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 in
