. wide, acutely caudate-acuminate, the smaller stipuliform,
2-several cm. long; pedicels few to several, about 1 (-2) cm. long,
little longer in fruit; calyx strigillose to glabrous, truncate, 2-2.5
(-4) mm. across, quite truncate, teeth lacking; corolla yellowish or
greenish white, pubescent, 1.5-2.5 cm. across, lobes 7 mm. long (or
longer); filaments subequal, 2 mm. long, anthers 3.5 mm. long; fruit
dark red, about 8 mm. in diameter. The Peruvian plant may prove
to be a variant at least of the type, which was collected near Quito;
specimens (except those of Mexia) determined by Morton with the
FLORA OF PERU 237
remark: it is either the same or very closely allied. Sterile specimens
of this and S. conspicuum Morton simulate each other.
San Martin: Juanjui, Klug 4261. Huanuco: Rio Cayumba,
Mexia 8259 (glabrate). Junin: Near La Merced, Killip & Smith
23940. Colonia Perene", Killip & Smith 25152. Near Peren6 Bridge,
Killip & Smith 25314. Balsapuerto, Killip & Smith 28446; 28507;
28555; 28557; 28572. Ecuador.
Solarium Jelskii Zahlbr. Ann. Naturh. Hofm. Wien 7: 7. 1892.
Lycianthes Jelskii (Zahlbr.) Bitter, Abhandl. Nat. Verh. Bremen 24:
399. 1920.
Densely divaricately and furcately branched shrub, the sparse
pubescence mostly simple; leaves thin, ovate or ovate-oblong, obtuse
or acutish, the larger 3-5.5 cm. long; flowers 2-4, between the leaf-
axils; pedicels 8-12 mm. long; calyx 3 mm. long, 10-costate, the 5
longer subulate teeth 1 mm. long, the 5 shorter often obscure; corolla
violet, the lobes 10-11 mm. long; filaments unequal, two 1.5-2 mm.
long, the others 3 mm. long, anthers all about 4 mm. long. Nearly
L. cutacensis but glabrate and calyx teeth shorter. F.M. Neg. 2580.
Cajamarca: Cutervo, Jelski 45; 47, type collections.
Solarium lineatum R. & P. Fl. Peruv. 2: 31, pi. 158, fig. &. 1799;
182. Lycianthes lineata (R. & P.) Bitter, Abhandl. Nat. Verh. Bre-
men 24: 442. 1920.
Leaves glabrous above, subappressed yellowish strigose beneath,
the larger to 5 cm. wide, about twice as long, oblong- to broadly
elliptic, coriaceous in age, the somewhat to much smaller similar;
flowers 1 or 2, white or faintly purplish, about 7 mm. long; pedicels
arcuate, in fruit 2-3.5 cm. long; calyx teeth subulate-setaceous, 2-
4 mm. long or 6-8 mm. long in fruit, the latter at maturity about
1.5 cm. thick, orange. My collections were unbranched shrubs a few
dm. tall on open grassy slopes. The description in part by Bitter
based on the Poeppig plate. After Ruiz and Pavon: nerves very
prominent beneath, 10-12 pairs, leaves ovate, acuminate, the larger
twice the size of the smaller, in plate about one-third, merely acute,
in age subrevolute, glabrous above, densely hirsute beneath, younger
leaves both sides; pedicels inter-axillary, one-flowered, filiform, en-
larged apically, pubescent as calyx, this with 10 subulate lobes, the
longer corolla white; berry included, brown.
Huanuco: Casapi, Poeppig 1537. Muiia, Ruiz & Pavdn, type;
3971. Rio Huallaga Canon, 4248.
238 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solarium luxurians Morton, Contr. U. S. Nat. Herb. 29: 66.
1944. Lycianthes Pearcei Bitter, Abhandl. Nat. Verb. Bremen 24:
355. 1920, not S. Pearcei Britton, 1895.
Aspect of S. glandulosum but with the slender longer calyx teeth
of S. Sprucei; younger parts stipitate stellate pubescent, the indu-
ment mostly lacking in age; leaves broadly ovate, rounded at base,
long-acuminate; flowers 4-6 in axils or often on short branchlets,
pedicels 9-12 mm. (in fruit 10-15 mm.) long; calyx 1 cm. long, 8 mm.
(in fruit 17 mm.) across, teeth alternately unequal, linear-subulate,
5-7 mm. long in flower; filaments unequal; anthers 5.5 mm. long;
berry globose, scarlet, to 1.5 cm. in diameter, seeds about 40, reticu-
late, granules 2. Type from Coroica, Bolivia, by Pearce. A shrub
or liana with white flowers, bright orange fruit.
Loreto: Iquitos, Killip & Smith 29849. Mishuyacu, King 1321;
1484 (det. Morton, ex char.).
Solanum lycioides L. Mant. 46. 1767; 161. S. lycioides L. var.
tomentosum Dunal, Hist. Sol. 174. 1813. S. candicans Dunal, Sol.
Syn. 23. 1816. S. lycioides L. var. parvifolium Wedd. Chloris And.
2: 107, pi. 55. 1857. S. Dombeyi Dunal in DC. Prodr. 13: 162. 1852?
Lycianthes lycioides (L.) Hassl. Ann. Cons. Jard. Bot. Geneve 20: 181.
1917; var. brachyphylla Bitter, Abhandl. Nat. Verh. Bremen 24: 327.
1920; subsp. tomentosa (Dunal) Bitter, I.e.; subsp. parvifolia (Wedd.)
Bitter, I.e. 328. L. candicans (Dunal) Hassl. I.e.
Shrub, a few dm. to rarely 2 meters tall, often intricately (some-
times virgately) branched, the short branchlets frequently spine-tipped;
leaves glabrous to stellate puberulent-tomentose, not unusually sub-
fasciculate, broadly obovate- to oblong-elliptic or narrower, rarely
subrotund (S. Dombeyi), commonly about 2-4 cm. long; flowers
ordinarily bluish-violet with yellow center, the 1-3 (5) pedicels soon
1.5-2 (2.8) cm. long; calyx teeth 5 (rarely with intervening smaller
teeth), (1) 2-2.5 mm. long; corolla usually about 1.5 cm. wide; fila-
ments of red or orange anthers unequal; fruit drupiform, to 8 mm.
thick with 8 sclerotically enclosed seeds. Variable especially in
leaves and degree of pubescence to complete lack; the var. tomento-
sum as to type was based on lack of spines as well as indument, a
not constant concomitance; S. Dombeyi, found only once, may be
recognizable, as suggested by Morton, Contr. U. S. Nat. Herb. 29:
56. 1929, on leaf shape, corolla 2-2.5 cm. wide, calyx teeth 1 mm.
long. F.M. Neg. 25701 (S. Dombeyi).
FLORA OF PERU 239
Morton, I.e. 54-55, showed clearly that the characters used by
Bitter to justify Lycianthes as a genus are invalid as they are found
at random in the genus in unrelated sections; the segregate there-
fore, if accepted, must rest on the single character of fruit; sclerotic
granules are present in Solanum without being tied together and,
as he remarks, the character is of doubtful taxonomic significance.
In 1917 Bitter himself referred to the type shrub as a Solanum.
Cajamarca: San Miguel to Asuncion, Ferreyra 7084- Huambos,
Prov. Chota, Soukup 4550; 4552 (det. Morton). La Libertad: Tru-
jillo to Suisicap, Rosa Scolnik 1271. Ancash: Below Ocros, Weber-
bauer 271+3. Lomas de la Chao, Goodspeed Exped. 205 (det. Johnston) .
Lima: Viso, Weberbauer 126 (det. Bitter); 166. Matucana, 332;
2890; 2948. Surco, Nunez 289 (var.). Huanuco: Near Huanuco,
2330; Ruiz & Pavdn (type, var. tomentosum) ; Seibert 2219. Chulque,
Mexia 04106. Tarma to Chinchao, Dombey 245 (type, S. Dombeyi) .
Junin: Huancayo, Soukup 3545. Tarma to Palca, Ruiz & Pavdn;
Weberbauer 1741; 176. Huacapistana, Velarde Nunez 785 (toward
S. Dombeyi, leaves rounded to abruptly acute base). Ayacucho:
Prov. Lucanas, Metcalf 30304. Puquio, Ferreyra 5463. Apurimac:
Oropeza Valley, Vargas 9769. Below Moyobamba, West 3734 (det.
Johnston). Saxaihuaman, H err era 846 (as S. pseudolycioides) .
Cuzco: San Sebastian, Pennell 13605. Ollantaitambo, Cook &
Gilbert 284; 431 A. Arequipa : Mum 15496; Pennell 13183. El Misti,
Sandeman 3810. Above Salamanca, Weberbauer 6855 (det. Bitter).
Puno: Near Puno, Soukup 448. Tacna: (Weddell, type, var. parvi-
folium). To Colombia, Argentina, Chile. "Upa-ttancar," "ttitir,"
"ttinquir" (Herrera); "misquitlancas" (Cook); "turu cashi" (Sei-
bert); "amacasa" (Dombey); "tin-tico" (West); "naranjito del cam-
po" (Mexia).
Solanum medusocalyx Bitter, Repert. Sp. Nov. 12: 549. 1913.
Lycianthes medusocalyx Bitter, Abhandl. Nat. Verh. Bremen 24:
437: 1920.
Pubescent with short and long spreading yellowish simple tri-
chomes most numerous on the branches and larger leaf veins; leaves
sessile or subsessile, very unequal, geminate, the larger broadly
lanceolate, subacuminate, 1-2 dm. long, the smaller obliquely
cordate-reniform, about 1 cm. long; flowers 4-9; pedicels 2-2.5 cm.
long; corolla stellate, violet or lilac, about 1.5 cm. wide, pubescent,
the lobes 2 mm. wide, 5 mm. long; stamens equal, the filaments
glabrous, 1.5 mm. long, anthers cordate-lanceolate, 4 mm. long;
240 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
calyx teeth 10, very unequal, subulate-filiform or setaceous, 6-7
mm. long; berry red, 8 mm. thick; seeds many, small, granules
absent. The recent specimen has shortly petioled leaves, longer
finely filiform calyx teeth. F.M. Neg. 2583.
Junin: Yapas, Pichis Trail, Killip & Smith 25446 (det. Morton,
with query). Loreto: Serra de Escaler, Ule 6804, type.
Solatium Poeppigii (Bitter) Morton, Contr. U. S. Nat. Herb.
29: 66. 1944. Lycianthes Poeppigii Bitter, Abhandl. Nat. Verh.
Bremen 24: 345. 1920.
