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BIOLOGY

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FLORA OF PERU

BY

DOROTHY N. GIBSON

CUSTODIAN OP THE HERBARIUM
FIELD MUSEUM OP NATURAL HISTORY

Univeisuy of Illinois
MAR 20 1968

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-A, NUMBER 2

NOVEMBER 10, 1967

PUBLICATION 1032

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FLORA OF PERU

BY

DOROTHY N. GIBSON

CUSTODIAN OF THE HERBARIUM
FIELD MUSEUM OF NATURAL HISTORY

BOTANICAL SERIES

FIELD MUSEUM OF NATURAL HISTORY

VOLUME XIII, PART V-A, NUMBER 2

NOVEMBER 10, 1967

PUBLICATION 1032

Library of Congress Catalog Card Number: 36-10426

PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS

FLORA OF PERU

DOROTHY N. GIBSON

HYDROPHYLLACEAE. Waterleaf Family

REFERENCES: Choisy, Me"m. Soc. Phys. Gen. 6: 106. 1833 and in
DC. Prodr. 10: 179. 1846. A. DC. Prodr. 9: 287 and 564. 1845.
Lindl. Veg. Kingd. ed. 3: 638. 1853. Benth. & Hook. Gen. PI. 2: 825.
1876. Peter in Engler & Prantl, Pflanzenf . 4(3a) : 54-71. 1892. Brand
in Engler, Pflanzenr. 4, 251: 1-210. 1913.

Plants usually herbaceous, sometimes suffruticose, annual or per-
ennial, often pubescent, scabrid, glandular, and sometimes armed.
Leaves alternate or opposite, sometimes rosulate, entire or pinnately-
(rarely palmately-) divided. Inflorescences usually cymose, often
helicoid, or may be reduced to a single flower ; flowers bisexual, actino-
morphic; calyx 5-lobed, imbricate; corolla 5-lobed, imbricate (rarely
contorted), rotate, campanulate, or funnel-form; stamens usually 5,
inserted on corolla tube near its base, alternate with corolla lobes,
anthers bilocular, dehiscing longitudinally, filaments often basally
dilated, sometimes subtended by appendages ; ovary superior to half-
inferior, usually unilocular with 2 parietal placentae meeting in the
center, styles 1 or 2 (rarely 3), ovules 4 to many; fruit of 2 carpels,
each 2 to many-seeded (occasionally 1-seeded due to abortion), de-
hiscing loculicidally, septicidally or rarely indehiscent; seeds vari-
ously rugose, reticulate or muricate; embryo small and straight;
endosperm copious or thin, cartilaginous or fleshy.

Hydrophyllaceae are of almost world-wide distribution, having
been reported from all continents except possibly Australia. Most
of the genera are highly variable; cytogenetic studies of the family,
such as those undertaken by Dr. Lincoln Constance and others in
recent years will no doubt be a great help in delimiting taxonomic
units.

The family is of little economic importance save for ornamentals.
Four of the 20 genera occur in Peru.

Ovary bilocular Hydrolea

Ovary unilocular

Style 1 Phacelia

Styles 2

101

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

Plant seldom more than 30 cm. tall ; herbaceous to partly woody,
annual or perennial Nama

Plant quite large, to several meters tall ; suffruticose to arbores-
cent, perennial Wigandia

HYDROLEA L.

Erect, often branching, sometimes becoming decumbent, herba-
ceous to partly woody, usually glandular-pubescent, sometimes spiny
in leaf -axils, annuals or perennials from taproots, usually in wet soil.
Leaves alternate, entire, elliptical-lanceolate to obovate. Flowers
usually few in lateral and /or terminal cymes; corolla usually blue,
sometimes white, broadly campanulate, usually exceeding the calyx,
calyx divided to base with lobes overlapping at base, persistent, ac-
crescent; stamens included, inserted on tube, partly adnate at base,
filaments dilated at base, anthers sagittate; styles 2 (rarely 3), elon-
gating, with clavate-capitate stigmas; ovary superior, globose to
nearly elliptic, bilocular, with 2 fleshy placentae containing numer-
ous ovules; capsule globose to nearly elliptic, dehiscing irregularly,
containing numerous, minute seeds.

The genus ranges from the southern United States through Mex-
ico, Central and South America, the East and West Indies, and in
Asia and Africa.

Sepals about equal to the mature, glabrous capsule

H. spinosa L. var. inermis
Sepals exceeding by 2-3 mm. the mature, pubescent capsule

H. zeylanica

H. spinosa L. var. inermis Spruce ex Bennett in Journ. Linn.
Soc. 11: 271. 1869. H. megapotamica Spreng. Syst. 4. Cur. poster.
114. 1827. H. spinosa L. var. megapotamica (Spreng.) Brand in
Engler, Pflanzenr. 4, 251: 182. 1913. Wigandia herbacea Choisy,
Me"m. Soc. Phys. Gen. 6: 117. 1833 and in DC. Prodr. 10: 184. 1846.

Erect herbs to 50 cm. tall, stem densely viscid-pilose, unarmed.
Leaves sub-sessile or on short petioles, lanceolate to oblong-elliptic,
acute, narrowed to the base, entire, usually 2-8 cm. long, glandular-
pubescent on margins and along veins on under surfaces with scat-
tered pubescence on upper surfaces; flowers usually short-peduncu-
late, few in axillary and apparently sub-terminal cymes; corolla
open-campanulate, blue, to 10 mm. long and 12-15 mm. across, ex-
ceeding the calyx by 2 or 3 mm.; calyx persistent, lobes ovate to

FLORA OF PERU 103

ovate-lanceolate, acute, glandular-pilose, 5-7 mm. long; stamens
equalling or slightly longer than corolla, adnate near base of corolla
tube, filaments dilated at base; ovary ovoid to nearly globose, gla-
brous; styles 2, elongating to 3 or 4 mm.; stigmas clavate-capitate;
mature capsule globose, glabrous; seeds numerous, minute, light
brown, rugose, longitudinally costate-undulate.

San Martin: Prov. Moyobamba, Rioja, 800 m., Woytkowski 6096;
Loreto: Ule 6567 (fide Brand).

Although the typical armed variety of H. spinosa L. may be ex-
pected in Peru as it and the var. inermis Spruce, both with varying
degrees of pubescence, are fairly widespread throughout most of
South America, the only two Peruvian specimens of the genus seen
were unarmed.

H. zeylanica (L.) Vahl, Symb. Bot. 2: 46. 1791; Choisy in
Mem. Soc. Phys. Gen. 6: 107. 1833 and in DC. Prodr. 10: 180. 1846;
Walker- Arnott in Hook. Bot. Mag. 2: 193, t. 26. 1836; Wight, Icon.
PI. Ind. Orient. 2, t. 601. 1843; Bennett in Journ. Linn. Soc. 11: 275.
1869; Brand in Engler, Pflanzenr. 4, 251 : 174. 1913. Nama Zeylanica
L. Sp. PL ed. 1: 226. 1753 and ed. 2: 327. 1762. Steris javana L.
Mant. PL 1: 54. S. 12(1). 1767. Steris aquatica Burm. Fl. Indica: 73,
L 39, f. 3. 1768. Hydrolea javanica Blume, Bijdrag. FL Nederl. Ind.
13: 725. 1825. H. arayatensis Blanco, Fl. Filipinas, ed. 1: 211. 1837;
Merrill, Rev. Ident. Blanco 61. 1905.

Erect to decumbent herbs of wet soil, to 50 cm. tall, stem glabrous
except for minute pubescence of extreme upper part, unarmed, leaves
short-petioled, lanceolate to elliptic, entire, acute, glabrous, to 7 cm.
long; flowers 2 to 7 in axillary and apparently sub-terminal cymes;
pedicels and peduncles glandular-pubescent; corolla small, deep blue,
broadly open-campanulate with petals becoming reflexed; calyx per-
sistent, glandular-pubescent, lobes lanceolate, acute, equalling the co-
rolla and exceeding the mature capsule by 2-3 mm.; stamens basally
adnate to corolla tube, filaments dilated at base; ovary globose, glan-
dular pubescent; styles 2, 1-1.5 mm. long; stigmas clavate-capitate;
mature capsule globose, glandular-pubescent; seeds numerous, minute,
rugose, longitudinally costate-undulate.

Peru: Matthews 3046.

NAMA L.

Erect to prostrate, simple or branched, herbaceous to partly
woody, pubescent, often glandular annuals or perennials from slen-

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

der taproots. Leaves usually few, mostly alternate and entire, rarely
dentate. Flowers single in axils of upper leaves or several in reduced
lateral or terminal cymes; corolla white to purple, sometimes pubes-
cent, tubular to funnel-form, exceeding the calyx; calyx divided
nearly to base, the lobes linear-lanceolate to linear-spatulate, sub-
equal, accrescent; stamens usually included, all basally adnate and
unequally inserted on the corolla tube or may be unequal in length,
filaments usually glabrous, their bases dilated or sometimes append-
aged; styles 2, usually free but sometimes partially united, with small
capitate stigmas. Ovary superior to semi-inferior, usually pubescent,
unilocular but appearing bilocular due to the ingrowing parietal pla-
centae. Capsule oblong to ovoid, cartilaginous, dehiscing loculici-
dally or septicidally, containing numerous brown, variously pitted,
reticulate, or smooth seeds.

Of the species reported from South America, only one, Nama di-
chotomum (R. & P.) Choisy, was seen from Peru.1 There is a broad-
leaved form of N. dichotomum in Peru, Argentina and Bolivia, which
Hitchcock (1933) separates as var. amplifolium (Brand) Hitchc., on
the basis of the wider leaves and "spatulate calyx lobes" as opposed
to linear-spatulate lobes of the typical variety. All specimens of this
form which were examined had linear-spatulate calyx lobes and the
leaf size was not constant. The specimen of Bang 958 (Bolivia) on de-
posit in the Field Museum of Natural History, cited by Hitchcock as
var. amplifolium, not only has linear calyx lobes, but only two leaves
are slightly more than 4 mm. broad; all the remaining leaves are con-
siderably less than 4 mm. As no other differences were found, this is
considered to be a growth form or ecological variant of N. dichotomum.

Two additional annual species are known from the Andean re-
gions of Argentina, Bolivia and Ecuador and might be expected to
occur in Peru. N. jamaicense L. (introduced in South America?) is a
slender plant with leaves usually broadly ovate, to 3.5 cm. wide, an
almost tubular corolla to 7 mm. long, and an elongate ovary contain-
ing 50 to 70 light brown seeds. Nama undulatum HBK. is robust,
branching from the base and with sessile cauline leaves; the corolla
is 6-9 mm. long, and the capsule contains 100 to 180 yellow seeds.

Nama dichotomum (R. & P.) Choisy, Me"m. Soc. Phys. Geneve
6: 113. 1833; Gray, Proc. Am. Acad. 5: 338. 1861; Brand in Engler,

Although N. rupicola Bonpl. ex Choisy, Mem. Soc. Phys. Geneve 6: 114.
(1833) is credited to Mexico and Peru, no South American specimens were seen,
and it is felt that the Pav6n collection referred to was probably from Mexico.

