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FLORA OF PERU
BY
GABRIEL EDWIN
ASSOCIATE PROFESSOR, BIOLOGY
ROOSEVELT UNIVERSITY, CHICAGO
BOTANICAL SERIES
FIELD MUSEUM OF NATURAL HISTORY
VOLUME XIII, PART V-B, NUMBER 3
MAY 21, 1971
PUBLICATION 1125
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UN.VERS.TY OP ,U,NO.S L.BRARY AT URBANA-CHAMPA.GN
FLORA OF PERU
BY
GABRIEL EDWIN
ASSOCIATE PROFESSOR, BIOLOGY
ROOSEVELT UNIVERSITY, CHICAGO
BOTANICAL SERIES
FIELD MUSEUM OF NATURAL HISTORY
VOLUME XIII, PART V-B, NUMBER 3
MAY 21, 1971
PUBLICATION 1125
Library of Congress Catalog Card Number: 36-10426
PRINTED IN THE UNITED STATES OF AMERICA
BY FIELD MUSEUM PRESS
FLORA OF PERU
GABRIEL EDWIN*
SCROPHULARIACEAE. Figwort Family
References: Benth. in DC., Prodr. 10: 186-188. 1846. Schmidt
in Martius, Fl. Bras. 8(1) : 230-231. 1863. Benth. & Hook., Gen PL
2: 913-980. 1876. Wettst. in Engler and Prantl, Pflanzenf. IV(3b) :
39-107. 1895. Kranzlin in Engler, Pflanzenf. IV, 257c: 1-122. 1907.
Pennell, Acad. Nat. Sci. Phila. Monogr. I: 1-45. 1935. Thieret,
Ceiba4: 164-182. 1954.
Annual or perennial herbs or less often shrubs, occasionally vines
(Calceolaria spp., Maurandya, Antirrhinum), glabrous to pubescent
with unbranched, usually unicellular hairs (occasionally multicellu-
lar in Calceolaria spp.), sometimes glandular or pulverulent. Leaves
usually opposite, sometimes alternate above and opposite below,
occasionally all alternate or ternate, rarely rosulate, estipulate, rarely
persistent (Hebe) , entire to usually toothed to occasionally pinnately
lobed or pinnate, rarely palmate, aciculate-linear to almost round,
minute to 25 cm. long and longer, sessile to long-petiolate; inflores-
cences various, determinate or indeterminate, simple to compound
spikes, racemes, dichasia, panicles or rarely umbellate, occasionally
flowers solitary or few-clustered, bracts or bracteoles usually present
and variously placed on the flower stalks; flowers bisexual, zygo-
morphic or rarely actinomorphic (Capraria); calyx 4-5-parted and
tube lacking or more often tube present and calyx 2 (4 -5) -toothed
or -lobed, sometimes the tube appearing to split only once and then
the calyx 2-lobed and usually at least partly one-sided on the flower,
the sepals when free occasionally valvate, all alike or one or two vari-
ously modified and wider (Bacopa), imbricate when united, all alike
or one longer (Mimulus, Calceolaria spp.) than the others; corolla
1 The author acknowledges support from the National Science Foundation,
grant GB 3560.
461
462 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
gamopetalous, the lobes 5, 4 or 2, rarely 3, usually bilabiate, the lips
equal to very unequal, lobes arched or spreading, tube sometimes
personate, long to almost lacking (Veronica), one or more anterior
petals sometimes saccate or spurred, rarely anteriorly with a fine
horn-like process at the base of a lobe (Angelonia) or the lips very
unequal and inflated (Calceolaria) or rarely sub-equal and inflated
(Calceolaria, Porodittia) ; stamens distinct, epipetalous, 5 or 4 or 2,
usually arising from the base of the tube, one or more stamens some-
times modified to a staminode, when 4, commonly didynamous, the
anterior pair frequently longer; anthers 2-celled or rarely one cell
obsolete (Buchnera, Calceolaria spp.), cells equal or unequal, contig-
uous or separated on stalks, dorsifixed or versatile, dehiscing longi-
tudinally, introrsely, sometimes splitting across the septum, rarely
by terminal pores; an annular or lateral, nectar-bearing disc usually
present, surrounding the ovary, or nectary on the reflexed edge of
the lower corolla lobe (Calceolaria) ; pistil one, ovary superior, locules
and carpels two (rarely unilocular), placentation axile; ovules numer-
ous, anatropous, placentae often conspicuous, style terminal; stigma
lobes 2, separate (Gratioleae) or united, punctiform, patelliform or
ligulate; fruit usually a many-seeded capsule, mostly septicidal,
sometimes loculicidal, occasionally indehiscent or poricidal (Antir-
rhinum, Porodittia), then locules unequal in size, the posterior smaller,
rarely a berry (Leucocarpus) ; seeds smooth or variously marked, often
angled, sometimes alate; endosperm soft, usually copious.
The family is cosmopolitan, containing over 200 genera and 3,000
species. The largest genera are Pedicularis with about 600 species
and Calceolaria with more than 500 species. The family has its great-
est concentration in South America, western North America, Africa,
and the Mediterranean regions. The limits of this family as well as
sectional and generic limits are still in dispute. A number of genera
separable only technically and with difficulty from some Solanaceae,
Gesneriaceae, Pedaliaceae, and Bignoniaceae.1
The family is of limited economic importance. Several species of
Calceolaria (Slipper flower), Antirrhinum (Snapdragon), Veronica
(Speedwell), Mimulus (Monkey flower), Penstemon (Beard-tongue),
as well as Cymbalaria, Russelia, Torenia, and Digitalis are garden
ornamentals. Digitalis yields alkaloids long used in medicine.
Thirty-two genera with 226 species occur in Peru. In addition, An-
1 The reader should remember that this and following descriptions are based
upon Peruvian material and may not be applicable elsewhere.
FLORA OF PERU 463
tirrhinum majus L. (Snapdragon) is commonly cultivated in Peru
but shows no tendency to escape.
KEY TO THE SUBFAMILIES, TRIBES, AND SUBTRIBES
1. Posterior lobes of the corolla external and overlapping in the bud.
Subfam. A. Scrophularioideae.
2. Corolla not enlarged at the base and not spurred anteriorly,
occasionally merely little saccate, with or without a fine horn-
like process at the base of the anterior lobe; capsule septicidal
or loculicidal by a simple median split, the septum breaking
from the capsule wall or rupturing, or the fruit indehiscent.
3. Stigma usually 2-lipped, when lips united plants subaquatic
and acaulescent or with repent or underground stems or with
sepals unequal in width; or stigma lips clavate; if tube al-
most lacking corolla subrotate, with 4 distinct lobes; herbage
most often glandular punctate Gratioleae.
4. Capsule usually septicidal, occasionally secondarily locu-
licidal, splitting to the base, when loculicidal the outer-
most sepal broader than the others; placenta simple; sepals
usually distinct or almost so (except Linaria spp., Limo-
sella), sometimes very unequal in width (Bacopa); leaf
blades and capsules usually glandular-punctate; inflores-
cences usually simple, racemose or the racemes sometimes
with fascicles of pedicellate flowers; anterior lobes of the
corolla usually smooth and glabrous within.
la. Gratiolinae.
4. Capsule loculicidal (secondarily barely if at all septicidal) ;
the outermost sepal about equal to the others in width;
or fruit an indehiscent berry; placenta branched; sepals
united over half their length; leaves and capsules glabrous
or pubescent but not glandular-punctate; flowers singly
pedicellate (in ours); anterior lobes of the corolla two-
ridged and pubescent within Ib. Mimulinae.
3. Stigmas united, capitate; plants terrestrial and caulescent;
sepals equal in width ; if corolla tube very short, corolla with
less than 4 distinct lobes; herbage with vesture various but
not glandular-punctate.
5. Capsule packed with hair internally; corolla weakly zy-
gomorphic, corolla- tube well developed; leaves in whorls
of threes on angled stems II. Russelieae.
464 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
5. Capsule without hair internally; corolla decidedly zygo-
morphic, the anterior lobes usually much exceeding the
posterior, tube often short; leaves opposite or the upper
alternate, occasionally ternate on terete stems.
6. A fine horn-like process developed at the base of the
corolla lobes anteriorly III. Angeloneae.
6. Corolla lobes without a horn-like process at base, lips
sometimes concave-saccate.
7. Stamens 4; corolla flattened, its lobes all evident;
the 5 sepals less than half the length of the capsule;
seeds blackish; inflorescences simply racemose;
stem quadrangular IV. Hemimerideae.
7. Stamens 2 or occasionally 3; lips of the corolla con-
cave-saccate (calceolate) or the posterior (upper)
much reduced, the individual lobes scarcely or not
at all evident; sepals 4, at least one-half the length
of the capsule, seeds brown; inflorescences usually
dichasiate or cymose; stem terete or nearly so, occa-
sionally squarrose above V. Calceolarieae.
2. Corolla anteriorly gibbose or saccate at base or with a spur at
the base of the anterior petal; capsule loculicidal, the septum
and adjacent cell wall persisting, the remaining wall splitting
irregularly VI. Antirrhineae.
1. Anterior lobes of the corolla external and overlapping in the bud.
Subfam. B. Rhinanthoideae.
8. Upper corolla lobes flattened or widely arched, often spread-
ing; anthers all distinct; stigma lobes distinct or united.
9. Stigmas distinct, plate-like; capsule septicidal and some-
times secondarily loculicidal VII. Digitaleae.
9. Stigmas united (except Gerardia sect. Dasistoma) ; capsule
loculicidal.
10. Anther cells divaricate; sepals united, 5.
VIII. Ouriseae.
10. Anther cells parallel; sepals 4 and free or united.
11. Stamens usually 2, less often 3-5 (8); corolla rotate
or subrotate, posterior lobes united; plants auto-
phytic IX. Veroniceae.
11. Stamens 4; corolla, salverform or campanulate; all
lobes evident and free; plants root-parasites or sap-
rophytes X. Buchnereae.
FLORA OF PERU 465
8. Upper corolla lobes narrowly arched, forming a definite galea
that encloses the frequently cohering anthers; pollen shed in
a mass; stigma lobes almost always united . . XL Euphrasieae.
KEY TO GENERA
Subfamily. A. Scrophularioideae (Antirrhinoideae of authors)
TRIBE I. Gratioleae, Sub tribe a. Gratiolinae
1. Leaves all alternate; corolla actinomorphic or weakly zygomor-
phic; stamens 5 or 4 Capraria.
1. Leaves opposite or verticillate or the upper sometimes alternate;
corolla basically zygomorphic, obscurely so in Scoparia', stamens 4
and didynamous or sometimes less than 4 (3 or 2) .
2. Plants acaulescent; anther cells wholly confluent. . . Limosella.
2. Plants caulescent; anther cells distinct.
3. Stem usually terete, or if quadrangular, without ridges or
wings; ridges to the anterior-lateral sinuses of the corolla
(due to adherence of the filaments) if developed low and not
projecting beyond; filaments simple; style without a semi-
persistent callosed base.
4. Anther-sacs proximate, stipes developed or not; seeds re-
ticulate; when anther-sacs stipitate, corolla 5-lobed, when
proximate corolla usually 4-lobed.
5. Sepals unequal, at least 1 or 2, usually the outer, wider
than the others; corolla glabrous within; anther-sacs
stipitate or not Bacopa.
5. Sepals equal ; corolla variously pubescent within ; anther-
sacs proximate.
6. Pedicels bibracteolate at apex (just below the calyx) ;
corolla pubescent within, usually only at the base of
the anterior lobes, occasionally on both lobes to the
base of the tube, plant repent-ascending to suberect or
erect; pedicels at least 3 times longer (up to 4 or more
times longer) than the calyx; corolla clearly zygo-
morphic, 5-lobed Conobea.
6. Pedicels ebracteolate; corolla densely hirsute within
at the bases of all the lobes; plant erect; pedicels often
less than 3 times longer than the calyx; corolla ob-
scurely zygomorphic, 4-lobed Scoparia.
466 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
4. Anther-sacs separated on stipes (connective arms); seeds
longitudinally striate and/or tuberculate-striate; corolla
4-lobed Stemodia.
3. Stem quadrangular, the angles ridged or with small wings;
two raised ridges to the anterior lateral sinuses of the corolla
(formed by the adherence of filaments) often projecting be-
yond these points as a knob-like process, the free part of the
filament appearing as a lateral outgrowth of the adherent
part; style semi-persistent, frequently enlarged and callosed
at the base.
7. Stamens 4, all antheriferous or 2 sterile (stami-
nodes) Lindernia.
7. Stamens 2 Micranthemum.
Sub tribe b. Mimulinae
8. Capsule dehiscing longitudinally; calyx tube longer than the cap-
sule; inflorescence with axillary, unbranched pedicels. .Mimulus.
8. Berry indehiscent; calyx tube shorter than the berry; inflores-
cence of much branched axillary cymes, borne on conspicuous
peduncles Leucocarpus.
TRIBE II. Russelieae
A single genus Russelia.
TRIBE III. Angeloneae
A single genus Angelonia.
TRIBE IV. Hemimerideae
9. Corolla orange, tube split to the base anteriorly; capsule second-
arily scarcely loculicidal Alonsoa.
9. Corolla white or blue or violet, split only part way anteriorly, if
at all; capsule secondarily, clearly loculicidal Basistemon.
TRIBE V. Calceolarieae
10. Anthers 2, opening by slits; corolla with two lips. . .Calceolaria.
10. Anthers 3, opening by pores; corolla with three lips. .Porodittia.
TRIBE VI. Antirrhineae
11. Herbs; corolla either with a definite spur or gibbous anteriorly
at the base, throat at least partly closed; calyx lobes clearly
imbricate.
FLORA OF PERU 467
12. Corolla spurred anteriorly at the base, throat closed by the
palate; anther-sacs parallel, distinct and oblong; stems erect
or spreading, procumbent Linaria.
12. Corolla gibbous anteriorly at the base, palate lacking (except
in M. antirrhiniflora) but throat partly closed by plaiting or
2 lines of hairs; anther-sacs confluent; plant twining (by
means of petioles and pedicels) ; perennial herbs . .Maurandya.1
11. Shrubs; corolla merely saccate anteriorly at base, throat not
closed by the palate; calyx lobes only slightly imbricate.
Galvesia.
Subfamily. B. Rhinanthoideae
TRIBE VII. Digitaleae
A single genus Digitalis.
TRIBE VIII. Ouriseae
A single genus Ourisia.
TRIBE IX. Veroniceae
13. Stamens more than 2, usually 3-5 (-8) ; sepals 4-5 (-8) united
into a campanulate calyx Sibthorpia.
13. Stamens 2; sepals 4, rarely 5, essentially free, calyx tube if de-
veloped, very short, not campanulate.
14. Herbs Veronica.
14. Shrubs or small trees Hebe.
TRIBE X. Buchnereae
15. Stamens with both anther cells developed ; corolla campanulate
or when salverform 8-15 cm. long or more; calyx campanulate or
when tubular, the tube 2.5-6.0 cm. long or longer.
16. Corolla salverform; calyx tubular; capsule ellipsoid; seeds
linear Escobedia.
16. Corolla and calyx campanulate or almost so; seeds usually
broader than linear, when linear borne in a subglobose cap-
sule Gerardia.
15. Stamens with only one anther cell developed; corolla salverform,
the tube less than 3.0 cm. long; calyx tubular, the tube not ex-
ceeding ca. 1.0 cm. long Buchnera.
1 Antirrhinum spp. widely cultivated in Peru treated at the end of this genus,
in text.
468 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
TRIBE XI. Euphrasieae
17. Anther-sacs equally placed; the leaves usually opposite (some
Bartsia spp. alternate).
18. Upper lip of the corolla reflexed-margined Euphrasia.
18. Upper lip of the corolla erect-margined
19. Lower lip of the corolla neither prominently 2-ridged nor
2-plaited ; seeds multistriate, testa close, reticulate . Bartsia.
19. Lower lip of the corolla prominently 2-ridged or 2-plaited;
seeds smooth within a loose, tuberculate and reticulate
testa Lamourouxia.
17. Anther-sacs unequally placed, the upper one attached by its
middle; leaves all or mostly alternate.
20. Lower lip of the corolla equal to or larger than the often
obtuse galea Orthocarpus.
20. Lower lip of corolla smaller than the sharper galea.
Castilleja.
ALONSOA R. & P.