Distinguished by the author from L. glandulosa (R. & P.) Bitter
by the sparser yellowish pubescence and longer trichome stipes;
pedicels 8-10 mm. long; calyx teeth 2.5-3 mm. long; fifth filament
5-5.5 mm. long, anthers 5.5-6 mm. long. The type, as observed
by Morton, has leaves glabrous above except veins, calyx teeth
equal, 1 mm. long, stamens larger; the discrepancy in characters
is probably close to individual variations or degree of development;
if not, some of the specimens cited under S. glandulosum may be
referable here. F.M. Neg. 23069.
Loreto: Yurimaguas, Poeppig 2406, type.
Solanum Rantonnetii Carr. ex Lescuyer in HeYincq, L'Hort.
Franc. seV. 2, 1: 197, pi. 16. 1859. Lycianthes Rantonnetii (Carr.)
Bitter, Abhandl. Nat. Verh. Bremen 24: 332. 1920.
Shrub, glabrous or nearly, with simple trichomes or these some-
times slightly branched early on leaves beneath and on margins,
the stems erect from base, branched, nearly angulate, about 5 mm.
in diameter below, internodes nearly 5 cm. long; petioles nearly
1-2.5 cm. long; leaves alternate or geminate, the larger mostly
5-7 cm. long, 3-4 cm. wide, to twice as large, all broadly lanceolate
to rhombic-lanceolate, cuneate to base, narrowed to acute or acu-
minate apex or this somewhat obtuse, pale green; lateral nerves 6-8;
flowers 5-7, subumbellate, pedicels 12-15 mm. long, stipitate glan-
dular; calyx teeth linear, the 5 larger 3-4 mm. long (to 6 mm. long
in fruit), the 5 smaller 1-2 mm. long or lacking; corolla violet,
rotate, 2-2.5 cm. across, lobes joined nearly to the cucullate, densely
pilose tip; 3 filaments 2-3 mm. long, 2 half as long, pubescent below;
anthers orange, equal, 3 mm. long, style to 5 mm. long; berry
subglobose, granules more than 25. After Bitter, Repert. Sp. Nov.
12: 458. 1913. The Peruvian specimens were probably cultivated
FLORA OF PERU 241
originally; perhaps all or mostly referable to S. acutangulum or the
similar, if distinct, S. stenolobum.
San Martin: Orroyo Branco, Allard 21764- Lima: Wawra 2584
(det. Killip); (Martinet 109}. Huanuco: Divisoria, Allard 21336
(det. Lyman Smith); Ferreyra 8140. Hacienda El Aguila, near
Tingo Maria, Ferreyra 4178; Allard 20412; 21145; 20660 A (all det.
Lyman Smith); Stork & Horton 9466 (det. Standley, S. stenoloba).
Tulumayo, Ferreyra 2147. Cuzco: Echarate, Goodspeed Exped.
10458 (det. Standley, S. stenoloba). Paraguay; Argentina; Bolivia
and Brazil. "Capaicayayi" (Argentina).
Solanum Sprucei Van Heurck & M. Arg. Obs. Bot. 67. 1870.
Lycianthes Sprucei (Van Heurck & M. Arg.) Bitter, Abhandl. Nat.
Verh. Bremen 24: 380. 1920.
A shrub, the branches, at least toward the tip, densely rusty
tomentose and usually glandular with simple and few to many
2-3-forked more or less stipitate trichomes intermixed; leaves softly
pubescent both sides, unequal, obliquely lanceolate, acuminate, the
larger 8-11 cm. long, about half as wide; flowers usually 2-3, 15-17
mm. long on pedicels soon 2 cm. long; fifth filament longer than
the equal 4, anthers all 5 mm. long; calyx densely pubescent, 10-11
mm. long, the 10 subequal narrowly subulate teeth 6-8 mm. long.
F.M. Neg. 23072.
San Martin : Tarapoto, Spruce 4352, type. San Roque, Wittiams
7135; 7653. Junin: Chanchamayo Valley, Schunke 7135. Bolivia?
Solanum stenolobum Van Heurck & M. Arg. Obs. Bot. 69.
1870. Lycianthes stenoloba (Van Heurck & M. Arg.) Bitter, Abhandl.
Nat. Verh. Bremen 24: 358. 1920.
Branches subcylindric, slender, often tortuous, early papillose-
puberulent, internodes 5-9 cm. long; petiole alate, 5 mm. long; leaves
membranous, glabrous or essentially, geminate, more or less unequal,
elliptic-lanceolate, the larger 5-8 cm. wide, about 1.5-2 dm. long,
the smaller often 3-5 cm. wide, 4-8 cm. long, or sometimes the
smaller similar to the larger; lateral nerves 9-10; flowers 1 or more
on pedicels about 1.5 cm. long; calyx teeth 10, subequal, linear-
subulate, about 5 mm. long, to 8 or 9 mm. in fruit; corolla violet,
stellate, 2 cm. across, lobes 4 mm. wide, glandular without and
puberulent, apically cucullate; filaments subequal (Bitter) or some-
times one longer(?); anthers 6.5 mm. long or longer; fruit red,
subglobose or ellipsoid, 1 cm. long, seeds about 15, the 2 subapical
242 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
stone cells larger. S. longidentatum (Bitter) Morton of Rio Acre,
probably in Peru, has filaments 0.6-1 mm. long, the fifth 2 mm.
long; trichomes 2-9-celled; in S. acutangulum the trichomes are
subequal in length. F.M. Negs. 33132; 2589 (Ule).
San Martin: Juan Guerra, Ule 6571, type. Tarapoto, Spruce
4210; Williams 6541; Ule 6873Ayd.cucho: Rio Apurimac Valley,
Killip & Smith 22989.
Solarium Ulei (Bitter) Morton, Contr. U. S. Nat. Herb. 29: 60.
1944. Lycianthes Ulei Bitter, Abhandl. Nat. Verh. Bremen 24:
437. 1920.
A shrub allied by author to L. medusocalyx Bitter but the slightly
larger leaves on petioles 6-9 mm. long and the smaller obliquely
ovate, 3-5 cm. long; pedicels 7-11 mm. long; calyx 3 mm. long
and broad, in fruit twice as large, the teeth only 1-2 mm. long;
corolla white, 1 cm. wide, the lobes 4-4.5 mm. long; filaments
0.6-0.8 mm. long, anthers 2 mm. long; seeds many, minutely re-
ticulate. Probably occurs within Peru; the Ecuador plant was des-
ignated subspecies dolichodontum Bitter, 438, calyx teeth 4-5 mm.
long. F.M. Neg. 9764.
Acre District: Seringal San Francisco, Ule 9764, type. Ecuador.
Subgenus LEPTOSTEMONUM
The subgenus Leptostemonum, to which all the following species
belong, is fairly well-marked by the more or less elongate and little-
to long-attenuate (to tip) anthers, which are dehiscent by terminal
minute or small posterior pores, or sometimes longitudinally; most
plants often aculeate, not always even in the same species. In
this group, especially, occur non-fertile flowers; they are usually
those with reduced style above base of inflorescence. The key ob-
viously has been devised as an aid to determination, not to suggest
relationships; thus variable or not understood species are sometimes
repeated. The same intent has been promoted in the keys of the
other subgenera; the exotic S. melongena L. and S. giganteum Jacq.
(in the text) are omitted.
Corollas rotate-stellate, sometimes deeply parted but segments some-
what ovate or ovate-lanceolate or, if narrow, flowers small
(-6 mm. long) and calyx teeth filiform.
FLORA OF PERU 243
Leaves green at least above but sometimes sparsely (minutely
or coarsely) pubescent with usually (rarely simple) stellulate
sessile or stiped trichomes.
Leaves at least in part pinnate or pinnately lobed, subsessile
or somewhat bipinnate, or very aculeate.
Leaves petioled, aculeate; calyx lobed; leaves more or less bi-
pinnate S. concisum, S. sisymbrifolium.
Leaves subsessile, little or not aculeate; calyx crenate.
Leaves pinnate S. Barbeyanum.
Leaves pinnately lobed or in part repand . . . . S. nemorense.
Leaves entire to repandly lobed; calyx lobed or early crenate.
Leaves sessile or nearly, or, if not, usually long-attenuate to
base.
Margins of large leaves spinulose or entire. S. yapacaniense.
Margins of small leaves repand-lobulate to entire.
S. leucopogon.
Leaves distinctly petioled, entire or rarely repand-lobulate
but not or sparsely prickly.
Calyx small, the lobes rounded; leaves soon glabrous above,
pallid or white tomentose beneath.
Indument compact; inflorescence pedunculate.
S. albidum.
Indument soft, open; inflorescence sessile. . . .S. coconilla.
Calyx lobes acute to caudate, rarely truncate-denticulate;
leaves fulvous beneath or subconcolor, often glabrate
or simply pubescent or scabrous above.
Calyx 4-5 (-8) mm. long; leaves 1-2 dm. long; flowers
to 1 cm. long.
Leaves acute or rounded at base.
Indument of branchlets and cymes sparse.
S. saponaceum.
Indument, at least of cymes, dense, hispid.
S. hispidum.
Leaves cordate S. rhamphidacanthum.
Calyx about 1 cm. long; leaves often 3 dm. long or longer;
flowers large (except sometimes S. Poeppigianum) .
Calyx lobes caudate, long echinate-aculeate.
S. kioniotrichum.
244 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Calyx lobes acute, tomentose-echinate.
S. Poeppigianum, S. grandiflorum.
Leaves more or less densely pubescent above (surface, unless S.
jamaicense, at least concealed), also beneath, unless in age,
entire to repandly lobed; cf. S. concisum; S. talarense may be
misplaced, ex char.
Flowers in sessile or subsessile lateral clusters; leaves softly
pubescent above as beneath or cuneate to base.
Leaves somewhat cordate; trichomes mostly or all simple.
Calyx lobes ovate, acute, 12-15 mm. long; leaves ample.
Calyx long-pilose S. quitoense.
Calyx tomentose S. topiro.
Calyx lobes subulate-filiform or if ovate-lanceolate, acutely
acuminate, shorter unless in fruit; leaves small or
ample.
Prickles all fine, terete or subterete, to 6 mm. long.
S. incarceratum.
Prickles in part longer, coarser or compressed.
Branches villous as 3-7-repand-lobulate leaves.
S. mammosum.
Branches long-hirsute as angulate-repand leaves.
S. pectinatum.
Leaves cuneate at base, entire; indument stellate.
S. jamaicense.
Flowers, unless early, usually in well-peduncled inflorescences;
indument at least in part forked or stellate, often stiped
trichomes.
Plants sordidly tomentose; calyx teeth subulate; corolla ro-
tate S. velleum.
Plants usually conspicuously orange or reddish stellate-
tomentose.
Indument developed more or less unevenly.
Calyx lobes caudate, more or less echinate.
S. kioniotrichum.
Calyx lobes acute or mucronate, somewhat tomentose
or stiped-stellate, rarely aculeate.
Flowers usually at least 2 cm. long and wide, few;
leaf indument in part stellate.