FLORA OF PERU 105

Pflanzenr. 4, 251: 150. 1913. Hydrolea dichotoma Ruiz & Pav. Fl.
Peruv. 3: 22, pi. 244, fig- b. 1802. N. tetrandra Pavon ex Choisy,
Me"m. Soc. Phys. Geneve 6: 113. 1833, in synon. N. stricta Phil. Fl.
Atac. 37. 1860. Marilaunidium strictum (Phil.) Kuntze, Rev. Gen.
PL 2: 434. 1891. N. dichotomum subsp. eu-dichotomum Brand in
Engler, Pflanzenr. 4, 251: 151. 1913 (in part). N. dichotomum subsp.
eu-dichotomum f. stricta (Phil.) Brand, I.e. 151, fig. 28. N. dichoto-
mum var. angustifolium Gray, Proc. Am. Acad. 8: 284. 1870. N. an-
gustifolium (Gray) A. Nels., Coult. and Nels. New Man. Rocky Mt.
Bot. 410. 1909. N. dichotomum subsp. angustifolium (Gray) Brand
in Engler, Pflanzenr. 4, 251: 151. 1913. N. dichotomum ft pauciflora
Choisy ex Gray, Proc. Am. Acad. 8: 284. 1870. Conanthus angusti-
folius (Gray) Heller, Bull. Torr. Bot. Club 24: 479. 1897. Marilau-
nidium dichotomum (R. & P.) Kuntze, Rev. Gen. PL 2: 434. 1891.
Marilaunidium tenue Woot. & Standl. Contr. U. S. Nat. Herb. 16:
162. 1913. N. dichotomum var. amplifolium (Brand) Hitchc. Am.
Journ. Bot. 20: 530. 1933.

Annual, slender, erect, simple or more or less dichotomously
branched herbs, 4-30 cm. tall from a slender taproot, hirsute and
glandular; leaves linear-spatulate to oblong-elliptic, 4-30 mm. long,
1-10 mm. wide, tapering to short petioles; flowers usually in pairs in
leaf axils, sessile or on short pedicels; calyx lobes linear-spatulate,
2-5 mm. long in flower, to 10 mm. in fruit, hirsute-ciliate; corolla
white to pale lavender, or white with blue or lavender lines, narrowly
campanulate, to 5 mm. long; stamens unequally placed about 1 mm.
from base of corolla, the glabrous filaments widened slightly just
above the adnate base and with minute free margins along the adnate
portions; styles to 2 mm. long; capsules pubescent, ovoid to oblong,
containing 20 to 60 brown, minutely reticulate seeds about 0.5 mm.
long.

Arequipa: Prov. Arequipa, cerros de Jesus, 2360 m., Vargas 7937;
southern slopes of Chachani Mtn., 3660 m., Hinkley 74; quebrado de
San Lazaro, 8500 ft., Munz 15485; Tingo, 2100-2300 m., Pennell
13108; Prov. Islai, Mollendo, Hitchcock 22406. — Cuzco: Prov. Calca,
2900 m., Vargas 723; Dist. Urubamba, Ollantaitambo, 3000-3100 m.,
Pennell 13679. — Huanuco: Prov. Ambo, about 7000 ft., Macbride
3192. — Junin: Prov. Huancayo, 3317 m., Soukup 2959. —La Lib-
ertad: Prov. Trujillo, Cerro Cabezon, 650 m., 'Ldpez M. 0678. — Lima:
Prov. Cajatambo, Ambar, 2110 m., Stork 11437; Prov. Huarochiri,
valley of Rio Rimac, 1800 m., Goodspeed & Metcalf 30246, San Barto-
lome", 1500-1600 m., Weberbauer 5291; Prov. Lima, Lurin, about

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

200 ft., Macbride 5975. — Moquegua: Prov. Moquegua, Lomas de
Mostacilla, 50-300 m., Vargas 8575; Torata, 2200-2300 m., Weber-
bauer 7409. — Piura: Dist. Mancora, Amotape Hills, Haught & Sven-
son 11550. — Tacna: Prov. Tacna, Lomas cerca al Morro Sama,
500-600 m., Ferreyra 12539. In Ecuador, and to Bolivia, Chile and
Argentina.

PHACELIA Juss.

Erect to decumbent, herbaceous, caulescent, branching, usually
pubescent and often glandular annuals or perennials from taproots or
creeping rootstocks. Cauline leaves usually alternate, entire, lobed,
pinnatifid or pinnately dissected ; basal leaves often rosulate. Flowers
few to numerous in usually helicoid, raceme-like terminal cymes, long
pedunculate to sessile; corolla blue, purple, pale lilac or white; nar-
rowly to widely campanulate; calyx divided nearly to base, per-
sistent, often accrescent; stamens usually exserted, all adnate and
equidistant from the base of the corolla- tube; a pair of scales, wholly
or partly adnate to tube, sometimes present at base of each filament;
style bifid. Ovary unilocular with 2 linear placentae adherent to
walls. Capsule ovoid, loculicidally dehiscent, containing 2 to many
(usually 4) minutely rugose-aerolate seeds.

A widely distributed and variable genus, well represented through-
out the Andes.

A considerable amount of work was done in an effort to find mor-
phological characters consistent enough to be of use in classification
of the so-called "P. magellanica" complex. A study of the scales at
the base of the filaments proved them to be inconstant; investigation
of the pubescence on the filaments revealed deviations in the flowers
of single plants — one or two stamens in a flower are sometimes gla-
brous while the remainder show varying amounts of pubescence;
plants with glabrous to slightly pubescent filaments occasionally have
stamens exserted well beyond the 3 mm. specification used by Mac-
bride (1917) to separate the typical variety of P. magellanica from
the var. robusta (Brand) Macbride; pubescence of leaves and stems
is equally variable. It is sometimes possible to separate certain An-
dean plants with glabrous to slightly pubescent filaments from those
with filaments obviously to densely pubescent, in support of the sep-
aration of P. secunda Gmel. and its subspecies pinnata (R. & P.)
Constance. However, other characters used by Macbride and by

FLORA OF PERU 107

Dundas1 do not seem to be consistent; i.e., the basal leaves of both
range from densely rosulate to not at all rosulate, and both exhibit
stamens exserted at various lengths past the corollas. Therefore, in
view of the fact that many specimens over a wide geographical range
exhibit numerous variations and forms suggesting intergradation, we
are treating this complex collectively as P. secunda Gmel., sens, lat.,
until satisfactory taxonomic units may be delimited.

Leaves entire to pinnate; when pinnate, pinnules entire. P. secunda
Leaves pinnately lobed to lacerate and parted; margins of lobes or
pinnules irregular and usually deeply crenate to dentate

P. pinnatifida

Phacelia secunda J. F. Gmel. L. Syst. Nat. ed. 13, 2: 330. 1791.
Hydrophyllum magellanicum Lam. Journ. Hist. Nat. 1: 373. 1792.
Heliotropium pinnatum Vahl. Symb. Bot. 3; 21. 1794. Aldea pin-
nata Ruiz & Pav. Fl. Peruv. 2: 8, t. 11 4-, fig- a. 1799. P. circinnata
(Willd.) Jacq. f. Eclog. 1: 135, t. 91. 1816. Hydrophyllum Aldea
Roem. & Schult. Sept. 4: 114. 1819. P. magellanica (Lam.) Cov.
Contr. U. S. Nat. Herb. 4: 159. 1893. Hydrophyllum magellanicum
pinnatum Macloskie. Rept. Princeton Patag. Exped. 8: 674. 1905.
P. magellanica (Lam.) Cov. f. amoena Brand in Engler, Pflanzenr. 4,
251: 97. 1913. P. magellanica ssp. barbata f. pinnata Brand, I.e. 99.
P. pinnata (R. & P.) Macbr. Contr. Gray Herb. 49: 37. 1917. P.
secunda J. F. Gmel. ssp. pinnata (R. & P.) Const. Univ. Calif. Publ.
Bot. 30(3): 241. 1959. P. secunda J. F. Gmel. ssp. plantaginea
(Brand) Const., L c.

Biennial or perennial, simple or branched, erect to semi-decum-
bent herbs, 4 cm. to 50 cm. tall from a taproot; more or less hirsute
and appressed-hispid throughout, foliage usually appearing greyish-
green; stem usually simple, more or less erect, additional shorter
stems often arising near its base; lower leaves usually long-petioled
and conspicuously veined, lanceolate, occasionally ovate, 1-10 cm.
long, 0.5-3 cm. broad, entire or pinnate, rarely trilobate, the one or
more pairs of lateral leaflets much smaller than the terminal ; cauline
leaves entire or pinnate, usually reduced upward; flowers numerous
in dense racemose to corymbose, often somewhat capitate cymes,
inflorescence elongating and spreading in fruit; calyx lobes lanceolate,
3-8 mm. or more long, hirsute-ciliate; corolla dingy- white, pink, blue-
lavender to purple, 4-6 mm. or more long; stamens exserted, fila-

1 Dundas, Frederick W. "A Revision of the Phacelia Californica Group (Hy-
drophyllaceae) for North America." Bull. So. Calif. Acad. Sci. 33: 152-178. 1935.

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

ments glabrous to pubescent, style well exserted, pubescent toward
base; capsule pubescent, ovoid; seeds to about 0.4 mm. long, brown.

Ancash: Prov. Bolognesi, Matarragra, 3520-3560 m., Ferreyra
5609; San Isidro entre Chiquian y Aquia, 3000 m., Ferreyra 7554.
— Arequipa: Prov. Arequipa, south slopes above Arequipa on trail to
Sumbay, 3665 m., Hutchison & Wright 724-0; Prov. Condesuyos,
Chuquibamba, among rocks, 10,000 ft., Stafford 1177. — Ayacucho:
Prov. Huancavelica, Orccon, near Conaica, 3550-3580 m., Tovar 80.
— Cajamarca: Prov. Celendin, 12 km. east of Encanada on road
from Cajamarca to Celendin, 3470 m., Hutchison & Wright 5121.
— Chasqui: Prov. Huanuco, about 10,500 ft., Macbride & Feather-
stone 1763. — Cuzco: Prov. Cuzco, Chuco, 3300 m., Vargas 135; Prov.
Cafias, slopes of San Andre's de Checca, 3860 m., Vargas 11020;
Prov. Espinar, 4200 m., Vargas 10544- —La Libertad: Prov. Hua-
machuco, summit above Aricapampa on road to Huamachuco, 3970
m., Hutchison & Wright 6262. —Lima: Prov. Canta, bushy slopes
of small canyon east of Canta, 3300-4100 m., Pennell 14619; Prov.
Huarochiri, between Rio Blanco and Chilca, 3650 m., Hutchison &
Tovar 4242', between Lima and Oroya, 3300-3400 m., Ferreyra 5292;
entre San Mateo y Casapalca, 3700 m., Ferreyra 6513; Matucana,
steep slopes, 8000 ft., Macbride 2939, Viso, rocky grassy talus slopes,
9000 ft., Macbride & Featherstone 639. — Pomopampa: Prov. An-
cash, Estacion 30 miles east of Huaraz, about 12,000 ft., Macbride &
Featherstone 2497. — Puno: Prov. Carabaya, cerca abra de anta-
pampa, 4150 m., Vargas 7003; Prov. Lampa, Palmpa, Pucara, 4200
m., Vargas 6401; Prov. Puno, from mountainside near Lake Titicaca,
3125 m., Shepard 42; Prov. Sandia, along road from Cuyocuyo to
summit, 3600 m., Metcalf 30708. To Bolivia, Chile and Argentina.