Branched herb or more often shrub, often glabrous except in the
inflorescence, occasionally pilose to glandular-pubescent, branches
mostly 4-angled at least above. Leaves sessile or petiolate, mostly
opposite or ternately verticillate, floriferous leaves alternate, bract-
like; flowers in racemose inflorescences, usually terminal, glandular-
puberulent, pedicels ebracteolate; calyx 5-parted, sepals narrow; co-
rolla zygomorphic, mostly orange, varying to scarlet-red, resupinate,
explanate-rotate, the short tube cleft to the base or almost so poste-
riorly; stamens 4, filaments longer or shorter than the divergent an-
thers that are confluent at apex; stigma lobes united capitate; capsule
ovoid or oblong, septicidal, and scarcely secondarily loculicidal, usu-
ally dehiscing apically to the middle; seed numerous, small, black or
almost so, most often longitudinally furrowed.
The genus is made up of about 12 species distributed from Mex-
ico to Peru, but not in the West Indies. Only one species in Mexico
and Central America. The area of greatest concentration is found
in the Andes Mountains. Eight species reported in Peru.
1. Leaves essentially linear and entire or distantly serrulate.
2. Anthers longer than the filaments; herbage, pedicels and cal-
yces subglabrous, puberulent or glandular-puberulent.
A. linearis var. linearis.
FLORA OF PERU 469
2. Anthers shorter than the filaments; herbage, pedicels and cal-
yces glabrous A. linearis var. glaberrima.
1. Leaves broader than linear, lanceolate to ovate or deltoid-ovate,
margins mostly more deeply and closely toothed, usually serrate
to incised-dentate (except linearis var. platyphylla) .
3. Anthers longer than the filaments.
4. Leaves sparsely serrulate or entire, not more than 8.0 mm.
broad.
5. Blade margins sparsely serrulate; calyx lobes usually
glabrous, up to 5 mm. long, less than fourth the mature
fruit; branches glabrous above; leaf blades mostly wing-
ing a short (ca. 1 mm.) petiole.
A. linearis var. platyphylla.
5. Blade sessile, margins entire; calyx lobes glandular-
pubescent, 7-8 mm. long, usually more than fourth the
mature fruit ; branches glandular-pubescent above.
A. integrifolia.
4. Leaves closely serrulate to serrate or dentate, sometimes
more than 8.0 mm. broad.
6. Leaf blades 3 or more times longer than wide (rarely
about 2.7 times longer than wide with glandular pedi-
cels), usually lanceolate or lance-ovate.
7. Corolla auriculate at the base of the posterior lobes;
herb; pedicels glabrous A. auriculata.
7. Corolla not auriculate at base of the posterior lobes;
shrub; pedicels usually glandular-pubescent.
A. acutifolia.
6. Leaf blades up to about 2.2 times longer than wide,
usually ovate or oblong-ovate; pedicels glabrous.
A. mathewsii.
3. Anthers shorter than the filaments ca. one- third to one-
fourth the length of the filaments.
8. Leaf-blades less than 2 cm. long, often sessile, if margins
toothed, teeth few or reduced, spine-like; petiole when
present not exceeding 2 cm.
9. Leaf blade margins with spine-like, very sharp teeth,
sinuses lacking, main blades lanceolate; stems glandu-
lar-puberulent, at least above; corolla red. . .A. minor.
470 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
9. Leaf blade margins few-toothed, the teeth broader,
sinuses present, main blades elliptic or elliptic-subovate;
stems glabrous; corolla white or almost so . . . .A. pallida.
8. At least some of the leaves more than 2 cm. long; at least
some petioles more than 2 mm. long, margins almost al-
ways beset with numerous teeth A. meridionalis.
Alonsoa acutifolia R. & P., Syst. Veg. Fl. Peruv. 153. 1798.
Shrub or subshrub, stems and branches usually pubescent or
glandular-pubescent, at least on upper half, less often pilose or
merely puberulent, occasionally wood glabrous, inflorescence ex-
cepted. Leaves often puberulent or pubescent, lanceolate to lance-
ovate, sometimes linearly lanceolate, up to 6 cm. long and 2 cm. wide,
usually about 2-3 cm. long and 0.7-1.0 cm. wide, sharply serrate,
serrations often hardened, sharp-pointed, sometimes distant, occa-
sionally irregular, acute to long-acute at apices, narrowed to usually
acute bases, secondary veins subobsolete, often only 3-pair visible,
tissue epunctate beneath; inflorescences racemose or racemes, termi-
nal on the branches; flowers axillary to reduced leaf -like bracts,
pedicellate and solitary or rarely 2 flowers in an axil; pedicels longer
than the mature flower and fruit, usually glandular-pubescent, occa-
sionally glabrous or almost so; calyx with 5, elliptical or oblong,
dimidiate, acute, usually glandular-pubescent lobes; corolla usually
glabrous, orange to red, anterior lip 2-3 times longer than the calyx,
posterior little exceeding the calyx; stamens about 6 mm. long, an-
thers about twice as long as the filaments; style twice the length of
the ovary, stigma lobes united, capitate or punctiform; capsule lance-
ellipsoid to lance-ovoid, up to 1.7 cm. long; seed broadly ovoid, black,
blunt. Common and widely distributed in Andean Peru; also found
in montane Bolivia.
Amazonas: Prov. Chachapoyas, Sierras, alt. 2,700 m., LI. Wil-
liams 7569. — Apurimac: Prov. Abancay, upper Rio Maranon, alt.
3,000 m., Stork & Horton 1064.6; Prov. Andahuaylas, Quebrada N. of
Chincheros, alt. 2,800 m., Stork & Horton 10763; Rio Pachachaca,
alt. 3,400 m., Weberbauer 5909 (possibly Prov. Abancay). — Caja-
marca: Prov. Celendin, Vicinity of Celendin, upper hills, Woytkowski
22. — Cuzco: Quebrada de Celiska-huara, alt. 3,400 m., Herrera 1503;
"Aya-maichia," Huancayo, C. Vargas C. 3130. — Huancavelica: Alt.
3,798 m., Soukup 27821; Prov. Huancavelica, 1 km. N. of Huan-
1 Pedicels and calyces glabrous.
FLORA OF PERU 471
cavelica, limestone boulder area, alt. 3,700 m., Stork & Horton 109211;
Surcubamba: edge of forest, roadsides and pastures at alt. 2,500 m.,
Stork & Horton 10364. — Junin: Prov. Tarma, 2-4 km. E. of Huasa-
huasi, alt. 2,400-2,500 m., Hutchinson 1126; Tarma, open hillside,
alt. 3,000-3,200 m., Kittip & Smith 21859.— Puno: Lake Titicaca.
alt. 4,000 m., Leg. ?; Soukup 574-— Sin Loc: Soukup 130; 1227; 1271,
Alonsoa auriculata Diels in Engler, Bot. Jahrb. 37: 426. 1906.
Perennial, subglabrous herb, stems minutely pilose or glabrous,
branched from the base. Leaves verticillate-ternate up to the in-
florescence, alternate in the inflorescence, lanceolate or lance-ovate,
4-7 cm. long and about 2 cm. wide, margins acutely serrate, acute at
apex, acutely narrowed to base, petioles 1.0-1.5 mm. long; inflores-
cence racemose; flowers axillary to reduced leaf -like bracts, pedicel-
late; pedicels 12-18 mm. long, erect; calyx of 5 sepals, united only
at base, lance-oblong, acuminate, glandular-pilose or glabrous, 2-5
mm. long; corolla glabrous, scarlet or red, posterior lip 4 mm. long
and 5 mm. wide,2 auriculate at base of the lobes, anterior lip with
lateral lobes ca. 4 mm. long and 7 mm. wide and median lobe broadly
triangular, 7 mm. long and 11 mm. wide; stamens 4, deflexed, fila-
ment 1 mm. long, anthers 3 mm. long, oval; capsule up to 10-12 mm.
long and about half as wide, ovoid or lance-ovoid ; seed black, oblong,
blunt, longitudinally furrowed; placenta columnar, naked, septum
soon lost after dehiscence.
Cuzco: Grassy slopes above Pillahuata, Cerro de Cusilluyac,
2,500-3,600 m., Pennell 13927.— Puno3: Sandia, Weberbauer 525.
Alonsoa integrifolia Edwin, Fieldiana: Botany 31 (8) : 228-229.
1967.
Branched shrub, stems terete, glandular-pubescent, at least above.
Leaf blades sessile, linear-lanceolate to elliptically lanceolate, 0.5-
2.0 cm. long and 0.1-0.3 cm. wide, entire, narrowed to apex and some-
what to base, decreasing in size from base to apex of stem, punctate
beneath, lower glabrous, upper sometimes puberulent or sparsely
glandular-puberulent beneath, costa little impressed above, flattened
and elevated beneath; inflorescence racemose; flowers mostly near
the apices of the branches and branchlets, axillary to a bract-like leaf,
1 Pedicels and calyces glabrous.
2 The lower lip due to 180° twisting of the pedicel.
* The type, collected at 2,100-2,300 m., destroyed at Berlin.
472 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
pedicellate; pedicels 10-12 mm. long, glandular-puberulent, usually
2 times longer than the subtending bract; calyx tube very short,
sparsely glandular-puberulent, calyx lobes lanceolate, 7-8 mm. long,
more than fourth the length of the mature capsule; corolla red1;
stamens 4, anthers confluent, longer than the filaments; ovary ovoid
or lance-ovoid, little longer or shorter than the style, united stigma
lobes subpatelliform; capsule lance-ovoid or ovoid, ca. 12 mm. long,
about 6 mm. wide at base, 1-2 mm. wide at apex; seed usually blunt,
or acute at one or both ends.
Cajamarca : Michiquillo ; shrub. Calcareous slope at alt. 3,000 m.,
Pennell & Reichlin 15033 — the type and only collection.
Alonsoa linearis (Jacq.) R. & P., Syst. Veg. Fl. Peruv. 154. 1798.
Celsia linearis Jacq., Ic. Rar. 3: 1. 1+97. 1786. Hemimeris linearifolia
HBK., Nov. Gen. et Sp. 2: 377. 1817. Alonsoa linearifolia Steud.,
Nomen. ed. 2. 60. 1840.
Branched shrub, occasionally subshrub2 to 1.0 or 1.5 m. tall, gla-
brous, or glandular pubescent in the inflorescence.3 Leaves linear,
to narrowly lance-ovate, not exceeding 8 mm. wide, acute to long-
acute both to base and apex, sessile or short-petiolate, 4 or more times
longer than wide, or 2 to 3 times longer than wide, margins entire or
serrulate; flowers in racemes, each subtended by a reduced bract and
usually 1 in each axil, pedicellate; pedicels longer to less often shorter
than the flower and fruit; calyx of 5 linear to narrowly lanceolate
sepals that are united only at the base, up to 5 (-7) mm. long and
1 mm. wide; corolla orange to scarlet-red, 8-10 mm. long, posterior
lip 2-lobed, due to the 180° twisting of the pedicel becoming the
lower lip, anterior lip 3-lobed, all lobes rounded ovate; stamens in-
serted at the base of the corolla tube, anthers longer or shorter than
the filaments, yellow, when anthers longer than the filaments, ca. two
times or more longer, when shorter (var. glaberrima) about a third
to a fourth as long as the filaments: ovary ellipsoid, much shorter
than the style that is slightly thickened at apex and base, stigma
lobes united capitate or coronate; capsule 5-16 (-18) mm. long, lance-
ovoid, or -oblong, placenta columnar. All collections so far seen are
from Peru.
1 Well pressed corolla lacking; but from the scant material, corolla resupinate,
2-lobed posteriorly and 3-lobed anteriorly.
2 Sometimes dying back to the ground but not an annual as indicated on some
labels, at least in Peru.
3 Gland-bearing hairs caducous and collections sometimes entirely glabrous in
fruit in var. linearis.
FLORA OF PERU 473
KEY TO THE VARIETIES OF Alonsoa linearis (Jacq.) R. & P.
1. Leaves linear, at least four times longer than wide; fruit usually
up to 12 mm. long, rarely up to 15 mm. ; pedicels usually less than
2 cm. long, when longer the bracts almost equaling the pedicels
in length.
2. Anthers longer than the filaments; plants glandular-pubescent
in the inflorescence at least in flower; fruit usually twice as long
as or longer than the basal width; corolla usually orange.
A. linearis var. linearis.
2. Anthers only a third or a fourth as long as the filaments; plants
glabrous even in flower; fruit often less than twice as long as
the basal width; corolla usually red or scarlet-red.
A. linearis var. glaberrima.
1. Leaves wider than linear, often narrowly lance-ovate, 2-3 times
longer than wide; fruit often more than 12 mm. long, up to
17 mm.; pedicels usually more than 2 cm. long; bracts only a
few mm. long A. linearis var. platyphylla.
A. linearis (Jacq.) R. & P. var. linearis
Ancash: Prov. Huaraz, Bafios de Chancos, alt. 8,500 ft., Sand-
man ^602. — Huanuco: San Rafael, alt. 2,697 m., Soukup 2243;
Carpish, 28 m., Stork & Horton 9884- — Junin: Mito, sunny black-
berry patch, alt. ca. 9,000 ft., Macbride & Featherstone 1413. — Lima:
Matucana, Rimactal alt. 2,500 m., Rauh-Hirsch P175. — Sin Dept.:
Yanano, sunny, grassy slopes, alt. about 6,000 ft., Macbride 3771.
—Sin. Loc.: Weberbauer 6326; Woytkowski 34134.
A. linearis var. glaberrima Edwin, Fieldiana: Botany 31 (8):
230-231. 1967.
Ancash: Cordillera Blanca, herb to 80 cm. tall. Kruzl P2279.—
Lima: Canta, rocky slopes, alt. 2,700-2,900 m., Pennell 14556;
Matucana, alt. about 8,000 ft., rocky slopes, Macbride & Feather-
stone 115.
A. linearis var. platyphylla (Diels) Edwin, comb, nov.; A. line-
arifolia var. platyphylla Diels in Engl., Bot. Jahrb. 37: 427. 1906.
Junin: Tarma, alt. 7,000 ft., Macbride & Featherstone 1023.
At present it seems better to recognize the varieties of linearis,
although future collections may blur the distinctions since all occur
in north-central Peru.
474 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Alonsoa mathewsii Benth. in DC., Prodr. 10: 250. 1846.
Perennial glabrous herb, stems terete, branched. Leaves petio-
late1 to sessile, usually ternate-verticillate, alternate in the inflores-
cence, ovate or oblong-ovate, up to 3.5 cm. long and 1.8 cm. wide,
serrate, acute at apex, narrowed to base and subacute, secondary
veins prominent beneath and subprominent above, subparallel, 5-
pair, forming acute angles with the costa, blades punctate beneath;
petioles up to 15 mm. long, usually only 7-8 mm.; inflorescences
racemose; flowers axillary, pedicellate; pedicels up to 2.2 cm. long;
calyx lobes elliptical, long-acute, about 5 mm. long; corolla red, ex-
ternally sparsely pubescent, posterior lip only little exceeding the
sepals, anterior about twice as long; anthers reniform to lanceolate,
3 times longer than the 1 mm. filament; ovary much shorter than the
style, stigma lobes united, punctiform; capsule ovoid, up to 11 mm.
long and 6 mm. wide; seed black, acute at one or both ends, placenta
columnar, naked.
Amazonas: Prov. Chachapoyas, Puente de Pipos, on Rio Sonche
opposite Daguas east of Chachapoyas. Banks, alt. 2,000 m., Pen-
nell 15750. — Cajamarca: Edwin 3557; 3561.
Alonsoa meridionalis (L.f.) Ktze., Rev. Gen. 2: 457. 1891.
Scrophularia meridionalis L. f., Suppl. 280. 1781. Alonsoa caulialata
R. & P., Syst. Veg. Fl. Peruv. 152. 1798. A. procumbens R. & P.,
Syst., Veg. Fl. Peruv. 153. 1798. A. incissaefolia R. & P., Syst. Veg.
Fl. Peruv. 154. 1798. Celsia urticaefolia Sims., Curt. Mag. 12: t. 417.
1798. Hemimeris urticaefolia Willd., Sp. PI. 2(1) : 282. 1800. H. pro-
cumbens and H. caulialata Pers., Syn. Fl. 2: 162. 1807. H. parviflora
and H. mutisii HBK., Nov. Gen. et Sp. 2: 376. 1817. H. intermedia
Lodd, Bot. Cab. t. 1456. 1829. Alonsoa parviflora and A. mutisii
Steud., Nomen. ed. 2. 60 and A. urticeaefolia 61. 1840. A. incisaefolia
var. latifolia Benth. in DC., Prodr. 10: 250. 1846. A. warscewiczii
Regel, Gartenfl. 3: 211. t. 91. 1854.