FLORA OF PERU 245
Branchlets finely aculeate S. talarense.
Branchlets stellate-tomentose S. gmndiflorum.
Flowers usually smaller, often many, if few, short;
leaf indument above in part simple.
S. Poeppigianum.
Indument even, short, compact, usually reddish unless
leaves above in age S. asperolanatum.
Corollas at least two-thirds parted, in any case buds as segments
suboblong-lanceolate (rarely narrowly ovate or ovoid), the latter
usually longer than 1 cm.; filaments at most 1 mm. long, free.
(Perhaps, in Peru, one collective and only two other species.)
Calyx and pedicels eglandular (typically); calyx lobes minute,
erect or reflexing.
Leaves glabrous or minutely pubescent above, often in part
geminate and about twice as long as wide; calyx minutely
denticulate S. subinerme.
Leaves finely but clearly pubescent above, in part ternate,
often suboblong, about three times longer than wide.
Calyx teeth and anthers slender S. heterophyllum.
Calyx teeth ovate, linear tips reflexing; anthers conical.
S. enoplocalyx.
Calyx and pedicels more or less glandular; calyx lobes early dis-
tinct, except S. torvum.
Leaves entire or subentire, rarely repandly lobulate.
Leaves cuneate to sessile or subsessile base . . . . S. jamaicense.
Leaves long-petioled.
Leaves in part ternate, suboblong S. heterophyllum.
Leaves often geminate, broadly ovate S. torvum.
Leaves pinnately lobed, very aculeate S. sisymbrifolium.
Solatium albidum Dunal, Hist. Sol. 206. 1813; 280. S. incanum
R. & P. Fl. Peruv. 40, pi. 175, fig. 6. 1799, not L., 1753. S. Mandonis
Van Heurck & M. Arg. Obs. Bot. 78. 1870 and subspecies cara-
bayense Bitter, Bot. Jahrb. 50, Beibl. Ill: 66. 1913. S. macrophyllum
Dunal, I.e. 199. 1913; 347?
A shrub or tree sometimes attaining 8 meters, the stems and
branches typically soon glabrous as the leaves above, but the former
often sparsely spinose, the prickles broad-based especially in age;
petioles terete, 2-15 cm. long, often, as the branchlets, purple,
246 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
rarely pubescent or aculeate; leaves solitary and geminate, broadly
ovate-oblong, very unequal at the rounded base, entire, repand,
or broadly lobed, somewhat acuminate, the larger to 5 dm. long,
nearly half as wide, those on the flowering branchlets much smaller,
all reticulate veiny above, strikingly and closely tomentose beneath
unless the 5 or 6 primary nerves, these usually not prickly; in-
florescence compound, often 5 to 10 cm. wide, the peduncles to
2 cm. long, finally glabrate as the primary branches, pedicels 6-8
mm. long, tomentose as the calyces and corollas, the former cyathi-
form, about 5 mm. across, 5-crenate or soon shortly 5-lobed, the
lobes rounded; corolla nearly 1 cm. long, white, glabrous within,
the lanceolate-acuminate segments to 6 mm. long; stamens equal,
6 mm. long; anthers fulvous below, oblong-linear with 2 apical
pores; ovary nearly glabrous, style basally articulate, flexuose,
pulverulent, the stigma with 2 filiform cusps; fruit globose, 6-8 mm.
in diameter, finally black. The Bolivian S. Mandonis Van Heurck
& M. Arg. and its variety apparently represented in Peru by West
7193 were united in Herb. Dahlem but indeed seem to differ in the
more persisting pubescence on the branches, the leaves shortly
aculeate on the lower portion of the midnerve beneath; probably
worthy of varietal indication. S. saponaceum Dunal varies in much
the same way. Probably the origin of the garden type of S. macro-
phyllum Dunal was Peruvian rather than Mexican and it apparently
pertains to S. albidum. F.M. Neg. 23129 (S. macrophyllum) .
Cajamarca: Cascas, Raimondi (det. Werdermann). Arenales,
Stork & Horton 10160 (det. Standley). Huanuco: Near Huanuco,
2363; Ruiz & Pavdn, type. Junin: San Ramon, Constance & Tovar
2230; Killip & Smith 24.745. La Merced, 5463; Soukup 2525.
Oxapampa, Soukup 2545. Ayacucho: Churubamba, Mexia 8129
(det. Morton, S. Mandonis). Aina, Killip & Smith 23126. Cuzco:
Huadquina, West 7193. Machupicchu, Herrera 3272. Santa Ana,
Cook & Gilbert 1654- Puno: Sandia, Weberbauer 582 (type, S.
Mandonis, subsp. carabayense). "Anti-Christo," "yurahuacta," or
"yurahuassa" (Ruiz & Pavon) ; "enves bianco," "huaritar" (Mexia) ;
"espina blanca."
Solanum asperolanatum R. & P. Fl. Peruv. 2: 39, pi. 174,
fig. b. 1799; 277. S. asperum Pers. Syn. 1: 229. 1805, not Vahl. S.
lanatum Dunal, Hist. Sol. 205. 1813.
A remotely prickly (only young stems below) or smooth shrub
or tree, to 10 meters tall; flowering branchlets and leaves especially
FLORA OF PERU 247
above early densely and shortly (evenly) reddish lanate with shortly
stipitate and sessile or subsessile stellate trichomes; young basal
or lower stems glabrous, aculeate, the upper espinose, pubescent;
petioles stout, 1 cm. long; leaves ovate or elliptic-ovate, entire
or subrepand or rarely shallowly lobulate, acuminate, the lower
solitary, the uppermost often subgeminate, 25 cm. long, 12 cm.
wide, or smaller, short stellate pubescent, somewhat denser beneath,
or sparsely stellate in age, the 4-5 primary nerves more prominent
below; corymbs subdichotomous, to 1 dm. long, about as wide;
peduncles to 2 cm. long, pedicels approximate, 6-8 mm. long; calyx
poculiform, 6 mm. across, the segments ovate, acute; corolla white,
lanate without, in the type shorter than 1 cm., plicate at base, the
segments lanceolate, acuminate, stellate-radiate; stamens half as
long as the corolla; filaments filiform, short, the anthers oblong-
acuminate, to 8 mm. long, dehiscing apically from two pores; style
straight, little longer than stamens, stigma bifid; fruit yellow, the
seeds lutescent. The specimens from San Martin have smaller
flowers, pubescence (in herb.) grayish. A shrub, seemingly char-
acteristic in pubescence, has notably larger flowers, and has been
referred here by Killip with doubt; it is worth-while to call it var.
Tovarianum Macbr., var. nov., corolla 5-fida, laciniis plus minusve
recurvatis, 1.5 cm. longis (type, Metcalf 30547). F.M. Neg. 12994.
San Martin: Juanjui, King 4258. Morales, Williams 5671. San
Roque, Williams 6972. Huanuco: Villcabamba, Rio Chinchao, 5166.
Pasco: Quillasu, Soukup 3323 (det. Lyman Smith). Junin:
Huasahuasi, Ruiz & Pavon, type; P. C. Hutchison 1134- Carpapata,
Killip & Smith 24389; 24427; 25761. Ayacucho: Ccarrapa, Fer-
reyra 3740; Killip & Smith 22275 (det. Morton) . Cuzco : Huad-
quina, West 7192. Puno: Sandia, Metcalf 30547 (type, var. To-
varianum).
Solatium Barbeyanum Huber, Bol. Mus. Goeldi 4: 605. 1906.
A glabrous scandent uncinate-aculeate shrub with yellowish-green
flowers distinguished from S. nemorense Dunal by the strictly solitary
and distinctly pseudopinnate leaves; leaves geminate, the larger
4-pinnate, pinnae 1.5 dm. long, 4 cm. wide or larger, alate midrib
about 2 cm. wide, the smaller usually a single pair and sometimes
a terminal leaflet often only a few cm. long; flowers few, the lower
falling; calyx crenate, 2.5 cm. long, 3 mm. across; corolla lobes
lanceolate-acuminate, 1 cm. long, white or greenish; stamens acu-
minate, subequal, 5-6 mm. long; pedicels 8 mm. long in flower,
248 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
becoming 2 cm. long and conical beneath; fruit bright orange or
reddish, 3 cm. in diameter. Character doubtfully an individual
variation; specimens referred by Morton to S. megistophyllidium
Bitter, Repert. Sp. Nov. 12: 554. 1913, this with 10-15 white
flowers, calyx 5 mm. across, subsessile anthers about 9 mm. long,
certainly related.
Loreto: Contamana, Huber 1335, type. San Antonio, Killip
& Smith 29^66. Soledad, Killip & Smith 29682; 29561. Itaya,
Williams 3517; Killip & Smith 29466. Brazil?
Solanum coconilla Huber, Bol. Mus. Goeldi 4: 604. 1906.
S. demerarense Dunal, var. inerme Dunal in DC. Prodr. 13, pt. 1:
257. 1852.
A shrub with broadly ovate-elliptic or nearly suborbicular sinuate-
angulate firm leaves (petioles 4-8 cm. long), unequal or usually
subcordate at base, glabrous above, pale beneath with a loose stellate
tomentum, 1-2 dm. long, often nearly as wide; cymes sessile or
subsessile, pseudolateral, 5-11-flowered; calyx 3 mm. long, 4 mm.
wide, minutely or obscurely denticulate, tomentose as the stout
1 cm. long pedicels and corolla without; calyx and corolla white,
9 mm. long, scarcely 1 cm. across; filaments very short; young fruits
stellate tomentulose, glabrous when mature, red, at least 2 cm. in di-
ameter. S. demerarense Dunal is S. stramonifolius Jacq. fide Bitter in
Herb. DC., surely a hasty conclusion as the East Indian species
has leaves stellate above, calyx lobes 2-3 mm. long; the Peruvian
shrub, however, may be only a variant of Dunal's, as he thought,
but that has aculeate branches and leaves, the latter membranous,
filiform pedicels, corollas 1.5 cm. across. F.M. Neg. 6812 (Poeppig).
Loreto: Leche, Pampa del Sacramento, (Huber 1532, type). Santa
Ana, Williams 1247. Rio Nanay, Williams 435; 693. Iquitos,
Williams 3555; Killip & Smith 27345. Santa Rosa, Killip & Smith
28713. Yurimaguas, Williams 4327; 4358. La Victoria, Williams
2577. Without locality, Poeppig 2225 (type, S. demerarense, var.).
"Coconilla."
Solanum concisum Dunal in DC. Prodr. 13, pt. 1: 327. 1852.