The annual P. simplicifolia Colla (P. clinopodioides Bertero)
from Chile, with entire, simple to trilobate leaves, is readily distin-
guishable, as the stamens are always included in the tubular-cam-
panulate corolla and the narrowly linear calyx lobes are more than
twice as long as the capsule.

Phacelia pinna tifida Griseb. ex Wedd. Chlor. Andin. 2: 85.
1857. P. pinnatifida (3 robusta Wedd., I.e. P. viscosa Phil. Flor. Ata-
cam. 37. 1860. P. pinnatifida var. elatior Griseb. Symb. Fl. Arg. 267.
1879. P. pinnatifida var. lobulata Hicken, Darwiniana 1: 134. 1924.

Biennial or perennial, erect, ascending, usually robust and often
much branched, to 50 cm. tall from stout taproot, glandular- viscid,
pilose and hirsute; leaves linear-oblong to oblong-elliptic, occasion-

FLORA OF PERU 109

ally to 16 cm. long, pinnately lobed to lacerate and parted, the mar-
gins of the lobes irregular and usually deeply crenate to dentate,
subsessile to long petioled; flowers numerous in dense, usually race-
mose cymes, inflorescence elongating and often becoming strictly
erect in fruit; calyx lobes linear to oblong-spatulate, obtuse, of un-
equal widths, to 7 mm. long or more, hirsute, considerably exceeding
the capsule; corolla lavender, blue, or white with blue limb, 5-9 mm.
long; stamens exserted, filaments glabrous; style exserted, hirsute
directly above and below the bifurcation, may be cleft from only
1 mm. above ovary to nearly half its length; capsule ovoid, pubes-
cent, 4-5 mm. long; seeds 2-3 mm. long, dark brown.

Arequipa: Prov. Camana, 4 km. from Caraveli on road to Atico,
1800-1900 m., Metcalf 30343. — Ayacucho: Prov. Huancavelica,
Jupas-pata, arriba de Conaica, 3700-3750 m., Tovar 245; Prov. Lu-
canas, entre Puquio y Nauca, 3400-3500 m., Ferreyra 5500. — Cuzco :
Prov. Calca, Pisac, 3200 m., Vargas 5075; Urubamba, open stony
slopes surrounding town, 2900 m., Vargas 2809; valley of the Uru-
bamba, 3300 m., Weberbauer 6913; Saxaihuanan, 3500 m., Herrera
3116; Prov. Canas, Asuncion Bridge, Apurimac River, rocky talus
slopes, 3725 m., Vargas 11035; Prov. Canchis, Aguas Calientes,
4150 m., Vargas 3325. —Lima: Prov. Huarochiri, Viso, 9000 ft.,
Macbride & Featherstone 764. — Moquegua: Prov. Moquegua, be-
tween Moquegua and Torata, 1900 m., Weberbauer 7425. — Puno:
Prov. Lampa, Andamarca, 13,300 ft., Sharpe 50; Prov. Puno, 4500
m., Soukup 86; vicinity of Lake Titicaca, 3125 m., Shepard 146;
3 km. from Chucuito on road to Puno, 4000 m., Metcalf 30685. Ex-
tending to Bolivia, Chile and Argentina.

P. nana Wedd. (P. sinuata Phil.; P. villosa Phil.), a perennial in
Chile, Argentina and Bolivia, is similar in appearance to P. pinnati-
fida, but the stamens and style of P. nana are never exserted.

P. artemesioides Griseb., found in Chile, Argentina, Brazil and
Uruguay, also closely resembles P. pinnatifida and may intergrade
with it; the foliage of P. artemesioides is more finely dissected, the
ultimate divisions sometimes appearing almost linear-filiform. Fur-
ther, the calyx-lobes of the few specimens examined were consider-
ably shorter than those of P. pinnatifida, exceeding the mature
capsule by only about 1 mm. or not at all.

A third species, P. cumingii (Benth.) A. Gray, with foliage much
like that of P. artemesioides, occurs in northern Chile and in Argen-
tina. The stamens and style are always included; the style is united
almost the entire length, being cleft only at the apex; and there are

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

usually 12 seeds instead of four as in P. artemesioides and P. pin-
natifida.

WIGANDIA HBK.

Large, erect, suffruticose to arborescent perennials, variously pu-
bescent (including stinging hairs) and glandular throughout. Leaves
often quite large but variable in size from about 5-40 cm. long, and
3-24 cm. broad, coriaceous, net-veined, ovate-cordate, alternate,
coarsely twice-dentate. Flowers few to many, sessile to sub-sessile,
often appearing densely spicate, in terminal cymes or panicles; co-
rolla greenish-white, yellow or lavender, open-campanulate, often
partially pubescent, equal to or usually exceeding the calyx; calyx
divided nearly to base, hispid-hirsute and ciliate, persistent, the lobes
linear-lanceolate to lanceolate, acute, accrescent; stamens exserted,
partly adnate to the base of the corolla tube, filaments usually pubes-
cent, anthers linear-oblong, more or less sagittate, dorsifixed ; styles 2,
elongate, with capitate-clavate, depressed stigmas; ovary superior,
oblong, pubescent, unilocular, with 2 laminiform placentae, the trans-
verse portions of each meeting in the center, and with the numerous
ovules borne chiefly along their revolute edges; capsule ovate-oblong,
pubescent, dehiscing loculicidally or septicidally, and containing more
than 200 minute, reticulate-rugose, brown seeds.

The genus is widely distributed, ranging from Mexico and Cen-
tral America to Argentina, weedy, and so variable that I see no reason
to change Sir Joseph Hooker's evaluation of the group of 18511 when,
in connection with his studies of W. caracasana HBK., he said,
". . . it must be confessed that other species of Wigandia present
great variations and intermediate gradations which render their
claims to specific identity extremely doubtful."

Corolla to 50 mm. long, greenish white to pale yellow W. crispa

Corolla to 18 mm. long, pale violet to purple W. urens

Wigandia crispa (R. & P.) HBK. Nova Gen. & Sp. 3: 129.
1819; Choisy in Me'm. Soc. Phys. Geneve 6: 116. 1833 and in DC.
Prodr. 10: 184. 1846. Hydrolea crispa Ruiz & Pav. Fl. Peruv. 3: 22,
t. 244. 1802. Wigandia pruritiva Spreng. Syst. 1 : 865. 1825. Ern-
stamra crispa (HBK.) 0. Ktze. Rev. Gen. PI. 2: 434. 1891. W. re-
flexa Brand in Engler, Pflanzenr. 4, 251: 135. 1913. "W. urens (L.)
Urban," ace. Urban in Fedde Rep. Spec. Nov. fasc. 15: 415. 1919.

1 Bot. Mag. ser. 3, 7: t. 4575. 1851.

FLORA OF PERU 111

Erect, suffruticose to arborescent plant to 6 m. tall; densely pu-
bescent throughout, overlaid with hispid, often glandular and sting-
ing hairs; leaves with 8 to 10 lateral veins, ovate, acute, subcordate
to cordate at base, irregularly biserrate, appearing deep green above,
greyish-white beneath, silky-pilose on both sides, usually noticeably
hispid along primary veins; flowers subsessile to sessile in scorpioid,
terminal panicles, corolla campanulate, greenish-white to pale yellow,
silky-hirsute and sometimes glandular without, large, to 5 cm. long,
exceeding the calyx by about 3 cm.; calyx persistent, lobes equal or
subequal, lanceolate, acute, glandular-pilose and hispid-ciliate, to
about 20 mm. long at an thesis, 30 mm. in fruit; stamens extending
7 to 10 mm. beyond sinuses but may be 4-7 mm. shorter than the
corolla lobes, subequal, basally adnate to corolla tube, filaments
pubescent at base but hairs soft and weak so that filaments may
appear glabrous if examined without a lens; anthers appearing more
sagittate than those of W. urens. Ovary ovate-oblong, silky-hirsute,
styles to 30 mm. long, pubescent only at base; stigmas clavate-
capitate; mature capsule oblong-ovoid, densely white-hirsute, ex-
ceeded by calyx lobes; seeds minute, reticulate-rugose, brown.

Junin: Prov. Tarma, 2000 m., Constance & Tovar 3586, Carpa-
pata, above Huacapistana, 2400 m., Killip & Smith 24363. — Piura:
Prov. Huancabamba, Stork 11370; Prov. Paita, Talara, Haught 86.
Also in Ecuador.

Known locally as "mano de leon."

Wigandia urens (R. & P.) HBK. Nova Gen. & Sp. 3 : 127. 1819 ;
Choisy, DC. Prodr. 10: 184. 1846, incorrectly as "W. urens (R. & P.)
Choisy." Hydrolea urens Ruiz & Pav. Fl. Peruv. 3: 21, t. 243. 1802.
W. peruviana W. Miller in Bailey, Cyclop. Am. Hort. 1975. 1902.
Ernstamra urens (R. & P.) 0. Ktze. Rev. Gen. PI. 2: 434. 1891.

Erect, branching, robust, suffruticose plant, to 4 m. tall, densely
sordid with white hispid to bristly-hirsute, sometimes glandular and
often stinging hairs; leaves with 5 to 7 lateral veins, ovate, obtuse to
sub-acute at apex, cordate to truncate at base, irregularly biserrate,
appressed pilose on upper surface and white-tomentose underneath ;
flowers subsessile to sessile in dense scorpioid terminal cymes or pan-
icles, corolla campanulate, pale violet to purple, lighter in throat,
silky pilose to tomentose without, to 18 mm. long, equalling or slightly
exceeding the calyx; calyx persistent, lobes subequal, acutely linear-
lanceolate, tomentose and hispid-ciliate, about equal to the mature
capsule; stamens included, subequal, filaments retrorsely hispid from

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

base to more than half to nearly two-thirds their length, basally ad-
nate to corolla tube. Ovary ovate-oblong, silky to hispid-hirsute,
styles 10-15 mm. long, pilose to more than one-third their length;
stigmas clavate-capitate; capsule oblong-conical, hispid-hirsute; seeds
numerous, minute, reticulate-rugose, brown.

Canta: Prov. Canta, Obrajillo, Wilkes Expedition. — La Liber-
tad: Chicama Valley, Smyth 72. —Lima: Prov. Canta, 1700-1800 m.,
Ferreyra 7261; Prov. Huarochiri, Matucana, 8000 ft., Macbride &
Featherstone 290; Olivo, 1140 m., Stadelman 2; Valley of Rio Rimac,
1900 m., Constance & Tovar 3583; Ruinas de Cajamarquilla, 300-
400 m., Ferreyra 2857. To Colombia and Ecuador.

Known locally as "Tabaquillo."