Herbs, subshrubs or occasionally shrubs, usually much branched,
stems and branches mostly 4-angled and often striate-ribbed, usually
erect, occasionally procumbent, glabrous or subglabrous, the pedicels
sometimes pubescent, rarely with the branchlet apices glandular-
pubescent. Leaves opposite, occasionally lowermost ternate-verticil-
late, usually petiolate, rarely sessile, ovate, lance-ovate, less often
1 Original description, "breviter petiolatis sessilibusive," and "lanceolatis ob-
longsive," in other respects the material at hand agrees with the protologue even
to the location.
FLORA OF PERU 475
lanceolate, occasionally broadly or triangular-ovate, up to 4-5 cm.
long and 3.0-3.5 cm. wide, usually about 3.0-3.5 cm. long and 1.0-
1.5 cm. wide, apices acute, bases very variable from obtusely or
rounded-cordate to subacute or acute, margins serrate, serrate-den-
tate or dentate, teeth close and sharp; petioles mostly 3-8 mm. long;
inflorescences terminal, racemes or racemose; flowers axillary, brac-
teate, pedicellate; bracts much shorter than the pedicels, linear to
lanceolate, serrate; pedicels about 1.5-5.0 times longer than the
flowers and fruit; calyx lobes lanceolate, 4-5 mm. long and 1.0-
1.7 mm. wide, entire, tube almost lacking; corolla orange to dark red,
lips very unequal, the 2-lobed posterior up to 1.5 times the length
of the sepals, anterior lip about 2-5 times longer than the sepals;
stamens exceeding the throat, the anthers a third to a fifth as long
as the thin filaments; style 2-3 times longer than the ovoid ovary,
stigma lobes united, punctiform; capsule 0.8-1.3 cm. long, ovoid to
lance-ovoid; seed shining black, oblong, blunt, longitudinally fur-
rowed, placenta columnar, septum caducous soon after dehiscence.
Amazonas: Llama, rain-forest (Cutervo) Sandeman 4136; Prov.
Hualgayoc, alt. 1,800 m., Soukup 3852. — Arequipa: lower slopes of
Misti, alt. 9,500 ft., Sandeman 3785; Prov. Condesuyos, Chuqui-
bamba, alt. 10,500 ft., Stafford 1203,—Cuzco: Paucartambo, alt.
between 2,500-3,600 m., C. Vargas C. 4-7. — Huanuco: open situation,
alt. 2,700 m., Woytkowski 80. — Junin: near Huancayo, alt. 3,300-
3,500 m., Weberbauer 6650; Prov. Tarma, Aqua Dulce, alt. 1,800 m.,
Woytkowski 35496.— Lima: Rio Blanco, alt. 3,000-3,500 m., Killip &
Smith 21537; Matucana, alt. about 8,000 ft., Macbride & Feather-
stone 105; Rio Chillon above Obrajillo, alt. 2,700-3,300 m., Pennell
14.360 and 14402; San Buenaventura, alt. 2,700-2,900 m., Pennell
14535; Rimac Valley, alt. 3,500 m., Rauh-Hirsch P214- — Sin. Loc.
et Leg: ex herb Ruiz & Pavon. — Moquequa: Carumas, in a moist
ravine, alt. 3,000 m., Weberbauer 7284- — Sin. Depto.: Yanano, alt.
about 6,000 ft., Macbride 3738; Valle de San Miguel, alt. 2,060-
2,260 m., Herrera2001.
At least three infraspecific categories could be presented based
upon differences in leaf-bases, pedicel length in relation to flower
length, relative sizes of corolla lobes, procumbent as opposed to erect
growth forms and kinds of leaf blade margins. However, this taxon
is so commonly collected that these characters, leaf-bases excepted,
overlap throughout most of its range. Therefore, it seems the better
course at least for the present to retain a single polymorphic taxon.
476 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
The commonest species of the genus, collected from Mexico through
Central America to Equador, Colombia, Venezuela and Chile as well
as Peru. Not known east of the Andean chain. The center of con-
centration is in Colombia.
Alonsoa minor Edwin, Fieldiana: Botany 31 (8) : 229-230. 1967.
Branched shrub, apices of branchlets glandular-pubescent. Leaves
very narrowly lanceolate, subsessile, up to 1.9 cm. long and 0.6 cm.
wide, margins few-serrate, serrations narrow, hard, attenuate to
apices and bases, sparsely black-punctate beneath; inflorescences
racemose; flowers axillary, solitary, pedicellate; pedicels glandular-
pubescent, up to 1.7 cm. long, longer than the flower, fruit and sub-
tending bract; calyces salverform in bud, tube very short, lobes 2.5-
3.5 mm. long, elliptical to narrowly-elliptical, acuminate; corolla red
to dark red, lobes shorter than the tube; stamens longer than the
corolla tube, exserted beyond the throat but shorter than the lobes,
anthers axe-shaped, densely pubescent, shorter than the filaments;
ovary little longer to little shorter than the style; immature capsule
9-10 mm. long and 4-5 mm. wide, ovoid; seed black.
Amazonas: Above Calcamar, in thin soil over limestone rocks,
alt. 2,600-2,900 m., F. W. Pennell 15609; Chachapoyas, Cerro Calla-
Calla, 8 km. W. of Leimebamba, Edwin 3702.
Alonsoa pallida Edwin, Fieldiana: Botany 31 (8) : 230. 1967.
Glabrous, erect, oppositely branching shrub, branchlets square.
Leaves elliptical or subovate elliptical, up to 1 cm. long and 0.6 cm.
wide, short-petiolate, margins few-toothed, serrate to laciniate, some-
times dimidiate at base, young blades sometimes puberulent above,
nerves elevated beneath; petioles up to 2 mm. long; inflorescences
racemes; bracts minute; pedicels glabrous, up to 9 mm. long, shorter
than the capsule; mature flowers lacking; calyx lobes about 3 mm.
long, lanceolate or narrowly elliptic, acuminate, 3-nerved, tube lack-
ing; corolla white; stamens included in the corolla tube; anthers
sagittate, shorter than the filaments; capsule 7-10 mm. long and 3-
5 mm. wide, ovoid; seed shining black. Known only from the type.
Cajamarca: Below Llama. Herb with whitish corolla. Banks,
alt. 1,900-2,100 m., Pennell 15918 (BM).
ANGELONIA H. and B.
Herbs or occasionally subshrubs, glabrous, viscid, glandular-
pubescent, stems and branches often 4-angled at the apices, round
FLORA OF PERU 477
below. Leaves all opposite or opposite below and becoming sub-
opposite and then alternate base to apex, blades narrowly lanceolate
to ovate, subsessile or sometimes sessile; flowers axillary, pedicellate,
1 or 2 in each axil or few-clustered in axils;1 calyx 5-toothed or more
deeply cut; corolla with short tube, subrotate, explanate, 2-lipped,
lobes 5, rounded, the 2-lobed posterior lip a little shorter than the
3-lobed anterior lip that is saccate at the throat and supplied with
a horn-like process at the base; stamens 4-didynamous, filaments
short, anther locules distinct, divaricate; ovary shorter than the
style, stigma lobes united, punctiform or capitate; capsule loculici-
dal, rarely secondarily septicidal; seed brown to yellow-brown with
raised wing-like reticula.
About 15-20 species, generally distributed in tropical America,
greatest concentration in Brazil.2
Two species collected in Peru. One of these only doubtfully an
Angelonia.
Leaves lanceolate, most or all, four or more times longer than wide;
plants viscid, glandular-pubescent A. angustifolia.
Leaves ovate, less than four times longer than wide; plants essentially
glabrous (wood and petioles sparsely puberulent) . .A. procumbens.
Angelonia angustifolia Benth. in DC., Prodr. 10: 254. 1846.
Viscid, glandular-puberulent herbs or subshrubs.3 Leaves up to
9 cm. long and 1 cm. wide, mostly much smaller, margins closely to
distantly serrate or serrulate, sometimes subentire, acute, narrowed
to base, petiole when present up to about 1 mm. long, costa visible,
remaining nerves obsolete or almost so; pedicels from 0.8-2.0 cm.
long; flowers usually 2, sometimes 1 in each leaf axil,4 calyx about
4-6 mm. long, sepals elliptical, acute; corolla purple or blue-purple,
about 2-3 times longer than the calyx; stamens short, filaments 1-
2 mm. long, little longer or shorter than the anthers; pistil about
3-4 mm. long; capsule much exceeding the calyx, globose, style per-
sistent; seed conical, interspaces of reticula deep, testa appearing
honeycomb-like.
1 Extra Peruvian material sometimes with flowers in terminal racemes.
2 In the last 30 years there has been a great proliferation of specific epithets in
this genus. A careful monograph would prove most useful.
3 Extra Peruvian material frequently glabrous.
4 Extra Peruvian material almost always one flower in each leaf axil.
478 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Loreto: Palta-Cocha on the upper Rio Nanay, LI. Williams 1279;
San Salvador on the Amazon River, LI. Williams 1550; Pebas on the
Amazon River, LI. Williams 1818; La Victoria on the Amazon River,
LI. Williams 2749; Iquitos and vicinity, alt. 120 m., LI. Williams
3580; Lower Rio Huallaga, alt. 155-210 m., LI. Williams 4490.
Collected from Mexico through Central America and the West
Indies to British Guiana, Venezuela, Colombia, and Peru.
This species is closely related to A. salicariaefolia H. & B. and
also to A. biflora Benth. It differs from the former where the leaf-
base is clasping and not narrowed and from the latter where the
vesture is eglandular, and the size of which is much greater. Con-
ceivably, intermediates may be collected uniting these three taxa.
Angelonia procumbens Nees ex Mart., Nova Acta Acad. Nat.
Cur. 11: 45. 1823. Physidium procumbens Schrad., in Got. Gel. Ang.
714. 1821. Schelveria arguta Nees ex Mart., in Bot. Zeit. 1: 299,
328. 1821.
Subglabrous branched herb, sparsely puberulent on the 4-angled
and winged branches, between the wings and on the very short,
thickened petioles. Leaves opposite, upper occasionally suboppo-
site, ovate, mostly subsessile, sometimes sessile, up to about 4.0 cm.
long and 1.5 cm. wide, acute or long-acute, obtuse or rounded at
base, margins serrate, especially upper three-fourths of blade, ser-
rations appressed, primary veins usually apparent; flowers short-
pedicellate, solitary or few-clustered, axillary; calyx enlarged at base,
drying black, sepals sparsely, very short-ciliolate; anthers divaricate
on very short filaments, style thickened, stigmas united, capitate.
San Martin: Juan Jui, Alto Rio Huallaga, forest, about alt. 400
m., King 3842. (Kew Herb.)
Good flowers and mature fruit lacking. Only provisionally de-
termined as an Angelonia. The stamens as usually occurring in this
genus as well as the thickened style with capitate stigma seem to
indicate this section of the family. The flowers are not resupinate
as in Alonsoa R. & P. However, Angelonia procumbens Nees ex
Mart, which this specimen so strongly resembles is a species of low-
land southeastern Brazil (Bahia).
BACOPA Aubl.
Erect, repent or creeping herbs, sometimes forming mats, when
erect up to about 60 cm. tall, glabrous to densely pubescent. Leaf
FLORA OF PERU 479
blades narrowly obovate to almost round, sessile or petiolate, from
less than 1 cm. long to 5 cm. or little longer, margins entire to irreg-
ularly coarsely dentate, bases attenuate to broadly obtuse, apices
long-acute to rounded, usually glabrous (except B. salzmanni Benth.
with long, golden hair on tissue beneath near the base) ; flowers axil-
lary, solitary, or occasionally few-clustered (B. stricta (Schrad.)
Britt.), pedicellate (the pedicels ebracteolate or bibracteolate at base
or apex, immediately below the calyx); sepals imbricate, 5, the 2
outermost (posterior) broadest, the lateral and anterior narrower,
sometimes only a fifth as wide as the posterior, sometimes the poste-
rior and lateral sepals much enlarged, often the larger sepals toothed,
calyx tube wanting, the sepals often only loosely enveloping the
flower and fruit; corolla tube cylindric to campanulate, lobes flat-
tened or rounded, the throat open, lips mostly subequal, the upper
(posterior) 2-lobed, the lobes united little higher than the 3-lobed
lower (anterior) lip; stamens usually 4, didynamous, included in the
tube, sometimes one or both of a pair (the smaller) with non-func-
tional anthers, rarely one or both of the lower pair wanting, anthers
stalked or sessile on the filament, when on stipes versatile, when ses-
sile sagittate, versatile or dorsifixed, usually very small, about one-
fourth to one-half the length of the filament; ovary ellipsoid, black
or brown when dry, style little dilated at apex, usually a little longer
than the ovary, stigma lobes at least partly distinct; capsule ellip-
soid to broadly ellipsoid or subglobose, frequently shorter than the
calyx, usually 2-valved, rarely 4-valved; seed numerous, testa retic-
ulate; placenta columnar, little if at all thickened.
The genus (Bacopa the aboriginal name for B. aquatica Aubl., the
type) is here treated in the broadest sense (See: Edwin, Fieldiana:
Botany 31: no. 8, 227. 1967), containing 50-60 or more species, many
very poorly collected and understood. Monographic revision em-
ploying all the means and methods at the disposal of today's worker
is most necessary. The generic limits as well as the specific are in
dispute. Distributed mostly in the warm regions (less in the tem-
perate) of the world. Rare in Europe, uncommon in Africa, and
sparsely represented in Australasia. Basically New World with
greatest concentration in Brazil and with many species also in north-
central and northeastern South America. A secondary center is pres-
ent in Mexico. Four species have been collected thus far in Peru,
two of which are rare.
1. Pedicels bibracteolate at apex.
480 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
2. Leaf blades with margins entire, often sessile, mostly not ex-
ceeding 1.5 cm. long, cuneate or obovate, with attenuate bases;
margins of the outer 3 sepals entire; anthers sessile on the fila-
ments, dorsifixed B. monnieri.
2. Leaf blades with coarsely toothed margins, petiolate, mostly
more than 1.5 cm. long, up to 5.5 cm., mostly ovate or ellipti-
cally ovate; margins of the outer sepals sometimes toothed;
anthers on short stipes, versatile B. stricta.
1. Pedicels ebracteolate or bibracteolate at base.
3. Stems densely pubescent; leaf blades sessile, clasping, almost
connate, widest at the base, pubescent beneath with golden
hair; pedicels ebracteolate; anthers sessile on the filaments,
dorsifixed B. salzmanni.
3. Stems essentially glabrous; leaf blades short-petiolate, nar-
rowed to base, glabrous; pedicels bibracteolate at base; an-
thers on short stipes, versatile B. procumbens.
Bacopa monnieri (L.) Wettst. in Engler and Prantl, Pflanzenf.
IV (3b): 77. 1891. Lysimachia monnieri L., Cent. Plant. 2: 9. 1756.
Gratiola monniera L., Amoen. Acad. 4: 306. 1759. Calytriplex obo-
vata Ruiz & Pav., Prodr. 84. 1797. Herpestis monnieri HBK., Nov.
Gen., et Sp. 2: 366. 1817. H. procumbens Spreng., Syst. Veg. 2: 802.
1825. Bramia monnieri (L.) Pennell, in Proc. Acad. Nat. Sci. Phila.
71: 243. 1920. Also Limosella calycina Forsk.; Septas repens Lour.;
Bramia indica Lam.; Monnieria brownei, M. africana, and M. pe-
dunculosa Pers.; Herpestis cuneifolia Michx.; Habershamia cuneifolia
(Michx.) Raf. and others.1
Repent or prostrate often matting, glabrous herb of wet places,
rarely floating on water. Leaves cuneate-obovate, attenuate to base,
usually not exceeding 1.5 cm. long and 5-6 mm. wide, entire, sessile
or almost so, apex rounded; internodes little longer or shorter than
the leaves; flowers axillary, solitary, pedicellate, small, up to about
1.0 cm. long and 1.5 cm. wide when open, pedicels bibracteolate at
apex, mostly not exceeding 1 cm. long, often only 6-7 mm. long
(rarely long-pedicellate, ca. 2.0-2.5 cm.), erect, rarely reflexed; brac-
teoles linear to narrowly ellipsoid; sepals free, 4-5 mm. long, poste-
rior (outer) pair about 2.5-3.0 mm. wide, lateral sepal 1.5-2.0 mm.
1 Full synonymy with extended citations for this taxon can serve no useful
purpose at this time. The synonyms supplied are believed to be sufficient to indi-
cate both the nomenclatural and taxonomic disputes still prevalent in the Gratiolae
generally and this taxon in particular.