Branches, at least, herbaceous, aculeate with unequal prickles
2-10 mm. long, and sericeous-velutinous with long-radiate stellate
trichomes; leaves pinnatifid, the lobes sinuate-repand, aculeate on
both sides and grayish-tomentose above, whitish-tomentose beneath,
7-10 cm. long; petioles stellate-hirsute and prickly, 2.5-4.5 cm. long;
FLORA OF PERU 249
peduncles 1-flowered, solitary or paired, with the calyces densely
echinate with yellow prickles; calyx lobes 6, long-acuminate; calyx
enlarged in fruit to nearly 2 cm. in diameter. Perhaps distinct
from S. sisymbrifolium Lam. in the more stellate pubescence; flower
parts 6. The Metcalf specimen is so glandular that the adhering
sand grains covered the stellate trichomes; its identity may be
questionable, as also the type locality, perhaps a similar name in
Ecuador or Mexico. F.M. Neg. 8581.
Amazonas: Chachapoyas, Metcalf 30806 (det. Killip)? Huanuco:
Chaucha, Ruiz & Pavdn, type. Ecuador?
Solanum enoplocalyx Dunal in DC. Prodr. 13, pt. 1: 222. 1852.
Branches flexuous, stellate-hispid, aculeate; leaves geminate or
ternate, entire or repand, ovate-oblong-lanceolate or broader to about
1 dm. long, 2-3 cm. wide, greenish but stellate-pilose above, stellate-
hispid beneath, the midnerve little if at all aculeate above, always
beneath; peduncles lateral, 8-12 mm. long, not aculeate; racemes
cymose, 4-5-flowered; pedicels subfiliform, 8-12 mm. long, aculeate;
calyx stellate-tomentose with many small prickles; calyx lobes ovate,
the narrow acuminate tips reflexed; corolla-lobes oblong-linear, 12-
15 mm. long, about 2 mm. wide; anthers conical, 7 mm. long, the
short filaments subcompressed. The leaves are unequal, the smaller
broader ones on petioles only 2-6 mm. long, the larger with petioles
1-2 cm. long. Not included by Standley in his "Shrubs and Trees
of Mexico," but probably a part of S. heterophyllum Lam. (S. sub-
inerme Jacq.) or closely allied.
Peru(?): (Pavdn, type). Mexico?
Solanum giganteum Jacq. Coll. 4: 125. 1790; 258.
A shrub or small tree with conical tomentose prickles on the
branches only; leaves elliptic-lanceolate, acute both ends, to 1.5 dm.
long, about 5 cm. wide, white-tomentose beneath, early farinose but
soon green and glabrous above; flowers purple, in pedunculate ter-
minal or lateral tomentose corymbs; calyx-lobes ovate-cuneate, acute;
corolla about 1 cm. broad, segments 5 mm. long, lanceolate, acumi-
nate. An African species, introduced into Peru if actually col-
lected there; the record is one of those collections in Herb. Boissier
(Geneva) mislabeled simply Pavon, many of which are known to be
species endemic to Mexico.
Peru(?): (Ruiz & Pawn).
250 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solanum grandiflorum R. & P. Fl. Peruv. 2: 35, pi. 168, fig. b.
1799; 338. S. astroites Jacq. f. Eclog. 97, pi 65. 1811-1816?; 346.
Becoming a small tree with erect stout trunk, the branches abun-
dantly foliose; petioles more or less prickly, to 6 cm. long, more or
less fulvescent stellate tomentose as the leaves beneath, peduncles
and few-flowered racemes, including the calyces and corolla without;
leaves broadly ovate-elliptic, and somewhat sinuately repand or lobu-
late, strongly unequal at base, acute, often 2-3 dm. long and half as
wide, the lower probably larger, in type sparsely (to densely) hispid-
ulous above with simple trichomes, the 6-8 primary nerves prominent
beneath as also the secondary; racemes 5 cm. long or longer, the
flowers cymose, pedicels stout, 6-12 mm. long (2 cm. in fruit) ; flower
bud ovate-acuminate, 2 cm. long, 1 cm. broad; calyx deeply parted,
the inordinately stiped-stellate (type) to subsessile or scurfy sessile-
stellate (var.) acute or subobtuse segments accrescent after anthesis;
corolla blue or white, to 7 cm. across, acutely plicate in the angles,
stellate, glabrous or nearly within but with a lanate oblong-lanceo-
late zone without about 1 cm. wide; stamens 10-12 mm. long, anthers
connivent at base, violet, with 2 minute apical pores; fruit lutescent,
pendent, about 5 cm. in diameter, the subtending 5-parted calyx
subcampanulate. It seems doubtful that the type actually had lan-
ate fruit (as described but not as to plate) but the Ruiz and Pavon
specimens at Berlin had a young fruit sparsely stellate, quite pos-
sibly glabrous at maturity. Corolla rich purple-blue, aging lavender-
blue; fruit to 4.5 cm. in diameter (Stork & Horton). The most
striking feature, apart from the size of the few flowers, is the radiate
pubescent zone on the corolla segments without; rather distinctive
is a less pubescent shrub which may be noted as var. aberrans
Macbr., var. nov., foliis supra glabris; pilis plerumque sessilibus vel
breviter stipitatis plus minusve cinereis (type, Soukup 2545}. The
plant of Jacquin f. described as an "annual from Chile and Peru" is
possibly S. grandiflorum. F.M. Neg. 2607.
Huanuco: Pampayacu, Poeppig 1243. Cuchero, Poeppig 1422.
Tingo Maria, Soukup 2279; Stork & Horton 9461 (det. Standley);
Allard 21156; 21626 (det. Lyman Smith). Pozuzo, 4654; Ruiz &
Pavdn (type, and type var.). Pasco: Oxapampa, Soukup 2545; 3344
(var.). Junin: La Merced, Soukup 2525 (var.). Puerto Yessup,
Killip & Smith 26362; 26396; 26398 (all det. Morton) . Loreto :
Pucallpa, Soukup 3035. "Mite," "balsa" (Soukup).
Solanum heterophyllum Lam. Illus. 2360. 1793; 215. S. asper-
rimum Bitt. & Moritz, Repert. Sp. Nov. 16: 393. 1919?
FLORA OF PERU 251
Branches often purple, as the stems often somewhat glandular,
toward the apex stellate-pilose and aculeate, the broad-based prickles
somewhat recurved at tip, the upper 4-6 mm. long; petioles about
0.5-1.5 cm. long, the leaves opposite or ternate, more or less repand
or sinuate, unequal, the larger sometimes subelliptic, all often sub-
oblong, oblique at acute base, acute at apex, to 1 dm. long, the
smaller two shortly petioled or subsessile to about 7.5 cm. and to
less than 4 cm. long, these often somewhat obovate, subobtuse or
even emarginate, all stellulate scabrous above, stellate tomentulose
beneath, the larger sometimes sparsely aculeate with subulate straight
yellowish prickles on both sides except sometimes the purple mid-
nerve; primary nerves 2-4; cymes 2.5-5 cm. long, not or sparsely
prickly, stellate-tomentose including the corollas without; calyx sub-
scarious, poculiform, 3-4 mm. across, with 5 mucronate subtruncate
or subulate teeth; corolla 2- nearly 3 cm. broad, the segments (10)
12-14 mm. long, oblong-linear, subacute, obscurely pilosulous with-
in; stamens equal or subequal, 8-12 mm. long, the very short fila-
ments affixed at tube apex, the slender linear equal to unequal anthers
orange with a purple dorsal line, dehiscing by 2 apical pores; ovary
hirsute as the straight style at base, this in fertile flowers exceeding
the stamens, clavate at apex; fruit globose, yellow (Mexia), 6-8 mm.
thick, the incrassate pedicels reflexed. This is S.heterophyllum, prob-
ably a form of S. subinerme Jacq. similar to S. juripeba Rich, but
differing in the ternately diverse leaves; strongly affine S. Hum-
boldtii Dunal, 216, of Venezuela but the prickles larger, often straight,
the racemes and calyx not spinose (Dunal) ; these are doubtfully ge-
netic. The plant of Richard from the Surinam apparently varies in
degree of prickles, these sometimes even absent, and in the size of
flowers, at least according to Dunal and others. The Bitt. & Moritz
plant from Venezuela, at least as to Peru, seems to be a part of the
Peruvian species, which possibly is highly variable; the authers indi-
cated no distinguishing characters. S. myrianthum Britton ex Rusby,
Bull. Torrey Club 26: 191. 1899, type from lower Rio Madre Dios,
fide author is allied and to be expected within adjacent Peru, but
cymes 2-3-forked, corolla lobes only 8 mm. long, anthers yellow,
distinct. Mexia noted her plant was scandent in tangle, the caustic
seeds used on skin spots. May be several species concerned but prob-
ably all variants of one; anthers may be equal to unequal and some-
times sublinear.
San Martin: San Roque, Williams 7332; 7678. Jepelacio, Klug
31+07. Huanuco: Divisoria, Swingle 119; 120. Near Tingo Maria,
252 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Swingle 8. Muna, 3940. Junin : Chanchamayo Valley, Schunke 136;
264; 1761. La Merced, 5710. Loreto: Boqueron, Swingle 100 (det.
Morton). Itaya, Mexia 6485 (det. Morton, S. asperrimum) ; Williams
3448. Hacienda Sinchoa, Swingle 116. French Guiana. "Ayoc
mullaca" (Mexia).
Solatium hispidum Pers. Syn. 1: 228. 1815; 275. S. stellatum
R. & P. Fl. Peruv. 2: 40, pi. 176, fig. b. 1799, not Jacq.; 275.
A shrub or becoming a small tree sometimes sparsely prickly on
the flowering branches, these only early rusty pubescent with long
stipitate and sessile or subsessile stellate trichomes, soon glabrate,
the same indument densely developed on the lateral and terminal
dichotomous corymbs (to 1.5 dm. wide, about 1 dm. long) including
the white flowers; prickles sometimes many, on lower ligneous stems
not or little enlarged, straight or nearly, 2-3 mm. long, broad at base;
petioles 2-2.5 cm. long; leaves broadly ovate-elliptic, more or less
oblique at the rounded-cordate base, acuminate, subentire to un-
evenly repand or 7-11-lobulate, sparsely (except undeveloped young-
est) green both sides with only scattered or few sessile or shortly
stiped stellulate trichomes, these in age asperous, especially on the
upper surface; peduncles about 2 cm. long, branches to 3 cm. long,
pedicels crowded, 5 mm. long (7 mm. in fruit); calyx lobes lanate-
stellate in type, usually ovate, acuminate, about 6 mm. long; corolla
stellate, about 1 (-1.5) cm. long, narrowed ovate lobes lanuginose
stellate without, medially puberulent within; anthers violet, 5.5-
6 mm. long; fruits globose, yellowish, about 1.5 cm. in diameter.