Wigandia caracasana HBK., found in Colombia and Venezuela
and which may be expected in Peru, I believe to be a form or variety
of W. urens, as the only distinguishing character appears to be its
constant short and velvety pubescence as opposed to the longer, his-
pid pubescence of W. urens. The "channelled" petioles of W. cara-
casana are also found in the other species; although it is true that the
calyx lobes of specimens identified as W. caracasana are sometimes
shorter than the mature capsule, they are also sometimes equal to
and may even exceed the capsule.

POLEMONIACEAE. Phlox Family

REFERENCES: Benth. in DC. Prodr. 9: 302-322. 1845. Benth. &
Hook., Gen. PL 2 : 820-824. 1876. Peter in Engl. & Prantl, Pflanzenf .
4(3a): 40-54. 1891. Brand in Engl. Pflanzenr. 4, 250: 1-203. 1907.
Wherry, A provisional key to the Polemoniaceae. Bartonia 20: 14-17.
1940. Grant, Natural History of the Phlox Family. 1: 1-273. 1959.

Plants usually herbaceous annuals or perennials, a few climbing
shrubs or small trees, usually somewhat pubescent, sometimes glan-
dular. Leaves alternate (lower ones sometimes opposite), entire,
serrate, or pinnately divided. Inflorescences usually cymose, the
flowers rarely solitary and axillary; flowers bisexual, actinomorphic
or weakly zygomorphic. Calyx 5-lobed, imbricate or valvate, per-
sistent, accrescent to subaccrescent ; corolla tubular (campanulate in
Cobaed), 5-lobed, lobes contorted, salverform to rotate; stamens 5,
inserted on corolla tube at various, often unequal heights, anthers
bilocular, introrse. Ovary superior, inserted on disc, trilocular (rarely
2 or 5 carpels), placentation axile, ovules 1-many on each placenta,
style filiform, stigmatic branches 3 (rarely 2 or 5); fruit a capsule

FLORA OF PERU 113

dehiscing loculicidally (septicidally in Cobaea) or rarely indehiscent;
seeds usually many but may be only 1 or 2 in each locule, usually
small, more or less angular, sometimes winged. Embryo spatulate;
endosperm usually firm, abundant.

The Polemoniaceae are naturally distributed throughout North
and South America and Eurasia. They occur as weeds in Africa and
Australia. Of the 317 reported species, 27 occur in South America;
five genera are presently known from Peru. The family is economi-
cally important for the ornamentals Cantua, Cobaea, Gilia, Linan-
thus, Phlox and Polemonium.

Plants herbaceous annuals or perennials

Leaves dissected, incised or lobed Gilia

Leaves entire or serrate Phlox

Plants suffrutescent or woody

Plants lianas, terminal leaflets modified to tendrils Cobaea

Plants small trees or shrubs

Leaves simple; capsule longer than calyx Cantua

Leaves pinnately divided; capsule shorter than calyx .... Huthia

CANTUA Juss. ex Lam.

Erect, branching shrubs (rarely becoming small trees), the
branches cylindrical, grooved to warty, usually at least partly pubes-
cent. Leaves coriaceous to subcoriaceous, alternate or fasciculate,
entire to sinuate-dentate; petioles short and usually pubescent; the
caducous juvenile leaves toothed and usually considerably larger than
adult leaves; adult leaves of all but two species nearly always entire
—rarely a few teeth appear; flowers usually few to many in terminal
corymbs, rarely solitary; corolla often rose to purple but may be
white, pink, yellow, greenish, or the tube may be white or yellow
with lobes yellow to purple; variously colored striations also occur;
corolla tubular to funnelform or salverform, often twice to three
times as long as the calyx, apex of corolla lobes from deeply bilobate
through retuse, retuse-apiculate, apiculate or even truncate-erose;
calyx purple to green, persistent, accrescent, tubular to cup-shaped or
campanulate, more or less zygomorphic, 3-5 (6) -lobed, lobes acute,
sometimes acuminate, one or all may be bifid, 2 lobes sometimes di-
vided to base so that calyx appears bilabiate; stamens included to
long exserted, partly adnate to and inserted on tube from very near
the base to a point one-fourth of the corolla length above the base,

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

filaments usually glabrous (rarely pubescent near or above point of
insertion), anthers versatile, sagittate or reniform; style filiform, in-
cluded or exserted, stigma 3-parted, branched or lobed; disc irregu-
larly 5-lobed; ovary superior, glabrous, ovoid to oblong-ovoid,
tricarpellate, trilocular with numerous ovules in each locule; ma-
ture capsule oblong, glabrous, dehiscing loculicidally by 3 valves,
containing numerous small, broadly-winged seeds.

The genus is South American, ranging through the Andean region
from Ecuador through Bolivia, Chile and to Brazil. All species are
variously known locally as "Ccantu," "Kantuta," "Decanto,"
"Ccellmo," "Flor del Inca," "Inca-Panccara," "Esfirna" and "Cha-
lagantu." C. candelilla is also known as "Candelilla"; C. flexuosa is
sometimes called "Turu," and C. quercifolia is known as "Pipiso"
or "Pepiso."

Leaves all obviously sinuate-crenate ................. C. quercifolia

Leaves entire to somewhat dentate

Calyx cup-shaped to campanulate, not more than 1.5 cm. long
Adult leaves 8-16 cm. long, 4-6 cm. wide .......... C. longifolia

Adult leaves usually less than 8 cm. long; never more than 10 cm.

long and 4 cm. wide

Corolla greenish-yellow, 2.5-3 cm. long, lobes usually apicu-
late .................................... C. pyrifolia

Corolla white, 2-2.5 cm. long, lobes retuse to deeply cleft

C. flexuosa

Calyx tubular to tubular-campanulate, 1.6-3.2 cm. long
Apex of expanded corolla truncate ............... C. candelilla

Apex of expanded corolla not truncate

Corolla 4-5 cm. long ........................ C. cuzcoensis

Corolla 5.5-8 cm. long ........................ C. buxifolia

C. bicolor Lemaire is omitted as there is no reason to believe that
it occurs in Peru. The single Peruvian specimen, Soukup 363, cited
by Infantes Vera1 has terminal and sub-terminal corymbose inflores-
cences of several flowers, with corollas to 6 cm. long, and is C. buxi-
folia Juss. According to her own description1 of C. bicolor, as well as
those of Lemaire,2 Paxton,3 Hooker,4 and Brand,5 the inflorescences

rQl. 1962.
2 Fl. des serres, ser. 1, 3: 242. 1847.
1 Mag. Bot. 15: 219. 1849.
4 Bot. Mag. ser. 3, 9: t. 4729. 1853.
8 Engler, Pflanzenr. 4, 250: 23. 1907.

FLORA OF PERU 115

consist of solitary flowers, each borne singly in leaf axils, with corollas
of only 4-5 cm. in length. This description fits Fiebrig 2155 from
southern Bolivia (Dept. Tarija, Prov. Arce, Huaya villa).

Cantua buxifolia Juss. ex Lam. Diet. Encycl. Bot. 1: 603. 1785;
Illustr. 1: t. 106, f. 2. 1823. Juss. Ann. Mus. Hist. Nat. Paris 3: 118,
t. 8. 1804; D. Don, Edinb. Phil. Journ. 7: 290. 1822; Benth. in DC.
Prodr. 9: 321. 1845; Hook, in Bot. Mag. t. 4582. 1851; Lemaire, Jard.
Fleur. 2:t. 115. 1852; Peter in Engler & Prantl, Pflanzenf. 4, 3a: 45.
1891; Brand in Engler, Pflanzenr. 270: 22. 1907; Infantes Vera, Lilloa
31: 84. 1962. C. ovata Cav. Icon. 4: 43, t. 363. 1797; Poiret in Lam.
Encycl. Me"th. Suppl. 2: 79. 1811; Roem. & Schult. Syst. 4: 366. 1819;
Benth. in DC. Prodr. 9: 321. 1845. C. tomentosa Cav. Icon. 4: 43,
t. 364- 1797; Persoon, Syn. PI. 1: 187. 1805. C. dependens Pers. Syn.
PI. 1 : 187. 1805. C. uniflora Pers. Syn. PI. 1 : 187. 1805. C. theaefolia
Don, Edinb. Phil. Journ. 7: 289. 1822. C. lanceolata Peter in Engler
& Prantl, Pflanzenf. 4, 3a: 45. 1891. C. buxifolia Juss. var. lanceolata
(Peter) Brand in Engler, Pflanzenr. 270: 22. 1907. C. buxifolia Juss.
var. ovata (Cav.) Brand in Engler, Pflanzenr. 270: 22. 1907. C. alu-
tacea Infantes Vera, Lilloa 31: 88. 1962. Periphragmos dependens
Ruiz & Pa von, Fl. Peruv. 2: 18, t. 133, 1799. P. uniflorus Ruiz &
Pavon, Fl. Peruv. 2: 18. 1799.

Erect, branching shrub or small tree, to 5 m. tall, more or less
pubescent throughout, although degree of pubescence quite variable,
glabrescent in part. Leaves ranging in size from only 0.6 cm. to
nearly 4 cm. long, sessile to short-petioled, usually elliptic but may
be obovate, oblanceolate, or linear-oblanceolate, acute, tapering to
base, usually entire, rarely irregularly dentate toward apex; inflores-
cence of few to many flowers on pedicels of varying lengths, in usually
lax terminal corymbs, rarely reduced to a solitary flower; corolla fun-
nelform, usually rose to purple but may be white, yellow, or the tube
yellow with lobes white or rose, 5.5-8 cm. long, the lobes bilobate,
retuse-apiculate or rounded and irregularly dentate; calyx tubular or
tubular-campanulate, 1.6-3.2 cm. long, lobes unequal, triangular,
acute, rarely acuminate, nearly glabrous to heavily pubescent, some-
times glandular, usually more pubescent at base and at apex of lobes,
inside as well as outside; stamens exserted or included, adnate to
tube at a point about 1-1.5 cm. from its base; ovary pyriform to
oblong, to 12 mm. long, glabrous; style usually exserted, rarely in-
cluded, to 9 cm. long; mature capsule oblong, to 2.5 cm. long, usually
containing 20-30 winged seeds, each 8-12 mm. long.

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

Without locality: Mathews 485 (1834); Lechler 1936 (only frag-
ments seen, along with FM photograph No. 27054) ; FM photograph
No. 14025, of Ruiz collection, without number or date. — Ancash:
Valley Santal, 3400 m., Rauh & Hirsch P2106; Prov. Bolognesi,
Callapata, 3850 m., Ferreyra 5804; Chupa, 3200 m., Ferreyra 7543.
— Arequipa: Prov. Arequipa, Rose 18978. — Apurimac: Prov. Anda-
huailas, Argama, 3300 m., West 3748; 2500 m., Stork & Horton 10727;
Prov. Apurimac, south of Chincheros, 3200 m., West 3688. — Aya-
cucho: Prov. Parinacochas, Coracora, 2900 m., Weberbauer 5790;
Valley of Rio de Ocona, above Calpamayo, 3650 m., Weberbauer
7177. — Cuzco: Prov. Cuzco, 3400 m., Ferreyra 2651; 3600 m., Her-
rera 1192. — Huanuco: Prov. Llata, 7000 ft., Macbride & Feather-
stone 2292; Prov. Dos de Mayo, Chavinillo, 8000 ft., Macbride &
Featherstone 1977. — Junin: Prov. Ambo, Chasqui, 10,500 ft., Mac-
bride & Featherstone 1908; Prov. Cerro de Pasco, Ambo, 3200 m.,
Seibert 2204; Prov. Jauja, 3300 m., Ochoa 108; Prov. Tarma, 3300-
3500 m., Ferreyra 3786; 10,600 ft., Saunders 539, 3500 m., Constance
& Tovar 2369, 2397. —Lima: Prov. Huarochiri, Obrajillo, Nee (July,
1802), Rio Blanco, 3500 m., Hutchison 579; San Mateo, 3200 m.,
Asplund 11492; 11,500 ft., Saunders 323; Viso, 9000 ft., Macbride &
Featherstone 640. — Puno: Prov. Huancare", Umuchi, 3125 m., Shep-
ard 103; Ushopata Valley, 3700 m., Cook & Gilbert 127; Prov. Sandia,
Limbani, 3200 m., Vargas 1284. Also in Bolivia.