FLORA OF PERU 481
wide, anterior pair less than 0.7 mm. wide, often not reaching 0.5
mm., outer and lateral ovate or elliptic-ovate, inner linear; corolla
zygomorphic, bilabiate, tube shorter than the calyx, included, yel-
lowish within, cylindric-campanulate, the white to pink to lavender
or blue lobes little shorter than the tube, posterior pair little longer
than the anterior three lobes, the posterior lobes united little more
than the anterior, entire corolla about 6-9 mm. long; stamens 4,
didynamous, anthers about one-half as long as the filaments, sessile,
sagittate, dorsifixed; ovary ellipsoid, shorter than the corolla tube
and the 3.5-4.5 mm. long style, stigma 2-lobed, lobes coronate or
punctiform; capsule ellipsoid or ellipsoid-ovoid, septically dehiscent.
Cajamarca: Osgood and Anderson 37. — Huanuco: Prov. Huanuco,
Huacho, 8 km. N. of Huanuco, 1,920 m. alt., Stork & Horton 9413;
7,000 ft., Macbride 3223.— Lima: Chancay, Seaside cliff, Stork 9364;
Callao, Macbride 5884; Miraflores, Pennell 14284; Vargas 1232; Sin
Prov.: near Viscas, Pennell 14446- — Piura: Negritos, Haught 126;1
E. of Cape Parinas, Haught 126.1
Greatest concentration is the Southern United States, not un-
common in Mexico, Central America, and the West Indies, sporadic
in South America to Paraguay and Brazil, rare in Malaya, Iraq, and
the Philippine Islands.
Bacopa procumbens (Mill.) Greenm., Field Mus. Bot. 2: 261.
1907. Erinus procumbens Miller, Gard. Diet. VIII (6). 1768 and
IX (13). 1797. Lindernia dianthera Sw., Prodr. 92. 1788; Miller
Gard. Diet. IX (2). 1797; Willd., Sp. PI. 3: 326. 1800. Herpestis
colubrina HBK., Nov. Gen. et Sp. 2: 368. 1818. H. chamaidryoides
HBK., Nov. Gen. et Sp. 2: 369. 1818; Benth. in DC., Prodr. 10: 393.
1846. H. peduncularis Benth. in Comp. Bot. Mag. 2: 56. 1836;
Benth. in DC., Prodr. 10: 394. 1846. Bacopa chamaidryoides Wettst.
in Engler & Prantl, Pflanzenf. IV (3b): 76. 1895. Also Monniera
dianthera Millsp. (comb. nov. only, not as to specimens); Mecar-
donia procumbens Small; M. viridis Small; M. tennis Small; Pagesia
peduncularis (Benth.) Pennell; P. procumbens (Miller) Pennell; P.
dianthera (Sw.) Pennell; Bacopa dianthera (Sw.) Descole & Borsini.
Glabrous, upright or prostrate herb, when prostrate sometimes
forming mats, usually blackening when dry. Stems terete, much
branched from the base; leaves mostly ovate or elliptically ovate,
1 The labels, data, and number of these collections are mixed or duplicated.
While these two sheets may comprise but one collection it is best to consider them
distinct. That the material was collected in Depto. Piura seems clear.
482 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
dimidiately-attenuate to the petiole, up to 3.4 cm. long and 1.2 cm.
wide, usually about 1.5-2.5 cm. long and 0.6-1.0 cm. wide; margins
dentate or serrate-dentate on upper three-fourths of blade, acute;
petiole up to 1 cm. long, narrowly winged; flowers axillary, solitary;
pedicels lax, up to 2.5 cm. long, usually at least 2 times longer than
the flowers, bibracteolate at base; bracteoles narrowly lance-ovate,
short-petiolate; sepals very little united, ca. 0.5 mm. or (usually)
completely free, 6-8 mm. long, three outermost ovate, broadest,
often with toothed margins, two innermost narrow, almost linear,
entire, all sepals acute, innermost occasionally acuminate; corolla
yellow, 8-10 mm. long, tube included in the calyx, the short lobes
exserted but only little exceeding the calyx, upper lip of 2 lobes, gla-
brous within, lower of 3 lobes or the lowermost 2 lobes united, inter-
nally with club-shaped hairs in or near the throat; stamens 4, didyna-
mous, anthers on very short stipes, about one-fifth to one-third the
length of the 2-3 mm. long filament, versatile; ovary ellipsoid, in-
cluded in or only little exceeding the frequently papery calyx, sep-
ticidal.
Sporadic in Central to Northeastern Peru.
Junin: Above San Ramon, Hacienda Schunke, 1,300-1,700 m.,
Schunke A9; About 4,000 ft., Macbride 5670; Mito, 9,000 ft., Mac-
bride & Feather stone 14.17. — Loreto: above Pongo de Manseriche,
alt. 200m., Mexia 6173.— San Martin: San Roque, alt. 1,300-1,500
m., LI. Williams 714-8. — Sin Dept. : in Montanis Peruviae, Haenke
1900.
Widely distributed in the warm temperate and tropical zones of
the New World, from sea-level to 9,000 ft. or more elevation. In
usually drier sites than the preceding species, sometimes a garden
weed. Most abundant in Central America and Mexico, common in
the southern United States. Rarer in the West Indies. Sporadic
in South America in Brazil, Ecuador, Colombia and one collection
seen from Paraguay.
There is reasonable expectation that further collecting, especially
from the Eastern slopes of the Andes and from lowland Peru will
result in uncovering at least some if not all of the following species,
known from Brazil, Venezuela, and Colombia: B. aquatica Aubl.,
B. lilacina (Pennell) Standl., B. reptans, and B. ranaria Benth.,
known also from widespread Central America collections. In addi-
tion, three others can be expected, at present well represented in
Central America: B. parviflora Standl., B. reflexa, and B. sessiliflora
Edwall.
FLORA OF PERU 483
Bacopa salzmannii (Benth.) Wettst. in Engler & Prantl, Pflan-
zenf. IV (3b): 76. 1895. Herpestis salzmannii Benth., in Comp. Bot.
Mag. 2: 58. 1836.
Erect or reclining herb, stems soft, densely beset with golden
hairs, the internodes often 2 or more times longer than the sessile
to clasping leaf blades (internodes much shortened near the apex).
Leaves ovate-orbicular, up to about 1 cm. long and wide, almost con-
nate, margins entire or microscopically distantly crenulate, densely
beset with golden hairs beneath, near the base and on the lower half
of the tissue; flowers solitary, axillary on long, recurved, golden-
pubescent, ebracteolate pedicels; pedicels up to 3-4 times longer than
the calyx; flowers up to about 7 mm. long, not as wide; calyx up to
5 mm. long, sepals free, outermost 2 broadly ovate, cordate at base,
acute to subobtuse at apex, inner sepals much narrower, little
shorter; corolla white or creamy white, only little exceeding the
calyx, tube much longer than the very short lobes; stamens 4, in-
cluded in the corolla tube, didynamous, filament pairs little to very
different in length, anthers sessile, subsagittate, dorsifixed, varying
from less than to more than one-half the length of the filaments,
ovary only about 1.5 mm. long, ellipsoid, style about 2 times as long
as the ovary, stigma lobes distinct, lamellate; capsule ellipsoid, in-
cluded in the calyx, septicidal.
Rare in Peru. Lima: Lima, Callao, alt. about 8 ft., Macbride
5987.
A Central American species, where it is polymorphic. Common
in boggy and other wet sites from British Honduras to Panama; not
infrequent in Mexico. Found sporadically in South America in Co-
lombia, Bolivia, and Brazil.
This taxon varies widely in habit; from erect and unbranched to
branched, repent, creeping, matted and prostrate. The latter con-
ditions would be expected in Peru.
Bacopa stricta (Schrad.) Edwall, Bol. Conn. Georgr. Sao Paulo
13: 176, 180. 1897. Herpestis stricta Schrad. in Link, Enum. 2: 142.
1822. H. domingensis Spreng., Syst. 2: 801. 1825. H. polyantha
Benth., Comp. Bot. Mag. 2: 57. 1836. Mella Vandelli, Fl. Lusit.
43, fig. 23. 1788. Generic description only, no generotype indicated.
This name is often a sectional or subgeneric designation of Bacopa
(or Herpestis} of authors, the section to which this species and
B. salzmannii Benth. pertain.
484 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Erect, subglabrous herb, stems terete or quadrangular, soft, usu-
ally sparsely and finely, very short-puberulent above. Leaves gla-
brous, petiolate, blades up to 6 cm. long and 2 cm. wide, usually
smaller, decreasing base to apex, ovate or elliptically ovate, narrowed
to long-acute bases and apices, occasionally apices subacuminate,
margins coarsely, irregularly dentate or serrate, venation pinnate,
apparent above and beneath; flowers small, up to about 8 mm. long,
usually solitary or few-clustered in leaf-axils, sometimes in short,
raceme-like inflorescences or racemes, pedicellate, dispersed often
from near the base of the plant to the apex; pedicels more or less
densely to sparsely pubescent, usually about as long as the calyces or
a little longer, shorter when the flower is immature, bibracteolate
near the apex; bracteoles linear, ca. 1 mm. long; calyx often dark red
in life, up to about 6 mm. long, usually 4-5 mm. posterior and lateral
sepals ovate, margins often soft-ciliate or ciliolate, acute, tube want-
ing; corolla up to 8 mm. long, not as wide, lobes only little exceeding
the calyx in length, tube and lobes white, tube internally marked
with yellow or blue near the throat; stamens 4, didynamous included
in the tube, anthers very small, often disposed on short stipes (visible
only at 20 X magnification or more) versatile, stipes sometimes sup-
pressed then anthers contiguous, dorsifixed ; style thick, much longer
than the 1-2 mm., ellipsoid ovary, stigma lobes wholly free; capsule
septicidal, short, ca. 3-5 mm. long, ovoid to ovoid-subglobose, style
frequently persisting.
Rare in Peru, only one collection seen, San Martin: San Roque,
1,300-1,500 m., LI. Williams 7165.
Sporadic and infrequent in South America; found also in Colom-
bia, Venezuela, Brazil, and Ecuador. Rare in the West Indies.
BARTSIA L.
Bartsia L., Gen. PL 262. 1754; Benth. in DC., Prodr. 10: 544.
1846. Trixago Stev., Mem. Soc. Mosc. 6: 4. 1823. Parentucellia
Viv., Fl. Lib. Spec. 31, t. 21, f. 3. 1824. Eufragia Griseb., Spicil.
Fl. Rumel. 2: 13. 1844. Alicosta Dulac., Fl. Hautes-Pyr. 381. 1867.
Annual or perennial herbs, sometimes subshrubs, usually vari-
ously pubescent and often glandular, erect to prostrate, usually
branched. Leaves usually opposite, occasionally alternate, the lower
often with axillary leaflets, mostly toothed, usually narrow (to ovate),
many times longer than wide, most often sessile and the apex obtuse,
upper frequently cordate-amplexicaul ; bracts similar to cauline or
FLORA OF PERU 485
modified, entire, subentire and narrower; flowers mostly 8-28 mm.
long, forming spikes or racemes, rarely solitary and axillary; calyx
tubular or campanulate, 4-lobed; corolla galeate, galea usually longer
than the 3-lobed lower lip, occasionally shorter, mostly glandular-
pubescent, sometimes glands wanting, rarely glabrous; lobes of the
lip erect or patent; throat occasionally with a small palate, tube
mostly longer than the lips; stamens 4, didynamous, with the style
included in the galea, filaments often with minute sessile glands, an-
thers usually long-pubescent, most often basally mucronate or apic-
ulate; stigma exserted, lobes united or rarely partially distinct, ovary
rarely glabrous, usually densely beset with stiff, long, golden hair;
capsule with vesture as the ovary, narrowly ellipsoid to ovoid, little
shorter to twice the length of the calyx, apically usually obtuse,
emarginate, apiculate; seeds numerous, minute, testa closely adher-
ent, surface many-striate-sulcatulate.
The genus has an unusual and disjunct distribution. One species
each in North America and Australia; between five and ten in Europe
and Africa. The center of concentration is in Andean South America
with about 50 species of which 32 are found in Peru. A number of
taxa most probably remain to be described, especially in Africa and
South America.
1. Corolla externally glabrous or sparsely hispidulous.
2. Galea equal to or longer than the lower lip of the corolla.
B. elongata.
2. Galea shorter than the lower lip of the corolla.
3. Leaf blade with entire margins B. integrifolia.
3. Most of the leaf blade margins grossly toothed. . . .B. aurea.
1. Corolla especially the galea, externally puberulent or pubescent
and often glandular.
4. Calyx 18-20 mm. long; corolla brown B. peruviana.
4. Calyx less than 18 mm. long; corolla yellow or red, less often
white or purple.
5. Anthers glabrous or very sparsely hispidulous.
6. Corolla tube shorter than the lips B. calycina.
6. Corolla tube equal to or longer than the lips.
7. Galea equal to or longer than the lower lip.
8. Pedicels of mature flowers 2-3 mm. long or longer.
9. Anthers blunt at the lower end, without mucra or
acumina. ..B.mutica.
486 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
9. Anthers mucronate or acuminate at the lower end.
B. pedicularioides.
8. Pedicels of mature flowers less than 2 mm. long.
10. Bract margins with few to many rounded ex-
trorse teeth or lobes; calyx usually glandular.
11. Plants erect, elongate, galea puberulent or
when glandular-pubescent, glands long-
stalked B. santolinaefolia.
11. Plants dwarf, often procumbent; galea gland-
ular-pubescent, glands subsessile . . B. diffusa.
10. Bract margins with small, sharp antrorse lobes;
calyx eglandular B. biloba.
7. Galea shorter than the lower lip.
12. Leaf blades most often lance-ovate, oblong or vari-
ously obovate; bracts ovate or obovate.
13. Blades mostly oblong to lance-ovate.
14. Stems suberect or ascending, clustered or branch-
ing from the base; plants dwarf, usually less
than 20 cm. tall B. diffusa.
14. Stems erect, solitary, branching above the base,
elongate, much more than 20 cm. tall.
B. mutica.
13. Blades and bracts variously obovate.
B. simulans.
12. Leaf blades linear; bracts linear, oblong-linear or
narrowly lance-subovate B. curtiflora.
5. Anthers distinctly pubescent with mostly long and curled or
matted hair.
15. Galea shorter than the lower lip of the corolla.
16. Corolla tube about as long as the lips.
17. Calyx lobes acute, shorter than the tube. . .B. aurea.
17. Calyx lobes obtuse or subobtuse, usually as long as
the tube or longer, rarely when shorter margins
crenate B. diffusa.
16. Corolla tube longer than the lips.
18. Calyx lobes obtuse; plants straggling, suberect,
or erect.
FLORA OF PERU 487
19. Galea little shorter than the lip; bracts ovate;
plants suberect-ascending, dwarfed . . E. diffusa.
19. Galea about one-half as long as lower lip ; bracts
oblong or elliptical; plant elongate, erect.
B. meyeniana.
18. Calyx lobes acute; plants usually erect.
20. Calyx lobes narrow, linear or little wider, mostly
as long as or longer than the tube.
21. Pubescence cinerous or canescent, appressed,
mostly eglandular; bracts usually entire, lin-
ear or narrowly lanceolate B. canescens.
21. Pubescence golden or white or brown, often
glandular; bracts usually toothed, broader
than linear B. bartsioides.
20. Calyx lobes lance-triangular, lance-ovate or nar-
rowly ovate or oblong, usually shorter than the
tube.
22. Tube of the 9-12 mm. long corolla more than
half included in the calyx B. curtiflora.
22. Tube of the 13-22 (-26) mm. long corolla
more than half exserted from the calyx.
B. anomala.
15. Galea equaling to 2-3 times longer than the lower lip of
the corolla.
23. Galea equaling to up to a fourth longer than the lip.
24. Corolla tube shorter than and included in the calyx,
usually equaling to sometimes shorter than the lips
of the corolla B. subinclusa.
24. Corolla tube exserted and longer than the calyx and
most often longer than the corolla lips.
25. Bracts entire or with a few pairs (up to 4) of dis-
tant often obscure teeth.
26. Calyx cleft more deeply medianly than laterally,
cleavages very short, lobes very unequal.
B. biloba.
26. Calyx cleft equally medianly and laterally, lobes
equal or almost so.
27. Flowers ca. 10 mm. long; calyx lobes deltoid-
triangular. . . ., B. parvifolia.
488 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
27. Flowers 13-18 (-20) mm. long; calyx lobes
linear or lance-linear B. bartsioides.
25. Bracts with several pairs of close, often obtuse or
rounded teeth.
28. Flowers up to 13 mm. long, rarely reaching
15 mm., usually 8-11 mm.; calyx lobes most
often much shorter than the tube.