The description of Dunal included S. Poeppigianum Sendt., collected
by Ruiz & Pavon at Muna. An exteme, perhaps distinct, is Pennell
14066, recordable as var. Cerrateae Macbr., var. nov., ramis fol-
liisque sparse sed longe stellato-hispidis. Equally aberrant, suggest-
ing in pubescence the unrelated S. selachophyllum Bitter but the
broad leaves not bullate, may be noted var. Ferreyrae Macbr., var.
nov., floribus racemosis circa 1.5 cm. latis; antheris flavis, 12 mm.
longis. Klug 3203, type (a 6-meter tree in old clearing, corolla dark
violet, anthers yellow) ; also apparently, Williams 2606, 2956, 2981 ,
all from La Victoria on the Amazon, branchlets glabrate; this may
well be an Amazonian species. F.M. Neg. 2609.
Huanuco: Panao and Pillao, Ruiz & Pav6n, type. Southeast of
Huanuco (near Panao), 2106 (det. Macbride, S. asperolanatum) .
Mito, 1485. Near Acomayo, Mexia 04118. Carpish, Stork & Hor-
ton 9886 (det. Standley, S. saponaceum). Cuzco: Near Rio Yana-
FLORA OF PERU 253
mayo, Pennell 14066 (type, var.). "Huircasan" or "campucasa"
(Ruiz & Pavon), "huachulla."
Solanum incarceratum R. & P. Fl. Peruv. 2: 40, pi. 176, fig. a.
1799; 249.
A suffrutescent or herbaceous plant 1-2 meters high, pilose even
to the flowers with simple trichomes and aculeate on the stems and
leaf -nerves, the prickles in part fine, in part stout; petioles to 2.5 cm.
long; leaves repand-angulate, cordate, to 1.5 dm. long, nearly as wide;
cymes lateral, to 5 cm. long, not prickly, the peduncles 4-6 mm. long;
pedicels in flower 1-1.5 cm. long, in fruit 2 cm. long or slightly longer,
strongly incrassate at apex; calyx cyathiform, the segments narrowly
linear, arcuate-recurved, in anthesis only about 5 mm. long, becom-
ing 1.5 cm. long and erect about the berry; corolla white or nearly,
perhaps about 2 cm. across, the ovate-lanceolate segments acutely
acuminate, spreading, finally revolute at tip, 8-10 mm. long; sta-
mens 5, equal, two-thirds as long as the corolla, the anthers ovate-
oblong, shortly attenuate, subconnivent; ovary glabrate, the filiform
glabrous style longer than the stamens, straight to the didymous
stigma; fruit white, to 2 cm. long, 1.5 cm. broad, glabrous, enclosed
in calyx lobes. Illustrated, Mart. Fl. Bras, pi 5, figs. 17-20. F.M.
Neg. 23120.
Huanuco: Yanano, 3803. Chinchao, Ruiz & Pavdn, type. Cas-
sapi, Poeppig 1659 (det. Bitter). Cuchero, Poeppig. Junin: Hua-
capistana, Killip & Smith 24102 (det. Morton). Ayacucho: Aina,
Killip & Smith 22699 (det. Morton). Bolivia; Brazil.
Solanum jamaicense Miller, Card. Diet. 8, no. 17. 1768; or
Swartz, Fl. Occ. Ind. 1: 154. 1797; 199.
A sparsely prickly shrub with geminate unequal angulate-lobed
broadly ovate cuneate-based leaves 1-1.5 dm. long, 6-10 cm. wide,
and subsessile umbel-like lateral racemes (to 2 cm. long in fruit) of
small white flowers; pubescence moderately dense, shortly stipitate-
stellate, somewhat whitening the leaves beneath; pedicels in fruit to
1 cm. long; calyx segments filiform, caudate in fruit, 6-7 mm. long,
usually with 1 or 2 prickles; berries 6-7 mm. in diameter, orange-red.
Determinations mostly by Standley; a common tropical weed.
Loreto: Iquitos, Ule 6241 (det. Bitter); King 335; 1204 (det.
Morton); Killip & Smith 26887; 26978; Williams 1345; 1482; 1484;
7978. Caballo-cocha, Williams 2180. La Victoria, Williams 2599.
Rio Itaya, Williams 129; 3225. To West Indies. "Coronilla."
254 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solatium kioniotrichum Bitter, in herb. S. grandiflorum R. &
P. var. leiocarpum Dunal in DC. Prodr. 13, pt. 1: 339. 1852.
Ut videtur ad 15 m. altum; foliis supra breviter stipitato-stellu-
latis vel demum aspero-hispidulis; junioribus plus minusve stipitato-
setosis, lobis breviter vel longe caudatis. A shrub or small tree,
sometimes coarsely spiny, the inordinately hispid-stellate branchlets
and inflorescence more often sparsely, the bristle-like trichomes 3-
4 mm. long, stellately branched at tip; leaves (upper) repand-mar-
gined, to few-lobulate, broadly elliptic, subsessile, the rounded base
oblique, subabruptly acuminate, about 2 dm. long and 1-1.5 dm.
broad, scabrous-hispid above with simple trichomes or these infre-
quently forked at tip, shortly grayish stellate pubescent beneath;
inflorescence lateral, the several-flowered raceme early crowded but
elongating as the mostly sterile flowers fall; pedicels in fruit to 8 mm.
long; calyx-lobes ovate-lanceolate, caudate-tipped, 1.5 cm. long or
longer including the 3-4 mm. tip, glabrous except for the bristle-
like sparse to very dense stiped stellate indument, the trichomes
typically 5-8 mm. long; corolla deep purple or white, the lobes ovate,
acute, 2 cm. long, medially stellate-tomentose without, the corolla
about 5 cm. broad; anthers 1.5 cm. long; berry subglobose, 4-5 cm.
in diameter. F.M. Negs. 2614; 33370 (var.).
San Martin: Zepelacio, Klug 3735. Junin: Puerto Yessup, Kil-
lip & Smith 26396. Chanchamayo Valley, Schunke 308. Above San
Ramon, Killip & Smith 24351; Schunke 612 (type, var.). Loreto:
Iquitos or near, Ule 6243, type; Killip & Smith 26905; Williams 1389;
1543; 3526; Mexia 6461 (type, var.). Balsapuerto, Klug 3071. Rio
Mazan, Jose Schunke 13 (det. Standley). Yurimaguas, Williams
4018; Poeppig 2223. Rio Nanay, Williams 296; 435. Pebas, Wil-
liams 1888. La Victoria, 2584- Brazil. "Siucahuito" (Schunke) ;
"simca-huito" (Quechua, Williams); "cocona."
Solatium leucopogon Huber, Bol. Mus. Goeldi 4: 604. 1906.
Shrub or clambering, the long branches (as leaf mid-nerve be-
neath) aculeate with short compressed recurved prickles and hispid
with long (4 mm.) spreading trichomes; leaves suboblong, long-atten-
uate to base, acutely acuminate, to 1.5 dm. long, 3.5-5 cm. wide,
subentire to repand-lobulate, typically membranous, sparsely hispid,
the trichomes on upper surface of leaves in part or both sides stellu-
late at least at base; flower clusters lateral or opposite leaves, sub-
sessile or peduncles 1 cm. long in fruit, pedicels 2 cm. long; calyx
lobes ovate, finely acuminate, 1 cm. long, in fruit 1.5 cm. long; corolla
FLORA OF PERU 255
about 1.5 cm. long, inordinately villous-pilose; fruits lustrous, yellow,
3 cm. in diameter. Allied to S. enoplocalyx Dunal but trichomes
rarely sessile, flowers 5-10, white-villous without, and calyx prickly
and hispid with long trichomes that are stellate at base. If Dunal's
plant is Mexican this may be distinct as a southern relative. Mexia
6335 (in fruit) has finely stellulate puberulent leaves both sides; Wil-
liams 1093 has firm stellulate scabrous repand leaves; there may be
an earlier name for this, probably extra-Peruvian, perhaps S. scabrum
Vahl, 216, to which the Williams and Mexia specimens could be re-
ferred except for the hispid branches, and they may be a variant of
that West Indian species. F.M. Neg. (Photo by Dahlgren).
Loreto: Canchahuaya, Huber 1370, type. Iquitos, Williams 8054-
La Victoria, Williams 2652. Leticia, Williams 3148. Lower Rio
Nanay, Williams 517. Pongo de Manseriche, Mexia 6335. Man-
finfa, Rio Nanay, Williams 1093. Brazil. "Cubunuchi," "intutu-
quiro" (Williams).
Solatium mammosum L. Sp. PI. 187. 1753; 250.
A bushy subherbaceous plant a meter or so high, densely villous
all over and armed on the stems and leaf-nerves with rather stout
yellow prickles, many 1.5-2 cm. long; leaves roundish, 3-4 (7) -lobed
or repand-dentate, sometimes 2 dm. long or longer, nearly as wide;
flowers subsessile, 1-few, pink-purple; calyx lobes long- villous with
articulated trichomes, not aculeate, linear, 4 mm. long; corolla to
3 cm. broad, the oblong-linear acuminate long-villous segments gla-
brous within, ciliate, 1-2.5 cm. long, 4 mm. wide; stamens less than
half as long as corolla, filaments 1.5 mm. long, anthers rounded at
base, conic-attenuate; ovary glandular punctate, glabrous, tapering
into the thick style, the stigma bilobed; fruits large, yellow with
white pulp, 3-6 mm. long, and purple-red punctate seeds. S. globi-
ferum Dunal, 251, known widely in cultivation, perhaps originally
from Martinique, has narrowly lanceolate glandular basally aculeate
calyx lobes about 5 mm. long, corolla 9-10 mm. long, fruits subglo-
bose; S. aculeatissimum Jacq., 244, a West Indies species, possibly
introduced, is similar but the ovate-acuminate calyx teeth are un-
equal, about 1.5 mm. long, aculeate; berry globose, 2-2.5 cm.
Cultivated at Hacienda Pampayacu for the poisonous fruits which
were used to exterminate rats. The seeds had been obtained from
plants said to be "wild" at Tingo Maria, Huanuco (Raimondi, det.
Werdermann). In the Ruiz and Pa von Herbarium from Guayaquil
(det. Werdermann) under name "Veneno."
256 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Huanuco: Pampayacu, Rio Chinchao, 5066. Loreto: Iquitos,
Raimondi; Williams 1483; 7977. La Victoria, Williams 2887. Con-
tamana, Tessmann 3094 (det. Werdermann) . Ayacucho: Kimpi-
tiriki, Killip & Smith 22915 (det. Morton). Northern South Amer-
ica and the West Indies. "Tintoma," "roconilla dulce," "veneno."
Solatium melongena L. Sp. PI. 186. 1753. S. esculentum Dunal,
Hist. Sol. 208, pi. 3. 1813; 355.