Dra. Infantes Vera retained as separate species C. ovata Cav. and
C. tomentosa Cav., and created a third, C. alutacea Infantes. Every
effort was made to find one or more characters that could be relied
upon to distinguish each of these from C. buxifolia Juss., without
success. In the case of C. ovata Cav., the original plate and descrip-
tion indicates flowers with apiculate corolla lobes, solitary in leaf
axils, leaves ovate-oblong, stamens exserted. Periphragmos uni-
florus Ruiz & Pavon, which Infantes agrees is synonymous with
C. ovata, is also described as having flowers borne singly in leaf axils,
with stamens exserted. Bentham states in part, "... floribus laxa
subcorymbosis v. in axillis superioribus solitariis. . . ." Infantes de-
scribes the inflorescences as ". . . pauci o multiflores, a veces flores
solitarias, nacen en la axila de una hoja. ..." and the corolla lobes as
"... enter os o bilobados . . ., mucronados, . . . pubescentes en los bordes."
However, no pubescence was found on corolla lobes of any of the
specimens which she cited and the lobes were not mucronate. A
Field Museum photograph of a Ruiz collection identified by Brand
as C. buxifolia var. ovata shows no flowers, but two leafy stems, obvi-

FLORA OF PERU 117

ously in the juvenile phase with ovate leaves and occasional, irregular
dentation toward the apices.

Although I was unable to see any of the specimens cited by In-
fantes as C. tomentosa, I did examine fragments of Lechler 1936, cited
by Brand as the type of his C. buxifolia var. lanceolata, which In-
fantes places in synonymy with C. tomentosa. The two calyces and
one leaf which were examined are not tomentose, in fact, they are not
as pubescent as are other specimens cited by her as C. buxifolia, i.e.,
Macbride & Featherstone 1069. Further, the single seed in the packet
of fragments is in no way different from those of C. buxifolia.

Her C. alutacea, with flowers described as white to yellowish, is
at best a color form of C. buxifolia; seeds of Herrera 2198 cited as
C. alutacea are identical with those of C. buxifolia, the apex of the
wing being irregular but not bilobate, and the seed measurement
9 mm. rather than 7 mm. One might be able to separate C. alutacea
from C. buxifolia by reading color notes on the labels if it were not
for the fact that Vargas 1284 was cited as C. alutacea, and his label
reads, "rojo y amarillo."

Cantua candelilla Brand in Engler, Pflanzenr. 4, 250: 22. 1907;
Infantes Vera, Lilloa 31: 95. 1962.

Erect, branching shrub or small tree to 4 m. tall, more or less
pubescent throughout. Leaves usually quite small but occasionally
to 3 cm. by 0.9 cm. wide, sessile to short-petioled, linear-oblanceolate
to oblanceolate-ovate, acute, tapering to base, usually entire, rarely
irregularly dentate toward apex, pubescent on both sides; flowers on
pedicels of varying lengths in terminal, usually lax corymbs; corolla
orange-pink to scarlet, 5-8 cm. long, incurved, often narrowing below
middle of tube, truncate at apex, occasionally very minutely pubes-
cent on the exterior at a point just below the lobes, the lobes retuse-
apiculate;1 calyx tubular, glandular-pubescent (heaviest pubescence
often at apex of lobes, inside as well as without), 1.6-2.7 cm. long;
stamens usually well exserted,2 often exceeding the corolla by as
much as 1.5 cm., adnate to tube at a point about one-fifth above its
base, filaments sometimes pubescent at point of insertion; ovary ob-
long-ovoid, glabrous; style exserted, exceeding the tube by as little

1 One exception, Hutchison 1252 (Dept. Ayacucho, Prov. Lucanas) was seen,
in which the retuse lobes usually lacked an apiculum. As the corolla limbs of this
collection were also more widely expanded than is usual in C. candelilla, this may
be a hybrid.

2 Stamens of all specimens seen were exserted to some degree with the excep-
tion of Weberbauer 7287 a.

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

as 0.2 cm. to as much as 2.8 cm.; mature capsule oblong-ovoid, gla-
brous, not more than 2 cm. long, about equal to or scarcely exceeded
by the calyx lobes; seeds brown, elliptic, winged, 6-8 mm. long.

Arequipa: Prov. Arequipa, 3200-3600 m., phototype, F. M. Neg.
No. 14024, of Weberbauer 1428', Chachani, 3300 m., Rank & Hirsch
P523; 3600-3700 m., Pennell 13264; 11,000 ft., Stafford 610. — Mo-
quegua: Prov. Moquegua, Carumas, 3200 m., Weberbauer 7266 and
7287-a. — Tacna: Prov. Tarata, 2800 m., Metcalf 30363; 3840 m.,
Pearson 32; Tarata Libre, Candarave, 3600 m., Weberbauer 7371.

Cantua cuzcoensis Infantes Vera, Lilloa 31: 102. 1962.

Erect, branching shrub or small tree to 5 m., of variable pubes-
cence.1 Leaves sessile to short-petioled, to 3.8 cm. long and 1.1 cm.
wide, oblanceolate to elliptic, acute, tapering to base, usually entire
but occasionally with a few teeth unequally placed toward apex,
usually with some pubescence at least along mid vein and /or near
base; inflorescence of many-flowered, terminal corymbs, flowers borne
on pedicels of vaying lengths (not more than 15 mm.) ; corolla 4-
5 cm. long, usually pink to crimson (tube sometimes cream or yellow-
ish tinged with pink), tubular with spreading lobes when completely
expanded, apex of lobes shallowly and unevenly cleft, borders of
lobes minutely ciliate to pilose; calyx tubular-campanulate, 1.6-2 cm.
long, lobes triangular and acute, pubescence usually minute and scat-
tered, heavier near base and at apex of lobes; stamens usually well
exserted,2 sometimes exceeding the corolla by as much as 1.8 cm.,
inserted about 4-5 mm. above base of tube; ovary oblong, to 12 mm.
long, glabrous; style exserted, to 6 cm. long; mature capsule oblong,
to 14 mm. long, usually containing 17-21 winged seeds.

Apurimac: Prov. Abancai, 11,000 ft., Balls B6903; Prov. Anda-
huaylas, 2500 m., Stork & Horton 10729. — Cuzco: Without locality,
Gay (Oct. 1839-Feb. 1840); Prov. Calca, 3150 m., Vargas 215;
3050 m., Vargas 9785. — Junin: Prov. Huancayo, Quebrada Occo-
pilla, 3345 m., Soukup 3627.

Cantua flexuosa (R. & P.) Pers. Syn. PL 1: 187. 1805; Infantes
Vera, Lilloa 31: 104. 1962. Periphragmos flexuosus Ruiz & Pa von,
Fl. Peruv. 2: 17, t. 131. 1799.

1 Without a lens, this species often appears glabrous.

2 Stamens were well exserted on all specimens seen except Soukup 3627, cited
by Infantes Vera in her original description but all other characters match the
description.

FLORA OF PERU 119

Erect, branching shrub or small tree, occasionally to 5 m. tall,
always somewhat pubescent, although degree of pubescence variable.
Leaves rarely to 9 cm. long and 3.6 cm. wide but averaging about
4 by 1.7 cm., short-petioled, elliptic, lanceolate to ovate, acute, taper-
ing to base, usually entire but occasionally with a few teeth toward
apex, usually somewhat pubescent, at least on underside along mid-
vein; flowers on pedicels of varying lengths, usually crowded in dense,
terminal or sub-terminal corymbs; corolla campanulate, somewhat
incurved, white, 1.5-2.3 cm. long, exceeding the calyx by 1-1.5 cm.,
tube sometimes pubescent on upper part, corolla lobes retuse to
deeply cleft, borders of lobes minutely ciliate to nearly pilose; calyx
0.8-1.5 cm. long, campanulate, minutely pubescent with heavier pu-
bescence at base and on tips of lobes, lobes triangular, unequal;
stamens exserted, exceeding the corolla by 1-1.5 cm., adnate to tube
at a point only 1 or 2 mm. above base; ovary oblong-ovoid, glabrous;
style exserted, to 4 cm. long; mature capsule oblong-ovoid, gla-
brous, to 2.5 cm. long, at least twice as long as the calyx; seeds
numerous, brown, winged, oblong to ovate, to 9 mm. long.

Apurimac: Prov. Abancai, Tilca, 2800 m., Rauh & Hirsch Pi 540;
3050 m., Vargas 1972. — Ayacucho: Prov. Huanta, Parcora, 3000 m.,
Killip & Smith 22182;Prov. La Mar, 3100-3200 m., Weberbauer 5554.
— Cuzco: Prov. Anta, 3100 m., Vargas 7281; Prov. Calca, 3200 m.,
Marin 2399; Prov. Paruro, 3200 m., Vargas 442; Prov. Paucartambo,
3000 m., Vargas 11184- — Huancavalica : Prov. Huancavalica, 2400
m., Stork & Horton 10895; Prov. Tayacaja, 3250 m., Stork & Horton
10262. — Huanuco: Type collection, without locality, Ruiz & Pavdn;
Prov. Ambo, 2800 m., Seibert 2215; Prov. Huanuco, Dunkafail, Sa-
wada P75; Muna, 7000 ft., Macbride 3923; 8000 ft., Pearce (May,
1863) ; Yanahuanca, 10,000 ft., Macbride & Featherstone 1182. — Ju-
nin: Prov. Cerro de Pasco, Huariaca, 3200 m., Asplund 11967; 7000
ft., Sandeman (June, 1938); Prov. Jauja, Mito, 9000 ft., Macbride &
Featherstone 1565. — Lima: Prov. Lima, Rose 18633. — San Martin:
Prov. San Martin, Uspayaca, 8500 ft., Macbride & Featherstone 1292.
Also in Bolivia.

Cantua longifolia Brand, Fedde Rep. Spec. Nov. 20: 46. 1924;
Infantes Vera, Lilloa 31: 83. 1962.