29. Dwarf, often less than 2 cm. tall, deep-
rooted, decumbent to suberect plants, usu-
ally branching from the base, frequently
stems clustered B. diffusa.
29. Elongate, more than 2 cm. tall, shallow-
rooted, erect plants, branching from well
above the base or unbranched, stems soli-
tary.
30. Foliage and fruit glabrous or subglabrous,
the few hairs when present on the fruit,
white; corolla whitish; stems unbranched.
B. glabra.
30. Foliage and fruit more or less densely
pubescent, the fruit beset apically with
golden hair; corolla yellow, pink or pur-
ple; stems most frequently branched.
31. Pubescence hispid, hard. . .B. hispida.
31. Pubescence pilose, soft.
B. santolinaefolia.
28. Flowers mostly 15-16 (-20) mm. long; calyx
lobes only little shorter to equalling the tube.
B. bartsioides.
23. Galea more than a fourth longer than the lip, up to 2-
3 times longer.
32. At least most of the flowers less than 14 mm. long.
33. Corolla tube equal to or shorter than the lips.
34. Calyx lobes subacute or obtuse.
35. Calyx lobes crenate B. filiformis.
35. Calyx lobes entire B. orthocarpiflora.
34. Calyx lobes acute, pubescence stiff (hispid).
B. elachophylla.
FLORA OF PERU 489
33. Corolla tube longer than the lips.
36. Bracts longer than the flowers.
37. Stem glandular-hispid above; calyx teeth tri-
angular; pedicels less than 3 mm. long.
B. brachyantha.
37. Stems with stiff, multicellular, pointed hair;
calyx teeth broadly ovate; pedicels often 3
mm. long B. duripilis.
36. Bracts shorter than the flowers.
38. Plants caespitose B. sanguined.
38. Stems not tufted, either branched or un-
branched.
39. Flowers and fruit on erect pedicels up to
7 (-8) mm. long B. pedicellata.
39. Flowers and fruit sessile or subsessile;
pedicels neither erect nor exceeding 3
mm. long.
40. Calyx lobes crenate B. filiformis.
40. Calyx lobes entire.
41. Flowers white or whitish, or yellow
or red; calyx lobes oblong, obtuse.
42. Corolla about 2 times as long as
the calyx, anthers sparsely, galea
little more, pubescent; galea up
to 2 times longer than the lip.
B. glabra.
42. Corolla about 3 times longer than
the calyx; anthers and galea
densely pubescent; galea about 3
times longer than the lip.
B. orthocarpiflora.
41. Flowers green; calyx lobes lance- tri-
angular, acute B. viridis,
32. At least most of the flowers more than 14 mm. long.
43. Corolla tube equal to or shorter than the lips.
44. Flowers on filiform pedicels, 5-10 mm. long;
galea up to about 2 times longer than the lip.
B. weberbaueri.
490 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
44. Flowers sessile or on thicker and shorter pedi-
cels, up to 4 mm. long; galea about 3 times
longer than the lip B. orthocarpiflora.
43. Corolla tube longer than the lips.
45. Leaves glabrous except for marginal articulated,
1 mm. long hair; stems solitary. .B. trichophylla.
45. Leaves usually pubescent on tissue and veins,
marginal hair present or lacking, when present
not articulated; when leaves glabrous, stems
clustered or tufted.
46. Galea usually deeply emarginate or obcor-
date; pubescence stiff, usually 2-celled.
B. altissima.
46. Galea entire or merely faintly emarginate at
apex; pubescence otherwise.
47. Stems caespitose, basally branched.
B. sanguined.
47. Stems solitary, not branching from the
base.
48. Flowers frequently 2 cm. long or longer;
galea up to 2 times the length of the lip.
B. inaequalis.
48. Flowers up to 16 mm. long; galea about
3 times the length of the lip.
B. orthocarpiflora.
Bartsia altissima Rusby, Phytologia 1: 75-76. 1934.
Upright (very) densely hispid herb, stems much branched and
sulcate, pubescence often clustered, the trichomes usually swollen at
the base and usually 2-celled, occasionally 1-celled, retrorse. Leaves
sessile, clasping, broadest at the base, lance oblong, up to more than
2 cm. long and almost or quite 1 cm. wide, coarsely crenate-dentate,
the teeth rounded-obtuse, the sinuses narrow, apex obtuse, veins
very apparent beneath, the costa often keeled, the pubescence fre-
quently clustered here and on the secondary veins, these 4-5 pairs;
spikes short, few-flowered, interrupted, short-pedicellate; pedicels ca.
3-4 mm. long; calyx 7-10 mm. long, tube campanulate, lobes un-
equally, broadly triangular, acute, shorter than the tube; corolla red
or dark brown-red, pubescent inside and outside, 15-18 mm. long,
tube a little longer than the galea, throat swollen, galea about 2 times
FLORA OF PERU 491
longer than the lip, usually deeply (for the genus) obcordate or emar-
ginate, lobes on the lips ovate; anthers lanate, mucronate; style very
short, puberulent apically, stigma lobes united, at least partly, most
often wholly, ovary densely beset with long, stiff, golden hair; cap-
sule densely pubescent as the ovary, ellipsoid or ovoid, blunt at apex,
a little longer than the calyx, style persisting.
Ayacucho: Between Huanta and Hacienda Pargora, alt. 4,200 m.,
Killip & Smith 23301; Huanta, Punas above Huanta, alt. 12,000-
13,000 ft., Pearce s.n. — Cuzco: Paucartambo to Tres Cruces, Cerro
de Cusilluyoc, thickets and rocky banks, alt. 3,400-3,600 m., Pen-
nell 13810; Pillahuata, Cerro de Cusilluyoc, thickets in valley of Rio
Pillahuata, alt. 2,300-3,000 m., Pennell 13926; Lares Prov., at Lares,
alt. 3,200 m., Marin 2307.
Type from Bolivia (Pongo) at 12,000 ft. The Peruvian material
differs in having unbranched hairs and a larger calyx but is clearly
the same taxon. Rusby's specimen apparently had no corollas or
fruit suitable for description.
Bartsia anomala Edwin, Phytologia 19: 362. 1970.
Perennial herb, usually pubescent, occasionally merely puberu-
lent on stems branches and foliage, erect, sometimes branching from
the base. Leaves linear to lanceolate, crenate, slightly bullate, ob-
tuse, narrowed to base, sessile; inflorescences spikes, usually lax and
basally interrupted; pedicels 1.0-2.5 mm. long; bracts usually as the
leaves, except margins distantly crenulate to almost entire, espe-
cially near the apices of the branches; flowers 13-22 (-26) mm. long,
corolla yellow or lower lip occasionally red to purple; calyx 8-11 mm.
long, tubular, the acute, narrowly lance-oblong or lance-triangular
lobes shorter than the tube; corolla tube longer than the lips, over
one-half exserted, galea shorter than the lower lip whose lobes are
oblong, rounded; filaments glabrous or with a few, minute, sessile,
scattered glands, anthers pubescent, mucronate at base; stigma lobes
united, style minutely, sparsely puberulent, ovary densely beset with
long, stiff, golden hair; capsule usually shorter than the calyx, occa-
sionally about equal to the calyx, ellipsoid-ovoid, obtuse, apiculate,
pubescent on the upper half, as the ovary.
Ancash: Cordillera Blanca, alt. 4,000 m., Kinsl. P2298.— Caja-
marca: S.W. of Paso de Gavilan, S. of Cajamarca, alt. 2,700-2,750 m.,
Pennell & Anderson 15104- — Cuzco: Limestone, open ground, alt.
11,300 ft., Gourlay 96; Prov. Anta, alt. 3,200 m., Herrera 3629, Cerro
de Colquepata, rocky slope on trail from Paucartambo to Vilcanota,
492 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
alt. 3,400-3,700 m., Pennell 14176.— Huancavelica : Prov. Tayacaja,
Hills N. of Pampas, alt. 3,200 m., Stork & Horton 10247.
This species differs from all other Bartsias by the combination
of the following characters : galea shorter than the lip and lanceolate-
oblong or lance-triangular, calyx lobes that are acute and clearly
shorter than the calyx tube as well as relatively long, much exserted
corollas.
The taxon appears sporadically over much of Andean Peru.
Bartsia aurea Edwin, Phytologia 19: 363. 1970.
Perennial (?) herb, up to 30 cm. tall, stems and branches lanate,
oldest stems excepted, branching from the base, main branches erect
or soon ascending. Leaves densely pubescent, sessile on the stems
and branches, oblong-linear, up to 4 mm. long, the bracteate blades
narrowly lance-ovate or linear, 5-7 mm. long, margins remotely
crenate, apex obtuse, base occasionally a little narrowed ; spikes ap-
ical, up to 4.5 cm. long, lax or subdense; bracts almost as long as the
calyx, with a few, distant pairs of marginal teeth; pedicels lacking or
up to 1.5 mm. long; calyx cylindric, 7-8 mm. long, densely pubescent,
eglandular or almost so, tube usually a little longer than the narrowly
lance-ovate, acute lobes; corolla yellow, 9-10 mm. long, tube a little
longer than the lips, galea glabrous to puberulent, shorter than the
lip (about one-half as long), lobes of the lip spreading, unequal, ellip-
tically ovate, the median much longer and a little narrower than the
lateral ; filaments inserted near the base of the corolla tube, apically
sparsely glandular with sessile glands, anthers sparsely to less often
densely long-pubescent, basally acuminate; style puberulent on upper
half, stigma-lobes united, punctiform, ovary densely beset with long,
stiff, golden hairs; fruit unknown.
Cuzco: Paucartambo, open, rocky slopes SSW. of Paucartambo,
alt. 3,200-3,400 m., Pennell 14171. The only collection.
Stems slender and with a grey cast when dry. The combination
of the galea much shorter than the lip and also glabrous or sparsely
puberulent, yellow corolla and unequal lobing of the lower lip, as well
as the small size of the corolla separates this taxon from all other
Bartsias.
Bartsia bartsioides (Hook.) Edwin, comb. nov. Lamourouxia
bartsioides Hook., Bot. Misc. 2: 234. 1831. Bartsia densiflora Benth.
in DC., Prodr. 10: 548. 1846; Wedd. Chi. And. Nov. 2: 130. 1860.
Bartsia thiantha Diels in Engler, Bot. 37: 432. 1906.
FLORA OF PERU 493
Perennial, stems erect and branched, often reflexed-hispid, occa-
sionally hair softer, plants pubescent and glandular-pubescent. Leaves
linear, linear-lanceolate, oblong or linear-oblong, up to more than
3 cm. long, most often between 1.0-2.5 cm. and up to 8 mm. wide,
frequently not exceeding 5 mm., crenate or dentate, often bullate,
margins plane or little revolute, obtuse, clasping at the sessile base;
flowers (13-) 15-18 (-20) mm. long, in usually dense spikes, yellow or
purple and yellow, very rarely white, sometimes different colored
flowers on one plant; bracts about as long as the calyx, margins en-
tire or subentire to closely toothed, the teeth rounded-obtuse or less
often acute; calyces 8-12 mm. long, inflated, lobes acute, little shorter
to most often about equal in length to the tube, linear to less often
very narrowly triangular; corolla tube usually a little exserted, some-
times equal to the calyx, longer than the lips, galea little shorter to
little longer than the lip, glandular-pubescent; anthers pubescent,
basally mucronulate, filaments sparsely glandular, the minute glands
with a bulbous base, abruptly narrowed to a linear apiculum; stigma
sparsely puberulent, ovary densely pubescent, hairs stiff, long and
golden; capsule ellipsoid-ovoid, little shorter to as long as the calyx,
pubescent as the ovary but less densely, apex obtuse, apiculate.
By far the commonest Bartsia in Peru and one of the most fre-
quently encountered in the central and southern Andes Mountains.
In Peru from Ancash to Puno at higher elevations.
Ancash: Above Laguna de Llanganuco, 25 km. E. of Yungay, alt.
4,000 m., Edwin 3806. — Apurimac: Cordillera Salcantay, alt. 4,300
m., Rauh-Hirsch P 1400; Prov. Andahuaylas, alt. 2,700 m., Stork &
Horton 10682. — Arequipa: Prov. Moquegua, Saylapa near Carumas,
alt. 3,600-3,700 m., Weberbauer 7347; Nevado de Chachani, alt.
3,200-3,600 m., Pennell 13256; Arequipa, alt. 3,100-3,300 m., Pen-
nell 1^276. — Ayacucho: Prov. Parinacochas, below Caucala, alt.
3,200-3,300 m., Weberbauer 5784- — Cuzco: Colinas de Laxaihuaman,
alt. 3,600 m., Herrera 2374; Cordilleras del Pachahuasan, alt. 4,400
m., Herrera 2575; Tambomachay, alt. 3,800 m., Marin 1417; Sacsa-
huaman, above Cuzco, alt. 3,500-3,600 m., Pennell 13697; Cuzco,
on trail to Pisac, alt. 3,500-3,600 m., Pennell 13697; Corapampa
between Cuzco and Pisac, alt. 3,500-3,600 m., Pennell 13704; Cerro
de Cusilluyoc, alt. 3,400-3,3900 m., Pennell 13811; Cordillera Ve-
ronica, Rauh-Hirsch P 966; Ausangate, alt. 4,700 m., Rauh-Hirsch
P 1150; Soukup 572; alt. 12,500 ft., Stafford 797; Parugo, alt. 3,800
m., Vargas 894; Canas, Asuncion Bridge, alt. 3,720 m., Vargas 11034;
Prov. Urubamba, alt. 2,850 m., Vargas 1156. — Junin: Prov. Huan-
494 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
cayo, alt. 3,400 m., Stork & Horton 10219. — Lima: Matucana, alt.
800 ft., Macbride & Featherstone 429; Canta, alt. 2,700-3,200 m.,
Pennell 14334; Rio Chillon, above Obrajillo, alt. 2,800-3,300 m., Pen-
mil 14428; San Buenaventura, alt. 2,700-3,700 m., Pennell 14621;
alt. 3,600-3,800 m., Pennell 14641 ; Huaros, alt. 3,200-3,600 m., Pen-
nell 14707. — Puno: Lake Titicaca, alt. 4,000 m., Mannheim M 70;
M 137; alt. 400 m., Soukup 103; 303; Putina, Azangaro, Soukup 970.
—Sin. Dept.: Haenke 1981; Sin. Loc.: Ruiz & Pawn s.n.; Soukup
1190.
Bartsia biloba Wedd., Chi. And. Nov. 2: 123. 1860.
Annual up to about 11-12 cm. tall, stems simple or branched from
the base, ascending, hispid, the hairs tending to aggregate in more or
less thick, definite lines. Leaves mostly oblong-lanceolate, up to
9 mm. long and about 1-2 mm. wide, hispidulous, crenate-bullate,
obtuse, at least the lower a little narrowed to the sessile base; spikes
more or less lax, appearing subdense in fruit, interrupted; pedicels
up to 2 mm. long, thickened; flowers up to 11 mm. long; calyx cam-
panulate-oblong, pubescent, 5-7 mm. long, tube much longer than
the lobes, cleft more deeply medianly than laterally; lobes unequal,
the lateral shorter than the median and than the tube, acute; corolla
"drab" purple, tube little longer than the calyx, and corolla lips;
galea concave at apex, densely tomentose, little longer than the lip
whose short lobes are ovate-rounded ; filaments glandular (30x) , an-
thers sparsely pilose or glabrous, mucronate at base; style glabrous,
stigma lobes united, ovary beset with long, stiff, golden hair; capsule
ovate, acute, pubescent as the ovary, about as long as the calyx.
Cuzco: Paso de Tre"s Cruces, Cerro de Cusilluyoc, alt. 3,800-
3,900 m., open grassy paramo, Pennell 13818.
The type also from Cuzco, not seen.
The unequally cleft calyx quickly and clearly delimits this taxon.
Bartsia brachyantha Diels., Bot. Jahrb. 37: 431. 1906.
Subshrub with stems up to 9.5 m. tall, angled, glandular-hispid
above. Leaves twining from the base, ovate or oblong, flattened,
crenate-serrate, hispid, 2-3 cm. long and 1.0-1.2 cm. wide; floral
leaves exceeding the flowers; spike interrupted; flowers very short-
pedicellate; calyx tube subscarious, 4-5 mm. long, teeth triangular,
very hispid, 4 mm. long; corolla purplish-blotched, pubescent, tube
5 mm. long and 3.5 mm. wide, galea 4 mm. long, lip 2 mm. long;
anthers pilose. (After original description.)