Annual or becoming a shrub, the stems aculeate or smooth; peti-
oles 3-3.5 (-13) cm. long; leaves weakly or not aculeate, subcordate
or not, entire or lobed, grayish rough or soft stellate, 10-16 cm. long,
8-11.5 cm. wide, 5 lateral nerves; inflorescence 4-5 cm. long, 3-6-
flowered, pedicels 2-3 cm. long (lowest) ; calyx (fertile) 1-1.5 cm. long,
with 6-7 (-8) unequal lanceolate stellulate lobes, 7-8 mm. long, 3 mm.
wide; corolla blue or violet (exceptionally white), campanulate, stel-
lulate, 2-3.5 cm. long, 3.5-5 cm. across; filaments glabrous, 1-2 (-4)
mm. long, the anthers 6.5-7 X 1-1.5 mm. thick, style 8-9 mm. long,
two-thirds of its length stellate; pedicels in fruit 5-7 mm. long, the
somewhat prickly berry sordid, white-yellow, 3-9 cm. long, 2.5-5 cm.
broad or larger, the subtending calyx to 7 cm. across; the seeds reni-
form, 3.5 X 2.5 mm. After Bitter; spineless or spiny in cultivation.
This vigorous annual of Asia is probably grown in Peru as in most
warm countries for the flowers and the curious ornamental but not
edible egg-shaped fruits.
Peru (probably in gardens and as an occasional escape). Asia.
Solatium nemorense Dunal, Hist. Sol. 194. 1813; 219. S. lacini-
atum R. & P. Fl. Peruv. 2: 40, pi 177 f fig. a. 1799, not Ait., 1789.
An entirely glabrous but prickly-stemmed more or less scandent
shrub with very long (to 3 dm.) racemes of smooth blue or white
flowers; leaves mostly geminate, unequal, sessile or nearly, the larger
rarely entire, ordinarily deeply angled or acuminately and pinnately
long-lobed, the opposite smaller one to about 1 dm. long, 4-4.5 cm.
wide (three-fourths the size of the larger), entire or repand, all un-
armed above, beneath aculeate on the midnerve; racemes cymose,
nearly opposite the leaves, solitary, minutely spinulose, mostly a dm.
long, some much longer, pendent in fruit, the filiform rachis cicatrose
from the fallen sterile flowers, pedicels 4-6 mm. long; calyx 4 mm.
across, subcyathiform, rather retusely 5-crenate; corolla white (Mexia)
or blue(?), 12-14 mm. broad, the ovate-lanceolate acuminate spread-
ing lobes 8 or 9 mm. long; stamens 8 mm. long, anthers white, basally
FLORA OF PERU 257
ovate, long-acuminate, equaling the style; ovary glabrous; stigma
suborbicular, concave; berry orange, subglobose, subumbonate at
base, in the enlarged calyx 3 cm. in diameter; seeds yellow. The
large yellow-orange berries are edible. The leaves are variable; many
are merely lobed but the rachis of the pinnate ones is broadly winged.
F.M. Neg. 2615.
Huanuco: Venenillo, 30 km. north of Tingo Maria, Ferreyra 4351
(det. Morton). Cayumba, Mexia 8304A (det. Morton). Chinchao
and Cuchero, Ruiz & Pav6n, type; Poeppig. Junin: Pichis Trail,
Killip & Smith 25531 (det. Morton). Loreto: Balsapuerto, Killip &
Smith 28387. Santa Rosa, Killip & Smith 28710. Yurimaguas,
Williams 5219; 4837; Killip & Smith 27618.
Solatium pectinatum Dunal in DC. Prodr. 13, pt. 1: 250. 1852.
Ex char, may be (fide Dunal) similar to S. mammosum L. in spines
(6-12 mm. long), size and shape of leaves but the pubescence harsher
(hirsute), the angulate-cordate leaves to about 10-angulate-repand or
-sinuate, and the very hirsute flowers subsessile, subcymose or glom-
erate; petioles (as aculeate branches, aculei 6-12 mm. long) densely
hirsute, the strong aculei 3-4 mm. long; corolla unknown; calyx after
an thesis 12 mm. long and wide, the ovate-lanceolate acutely acumi-
nate lobes membranous, glabrate within, 6 mm. long; ovary obtuse,
densely hirsute; fruit unknown. Not seen since first draft of this
account; since apparently it has not been collected again in the much
collected area the label may be an error or quite possibly the speci-
mens cited under S. mammosum L., at least in part, belong here.
Loreto: Yurimaguas, Poeppig 2224, type.
Solatium Poeppigianum Sendt. in Mart. Fl. Bras. 10: 103.
1846; 231.
Shrub with flexuose or scandent branches densely to sparsely red-
dish stellate-tomentose and sparsely aculeate; prickles short, recurved;
petioles about 2 cm. long; leaves ovate, elliptic, usually entire, rarely
lobed, around 12 cm. long and 8-10 cm. broad, green, typically ap-
pressed pubescent above (indument long simple trichomes or tri-
chomes rarely 1-2-branched at base) or becoming glabrate, densely
stellate-tomentose beneath and often remotely aculeate on the mid-
nerve; inflorescence reddish stellate-tomentose; cymes in type ter-
minal, about 2.5 cm. long, peduncle half as long; pedicels 2-6 mm.
long, stellate-lanuginose (not prickly) as also the calyx and corolla
without; calyx poculiform, 10-12 mm. broad, the ovate-lanceolate
258 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
acute segments unequal, marginally hirsute, glabrous within, to 1 cm.
long, 2-4 mm. wide; corolla 5-parted, segments oblong-linear, sub-
obtuse, glabrous within, reticulate-veined, (8) 12-14 mm. long; sta-
mens equal, scarcely shorter than corolla, inserted at base of tube,
the anthers subconic, attenuate and sublinear above, 8-10 mm. long;
ovary very hirsute with long erect yellowish trichomes; style 1 cm.
long, sulcate, glabrous and slightly recurved toward the bifid stigma;
younger fruit hirsute, subtended by the persisting calyx. The type
has mostly stipitate-stellate trichomes, the trichome branches long
and weak while the trichomes on leaves above are mostly simple or
furcate; also, they are long and weak or sessile and more or less stel-
late at the very base; sometimes stellate, usually stiped trichomes
dominate the upper side of the ovate-lanceolate leaves (entire or
lobed), and the subcampanulate corollas are larger, var. Vargasii
Macbr., var. nov., pilis plerumque stipitato-stellatis; corollis 2 cm.
latis. Perhaps the species includes other varieties, more likely indi-
vidual variants, especially in pubescence; type young, corolla noted
by Poeppig as green, by me as greenish-yellow; these specimens have
petals only 8 mm. long, stamens 5 mm. long. Perhaps the name
should be restricted to this plant, using another name (by Bitter, in
herbarium) for the rest. Herrera 1799 (not seen recently) has forked
racemes several cm. long, the flowers few by abortion, so may be
S. velleum Swartz; size of flowers may be a more basic character;
Cook & Gilbert 1273, Soukup 538, and Seibert 2005 have corollas
nearly as large as those of var. Vargasii Macbr. but pubescence in
part, especially on the leaves above, is simple or mostly simple. For
convenience this seemingly intermediate plant may be named var.
medians Macbr., var. nov., corollae valde partitae, laciniis ovato-
lanceolatis, 12-15 mm. longis (Soukup 538, type). Solution of the
problem awaits determination of the genetically salient characters.
F.M. Negs. 6822; 2596 (Weberbauer 4534).
Cajamarca: Monte Seco, Prov. Hualgayoc, 3883. Amazonas:
Moyobamba, Weberbauer 4534. San Martin: Tarapoto, Ule 31 in
part (new species in herb, by Bitter). Jepelacio, Klug 3284 (det.
Standley). Ancash: Marcara, Velarde Nunez 3228. Lima: Rio Ri-
mac, Soukup 3557; 1047 (toward S. hispidum in leaves). Huanuco:
Muna, 4083 (det. Standley) ; 4909; Chavanillo, Rio Maranon, 2305.
Junin: La Oroya, Kalenborn 147. Chongos Bajo, Soukup 3617.
Palca, Velarde Nunez 619. Parco, near Huancayo, West 3631 (type,
var. Vargasii). Near Tarma, Constance & Tovar 2212 (var. Vargasii) ;
Isern 2533; Killip & Smith 21787. Loreto: Yurimaguas, Poeppig
FLORA OF PERU 259
1231 (2131 in Fl. Bras, type). Huancavelica: Ancco, Soukup 4017.
Pampas, Stork & Horton 10257 (det. Standley). Ayacucho: Near
Huanta, Killip & Smith 23313. Cuzco: Machupicchu, Hen era 3233;
3238. Lares Valley, Herrera 1799. Prov. Cuzco, near Cuzco, Var-
gas 217. Rio Churumayo, Soukup 538. Lucumayo Valley, Cook &
Gilbert 1273 Madre de Dios: Maldonado, Seibert 2005. "Urmish"
(West).
Solanum quinquangulare Willd. ex Roem. & Schult. Syst. 4:
669. 1819; 239.
Vigorous, the branchlets densely prickly with somewhat refracted
very unequal slender subulate spines, the longest 6 mm. long (in Peru
some 2 cm.) and in typical form also long-hirsute with simple pellucid
trichomes as the petioles (to 2 cm. long) and peduncles (8-12 mm.
long) ; leaves smooth, at the subcuneate or subtruncate base, trian-
gular, appressed hirsute, ciliate and aculeate both sides, 1-1.5 (2) dm.
long, 8-10 cm. wide, the acutely acuminate 3-5 lateral lobes divari-
cate, the intermediate lobe largest; cymes to 2.5 cm. long, the 4 or
5 pedicels filiform, 1 cm. long; corolla 8 mm. across, 5-parted, the
narrowly oblong acute segments 8 mm. long; calyx 4 mm. broad,
accrescent in fruit, the ovate lobes not aculeate; stamens two-thirds
as long as corolla, filaments capillaceous, 2 mm. long, anthers ovate-
lanceolate, ventricose at base, attenuate, 2-pored, about 6 mm. long;
ovary glabrous, style filiform, 5-6 mm. long. The Peruvian plant
is sparsely hirsute, with much longer prickles and subcordate leaves
and so it resembles S. ciliatum Lam., 241; otherwise it is more like
the plant of Willdenow; may have been introduced.
Cuzco: Along railroad, Lucumayo, Vargas 2788 (det. Killip).
Maquina, in riverside brush, West 8030. Venezuela; Colombia.
Solanum quitoense Lam. Illus. 2: 16. 1797; 352. S. angulatum
R. & P. Fl. Peruv. 2: 36, pi. 170, fig. a. 1799.
A very pilose-stemmed, often unarmed 1-2-meter shrub, the up-
per branches, leaves on both sides and racemes including the corollas
in type densely and softly lanuginous; petioles 5-10 cm. long; leaves
ample, the lower 4.5 dm. long and about two-thirds as wide, the upper
geminate, subequal, 11-14-sinuate-lobate, mostly 1.5 dm. long, mid-
nerve and the 5 lateral nerves obvious both sides but scarcely promi-
nent; racemes 4-5-flowered, peduncles 6 mm. long, pedicels crowded,
1-1.5 cm. long, apically incrassate; calyx campanulate, 14-16 mm.
across, the ovate-lanceolate segments acute; corolla about 2 cm.