Erect, branching shrub, to 6 m. or perhaps taller, somewhat pubes-
cent throughout except on larger stems, probably becoming glabres-
cent with age. Leaves 8-16 cm. long, 4-6 cm. wide, short-petioled,
elliptic to obovate, acute, tapering to base, with 7-20 teeth on the
upper two- thirds of the leaf margins; flowers borne on pedicels of

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

varying lengths (1-5 cm.) in lax, terminal corymbs; corolla greenish-
yellow, funnelform, 2-2.5 cm. long, exceeding the calyx by 1-1.5 cm.,
with some scattered pubescence on upper part of tube, lobes irregu-
larly bilobate; calyx cup-shaped to campanulate, to 1.3 cm. long,
minutely pubescent, lobes triangular, unequal ; stamens exserted, ex-
ceeding the corolla by 1-1.5 cm., adnate to tube at a point 5-12 mm.
above the base; ovary oblong-ovoid, glabrous; style exserted, to 3 cm.
long, mature capsule oblong, glabrous, to 2.5 cm., about twice as
long as the calyx; seeds numerous, brown, winged, suborbicular to
pyriform, to 4 mm. long.

Libertad: Prov. Pataz, 2600 m., Weberbauer 70381 and 7039.
-Without locality: Ruiz & Pavon (1778-1788).

Cantua pyrifolia Juss. ex Lam. Encycl. Me"th. 1: 603. 1783;
Illust. 1, t. 106, f. 1. 1791; Willd. Sp. PL 1(2): 879. 1797; Juss. in
Ann. Mus. Paris 3: 117, t. 7. 1804; HBK. Nova Gen. & Sp. 3: 161.
1818; Benth. in DC. Prodr. 9: 320. 1845; Hook. Bot. Mag. 74: t. 4386.
1848; Lemaire, Fl. des serres 4: t. 383. 1848; Infantes Vera, Lilloa 31:
97. 1962. C. peruviana Gmel. Syst. 347. 1791. C. loxensis Willd.
in Roem. & Schult. Syst. Veg. 4: 369. 1819. C. ochroleuca Brand in
Engler, Pflanzenr. 4, 250: 23. 1907.

Erect, branching shrub or small tree to 6 m. tall, always some-
what pubescent, although degree of pubescence is quite variable.
Leaves 3-10 cm. long, 1.5-4 cm. wide, short-petioled, elliptic to obo-
vate, acute, tapering to base, usually entire but occasionally with a
few teeth toward apex, usually somewhat pubescent, at least on un-
derside along mid- vein ; flowers borne on pedicels of varying lengths,
in lax, terminal corymbs; corolla funnelform, greenish-yellow, 2.5-
3 cm. long, exceeding the calyx by 1-1.5 cm., tube often pubescent,
lobes usually apiculate but rarely somewhat retuse, borders of corolla
lobes minutely ciliate; calyx about 7 mm. to not more than 1.5 cm.
long, cup-shaped to campanulate, minutely pubescent, lobes triangu-
lar; stamens exserted, exceeding the corolla by at least 1 cm., adnate
to tube at a point 4-9 mm. above base, filaments often pubescent at
point just above insertion; ovary oblong-ovoid, glabrous; style ex-
serted, to 3.5 cm. long; mature capsule oblong, glabrous, at least
twice as long as the calyx; seeds numerous, brown, winged, pyriform
to elliptic, to 6 mm. long.

1 The specimens cited by Brand, in the Berlin Herbarium, are presumed de-
stroyed; duplicate specimen of Weberbauer 7088, in the Field Museum Herbarium,
is here designated as lectotype.

FLORA OF PERU 121

Amazonas: Prov. Bongara, 2400 m., Wurdack 977; Prov. Chacha-
poyas, Mathews 3046. Also in Ecuador.

Cantua quercifolia Juss. Ann. Mus. Hist. Nat. Paris 3: 118.
1804; Poiret in Lam. Encycl. Me"th. Suppl. 2: 80. 1811; Benth. in
DC. Prodr. 9: 320. 1845; Brand in Engler, Pflanzenr. 4, 250: 20. 1907;
Infantes Vera, Lilloa 31: 80. 1962.

Erect, branching shrub to 3 m. tall, glandular-pubescent through-
out. Leaves subsessile (occasionally clasping) to short-petioled,
elliptic to lanceolate-ovate, acute, sinuate-crenate, cuneate at base
or lyrate (rarely auriculate), usually 3-9 cm. long, viscid, densely
glandular-pubescent; flowers short-pedicellate, few to many in ter-
minal corymbs; corolla salverform, white, or tube may be yellow
with white lobes, 3^4 cm. long and 2.5 cm. across, exceeding the
calyx by about 2 cm. ; calyx tubular-campanulate, glandular-pubes-
cent, 1.5-2.8 cm. long, lobes narrowly lanceolate, acute, 3-nerved;
stamens exserted, exceeding the corolla by 1 cm. or more, inserted
near base of tube; ovary oblong-ovoid, glabrous; style exserted, 3.5-
5 cm. long; mature capsule oblong, glabrous, to 2.5 cm. long, exceeded
by calyx lobes; seeds numerous, brown, winged, the wing usually
notched on one side, orbicular or suborbicular, to 3 mm. long.

Amazonas: Prov. Chachapoyas, 2000-2400 m., Wurdack 575;
7000 ft., Sandeman 27. — Cajamarca: Santa Cruz, Soukup 4825.
— Piura: Prov. Huancabamba, 2000-2100 m., Ferreyra 15662; 2500
m., Scolnik 1429. — Without locality: Mathews 1500, 3047. Also re-
ported from Ecuador.

COBAEA Cav.

Liana, usually somewhat pubescent. Leaves alternate, pinnately
compound, leaflets entire, lateral leaflets broad, short-petioled, the
terminal leaflet usually modified as a tendril; flowers solitary on long
peduncles in leaf axils; corolla green, greenish- white, yellow, pink,
violet or purple (rarely striped), large, campanulate, lobes orbicular
to ovate, sometimes caudate; calyx persistent, herbaceous, the 5 se-
pals divided almost to base, regular; stamens inserted near base of
corolla tube, filaments slender, anthers versatile, sometimes appear-
ing sagittate; style included or exserted, stigma 3-lobed or branched;
disc 5-lobed; ovary superior, glabrous, elliptic-ovate, tricarpellate,
trilocular; mature capsule exceeding calyx, elliptic or elliptic-ovoid,
dehiscing septicidally, each locule containing 2 to several large, flat,
winged seeds.

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

Of the 19 described species, only three are known from Peru. The
genus ranges from Mexico to Venezuela and in the Andes from Co-
lombia to northern Chile, always in humid, montane forests. Cobaea
scandens is in cultivation in Europe as well as the Americas ; it is vari-
ously known in South America as "Enredadera," "Ccoto-ccoto," and
'Trinitaria."

Corolla lobes not caudate

Calyx lobes broadly rounded at apex, mucronate C. scandens

Calyx lobes lanceolate, acute C. campanulata

Corolla lobes linear-caudate C. penduliflora

Cobaea campanulata Hemsl. The Garden 17: 352. 1880. Rosen-
bergia campanulata House, Muhlenbergia 4: 24. 1908.

Liana; stems largely glabrous. Leaves subsessile to short-peti-
oled, petioles glabrous or occasionally somewhat pubescent, leaflets
entire, glabrous or minutely ciliate, elliptic to obovate, acuminate,
the lower pair usually rounded at base, subsessile, others oblique at
base with longest cilia often near base, short-petioled, mature leafiest
averaging 4-8 cm. long, 2-3 cm. wide; flowers solitary on peduncles
12-18 cm. long; corolla tubular-campanulate, white or yellowish,
4-6 cm. long, exceeding the calyx by 2.5-4 cm., glabrous without,
lobes ovate; calyx segments united only at base, sepals linear-lanceo-
late, acute, long-ciliate; stamens exserted or about equalling the co-
rolla lobes, inserted at a point ca. 1 cm. above base of tube, filaments
wooly above point of insertion, pubescence extending down onto the
tube; ovary elliptic-ovoid, glabrous, style exserted or included; ma-
ture capsule elliptic-ovoid, glabrous, 4-6 cm. long, exceeding calyx
lobes by 2-3 cm.; each lobe of disc shallowly bilobate; seeds light
brown, ca. 10 mm. long, 6 mm. wide, broadly winged.

Piura: Prov. Ayavaca, valley of Quiros river, 1000-1100 m., Weber-
bauer 6425; 1500 m., Weberbauer 6397; Prov. Huancabamba, west of
Canchaque, 1200 m., Stork 11395. Also reported from Chile.

Cobaea penduliflora (Karst.) Hook, in Bot. Mag. 95: t. 5757.
1869; Brand in Engler, Pflanzenr. 4, 250: 28. 1907; Pittier, Bol. Soc.
Venez. 3(26): 303-306. 1936. Rosenbergia penduliflora Karst., Fl
Columb. Sp. Sel. 1: 27, t. 14- 1858. C. hookeriana Stand!., Contr.
U. S. Nat. Herb. 17: 451, t. 26. 1914.

Liana, usually somewhat pubescent, at least at stem nodes and
on petioles. Leaves sessile or subsessile, the 4-6 petiolulate leaflets,

FLORA OF PERU 123

glaucous, sometimes with scattered pubescence, entire, obovate to
elliptic, acute to somewhat acuminate and mucronate, oblique at
base, mature leaflets 3-11 cm. long, 2.5-5 cm. wide; flowers solitary
on peduncles 20-28 cm. long; corolla greenish, the tube campanulate,
said to be 1-5 cm. long, minutely pubescent, the lobes becoming nar-
rowly linear, 4-5 cm. long, 1-2 mm. wide, obtuse at apex or obscurely
bilobate; calyx segments united only at base, linear-lanceolate, nar-
rowly apiculate, minutely puberulent, 2.5-3 cm. long, 0.7-0.8 cm.
wide; stamens exserted at anthesis, 9-12 cm. long, filaments wooly
above point of insertion near base with pubescence extending down
onto the tube; ovary ovoid-acuminate, glabrous, style exserted, 10-
13 cm. long; mature capsule ovoid-trigonal, glabrous, exceeding calyx,
4.5-5 cm. long; each lobe of disc bilobate; seeds light brown, 2.5-3 cm.
long, 1-1.3 cm. wide, emarginate at base, broadly winged.

Although I saw no Peruvian collections, Brand's citation reads:
"Ost-Peru: Tarapoto (Spruce n. 4353, Herb. Boissier)." It is cer-
tainly to be expected in Peru as it is known from the Andean region
of Venezuela and Ecuador.

Cobaea scandens Cav. Icon. PL 1: 11, 1. 16, 17. 1791; Benth. in
DC. Prodr. 9: 322. 1845; Brand in Engler, Pflanzenr. 4, 250: 24. 1907.
Rosenbergia scandens House, Muhlenbergia 3: 23. 1908.