FLORA OF PERU 495
Sin. Dept.: Above Cuyocuyo, alt. 3,600-3,800 m., Weberbauer
919. (Material not seen.)
Bartsia calycina Diels., Bot. Jahrb. 37: 432. 1906.
Laxly branched subshrub, up to 50 cm. tall, branches dark purple,
with the leaves glandular-hispidulous above. Cauline leaves patent,
dilated at base, lance-ovate, conspicuously bullate-crenate, 18-22
mm. long and 2-4 mm. wide, none smaller; floral leaves a little
smaller, acute, usually entire, exceeding the flowers; calyx glandular,
teeth linear-lanceolate, very acute, 2 times longer than the tube,
about 10 mm. long; corolla purple, tube about 6 mm. long, galea
7 mm. long, lip 8 mm. long, internally deep yellow; anthers glabrous.
(After original description.)
Ancash: Between Samanco and Caraz near Cajabamba, alt.
3,000-3,500 m., Weberbauer 3166. (Material not seen.)
Bartsia canescens Wedd., Chi. And. Nov. 2: 123. 1860.
Perennial or annual herb, with usually much branched, clustered,
canescent-hispidulous stems. Leaves linear-oblong, hispid, sessile,
4-8 mm. long and 1-2 mm. wide, eglandular as the stems and
branches, crenate and plane to entire and revolute, obtuse, not nar-
rowed to base; spikes few-flowered, lax; flowers sessile or subsessile;
calyx 9-11 mm. long, ovate-oblong, glandular-hispid, lobes linear,
entire, acute almost equaling the length of the tube; corolla 12-18
mm. long, tube exserted, longer than the lips, galea yellow, lip red-
dish; galea glandular-hispid, usually a little shorter than the sub-
glabrous or glabrous lip, rarely equaling its length, lip erect, lobes
unequal, median bigger than the lateral, rounded ; filaments glabrous,
anthers pilose, basally acuminate; style pubescent above, stigma
lobes united, punctiform, ovary narrowly ellipsoid, beset with long,
stiff, golden hair; capsule apically pubescent, about as long as the
enveloping calyx.
Apurimac: Prov. Andahuaylas, Pincos, alt. 2,700 m., Stork &
Horton 10686. — Ayacucho: Weberbauer 5516. — Cuzco: Gay s.n. (not
seen). — Lima: Dombey s.n.
Bartsia curtiflora Edwin, Phytologia 19: 363. 1970.
Erect or ascending, dwarf to 35 cm. tall, usually perennial, rarely
annual herb, usually branching from the base, densely pubescent,
the inflorescence often glandular-pubescent on stems, calyx and co-
rolla. Leaves linear to narrowly lance-subovate, only rarely exceed-
496 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
ing 1.2 cm. long, usually less than 1.0 cm. long and l-3(-4) mm.
wide, subentire to crenulate, plane or little revolute, obtuse, narrowed
or not to the sessile, clasping base; inflorescences long or short, usu-
ally dense, occasionally lax, often interrupted below, spikes or spike-
like; pedicels up to 2 mm. long; bracts linear and few- toothed to
lance-subovate and more or less closely crenulate; calyx tubular-
funnelform, 7-8 mm. long, lobes lanceolate or narrowly lance-oblong
or triangular, a little shorter to about equaling the length of the tube,
acute; corolla usually yellow, 9-12 mm. long, tube from more than
one-half to almost entirely included, longer than the lips, galea a
little shorter than the lip; filaments sparsely glandular on upper one-
half with sessile glands, anthers glabrous to more or less densely
long-pubescent, basally acuminate; stigma lobes united, style puber-
ulent over its entire length, ovary densely stiff, golden pubescent;
capsule longer than the calyx, oblong to ellipsoid-ovoid, obtuse with
a sharp apiculum, the style persisting or deciduous.
Arequipa: Pichu Pichu, alt. 13,500 ft., Stafford 810.—Cuzco:
Prov. Anta, alt. 3,200 m., Herrera 3625; 3626; 3627b; alt. 3,450 m.,
Vargas 296; Cuzco: Ollantaitambo, open rocky slope, alt. 3,000-
3,100 m., Pennell 13643.— Lima: Rio Blanco, alt. 3,000-3,500 m.,
Kittip & Smith 21633.— Puno: Araranca, alt. 4,100-4,300 m., Pen-
nell 13433.
This species differs from all other Bartsias by the combination of
the following characters: galea shorter than the lip, bracts linear to
narrowly lance-subovate as are the leaves, corolla tube largely in-
cluded in the calyx, the anthers sometimes glabrous and the narrow
acute calyx teeth that are as long or almost as long as the calyx tube.
Bartsia diffusa Benth. in DC., Prodr. 10: 546. 1846. B. pumila
Benth. I.e. 546.
Subshrub, dwarf to about 20 cm. tall, decumbent to ascending,
diffusely branched from the base, sometimes stems clustered, pubes-
cent to glandular-pubescent and often viscose. Leaves oblong to
lance-ovate or narrowly ovate, 4-11 mm. long, bullate-crenate, ob-
tuse at apex, little narrowed to the sessile base; inflorescences dense,
usually glandular-pubescent spikes; bracts ovate or obovate, margins
obtusely toothed; pedicels only rarely reaching 2 mm. long; flowers
8-11 (-13) mm. long, yellow, less often yellow and red or red; calyx
tubular-campanulate, lobes obtuse, usually longer than the tube,
entire, occasionally shorter than the tube, crenulate; corolla tube
usually exserted, a little longer than the lips, rarely only equal to the
FLORA OF PERU 497
lips, galea most often shorter than the lower lip, occasionally vary-
ing to a little longer, glandular-pubescent or eglandular or puberu-
lent; anthers glabrous to sparsely pubescent, mucronate basally;
ovary densely pubescent, hair stiff and white; capsule up to 8 mm.
long, exceeding the calyx, pubescent as the ovary, ellipsoid, obtuse.
Arequipa: Nevado de Chachani, alt. 3,900-4,000 m., Pennell
13287', alt. 4,000 m., Rauh-Hirsch P 616; 617; Pichu Pichu, alt.
13,500 ft., Stafford 810.—Cuzco: Cerro de Colquipata, alt. 4,000-
4,200 m., Pennell 13734; Aricoma Pass above Crucero, alt. 16,000 ft.,
Stafford 1099. — Dept. and Prov. Huancavelica: in puna grass, alt.
3,900 m., Stork & Norton 10848. — Junin: Cobracancha Valley, alt.
4,200 m., Grant 7540; near Morococha, alt. 4,350-4,800 m., Grant
7559; Yauli, alt. 13,500 ft., Macbride & Featherstone 912; 924; Huan-
cayo: In puna grass, alt. 4,500 m., Stork 10938; alt. 4,400 m., Stork
10941. — Lima: Cerro Colorado, E. of Canta, paramo slope, alt.
4,000-4,100 m., Pennell 14675; alt. 4,500 m., Rauh-Hirsch P 258.—
Pasco: Cerro de Pasco, Mathews s.n. — Puno: Santa Lucia, alt. 14,000
ft., Sharpe 150; Capachica Peninsular, alt. 12,700 ft., Tutin 1018 —
Sin. Depto: Huaron, alt. 14,000 ft., Macbride & Featherstone 1117.
— Sin. Loc.: Weberbauer 6480; Weddell s.n.
Bartsia duripilis Edwin, Phytologia 19: 364. 1970.
Subshrub, erect and branched, wood and foliage hispid, the pu-
bescence thickened at base, stem 4-angled and striate-sulcate. Leaves
amplexicaul and cordate-subhastate, widest at the base, mostly
shorter than the internodes, oblong, the largest about 4.0 cm. long
and 1.5 cm. wide, frequently 3.0 cm. long and 1.0 cm. wide, very
coarsely crenate, obtuse; flowers axillary, pedicellate, in upper leaf-
axils; bracts longer than the flowers, subovate; pedicels up to 5 mm.
long, densely pubescent; calyx obliquely campanulate, 8-9 mm. long,
tube much longer than the broadly ovate, obtuse, 1 -veined lobes,
pubescent; corolla up to 12 mm. long, dark rose colored with the
lower lip greenish bordered, tube longer than the lips, included in the
calyx, galea dorsally tomentulose, obtuse, at least 2 times longer than
the lower lip, barely emarginate, lip at least partly included in the
calyx, lobes ovate, subequal; filaments with sessile glands on upper
part, anthers pubescent with long, matted hair, acuminate at base;
style puberulent, stigma lobes united, ovary densely beset with long,
golden, stiff hair; fruit unknown.
Ayacucho: Prov. Huanta: Putis, Choimacota Valley, in evergreen
shrub woods, alt. 3,200-3,300 m., Weberbauer 7533.
498 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
A clearly distinct taxon differing from all other Bartsias by the
following combination of characters: stiffly hispid pubescence, large
leaves, included corolla tube and partly included lower lip, oblique
calyx with short, broad, obtuse lobes, longish pedicels, flowers borne
in axils of leaves, not forming a definite spike or raceme and of small
size. The lack of a definite inflorescence is rare in Bartsia.
Bartsia elachophylla Diels., Bot. Jahrb. 37: 431. 1906.
Perennial herb or subshrub, branches ascending, stem hispidulous
above, about 50 cm. long. Leaves minute, sessile, suboblong-crenate,
bullate, 4-6 mm. long and 1 mm. wide, pubescent; spike few-flowered,
short, lax; flowers subsessile or short-pedicellate; calyx tube 4 mm.
long, teeth broadly triangular or triangular, short, 1.5 mm. long;
corolla tube about 4 mm. long, galea 5 mm. long, externally villulose,
light purple, lip erect, 3.5 mm. long, dark yellow or purple; anthers
pilose. (After original description, a few details added.)
Huanuco: Pillao, dry slope, alt. 2,700 m., Woytkowski 34-029.—
Junin: Weberbauer 2087, the type, not seen.
Bartsia elongata Wedd., Chi. And. Nov. 2: 127. 1860. B. elon-
gata subvar. pusilla Wedd., I.e.
Erect, annual, branched, glandular-pubescent herb. Upper leaves
linear-oblong and lanceolate, 5-15 mm. long, lower often wider and
amplexicaul, bullate-crenate, apex elongate; flowers short-pedicellate
or sessile, longer than the bracts; calyx oblong-campanulate, 5-6 mm.
long, glandular-hispid, calyx-teeth linear or linear-oblong, entire,
obtuse, a little longer than the tube; corolla up to 8 mm. long, the
tube as long as the calyx, galea hispidulous or glabrous, subequaling
the lip in length; anthers pubescent, with a very short barb or mucron
at the base, capsule oblong, apiculate (after the original description) .
Only a fragment of Weddell 3665 from the high elevations near
Carabaya seen. Collected (ace. Wedd.) also from the mountains
around Cuzco.
This species is close to B. hispida Benth and B. subinclusa Benth
differing from the former in having a much shorter, almost glabrous
corolla with tube included in the calyx and from the latter in having
a smaller calyx, non-tomentose corolla and more hispid stem. Study
of further collections when available may prove these taxa to be
conspecific.
FLORA OF PERU 499
Bartsia filiformis Wedd., Chi. And. Nov. 2: 126. 1860.
Perennial herb, stems elongate, thin, erect, subsimple, 4-angled,
puberulent. Leaves linear-oblong and linear, 8-12 mm. long, obtuse,
bullate-crenate, not or a little dilated at base, puberulent and gla-
brate; spike elongate, lax; flowers short-pedicellate or subsessile;
calyx 5-6 mm. long, oblong-campanulate, sparsely glandular-hirtus
or almost glabrous, lobes triangular, obtuse, crenate, a little shorter
than the tube; corolla 6 mm. long, tube included, galea ovate, ob-
tuse, pubescent, about 2 times as long as the lip; anthers sparsely
pilose; capsule oblong, subemarginate, apiculate; corolla yellow-
green. (After original description and comments.)
Sin. Dept. : Near Ayapata. Lechler exsicc. n. 1818 (Type K).
Bartsia glabra Edwin, Phytologia 19: 365. 1970.
Herb, stem simple, sparsely puberulent, rarely pubescent with the
short-pubescence subappressed or appressed, foliage and fruit gla-
brous or the capsules occasionally with scant, white pubescence api-
cally. Leaves linear or aciculiform to narrowly oblong-linear, up to
11 mm. long and 1-2 mm. wide, often 6-8 mm. long, margin bullate-
crenate and revolute, obtuse and hammate at apex, slightly broad-
ened and clasping at the sessile base, the costa elevated beneath;
spikes dense above, interrupted below; bracts shorter than the
flowers, similar to the stem leaves, except margins crenate on upper
part only and a little wider in relation to the length; pedicels 1-
2 mm. long; flowers up to 12 mm. long, usually 8-10 mm.; calyx
4-6 mm. long, tube longer than the lobes, pubescent on the veins,
lobes narrowly ovate, acute, pubescent on the veins, margins ciliate;
corolla whitish (in life), tube exserted, longer than the lips, galea
pubescent, a little longer than the lower lip; filaments apically
sparsely glandular and sparsely, very short-puberulent just below
the summit, anthers glabrous to sparsely long-pubescent, acuminate;
style puberulent above, stigma lobes united, ovary glabrous or api-
cally with scant, stiff, white pubescence; capsule ellipsoid-ovoid,
shorter than the calyx.
Amazonas: Cerro de Fraijaco (Huaui-Huni), N.E. of Tambo de
Ventilla, dry jalca, alt. 3,500 m., Pennell 15878.
This species differs from all other Bartsias by the following com-
bination of characters: corolla whitish, foliage, fruit, and ovary
glabrous or almost so, eglandular galea and appressed puberulence
on most of the stems.
500 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Bartsia hispida Benth. in DC., Prodr. 10: 547. 1846.
Perennial, erect, paniculately branched herb, the stems hirsute
with erect trichomes. Leaves lanceolate to linear-lanceolate, up to
about 12 mm. long, scabrous-hirtus crenate-reflexed, upper caudate-
amplexicaul at base; spikes numerous, lax; bracts reaching to about
the calyx, crenate, hispid; flowers subsessile; calyx about 6 mm. long,
glandular-pilose, teeth narrow, shorter than the tube; corolla about
12 mm. long, tube exserted; galea pubescent; equaling to little longer
than the lip ; anthers very sparsely pilose. (After original description.)
Chachapoyas: Mathews 778 (K). — Junin: Tarma, 1-3 km. E. of
Palca, near Carretera Central, alt. ca. 2,000 m., Edwin 3924-
Bartsia inaequalis Benth. in DC., Prodr. 10: 547. 1846. B. cam-
porum and B. thiantha Diels., Bot. Jahrb. 37: 432-433. 1906.
Perennial herb or subshrub, much branched, variously pubescent
or glandular-pubescent, frequently hispid, stems and branches usu-
ally 4-angled, especially above. Leaves variable in size and shape,
narrowly lanceolate to lance-oblong, rarely almost linear, occasion-
ally lance-ovate, up to 3 cm. long or little longer, often the main
leaves exceeding 2 cm. long, frequently clustered beneath, apex acute
to more often subobtuse, margin crenate-bullate, sessile, upper leaves
cordate-amplexicaul ; inflorescences racemose or spike-like, long or
short, few- (6 -8) to many-flowered (up to 30 or more), dense to lax,
pedicels up to 5-6 mm. long, especially the lower; bracts shorter than
the flower, lower similar to the cauline leaves, varying to almost or
rarely quite entire above; calyces 6-11 mm. long, subcampanulate
to subfunnelform, teeth narrowly triangular to lanceolate, acute,
usually shorter than the tube; corolla dark red to purple, large for
the genus, up to 2.7 cm. long, frequently exceeding 2.0 cm., tube sub-
conical, longer than the lips, throat a little inflated, galea about 2
times longer than the lip, densely pubescent or glandular-pubescent
externally, notched at apex, lobes of the lip ovate, obtuse, sometimes
the median apically notched, filaments sparsely, microscopically pu-
berulent on upper half, stigma lobes united, ovary densely matted
with golden hair; capsule ovoid, little shorter or longer than the
calyx, the tube of which is greatly inflated, obtuse apically pubescent
as the ovary, the style persisting or early deciduous.
Apurimac: Cord. Salcantay alt. 3,500 m., Rauh-Hirsch P 1454-
— Arequipa: Pichu Pichu, alt. 11,500 ft., Stafford 811. — Cuzco: Prov.
Urubamba, Macchu Picchu, alt. 2,000-3,000 m., H err era 2000; Rauh-
Hirsch P 785; Saunders 426; Stafford 795; San Gabon, Chichacori
FLORA OF PERU 501
Valley, 10,500 ft., Fisher 18; Allantaitambo, alt. 2,800 m., Herrera
34.11; Yanamancha, alt. 3,799 m., Marin 1783; Sacsahuaman, alt.