260 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
broad, the oblong linear segments 6 mm. wide, subobtuse; anthers
yellow-orange, ventricose at base, 3-4 mm. long, the 2 terminal pores
minute; style filiform, scarcely longer, little clavate toward the oblique
stigma; ovary subglobose, hirsute; fruit globose in size, the color and
taste resembling that of citrus, the younger lanuginose, at maturity
glabrous, lustrous, fragrant. According to Feuill^e, Arzn. Peru 2:
85, pi. 46- 1766, the flowers are white within, violet without. Sup-
posedly native to Ecuador, the fruit is called "naranjitas de Quito."
This is the var. septentrionale R. E. Schultes & Cuatr., Bot. Mus.
Leaflets, Harvard 16: 100. 1953, devoid of spines and spinules, not
known from Peru. Illustrated (the variety) photo of flower and fruit
by Cuatrecasas, pi. 17; Schultes, I.e. 14. pi. 10 (plant). Ruiz and
Pavon noted that the fragrance and flavor of the fruits contributed
to the pleasure of the mate tea. Schultes and Cuatrecasas quoted
from notes of J. V. Sigvold Muller on a specimen at Kew, in part:
a climber, the bright orange fruit to 5 cm. in diameter, pulp bright
green, very juicy and aromatic; the pulp eaten with a little sugar is
exceedingly pleasant and cooling; with seeds strained out it gives an
excellent flavor to ices. The authors of the variety give a bibliog-
raphy that includes a good account by Hodge, Journ. N. Y. Bot.
Gard. 48: 155-157. 1947.
Amazonas: Chachapoyas, (Mathews). Lima: Especially in gar-
dens, (Feuille"e) ; Ruiz & Pavon; Nee. Junin : Pichis Trail, Killip &
Smith 25783? Graves less pubescent). Loreto: Cultivated at Yuri-
maguas, Poeppig 2222. Ecuador; Colombia; Venezuela. "Lulo."
Solatium rhamphidacanthum Bitter, Repert. Sp. Nov. 12:
436. 1913.
A 1-5-meter shrub apparently supported by the hooked com-
pressed prickles (to 4 mm. long) on branches, petioles (3-5 cm. long)
and leaves, especially on the midnerve beneath; pubescence, at least
on the branches, consisting of long (4-5 mm.) red trichomes and some
minute stellate ones, that of the leaves and inflorescence in part
glandular; branches 4-5 mm. thick, internodes often 5-7 cm. long;
leaves ovate-cordate, long-acuminate, about 1-1.5 dm. long, 8-12 cm.
wide, the pubescence above mostly on the 7 or so lateral nerves and
veins; peduncles extra-axillary, slender, 3 cm. long, the many (12)
pedicels about 8 mm. long, subsecund; calyx as corolla stellate, the
lanceolate acute or acuminate lobes 7-8 mm. long; corolla bright
yellow-green, the subobtuse stellulate and glandular lobes 12 (-14)
mm. long; stamens equal, filaments glabrous, anthers long-lanceolate,
FLORA OF PERU 261
subcordate at base, attenuate, 1 cm. long; ovary trichomes very
unequal, mostly glandular, style incurved, 12 mm. long, pubescent
except above with simple glandular or furcate trichomes, the stigma
incrassate. Probably occurs in adjacent Peru. F.M. Neg. 2635.
Rio Acre: Seringal Auristella, Ule 9763, type.
Solatium saponaceum Dunal, Hist. Sol. 206. 1813; 274. S.
scabrum R. & P. Fl. Peruv. 2: 39, pi 175. 1799, not Vahl, 1796.
A shrub or small openly branched tree, sometimes 4 meters high,
with smooth to more or less sparsely aculeate stems and branches;
petioles 2-4 cm. long; leaves ordinarily alternate, ovate-oblong, sin-
uate- or angulate-lobed, rarely deeply, notably venose, often 1.5-
2 dm. long, 1-1.5 dm. wide, sometimes much smaller, unequal at
base, acute, usually green and soon glabrous but typically stellulate
scabrous on both sides, especially above; corymbs terminal and lat-
eral, dichotomous, sometimes a dm. wide, often much smaller, some-
what glandular puberulent, the trichomes mostly or all simple on
the ascending-erect pedicels, these incrassate above, early 8-14 mm.
long, to 2 cm. long in fruit; calyx cyathiform, 4-6 mm. across, the
(early) narrowly ovate-lanceolate acutely acuminate lobes 3-5 mm.
long, in fruit little longer but ovate at base; corolla blue or white, to
about 1.5 cm. broad, 1.5-2 cm. long, deeply divided, the ovate-lanceo-
late apiculate segments at least 6 mm. long, 4-7 mm. wide, finely
stellulate without; stamens 8 mm. long, slightly unequal, the some-
what connivent anthers oblong-ovate below, narrowed to the minute
terminal pores; style longer than stamens; fruit orange, 2-celled,
12 mm. thick. Sometimes the petioles and leaves beneath are
prickly and the latter also evanescently grayish-white pubescent as
in the Raimondi collection from Piura. Dunal drew his description
from Ruiz and Pavon and their plate. The stamens are not equal
and usually exceed 6 mm. ; flowers of plants in central Peru are often
somewhat smaller than those from Cuzco but not always; the leaves
are variable in size, sometimes glabrous and more or less lobed in
both regions, rarely almost binnate (my 5291}; inflorescences are
often reduced but the rachis is ultimately racemose, sometimes nearly
1 dm. long, usually shorter even with mature fruit. It is still not
certain that there is only one variable entity but it seems so to me.
Probably an earlier name from one of the extra-Peruvian similar
species will be shown to apply; Bitter referred Spruce 1+615 from
Tarapoto to S. caricaefolium Rusby, Bull. N. Y. Bot. Gard. 8: 118.
1912, with the remark, "spines more numerous, longer." The fruit,
like that of S. albidum Dunal, is saponaceous.
262 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Piura: Chalaco, Raimondi (det. Werdermann, affine). Caja-
marca: Cutervo, Raimondi (det. Werdermann). Cajabamba, Fer-
reyra 3057. Tambillo, Jelski 57 (det. Zahlbruckner). San Martin:
Moyobamba, Woytkowski 35304 (det. Cuatrecasas, S. caerulescens
Benth.). San Roque, Williams 7263. Tarapoto, Williams 5509.
Rioja, Woytkowski 5. Huanuco: Yanano, 3731. Muna and Chin-
chao, Ruiz & Pavdn, type. Pozuzo, 4635. Cushi, 4823. Junin: La
Merced, 5257; 5291; Killip & Smith 23765; 23767; Soukup 2544;
2543; 2548. Huacapistana, Killip & Smith 24100; 24291; Velarde
Nunez 3748 (det. Morton) ; Weberbauer 2036. Palca, Soukup 3472.
Chanchamayo, Schunke 543; 544- Rio Perene", Killip & Smith 24986;
25066; 25203. Rio Pinedo, Killip & Smith 23571 . Carpapata, Kil-
lip & Smith 24365 (det. Morton) ; 3468. Ferreyra 3640 (det. Mor-
ton). Tarma, Velarde Nunez 748. San Ramon, Constance & Tovar
2220. Loreto: Rio Nanay, Williams 1131. Iquitos, Williams 1308.
Ayacucho: Aina, Killip & Smith 22556. Ccarrapa, Killip & Smith
23224- Apurimac: Abancay, Vargas 461 (det. Standley, S. nitidum) ;
462Cuzco: San Miguel, Urubamba Valley, Cook & Gilbert 1170
(det. Morton) ; Herrera 2023. Quellomayo, Prov. Convention, West
8015. Machupicchu, Herrera 3204 (det. Standley); 3276 (det. Mor-
ton). La Maquina, West 8029. Madre de Dios: Maldonado, Sei-
bert 2004 (det. A. C. Smith, vel aff.). Bolivia? "Casiamuru," "tulu-
luque," "ama mullaca" (Williams).
Solanum sisymbrifolium Lam. Illus. 2: 25. 1797; 326.
The only (cf. S. concisum) Peruvian species prickly and viscid-
villous at least in inflorescence and with pinnatifid or bipinnatifid
leaves, these sometimes glabrate, the largest 3 dm. long, segments
1 dm. long, petioles 1 dm. long; a weedy plant, even the calyx often
armed with yellow prickles, the lobes in fruit to 1.5 cm. long; berry
finally yellow, 12 mm. in diameter; flowers white. For an illustration
of the inflorescence and fruit see Bitter, Bot. Jahrb. 55, Beibl. 121:
pi. 2. 1917, showing the berry without markings or essentially.
Probably introduced. In Bolivia called "Alco-chilco;" in the Ruiz
and Pavon herbarium from Guayaquil labeled "Chancolet."
Piura: Ayavaca, (Bonpland). Cajamarca: Chola and Nancho,
Raimondi. Huambos, Soukup 4459 (det. R. S. Cowan). La Liber-
tad: Raimondi. San Martin: San Roque, Williams 7640. Bella-
vista, Ferreyra 4752. Loreto: Boca de Yarina, Tessmann 3490.
Iquitos, Killip & Smith 27095; Williams 4993; Klug 321; 957. Yuri-
maguas, Williams 4280. Cuzco: Echarate, Goodspeed Exped. 10469
FLORA OF PERU 263
(det. Standley). Santa Ana, Cook & Gilbert 1441,' 1488. Argentina
to southern United States. "Ocote mullaca," (Ferreyra), "misqui-
corrota" (Cook & Gilbert).
Solanum subinerme Jacq. Stirp. Amer. 50, pi. 40, fig. 3. 1760;
259. S. juripeba Rich. Act. Soc. Nat. Paris 1: 107. 1792; 214, fide
Urban.