Liana with some scattered pubescence on stem and petioles.
Leaves sessile, leaflets entire, glabrous (rarely minutely ciliate) , acu-
minate, the upper two pairs short-petioled, elliptic to obovate, oblique
at base, the lower pair usually auriculate, sessile or subsessile, mature
leaflets averaging 7-8 cm. long, 3-3.5 cm. wide, but may attain 10 by
5 cm. ; flowers solitary on peduncles 15-22 cm. long; corolla campanu-
late, green, greenish- white, pink, rose, purple, or variously striped,
5-6 cm. long, exceeding calyx by 2.5-3.5 cm., sometimes with scat-
tered pubescence on outside of tube beneath lobes, lobes orbicular to
obovate; calyx broadly campanulate, glaucous without but minutely
short-pubescent within, the lobes 2.5-3.5 cm. long, 2-3 cm. wide,
undulate, mucronate; stamens exserted, filaments wooly above point
of insertion near base of tube with pubescence extending down onto
tube; ovary elliptic-ovoid, glabrous, style slightly exceeding or about
equalling corolla tube; mature capsule elliptic-ovoid, glabrous, 5-
7 cm. long, exceeding calyx lobes by 2.5-3 cm.; each lobe of disc
deeply bilobate; seeds light brown, ca. 16 mm. long, 10 mm. wide,
broadly winged.

Amazonas: Prov. Chachapoyas, Rio Utcubamba, 2150-2250 m.,
Hutchison & Wright 4915. — -Apurimac: Prov. Apurimac, Abancai,

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

2300 m., Soukup 723; 7500 ft., Saunders 763; Prov. Andahuaylas,
2700 m., Stork & Horton 10773. — Junin: Prov. Tarma, Lucumayo
Valley, 1800-3600 m., Cook & Gilbert 1377; Monte Rico La (Merced),
Soukup 3509 ; Utcuyacu, 1800 m., Woytkowski 35385. Western South
America, Venezuela to northern Chile.

GILIA Ruiz & Pavon

Annual, biennial or perennial herbs, usually somewhat pubescent,
often glandular. Leaves alternate, entire and linear or pinnately dis-
sected or incised ; inflorescences usually cymose, loose cymes or dense
heads, or solitary flowers; corolla white, blue, violet, pink or rarely
yellow, showy to small and inconspicuous, regular, usually funnel-
form but sometimes nearly salverform; calyx persistent, accrescent,
regular, the 5 sepals united by a hyaline membrane for one-third to
one-half their length; stamens inserted on tube, usually slightly un-
equal in length, exserted or included, anthers oval; style included or
only slightly exserted, usually slender with 3 stigmatic branches;
ovary elliptic-ovoid to pyriform; mature capsule ovoid, trilocular,
dehiscing loculicidally, each locule containing 1-many small brown
seeds appearing winged when immature but ovoid to trigonous, with-
out wing, when mature; seeds mucilaginous when wetted.

This extremely heterogenous genus ranges from southern British
Columbia through the mountains, southwestern plains and deserts of
North America, eastward through Texas to the south Atlantic coast,
and recurs in mountains, deserts, and coastal lomas of western South
America. Of the 56 reported species, three are known from Peru.
A few species have been grown as garden ornamentals.

Stamens inserted in sinuses of corolla lobes

Plants erect, to 20 cm. tall G. laciniata

Plants with short central stems, either decumbent or developing

decumbent lateral stems
Plants 1-3 cm. tall; leaves pinnately dissected with segments

narrowly linear G. laciniata var. alpina

Plants to 10 cm. tall; leaves irregularly incised, broad

G. lomensis

Stamens inserted at a point between base of corolla tube and midway
of throat G. glutinosa

Gilia glutinosa Phil. Linnaea 30: 196. 1859 and Anal. Univ.
Chile 90: 212. 1895; Johnston, Contr. Gray Herb. 85: 96. 1929.

FLORA OF PERU 125

G. ramosissima Phil. Cat. PL Itin. Tarapacd 53. 1891. G. glabrata
Phil. Anal. Univ. Chile 90: 213. 1895. G. ramosissima var. gla-
brata Reiche, Anal. Univ. Chile 120: 194. 1907 and Fl. Chile 5: 153.
1910. G. cobijanensis Brand, Pflanzenr. 4, 250: 98. 1907. G. cha-
chanensis Johnston, Contr. Gray Herb. 70: 82. 1924.

Annual or perennial, 5-20 cm. tall, glandular-pubescent, viscid
throughout. Leaves narrowly linear, 0.5-1 mm. wide, 0.5-2 cm.
long, not more than once dissected ; flowers solitary or 2 in leaf axils,
on pedicels 1-3 cm. long, corolla 6-12 mm. long, rotate to campanu-
late, white, blue, violet, or tube white with blue or violet lobes, lobes
obtuse to obovate or apiculate; calyx 5-8 mm. long, lobes lanceolate,
acute to acuminate; stamens included, filaments inserted near base
of corolla to about midway of throat; ovary elliptic-ovoid, glabrous;
mature capsule ovoid, glabrous, 5-8 mm. long, equal to or slightly
exceeding calyx; each locule containing numerous small, brown seeds.

Arequipa: Prov. Arequipa, Isern 2138; Misti Volcano, Cardenas &
Rodriguez 6; 2300-2400 m., Pennell 13055; 2500 m., Rauh & Hirsch
P569; 2600 m., Vargas 7964; Tingo, 2100-2300 m., Pennell 13113;
Chachani Mountain, 3300 m., Hinkley 25. — Moquegua: Prov. Mo-
quegua, 1900-2000 m., Weberbauer 7428. — Tacna: Prov. Tacna,
500-600 m., Ferreyra 12510, 12526, 12536.

Gilia laciniata Ruiz & Pavon; Fl. Peruv. 2: 17, t. 123. 1799;
Benth. in DC. Prodr. 9: 312. 1845; Grant, Aliso 6(2): 73. 1966.
G. erecta Hieron. Bol. Acad. Nac. Cordoba 3: 368. 1879; G. laciniata
var. erecta Brand, Pflanzenr. 4, 250: 106. 1907. Thouinia multifida
Dombey ex Juss. Ann. Mus. Paris 3: 119. 1804. Cantua breviflora
Juss. Ann. Mus. Paris 3: 119. 1804; C. laciniata Poir. Lamarck,
Encycl. Me"th. Bot. Suppl. 2: 81. 1811. Ipomeria albida Nutt. Gen.
N. Amer. PL 1: 125. 1818.

Annuals to 20 cm. tall, erect, glandular-pubescent. Leaves pin-
nately dissected 1-3 times, 1-5 cm. long, the segments narrowly linear
to 2 mm. wide, the mid-section of blade rarely to 4 mm. wide; flowers
1-several in leaf axils on pedicels 0.5-7 mm. long, corolla funnelform,
white, pink, lavender or purple, or tube white with colored lobes,
lobes obtuse to obovate, tube included in calyx; calyx 6-8 mm. long,
lobes lanceolate, acute; stamens usually included but may be short
exserted, filaments inserted in sinuses of corolla lobes; ovary elliptic-
ovoid, glabrous; mature capsule ovoid, glabrous, 5-7 mm. long, ex-
ceeded by calyx lobes, each locule containing numerous small, brown
seeds.

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

Apurimac: Prov. Grau, 3000 m., Vargas 5728. — Arequipa: Prov.
Arequipa, 2600-2700 m., Pennell 13190, 13240; 3355 m., Hinkley 19;
8540 ft., Sandeman 3951, 3999; Tiabaya, 2100-2200 m., Pennell
13097; Rio Blanco, 12,000 ft., Macbride & Featherstone 737; Prov.
Islay, lomas de Mollendo 500-600 m., Ferreyra 6376. — Cuzco: Prov.
Cuzco, Sicuani, 13,500 ft., Stafford 521-A. —La Libertad : Prov. Tru-
jillo, 550 m., Lopez M. 1527. — Lima: Prov. Chancai, Huaura, Ruiz
(1778-1788); Prov. Lima, Lurin, 200 ft., Macbride 5964; San Bar-
tolo, 500-800 ft., Saunders 166. — Moquegua: Prov. Moquegua,
lomas de Mostacilla, 50-300 m., Vargas 8595; Carumas, 3200 m.,
Weberbauer 7302; Torata, 2200-2300 m., Weberbauer 7400. — Puno:
Prov. Puno, Araranca, 4100-4300 m., Pennell 13435.

Gilia laciniata var. alpina Weddell, Chlor. And. 2: 81, t. 58,
1859. G. alpina Brand, Pflanzenr. 4, 250: 107. 1907.

Annuals, 1-3 cm. tall, with short central stems and decumbent
lateral stems, more or less glandular-pubescent throughout. Leaves
pinnately dissected 1-3 times, 0.5-2.5 cm. long, the segments nar-
rowly linear to 2 mm. wide; flowers 1 -several in leaf axils, short pe-
dicillate, corolla funnelform, pink, violet or purple, tube included in
calyx; calyx 3-6 mm. long, lobes lanceolate, acute; stamens usually
included but may be short exserted, filaments inserted in sinuses of
corolla lobes; ovary elliptic-ovoid, glabrous, mature capsule ovoid,
glabrous, 3-4 mm. long, exceeded by calyx lobes, each locule contain-
ing numerous small, brown seeds.

Junin: Prov. Yauli, 13,500 ft., Macbride & Featherstone 917.
—Puno: Prov. Puno, Santa Rosa, 13,500 ft., Stafford 517; Chuqui-
bambilla, 3850-3900 m., Pennell 13410.

Although Dr. Grant1 placed the variety alpina in synonymy with
G. laciniata, I have returned it to varietal status, not only because of
its short, decumbent growth habit but also because its calyces are
3-6 mm. long, whereas those of G. laciniata are 6-8 mm. long.

Gilia lomensis Grant, Aliso 6(2) : 72, fig. 4- 1966.

Annuals to 10 cm. tall, with short central stem and decumbent
lateral stems villous to glandular pubescent; leaves of type specimen
(garden progeny) irregularly incised, 0.5-2 cm. long, 0.3-1 cm. broad;
leaves of specimens cited as representative pinnately dissected, the
segments usually narrowly linear, 1-3 cm. long, either entire or in-
cised, rarely broadened to 2 mm.; flowers 1-several in leaf axils, on

1 Aliso 6(2): 73. 1966.

FLORA OF PERU 127

pedicels 0.5-4 mm. long, corolla funnelform, white or pink, tube in-
cluded in calyx; calyx 5.5-7.5 mm. long,1 lobes acute, stamens included
or short-exserted, filaments inserted in sinuses of corolla lobes, ovary
elliptic-ovoid, glabrous; mature capsule ovoid, glabrous, 5-6 mm.
long, exceeded by calyx lobes; each locule containing numerous small,
brown seeds.

Arequipa: Prov. Camana, 125 m., Worth & Morrison 15650.
—Lima: Prov. Chancai, south slope of Cerro, 80 m., Stork, Beetle &
Morrison 9093; V. & A. Grant 2244, TYPE, grown in California from
seeds collected on southwestern slope of Cerro San Geronimo by
0. Velarde and M. Reiche.