3,500-3,600 m., Pennett 13551; below Colquipata, alt. 3,200-3,300
m., Pennell 13775; suburbs of Cuzco, Soukup 571; Paucartambo,
alt. 3,000-3,100 m., Vargas 314; Colinas del Rodadero, Vargas 3131.
— Huancavelica: Prov. Tayacaja, Pampas-Salcabamba Trail, alt.
2,500 m., Stork & Horton 101+38. — Huanuco: several km. SE. of
Carpish, alt. 2,800 m., Stork & Horton 9904- — Lima: Prov. Huaro-
chin, km. 85, Carretera Central, high mountain above Matucana,
alt. 10,500-11,000 ft., Saunders 289.— Sin. loc.: Diehl, 2556.
This common species of southern Peru has the largest flowers
found in the genus in the country. These are uniformly dark red
to purple.
Also collected in Bolivia and Ecuador where the leaves are larger,
wider in relation to their length, and more coarsely crenate.
Bartsia integrifolia Wedd., Chi. And. Nov. 2: 131. 1860.
Minutely puberulent, thin stemmed, usually branching subshrub.
Leaves narrowly linear, 5-15 mm. long, margin entire, revolute, apex
acute, base not or little dilated; flowers short-pedicellate; calyx ob-
long-campanulate, 5-9 mm. long, puberulent or very finely tomen-
tose, calyx lobes linear-triangular, entire, acute, little shorter to
equaling the tube; corolla 10-15 mm. long, tube exserted, galea ovate,
obtuse, glabrous, little shorter than the tube; anthers densely pilose,
mucronulate at base; capsule oblong, obtuse, apiculate. (After the
original description.)
Only a fragment of Castelenau,s.n. (the type) from Depto. Lima
seen. The only other collection cited is Cay, s.n. from Cuzco.
Entire-leaved species of Bartsia are not common.
Bartsia meyeniana Benth. in DC., Prodr. 10: 546. 1846.
Erect or ascending soft-pubescent and glandular-pubescent per-
ennial herb. Leaves oblong-lanceolate, up to about 1.6 cm. long,
usually shorter, obtuse, sessile and sometimes clasping, broadest at
the base, crenate, crenations bullate; spikes short, interrupted below,
subdensely few-flowered; bracts smaller than but similar to the
leaves, elliptic or oblong; flowers subsessile; calyx 7-9 mm. long, the
narrowly triangular, obtuse teeth almost as long as the tube; corolla
red, 16-19 mm. long, tube longer than the lips, galea sparsely to
densely pubescent, much shorter than the spreading, reflexed lip (ca.
one-half the length of the lip) ; filaments short for the genus, inserted
502 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
at the apex of the tube, dotted with a few, sessile, red glands on the
upper part; anthers pubescent with long, white hair, with a short,
sharp acumin at base; style pubescent on the upper half, stigma
lobes united, ovary densely beset with long, golden hair; capsule
ex desc., oblong, acute, pubescent.
Junin (Pisco) : Castroverreina, Pisco Valley, alt. 3,600 m., Rauh -
Hirsch P 372. — Also reported, Sin. Dept.: near Tacora, Meyen s.n.
Found at high elevations in Bolivia.
Although the Peruvian specimen is erect and 40 cm. tall, the
taxon elsewhere contains dwarfs with ascending branches.
Bartsia mutica Benth. in DC., Prodr. 10: 548. 1846. Euphrasia
mutica HBK., Nov. Gen. et Sp. 2: 334. 1817. Bartsia chilensis Benth.
in DC., Prodr. 10: 547. 1846.
Based upon an unpublished name of Bertero, "Lamourouxia eu-
phrasioides," not conspecific with Bartsia euphrasioides Wedd. of
Ecuador.
Annual or perennial, erect, much branched, glandular, hispid
herb. Leaves hispid, oblong to linear-lanceolate, obtuse, up to 2 cm.
long or longer, grossly crenate, plane or a little revolute, cordate-
amplexicaul at base; floral leaves smaller, otherwise as the cauline;
spikes lax; pedicels mostly 2-3 mm. long; calyx 7-9 mm. long, pubes-
cent, lobes linear, subacute to subobtuse, often unequal, about as
long as the tube, entire to less often sparsely denticulate or serrulate;
corolla purple, 10-16 mm. long, the tube longer than the lips, ex-
serted, galea pubescent or more often glandular-pubescent, longer to
shorter than the lip with lobes elliptical to narrowly ovate; filaments
glabrous, the pairs subequal in length, anthers glabrous, blunt to
acute at base, neither mucronate nor acuminate; ovary densely pu-
bescent, style elongate, stigma unequally bifid; capsule longer or
shorter than the calyx, pubescent.
Lima: Along Rio Chillon, near Obrajillo, alt. 2,200-2,500 m.,
Penned 14326. — Sin. Dept.: Between Lucarque and Ayavaca, Bon-
pland 3466. (Fragment seen.)
This taxon is not uncommon in Chile. The description has been
based largely on specimens collected there. Both Benthan and Wed-
dell found difficulty in distinguishing B. mutica from B. chilensis. It
seems the better course to unite the specimens under one appellation
at least until further Peruvian specimens become available.
FLORA OF PERU 503
Bartsia orthocarpiflora Benth. in DC., Prodr. 10: 545. 1846.
Perennial herb, occasionally annual, variously pubescent, stems
erect, branched, long- and short-pubescent. Leaves hirtellus, sessile,
bullate-crenate, linear-oblong to narrowly lance-ovate, widest at the
base, obtuse apically; spike usually short, basally interrupted; pedi-
cels 1-2 mm. long, calyx oblong-campanulate, 5-6 mm. long, tube
much longer than the 1.5 mm., frequently obtuse teeth; corolla red
to green-yellow, up to 16 mm. long, mostly 9-14 mm., most often
narrow, tube much exserted, densely puberulent and often glandular,
longer or shorter than the galea, galea about 3 times as long as the
lip, apically barely emarginate; lobes of the lip about 1 mm. long;
filaments glabrous or under 30 X, apically, sparsely glandular or
puberulent, anthers densely long-pubescent, basally mucronate; style
puberulent, stigma lobes wholly united, punctiform to subulate;
ovary densely beset with long, stiff, golden hair; capsule ellipsoid-
ovoid, apically long and stiff pubescent, longer than the calyx.
Apurimac: Rio Pinco, alt. 3,500-3,600 m., Weberbauer 5904.—
Cuzco: Cerro de Cusilluyoc, open grassy slope, alt. 3,400-3,500 m.,
Pennell 13808.— Huanuco: Pillao, 2,700 m. alt., Woytkowski 50 —
Junin: Prov. Huancayo: between Huancayo and Acopalca, alt.
3,700 m., Stork 10924— Sin. Loc.: Pavon s.n.
A basically Bolivian species but reaching to South-Central Peru.
The galea that is about three times longer than the lip helps to de-
limit this taxon.
Bartsia parvifolia Benth. in DC., Prodr. 10: 545. 1846.
Perennial herb, stems minutely puberulent, branched, the branch-
ing opposite, at least the lower and middle nodes longer than the
leaves on the main stem and branches. Blades linear-oblong to nar-
rowly oblong-subovate, reaching 2.0 cm. long, mostly less than
1.0 cm. long and (a few) up to 5 mm. wide, frequently narrower,
hispidulous, sessile and often clasping, bullate-crenate, obtuse, base
narrowed or a little wider; spike lax; flowers sessile or subsessile;
calyx oblique, ovate-campanulate, pubescent, especially on the main
veins, tube 4.5-5.0 mm. long, lobes 1.5-2.0 mm., broadly triangular;
corolla about 10 mm. long, tube a little exserted; galea red, tomen-
tose, little longer than the yellow or green lower lip ; anthers pilose,
mucronate at base; stigma lobes united, punctiform; capsule oblong,
about as long as the calyx.
504 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Huancayo: Huaytapallana, alt. 4,900 m., Rauh-Hirsch P 1737.
— Junin: Prov. Tarma: Between Palca and Carpapata, alt. 2,900 m.,
Stork 10980.
This taxon described from Venezuela differs from Bartsia bart-
sioides (Hook) Edwin in the shape of the calyx lobes and their size
in relation to the tube, the size of the flower, and the lax inflorescence.
Bartsia pedicellata Edwin, Phytologia 19: 366. 1970.
Erect annual, simple or with a few, short rosulate branches, stems
pubescent-puberulent with flattened, somewhat curly, multicellular
hair. Leaves glabrous or sparsely pubescent on the margins and main
veins with multicellular hair, linear or narrowly oblanceolate, up to
about 1.0 cm. long and 0.2 cm. wide, crenate-revolute, apex obtuse,
hammate, base sessile subclasping; floriferous over most of the length
of the stems in lax racemes; bracts wider than the cauline leaves;
pedicels 7 (-8) mm. long, erect, almost parallel to the stems; flowers
8-11 mm. long; calyx oblique, tube longer than the crenate, obtusely-
rounded lobes ; corolla purple, tube longer than the lips, galea tomen-
tose with multicellular hair, about twice as long as the lip, lobes of
the lip elliptic-oblong; filaments sparsely glandular above with ses-
sile glands, anthers tomentose, acuminate at base; style glabrous,
stigmas united, ovary glabrous; capsule glabrous, oblong, almost
twice the length of the calyx, mucronulate at the obtuse apex.
Cuzco: Paucartambo to Tres Cruces, Cerro de Cusilluyoc, moist
gravelly banks and swales, alt. 3,500-3,800 m., corolla "dull magenta
purple." Annual herb. Pennell 13815.
This taxon differs from all other Bartsias by the combination of
multicellular, flattened pubescence on the stems and branches, erect,
strict, 7-8 mm. long pedicels, crenate calyx lobe, and glabrous cap-
sule twice the length of the calyx.
Bartsia pedicularioides Benth.. in DC., Prodr. 10: 546. 1846.
Perennial, root shoots short, thin ascending, leaves minute to
about 8-10 mm. long, short-petiolate. Stems erect, much branched,
viscose, pilose; leaves oblong, subsessile, larger than root shoots,
bullate-crenate, obtuse; spike glandular-pilose, interrupted at base;
flowers violet, at least the lower pedicellate; calyx 5-6 mm. long,
ovate-oblong, lobes triangular-ovate, shorter than the tube; the co-
rolla 8-9 mm. long, little exceeding the calyx, lips subequal, galea
hirsute or almost lanate, oblong, obtuse, lobes of the lip short, ob-
tuse; anthers glabrous; capsule ovate, 8-11 mm. long, hirsute at apex.
(After original description and Weddell I.e. 121-122.)
FLORA OF PERU 505
Cuzco: Mountains of Cuzco. Gay s.n. (Material not seen.) Also
reported from Venezuela and Ecuador.
Weddell includes Bartsia parvifolia Benth. in the synonymy of
this taxon. It is retained as a species in this paper.
Bartsia peruviana Walp., Nov. Acta. Acad. Caes. Leopold.
Carol. Nat. Cur. 19: suppl. 1, 400. 1843; Rep. 3: 408. 1844; Benth.
in DC., Prodr. 10: 357. 1846; Wedd. Chi. And. Nov. 2: 128. 1860.
Herb with opposite branching, erect, glandular-tomentose stems,
blackening in drying; leaves linear, 13-20 mm. long and 3-6 mm.
wide, crenate-dentate, obtuse, glandular-tomentose, sessile; inflores-
cences spicate or flowers axillary, subsessile, solitary; calyx 18-20
mm. long, campanulate, equally 4- toothed, teeth linear, obtuse; co-
rolla brown, little longer than the calyx; seeds very small, subsmooth.
(Description after Walpers, 1844.)
Arequipa: Chuquibamba Prov.: Condesuyos, under rocks, open
hillside, alt. 10,500 ft., Stafford 1214.—Puno: Laguna de Titicaca &
Tisslaoma, Meyen 3900 (material not seen) .
Type (K) from Bolivia (Sorata).
Bartsia sanguinea Diels., Bot. Jahrb. 37: 433. 1906.
Caespitose dwarf, stems 5-8 cm. tall, sparsely pilose or glabrate,
blackening on drying. Leaves thickened, almost glabrous, obovate,
conspicuously bullate-crenate above, almost entire below; spike few-
flowered; calyx subinflated, subhyaline, pilose, 7 mm. long and 4.5
wide, teeth triangular; corolla blood-red, tube about 6-8 mm. long;
galea densely short- tomentose, 6-7 mm. long, lip 5 mm. long; anthers
pilose; capsule glabrous at the apex. (After original description.)
Sin Dept.: Roadside between Cuzco and St. Anna, alt. 4,000 m.,
Weberbauer ^9^5. (Not seen.) The type seen, from Bolivia (So-
rata), Rusby 1092 (K).
Bartsia santolinaefolia Benth. in DC., Prodr. 10: 546. 1846.
Euphrasia santolinaefolia HBK., Nov. Gen. et Sp. 2: 333, pi. 166.
1818.
Perennial herb to subshrub, from almost unbranched to profusely
branching from the base, erect, elongate, frequently rigid, usually
glandular-pilose, often (in life) viscose. Leaves oblong-linear to lan-
ceolate or lance-ovate, up to almost 3 cm. long and 4-5 mm. wide,
usually considerably smaller; margins coarsely crenate-bullate, apex
506 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
obtuse, the sessile base usually not narrowed, often clasping; spikes
dense to lax and often interrupted basally; bracts ovate or obovate,
closely to subdistantly, coarsely crenate, subsessile, about as long as
or little longer than the calyx, uppermost often only with 1-2 pairs
of teeth; calyx tubular-campanulate, 6-8 mm. long, lobes oblong,
obtuse, entire and longer than the tube, rarely crenate-crenulate
and shorter than the tube; corolla yellow to purple, 9-13(-15) mm.
long, tube exserted, longer than the lips, galea little longer than the
lip, puberulent or glandular-pubescent, glands on long hair; anthers
glabrous or pubescent, mucronulate at base, filaments puberulent;
ovary densely long, stiff, golden pubescent; capsule pubescent on up-
per half as the ovary, ellipsoid, not filling the inflated calyx and about
equal in length, emarginate and frequently apiculate.
Amazonas: Daguas on Rio Sonche, E. of Chachapoyas, alt. 2,400
m., Pennell 157^5; Mathews s.n. — Ancash: Cordillera Blanca, alt.
3,900 m., Rauh-Hirsch P2057. Cajamarca: S.W. of Cajamarca,
alt. 2,800-2,900 m., Pennell & Reichlin 1^969— Cuzco: near Rio
Paucartambo, below Paucartambo, alt. 2,900-3,200 m., Pennell
13796. — Huanuco: slope of Hill, western slope of Torre-huase, alt.
3,300 m., Woytkowski 329.
This taxon forms a close alliance with B. diffusa Benth. and B.
hispida Benth.
Bartsia simulans Edwin, Phytologia 19: 367. 1970.
Perennial shrublet up to about 20 cm. tall, branching from the
base. Stems terete, drying light to dark brown, the soft, glandular-
pubescence mixed with soft and also scattered, stiff, eglandular hairs;
leaf-blades thickened, sessile, pubescent above, beneath and on the
margins as the stem, except more stiff hairs present, obovate or
spathulate-obovate or oblong-obovate in outline, up to about 1.0 cm.
long and half as wide, margins with 3-5 pairs of rounded, close or
distant lobes, these increasing in size base to apex, the apical lobe
largest, narrowed to base, venation obscure above, apparent beneath ;
flowers in short, apical, dense, spikes; bracts similar to the leaves,
except a little smaller and the 2-3 pairs of lobes usually more or less
distant, shorter than the flowers; calyx glandular, 9-11 mm. long,
tube 4-5 mm. long, little shorter than the 5-6 mm. long, lance-
oblong or oblong, subacute lobes; corolla 10-13 mm. long, yellow
and red, almost wholly included in the calyx to little exserted, the
corolla tube entirely included, narrow, longer than the lips, galea
3.0-4.5 mm. long, dorsally densely glandular, apically notched, little
FLORA OF PERU 507
shorter than the lip the lobes of which are very broadly ovate to sub-
orbicular, the median little larger than the lateral; stamens didyna-
mous, the pairs very unequal in length, anthers glabrous, basally
mucronulate, filaments sparsely puberulent (under 20 X); style ex-
ceeding the stamens, stigma exserted punctiform, ovary densely
golden pubescent, the hairs stiffish; capsule elliptical or elliptically
obovoid, little shorter than the calyx, pubescent at least apically with
long, stiffish, golden hairs and apiculate style tardily deciduous or
persistent, placenta thin; seed numerous, ovoid, narrowed to both
ends, testa striate-striatulate, closely adherent.