Not surely specifically distinct from S. torvum Sw. or at least from
S. heterophyllum Lam.; branched, about 1 meter tall, lightly fari-
nose tomentose above, not aculeate or sparsely; leaves oblong-ovate
(broadly elliptic or ovate-elliptic in Peru), acuminate each end (in
Peru acute or rounded and usually obliquely at base), entire or 3-4-
repand-lobulate, glabrous above or the midnerve and 5-7 lateral
nerves somewhat puberulent, often 1-2 dm. long, half as wide, more
or less tomentulose or puberulent beneath; petioles often geminate,
the shorter 1-1.5 cm., the longer 3-4 cm. long, the smaller leaves
about a fourth as large as the opposite one; racemes 2.5-7 cm. long,
lateral or axillary, pedicels early crowded, recurved to spreading-
erect, finally to 2 cm. long, little incrassate; calyx lobes minute or
short, ovate, obtuse; corolla purple, parted to base or nearly, the
oblong lanceolate segments puberulent without, acuminate, 2-2.5 cm.
long; stamens about 1.5 cm. long, filaments 0.75 mm. long, as wide
as long, anthers 11-12 mm. long-attenuate; ovary pubescent, style
13-17 mm. long, stigma bilobed; berry globose, about 1 cm. thick,
yellowish, perhaps also dark in color. Very probably should include
S. heterophyllum Lam., its leaves varying from oval to oblong but
more usually geminate or ternate and pubescent above. The plant
of Richard (Guiana), to which specimens have been referred, has
glabrous ovary and ovate, mucronulate calyx lobes, characters prob-
ably not significant; many species exhibit variation in amount of
indument (to lack) on ovary and development of calyx lobes. Allard
collections det. Lyman Smith; others mostly det. Morton.
San Martin : Pongo de Cainarachi, Klug 2725. Toward Juanjui,
Ferreyra 4478; Arroyo Bravo, toward Pucallpa, Allard 20406.
Huanuco: Tingo Maria to Divisoria, Ferreyra 977; Allard 21305;
Woytkowski 34544, Swingle 8; 100; 120. Loreto: Prov. Coronel
Portillos, Ferreyra 4301. Toward Pucallpa, Ferreyra 8107. Man-
finfa, Williams 1131. Rio Nanay, Williams 1248. Colombia; West
Indies. "Chupusacha" (Ferreyra), "sinca-huito," "sinca-huitello"
(Williams).
264 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Solanum talarense Sven. Amer. Journ. Bot. 33: 485. 1946.
Suffruticose, the procumbent or suberect stem about half a meter
long, the few branches (these 5 mm. thick) yellow, densely aculeate
and pubescent; petioles to 3 cm. long, little aculeate; leaves entire,
ovate, subtruncate or unequally decurrent at base, acute, 3-8 cm.
long, 1-5 cm. wide, pubescent both sides with sessile stellate or simple
septate and glandular stipitate trichomes; peduncles terminal (?),
mostly bi-flowered (or obsolete and pedicels forked?) ; calyx deeply
5-lobed, aculeate; corolla greenish-yellow, 3 cm. across, deeply parted,
the 5 ovate lobes 3-4 mm. wide, 8-10 mm. long, acuminate, stellate-
pubescent without; filaments glabrous, 1-1.5 mm. long, hyaline-alate,
anthers linear, narrowed apically, the terminal chinks finally dehisc-
ing laterally; style filiform, 9 mm. long, glandular toward base, the
stigma little larger; young fruit glandular. Apparently resembles
most the purple-flowered S. crinitipes Dunal, 317, of Ecuador, but
entirely different in the membranous leaves and type of pubescence
(Svenson) ; the plate shows one flower on pedicel about 1 cm. long,
two flowers apparently from the same pedicel (or peduncle?); the
anthers are scarcely attenuate, but minutely bipored. Illustrated,
Svenson, 484, pi. 19, fig. 3.
Piura: Quebrada Mogollon, Amotape Hills, (Haught & Svenson
11534, type).
Solanum topiro HBK. Nov. Gen. & Sp. 3: 25. 1818; 353.
Type described by the collector as a much-branched unarmed
shrub 2 meters high, the stem tomentose, the flowers extra-axillary,
2-several clustered; branches, leaves beneath, petioles, these nearly
4 (-12) cm. long, and flowers without stellulate-tomentose, the
trichomes of the branches obviously stiped, shortly so or subsessile
on the leaves and flowers; leaves pubescent above, strongly oblique
at base, acutely angulate, subacute, to 3 dm. long, 2 dm. wide
(to 48 X 36 cm., Schultes), with membranous appressed pilose
simple or long-furcate trichomes above, reticulate-veined, the 7
primary nerves prominent beneath; pedicels about 5 mm. long;
calyx deeply 5-parted, the ovate acute equal segments little shorter
than the rotate corolla, this with very short tube, ovate-oblong
acute somewhat unequal segments 1.5-2 cm. long; stamens 5, shorter
than the corolla, subsessile, the erect oblong anthers connivent,
acuminate, yellow, glabrous, the 2 subrotund pores minute; ovary
ovoid-trigonous, hirsute; fruit finely tomentose, the indument cadu-
cous (Schultes), edible, the seeds sublenticular, 3-4 mm. long,
FLORA OF PERU 265
2-2.5 mm. wide (Schultes). Type by Bonpland from the con-
fluence of the Rios Orinoco, Atabaju and Quariares. After writing
the above an informative account by Schultes appeared, Bot. Mus.
Leaflets 18: 229-244. 1958, with three excellent photographs by
the author and a beautiful drawing by L. W. Smith. The author
observes that the plant apparently has never been collected except
about cultivated areas, often abandoned, and that the reddish ripe
fruit is eaten fresh or used to make a rather acidulous beverage.
Schultes quotes Fennell, For. Agric. 12: 181. 1948, ". . . . the cocona
is a permanent acquisition as a valuable horticultural plant."
Loreto: In clearing, Iquitos, Killip & Smith 27367. Puerto
Yessup, (Killip & Smith 26363). Puerto Arturo, (Killip & Smith
57823}. Rio Nanay, Williams 1254- Yurimaguas, Killip & Smith
27999. Caballo-cocha, Williams 2490. Colombia; Venezuela.
"Topiro," "cocona," "coconilla," "lulo" (Colombia).
Solanum torvuni Sw. Prodr. 47. 1788; 266.
Shrub or small tree (-4 meters) mostly stiped stellate-pubescent
but branchlets becoming smooth; petioles 1.5-4 cm. long, usually
not prickly; leaves often geminate on branchlets, ovate or oval-
rhombic, acute or subcordate at base, 6.5-13.5 cm. long, 5-12.5
cm. wide or larger, repand or lobulate with 2-3 obtuse teeth, slightly
rough and sordid white above, prickly or with a few prickles on
midrib, rarely elsewhere; inflorescence early lateral, 2-3-forked, 50-
100-flowered; pedicels 3-10 mm. long, sparsely stipitately glandular;
calyx 3.5-4 mm. long, 5 mm. across, lobes apiculate, 2-3 mm.
long, stellate-glandular; corolla white, stellate, 2.5 cm. across, the
5 lanceolate lobes acute or obtuse, 1 cm. long, half as wide, stellate;
stamens subequal, 0.5-1 mm. above base, filaments glabrous, 1 mm.
long, anthers lanceolate-ellipsoid, 6-7 mm. long, 1 mm. broad,
attenuate to apical small pores; ovary globose, stellate, style 8-10.5
mm. long, curved only at tip, stigma obscurely bilobed; fruiting
pedicels 15-18 mm. long, incrassate below calyx, the lobes of this
4 mm. long, 1.5 mm. wide; berry sordid yellow, 11-15 mm. thick,
the many flat kidney-shaped seeds yellow-brown, reticulate, 2.5
by 2 mm. After Urban, Symb. Ant. 6: 233. 1909, and Bitter,
Bot. Jahrb. 55: 87. 1919. Apparently the Williams specimens with
somewhat repand-lobulate but nearly glabrous leaves and small
flowers belong here; however, they may represent S. hyoscy ami-
folium Rusby, Bull. Torrey Club 26: 190. 1899, from lower Rio
Madre Dios, adjacent Bolivia, the leaves green but scabrous stellate
above, calyx 3-4 mm. long, corolla lobes 7 mm. long.
266 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Loreto: Caballo-cocha, Williams 2478; 2479. Northern to Cen-
tral America and the Caribbean; apparently also south to Bolivia.
Solatium velleum Swartz ex Roem. & Schult. Syst. 4: 643. 1819;
307. S. leptostachys Dunal in DC. Prodr. 13, pt. 1: 306. 1852?
A sordid or yellowish somewhat glanduliferous lanuginose-
tomentose sparsely aculeate shrub, the spines on the short or elongat-
ing branches mostly compressed, those on the leaf-nerves sub-
acicular; petioles about 2-3 cm. long; leaves solitary, ovate, rounded-
subcordate at base, acute, to 7 or 8 cm. long, about half as wide,
subcanescently pilose above, more tomentose beneath, with not more
than 4 pairs of lateral nerves; cymes soon opposite the leaves, not
aculeate, bifid, early scorpioid, finally strict, erect, elongated, the
pedicels typically only a few mm. long even in fruit; calyx open-
campanulate, with thin rounded sinuses between the 5 subulate
teeth; corolla obtusely rotate-angulate, sericeous without, 1.5-2 cm.
across; anthers oblong-lanceolate, 4 mm. long, apical pores finally
extending as a chink to base; ovary pilose, style curved toward tip.
Seems to be nearer the more widely distributed S. velleum Sw.
than the scarcely distinct segregate of Dunal, as it has the same
indument, rounded calyx sinus and corolla, but it differs at least
as follows and may bear the name of an enthusiastic student of
the plants of his country: var. Tovarii Macbr., var. nov., pedicellis
circa 1.5 cm. longis, demum ad 2 cm. longis, recurvatis.
Puno: North (8 km.) of Limbani, Metcalf 30545, type (det.
Killip, S. leptostachys, aff.). Brazil.
Solanum yapacaniense Kuntze, Rev. Gen. 3, 2: 228. 1898.
Erect, shrubby, 1 meter high, glabrous, terete, remotely aculeate
(also on midnerves), stout, straight or curved, to 7 mm. long, 3 mm.
wide at base; leaves entire or remotely spinulose dentate, lanceolate,
attenuate to sessile base, to 3 dm. long, half as wide; peduncles
opposite leaves, aculeate, to 6 cm. long; flowers 3-8, approximately
racemose; calyx cyathiform, 3 mm. wide, 2 cm. long, with 5 broad
acute lobes; corolla white, 5-parted, to 1 cm. long, the segments
oblong-lanceolate; stamens equal, shorter than corolla, anthers acu-
minate, 3 times longer than filaments; style long as stamens, stigma
obtuse, ovary globose. Near S. lancifolium Jacq., larger leaves,
small, aculeate inflorescence (author), but in Peru at least resembles
most S. nemorense Dunal but the remotely aculeate leaves larger,
corolla greenish-white or yellowish, fruit bright orange, to about 4
FLORA OF PERU 267
cm. in diameter, the light brown reniform seeds about 5 mm. long
(Peruvian specimens); perhaps it will prove to be a variant of
Dunal's species. Type from Rio Yapacani, Bolivia, at 400 meters.
Determinations by Morton.
San Martin: San Roque, Williams 6951. Junin: Dos de Mayo,
Pichis Trail, Killip & Smith 25854- Ayacucho: Kimpitiriki, Killip
& Smith 23019. Loreto: Balsapuerto, Killip & Smith 23019.
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