In order to avoid confusion, it should be made clear that the type
material, which is garden progeny, differs markedly from the speci-
mens collected in Peru and cited as representative. The garden plant
is much larger, more robust, more profusely branched, more decum-
bent, with broad leaf blades irregularly incised, not pinnately dis-
sected. The leaves of the two small Peruvian specimens show the
pinnate dissection typical of G. laciniata, although a few leaves dis-
play a slight broadening (to 2 mm.) of mid-blade and segment. Of
the ten small plants on the two specimen sheets seen, eight were erect
and only two showed evidence of developing decumbent lateral stems.
Further, slightly broadened leaves were observed on several collec-
tions of G. laciniata, including two from lomas, Vargas 8595 and
Ferreyra 6376; these plants are quite tall and erect, with calyces 7-
9 mm. long in fruit. Unusually broad leaf segments occasionally
occur on specimens of G. laciniata var. alpina, i.e., Stafford 517 and
Pennell 13410. As these plants are decumbent with mature calyces
shorter than those of the typical variety, G. lomensis may prove to
be even more closely related to the var. alpina than to G. laciniata
sens. str. It would be helpful if we knew how the two latter plants
would develop in garden environment. It is hoped that additional
collections and continued investigations by such workers as Dr. Grant
and his associates will eventually clarify the complex inter-relation-
ships that are apparent in Gilia.

HUTHIA Brand

Erect, branching shrubs, the branches cylindrical, glandular-
pubescent throughout. Leaves linear-lanceolate to narrowly linear,

1 According to the original description, length of fruiting calyx is only 6 mm.
As a few calyces to 7.5 mm. were seen on the type material, this may indicate a
closer relationship with G. laciniata.

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

alternate or fasciculate, always deeply pinnatifid, the segments
fleshy, obtuse to obovate, revolute; flowers usually few (sometimes
solitary) in terminal or axillary corymbs; corolla blue or violet, tubu-
lar or funnelform, two to five times as long as the calyx, apex of
corolla lobes obovate-irregular to truncate-erose; calyx persistent,
somewhat accrescent, tubular to tubular-campanulate, more or less
zygomorphic, 3-5 (6) lobed, the lobes 3-nerved, subulate to acumi-
nate, 2 lobes sometimes more deeply divided than others so that
calyx appears sub-bilabiate; stamens included, partly adnate to and
inserted on tube at a point one-third to one-half of the corolla length
above the base, filaments somewhat dilated at base, pubescent near
point of insertion, or papillose-hirsute for entire length, anthers ver-
satile; style included, filiform, stigmas 3-branched; disc irregularly
5-lobed; ovary superior, glabrous, oblong-ovoid, 3-carpellate, 3-locu-
lar, with numerous ovules in each locule; mature capsule oblong,
glabrous, dehiscing loculicidally by 3 valves, containing 60-80 small,
trigonous, narrowly winged seeds.

The genus is obviously very close to Cantua, but may be distin-
guished by its characteristic foliage, blue to violet corolla, and its
trigonous seeds which are never more than 2.5 mm. long (the more
broadly winged seeds of Cantua ranging in length from 3-11 mm.),
although these characters hardly seem of generic worth.

Only two species of Huthia are known, both reported only from
the Department of Arequipa in Peru.

Corolla 2-3 cm. long H. coerulea

Corolla 4-5.5 cm. long H. longiflora

Huthia coerulea Brand, Bot. Jahrb. 42: 175. 1908.

Erect, branching shrub to 1 m. tall, glandular-pubescent through-
out. Leaves alternate and usually fasciculate, 0.5-3.5 cm. long, 1-
6 mm. wide, sessile to short-petioled, linear to linear-lanceolate,
deeply pinnatifid, the segments obtuse to obovate, revolute; the up-
permost bract-like leaves sometimes entire, or the upper one entire
and the lower partly pinnatifid; flowers short-pedicellate, usually
few in terminal or axillary corymbs, sometimes solitary in leaf axils;
corolla funnelform, blue or blue-violet, glabrous, 2-3 cm. long, usu-
ally 2-3 times longer than the calyx, corolla lobes irregularly obovate,
almost half the length of the corolla; calyx tubular to tubular-cam-
panulate, glandular-pubescent, 7-12 mm. long, the lobes acuminate;
stamens included, inserted on tube at a point about 5-7 mm. above
its base, filaments densely papillose-hirsute with longer pubescence

FLORA OF PERU 129

at base; ovary oblong-ovoid, glabrous; style included, 1.5-2.5 cm.
long; mature capsule oblong-ovoid, glabrous, 0.5-1 cm. long, ex-
ceeded by the calyx lobes; seeds numerous, brown, narrowly winged,
trigonous, to 2.5 mm. long.

Arequipa: On slopes of Misti Volcano, Cdrdenas & Rodriguez 4',
8300 ft., Saunders 345; 9000 ft., Sandeman 3754; arid plains, over
3200 m., West 7139; on Mt. Chiwata, 2200 m., Eyerdam & Beetle
22110; cerros de Arequipa, 2350 m., Ferreyra 2576; 8000 ft., Staf-
ford D-8.

Huthia longiflora Brand, Bot. Jahrb. 50, Beibl. Ill: 51. 1913.

Erect, branching shrub to 1 m. tall, densely glandular-puberulent,
canescent. Leaves alternate, often fasciculate, sessile to short-peti-
oled, narrowly linear, pinnatisect, 1-20 mm. long, 1-3 mm. wide, the
short segments rounded, revolute; flowers long-pedicillate , solitary
or few in lax, terminal corymbs; corolla tubular, violet, glabrous,
4-5.5 cm. long, 3-5 times longer than the calyx, corolla lobes obtuse
to truncate and erose, 1-1.5 cm. long; calyx tubular, glandular-
pubescent, 1-1.5 cm. long, the lobes lanceolate, subulate to acumi-
nate; stamens included or equalling the corolla length, glabrous ex-
cept becoming wooly near point of insertion, about 1 cm. above base
of tube; ovary oblong-ovoid, glabrous; style about equalling corolla
length; mature capsule pyriform to oblong, glabrous, 7-10 mm. long,
exceeded by calyx lobes; seeds numerous, brown, trigonous, narrowly
winged, to 2.5 mm. long.

Arequipa: Near confluence of Rio de Lomas and Rio Yauca,
1900-2000 m., Weberbauer 5752.

PHLOX L.

Erect, branching, diffuse or caespitose perennials or annuals, her-
baceous to suffrutescent (not in ours) . Leaves entire, usually oppo-
site, sessile to petioled; flowers in terminal corymbs or cymes or in
pairs in leaf axils (rarely solitary) ; corolla salverform, tube slender
and throat usually constricted, often showy but may be inconspicu-
ous, red, purplish, blue, cream or white; calyx tubular or campanu-
late, 5-lobed, persistent, accrescent, the lobes acute to acuminate,
with a membrane below the sinuses; stamens short, included, un-
equally inserted; style usually slender, stigmas 3-branched or lobed;
ovary superior, 3-loculed, ovoid to oblong; mature capsule ovoid to
oblong, dehiscing loculicidally, seeds oblong, 1-few in each locule.

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

The genus ranges throughout temperate North America and
northern Asia, but only one species, Phlox gracilis, is known from
South America. There are certain differences between this poly-
morphic species and most of the remaining species of Phlox (P. gra-
cilis has a smaller corolla with slightly flaring base, and seeds that
become somewhat mucilaginous when wetted), which to some bot-
anists justify its segregation as a minor genus Microsteris.1 However,
I agree with Mason2 that the differences are minor and that its rela-
tionship within the family is more clearly shown by including it in
Phlox.

Phlox gracilis (Dougl. ex Hook.) Greene, Pittonia 1: 141. 1887.
Gilia gracilis (Dougl.) Hook. Bot. Mag. 56: t. 2924- 1829; Brand in
Engler, Pflanzenr. 4, 250: 88. 1907. G. gracilis (Dougl.) Hook. ssp.
eu-gracilis Brand and varieties stricta (Greene) Brand, eritrichoides
(Griseb.) Brand, glabella (Greene) Brand, micrantha (Kellogg) Brand,
and subsp. humilis (Dougl.) Brand, I.e. 89-91. G. gracilis var. minu-
artioides (Franchet) Borsine, Lilloa 8: 212. 1942. Collomia gracilis
(Dougl.) Lindley, Bot. Reg. 19: 1. 1622. 1833 and DC. Prodr. 9: 308.
1845. C. gracilis var. congesta Wedd., Chlor. And. 2: 80, pi. 58-A.
1859; C. gracilis var. andicola Wedd., I.e. C. micrantha Kellogg,
Proc. Calif. Acad. 3: 18. 1863. C. eritrichoides Griseb. Abh. Konigl.
Ges. Wissensch. Gb'ttengen 6: 129. 1854. Navarretia gracilis (Dougl.
ex Hook.) Kuntze, Rev. Gen. PI. 1: 433. 1891. Microsteris gracilis
(Hook.) Greene, Pittonia 3: 300. 1898. M. gracilis ssp. humilis
(Greene) V. Grant in Munz, A California Flora 478. 1959. M. hu-
milis (Dougl.) Greene, Pittonia 3: 301. 1898. M. micrantha (Kell.)
Greene, I.e. 303. Polemonium morenonis 0. Ktze. Rev. Gen. PL
3(2): 203. 1898.

Small, branching annual, more or less glandular-pubescent through-
out, diffuse or erect, 1-12 cm. tall. Leaves entire, linear or linear-
oblong to oblanceolate or spatulate, 1-3 cm. long, predominantly
opposite (the lower always opposite, only the uppermost alternate),
sessile to short-petioled ; flowers usually in pairs (rarely 1 or 3) in leaf
axils; corolla salverform, 5-12 mm. long (usually less than 8 mm.),
the tube white or cream, the lobes bilobate, white, cream, rose, laven-
der or purple, exceeding the calyx lobes by only 1-3 mm.; calyx 4-

1 Wherry, Edgar T. Brittonia 5(1) : 60-63. 1943; "The Genus Phlox" in Morris
Arb. Monogr. 3: 1-174. 1955.

Grant, Verne E. Natural History of the Phlox Family, Vol. 1: 43, 46, 63,
77. 1959.

2 Mason, H. L. Madrono 6: 122-177. 1941; Madrono 10: 202-203. 1950.

FLORA OF PERU 131

10 mm. long, cylindrical, the lobes herbaceous, subequal, 3-nerved,
acute, united by a hyaline membrane, the lobes equalling or longer
than the tube; stamens included, the short filaments partly adnate
to and unequally inserted on the tube at a point a little above the
middle, anthers sagittate; style included, slender; ovary glabrous,
ovoid; mature capsule glabrous, ovoid, rupturing the calyx in age,
each locule containing 1 brown, lenticular seed, the seeds becoming
somewhat mucilaginous when wetted.

Plants with a small corolla (5-8 mm. long) were first thought to
be ssp. humilis (Greene) V. Grant. However, the corollas of all those
examined from Peru were well exserted, and the larger ones (8-
12 mm.) were often only barely exserted.

Arequipa: Arequipa, 3000-3200 m., Pennell 13232; Chachani,
11,000 ft., Stafford 618; Prov. Condesuyos, between Chuquibamba
and Salamanca, 4000 m., Weberbauer 6851. — Cuzco: Prov. Anta,
Rio Blanco, 15,000 ft., Macbride & Featherstone 815. — Huanuco:
Huillacachi, Matucana, 3300 m., Weberbauer 5725. — Puno: Prov.
Sandia, Hill (Jan.-March, 1903). Also in Bolivia, Chile and Ar-
gentina.

HECKMAN

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