Puno: San Antonio de Esquilache in clefts in face of rock at alt.
15,500 ft., May 16, 1937. Stafford 743.
Most closely resembling Bartsia diffusa Benth. which has ovate
bracts with lobes decreasing in size base to apex, exserted corollas,
wider calyx lobes that are usually not longer than the calyx tube,
and different pubescence.
Bartsia subinclusa Benth. in DC., Prodr. 10: 547. 1846.
Hispidulous-scabrous plant, stems glandular-hirtus above, soft-
pubescent below. Leaves sessile below, cordate-clasping above, pu-
bescent with stiffish, white hair, especially on the margins and main
veins, linear-lanceolate, up to about 1.8 cm. long and 3-5 mm. wide,
crenate, crenations recurved, obtuse, spike few-flowered ; calyx about
10 mm. long, teeth linear, obtuse, shorter than the narrow tube,
exceeding the corolla tube and throat; corolla tomentose, only the
apices of the lips exserted ; galea about equal to the lip, tube equal to
or shorter than the galea; anthers sparsely long-pubescent, basal
acumin short. (After original description.)
Sin. Dept.: Near Tissacoma, alt. 15,000 ft., Meyen s.n. (Type K.)
Bartsia trichophylla Wedd., Chi. And. Nov. 2: 122. 1860.
Small perennial (subshrub?) with the stems profusely branching
from the base, primary branches ascending, to 10 cm. tall, apically
hispid-pilose. Leaves densely crowded, especially below, lower 4-
5 mm. long and 1.0-1.5 mm. wide, size increasing to the middle of the
plant to about one-half bigger, oblong or spathulate-lanceolate, ob-
tuse, bullate-crenulate, attenuate to a short petiole, glabrous except
for the margin (that is) ciliate with articulated 1 mm. long hair;
flowers about 6-8, in sometimes lax racemes; pedicels almost as long-
as the 5 mm. calyx but shorter than the bracts; calyx oblong-cam-
panulate, sparsely pilose, lobes oblong, obtuse, crenate, shorter than
508 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
the tube; corolla tube glabrous, 8-9 mm. long, little exserted, galea
densely tomentose, oblong, obtuse, about 2 times longer than the
lip; lobes of the lower lip about 1 mm. long, rounded, glabrous; fila-
ments sparsely, retrorsely barbed on one side, anthers sparsely
barbed ; ovary glabrous. (From original description and comments.)
Sin. Dept. : South Peru. Rocky hills between Puno and Arequipa,
alt. 4,000 m., Weddell s.n.
Bartsia viridis Edwin, Phytologia 19: 367. 1970.
Annual, erect herb, old stems glabrate, younger with definite lines
of soft pubescence, all stems 4-angled, sulcate, drying very dark red
to black. Leaves linear or aciculiform, usually minute to 6 mm. long,
occasionally reaching 2.0 cm., bullate-crenate and revolute, apex ob-
tuse, hammate, often a little widened to the clasping base, sessile,
glabrous or sparsely pubescent, main vein elevated beneath, other
venation not visible ; bracts crenate, at least the upper ellipsoid-ovate,
about as long as or barely exceeding the calyces; flowers mostly ag-
gregated into short, lax, interrupted spikes, sessile or subsessile on
pedicels reaching 2 mm. long, usually not exceeding 1.5 mm., and
very small for the genus, 8-10 mm. long, occasionally up to 11 mm.
long; calyx narrow, usually glandular-puberulent, 4.0-5.5 mm. long,
tube longer than the lance-triangular, acute lobes; corolla green,
tube longer than the lips, galea 2 times as long as the lower lip,
usually 1.5 times longer, eglandular, pubescent, lobes of the lip un-
equal, the median longer than the lateral, all ovate, rounded; fila-
ments sparsely dotted with small, sessile glands, anthers pubescent,
mucronate at base; style puberulent above, stigma lobes united,
ovary beset with long, stiff, golden hair; capsule ovoid to broadly
ovoid, obtuse to rounded at apex, with long, stiff, golden hair api-
cally, a little shorter than the calyx and enveloped by it.
Cuzco: Paso de Tr£s Cruces, Cerro de Cusilluyac, mossy banks
and knolls on paramo, alt. 3,700-3,900 m., Pennell 13826.
The combination of very small, green flowers with included co-
rolla tube and partly included lower lip as well as the arrangement
of the pubescence on the stems and branches separates this taxon
from all other Bartsias.
According to the collector the corolla is "mignonette green."
Bartsia weberbaueri Diels., Bot. Jahrb. 37: 431. 1906.
Vine, the stem twining among shrubs, up to 2 m. long, hispidulous
above. Leaves patent, sessile, cordate at base, oblong or lanceolate-
FLORA OF PERU 509
oblong, stem leaves 15-20 mm. long and about 4-5 mm. wide, cre-
nate, leaves of the lateral branches smaller; flowers on slender pedi-
cels, 5-10 mm. long; calyx glandular-hispid, tube 10 mm. long and
6-7 mm. wide, teeth triangular, about 7 mm. long; corolla tube
10 mm. long, galea wide, externally villose, 10 mm. long, dark pur-
ple, lip erect, 6-7 mm. long, bright green; anthers pilose, exserted.
(After original description.)
Ancash: Cajatambo Prov., near Ocros, in shrub snag, alt. 3,500-
3,700 m., Weberbauer 2692. (Material not seen.)
BASISTEMON Turcz.
Basistemon Turcz., Bull. Soc. Imp. Nat. Mosc. 36(2) : 214. 1863.
Hassleropsis Chod., Bull. Herb. Boiss. ser. 2, 4: 285. 1904. Saccan-
thus Herzog., Meded. Rijks Herb. Leid. 29: 47. 1916.
Reference: Baehni and Macbride, Candollea 5: 345. 1934. and
7: 185-188. 1936. Junell, Symbol. Bot. Upsal. 4: 1934. Moldenke,
Bull. Torr. Bot. Club 63(6): 345. 1936.1
Shrubs or small trees, usually subglabrous or new wood sparsely,
minutely puberulent; leaves opposite, subsessile to sessile, ovate or
broadly ovate, long-acute to acuminate, acumin sharp, 4-15 cm.
long and 1.8-8.0 cm. wide, base subacute to obtuse, margin entire,
or minutely serrulate, glabrous, punctate or puncticulate on both
surfaces; petioles when present only up to 2-3 mm. long, swollen at
base, margins sparsely puberulent; flowers axillary, 2-5 fasciculate,
pedicellate; pedicels 3-10 mm. long, glabrous, or sparsely, minutely
puberulent, bracteolate at base; calyx about 3 mm. long, ovate;
corolla bilabiate, 5-8 mm. long, tube short to long, saccate at base,
lips subequal, lobes 5, all about equal or the 2 anterior little shorter;
stamens 4, didynamous, the pairs little to very unequal, all inserted
at or near the base of the tube, anther connective much enlarged at
maturity, the thecae at length divergent and dehiscing by longitudi-
nal slits; ovary shorter than the erect style, stigma lobes united; cap-
sule ex desc. loculicidal; seeds few to numerous, axile and pendulous,
albumen lacking. Two species collected in Peru.
A genus of six species at present wholly restricted to South Amer-
ica in Peru, Colombia, Argentina, and Paraguay, from 30-1,500 m.
1 The details of the checkered taxonomic and nomenclatural history of this
genus are beyond the scope of this flora. Although only six species are known at
present, the genus has been placed in the Verbenaceae and the Bignoniaceae as well
as in the Scrophulariaceae, where it provisionally appears here.
510 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
alt. Very few collections known, perhaps not more than 40. The
specific variation displayed by and the altitudinal spread of the few
collections would seem to imply a large genus (in species number)
yet poorly collected.1
Leaves 10-15 cm. long and 5-6(-8) cm. wide; petioles 2-3 mm. long.
B. peruvianus.
Leaves 3.5-6.0 cm. long and 1.5-3.3 cm. wide; petioles lacking or up
to 0.8 mm. long B. intermedius.
Basistemon intermedius Edwin, Phytologia 19: 368. 1970.
Unarmed shrub with glabrous leaves, otherwise mostly minutely
puberulent. Leaves ovate, acuminate, 3.5-6.0 cm. long and 1.5-
3.3 cm. wide, bases obtuse, margins minutely serrulate, punticulate
on both surfaces; petioles up to 0.8 mm. long or lacking, minutely
puberulent, gibbose at base and at least partly surrounding the stem ;
flowers 2-3-fasciculate, in axils of unmodified leaves, pedicellate;
pedicels 3-5 mm. long, puberulent, with a pair of minute, hard, pu-
berulent bracteoles at the bases; calyx with 5 ovate, puberulent
sepals, 3.0-3.5 mm. long, apices long-acuminate, margins ciliolate,
united about one- third their length into a campanulate tube; corolla
weakly saccate at base, white, 7-8 mm. long, bilabiate, lips subequal,
the 2 anterior lobes little shorter than the 3 posterior, tube about
6 mm. long; stamens 4, didynamous, anterior pair about 2 mm. long,
posterior 3 mm. long, anthers sub ovoid, divergent, dehiscing by longi-
tudinal slits, connective enlarged, hardened at maturity, filaments
gibbose at base, inserted about 1 mm. above the base of the corolla
tube; ovary narrowly ellipsoid-ovoid, much shorter than the style,
stigma lobes united punctiform; fruit unknown.
Huanuco: Ganso Azul, Agua Caliente on Rio Pachitea, alt. 1,000
ft., dense shade; Oct. 17, 1942, Sandeman 3380. (Type K; Isotype
OXF.)
Near B. peruvianus Benth. & Hook, f . apud Hook f . & Jacks, but
differing in the length of corolla lobes in relation to the tube, distri-
bution and amount of vesture and size of the leaves. The taxon
appears to stand between B. peruvianus and B. rusbyi Moldenke.
Basistemon peruvianus Benth. & Hook, f . apud Hook f. &
Jacks. Ind. Kew. 1: 277. 1895.'
1 An opinion of Pennell in correspondence to Moldenke.
2 Basistemon in Benth. & Hook, f., Gen. PI. 2: 1,224-1,245. 1876, barely re-
ferred (without specific diagnosis) to the species.
FLORA OF PERU 511
Bracteoles and sepals ciliolate, petioles minutely puberulent,
otherwise a glabrous shrub or small tree. Pedicels 5-7 (-8) mm. long,
bracteoles minute; calyx lobes 2.5-3.5 mm. long, narrowly ovate,
connate about one-third their length into a flaring tube; corolla (ex
coll.) lilac, white and cream, tube ca. 1.5 mm. long, lobes ca. 4.5 mm.
long; stamens included or little exserted, filaments 1.5-2.5 mm. long,
flattened or terete, anthers separated on a bulbous connective, diver-
gent; ovary narrowly ellipsoid. Fruit not seen.
San Martin: Juan Jui, Alto Rio Huallaga at 400 m. alt., forest,
King 3780 (K) ; near Tarapoto, Spruce 4515, the type (K) . The Klug
specimen is in close agreement with the type seen at Kew.
BUCHNERA L.
Annual or perennial, probably hemi-parasitic herbs, becoming
black upon drying. Stem erect, simple or branched, glabrous to
hispid-scabrid, hairs often callus based; leaves all opposite, sometimes
subopposite above or all alternate, simple, usually sessile, occasion-
ally short-petiolate, rarely forming small, basal rosettes, filiform to
linear-lanceolate, less than 1 mm. to a few mm. wide, apex acute or
obtuse, margin entire or with a few, scattered, small teeth, 1-5-
nerved; inflorescence terminal, spicate, compact to lax; flowers sub-
tended by one bract and two bracteoles; calyx tubular, 10-nerved,
regularly to irregularly 5-dentate, glabrous to hispid-scabrid or ver-
rucose; teeth erect, spreading or reflexed; corolla white, blue or
violet, salverform, tube longer than the calyx, externally glabrous to
softly pubescent, lobes subequal, ovate to subrotund, entire to emar-
ginate, throat pilose; stamens 4, included, slightly didynamous; an-
thers monothecous; stigma linear-cylindric, style apex clavate, en-
tire; capsule ovoid, oblate to cylindric, shorter to longer than the
calyx, loculicidal, seeds numerous.1
A genus comprising about 100 species which are almost wholly
tropical and chiefly Old World. Sixteen species are found in the New
World. A single collection is known from Canada. Taxa are found
more or less abundantly in the United States, Central America and
the West Indies and South America, Chile excepted. Four species
have been found in Peru.
1. Corolla externally glabrous; annual herbs.
1 This generic description and the following species descriptions after Philcox,
Kew. Bull. 18 (2): 275-315. 1965.
512 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
2. Calyx teeth lanceolate — or long — acuminate, more or less
spreading; bracteoles linear or linear-lanceolate.
3. Leaves alternate; calyx glabrous; stems glabrous or sparsely
pubescent below with adpressed, upward directed hairs.
B. palustris.
3. Leaves opposite or subopposite; calyx tube hispid-scabrid
on nerves; stems pubescent throughout with callus-based
hairs B. pusilla.
2. Calyx teeth triangular, erect; bracteoles narrowly ovate-lance-
olate B. weberbaueri.
1. Corolla externally soft-pubescent; perennial herbs B. rosea.
Buchnera palustris (Aubl.) Spreng., Syst. 2: 805. 1825. Piripea
palustris Aubl., PI. Guian. 2: 628, t. 253. 1775.
Annual herb, up to 65 cm. tall, usually not exceeding 45 cm., sub-
glabrous. Stem simple or branched, terete, glabrous above to
sparsely pubescent below with upwardly directed hairs; cauline
leaves alternate, narrowly linear, 1.0-3.5 cm. long and 0.5-1.5 mm.
wide, entire, acute, 1-nerved, minutely scabrous-verrucose, finely
hispid on the margin basally; lowest leaves (basal) minute, obovate,
entire; inflorescence lax, up to 15 cm. long; bracts ovate-lanceolate,
subacuminate, 3.0-5.5 mm. long, scabrous-verrucose, ciliate, cari-
nate; bracteoles linear-lanceolate, setaceous-ciliate, little shorter than
bracts; calyx 7-10 mm. long, not reticulate between main nerves,
glabrous, teeth erect, lanceolate-acuminate, subciliate; corolla blue,
pink to purple, tube slender, glabrous, incurved, up to twice as long
as the calyx, lobes obovate, emarginate, subequal; capsule oblate-
cylindric, obtuse, equaling to slightly longer than the calyx; style
base deciduous.
San Martin: Tarapoto, Spruce 4397 p.p.
Riversides and marshy places from Colombia through Venezuela
and the Guianas to Brazil and westward to eastern Peru.
Buchnera pusila HBK., Nov. Gen. et Sp. 2: 340. 1818. B. tine-
toria Bertol., Fl. Guatim. 26. 1840; B. pilosa Benth., Bot. Voy. Sulph.
144. 1845. B. lithospermifolia sensu Benth. in DC., Prodr. 10: 497.
1846, non HBK. B. major Polak, Linnaea 41: 588. 1877. B. elon-
gata sensu Hemsl., Biol. Centr. Am. 2: 457. 1881 et sensu Goyena,
Fl. Nicar. 619. 1911, non Sw.
FLORA OF PERU 513
Annual herb, usually 15-30 cm. tall, rarely up to 70 cm. Stem
slender, simple or branched, hispid with callus-based hair; leaves
subopposite, upper linear, entire or subentire, up to 2 mm. wide,
1-nerved, densely hispid-scabrid, lower obovate-oblong to oblong,
entire to deeply dentate, obtuse or acute, 1.5-5.0 mm. wide, hispid-
scabrid, often similar to upper leaves; inflorescence up to 12 (-24)
cm. long, lax to congested; bracts lanceolate, hispid-scabrid, up to
5.5 mm. long; bracteoles linear, shorter than bracts; calyx up
to 8 mm. long, nervation between the hispid-scabrid main veins
lacking, otherwise glabrous, teeth long-acuminate, spreading to re-
flexed in fruit; corolla usually white, occasionally pink to lilac or
purple, tube slender, externally glabrous, little to twice as long as
the calyx, lobes 4-10 mm. long, ovate to obovate, entire, throat
pilose; capsule oblong, compressed, equal to the calyx.
Cuzco: Prov. Urubamba, petals pink, open grass slope, alt. 2,900
m., Vargas 2702. — Junin: Mito, northeastern grasslands, flowers
pink, alt. about 9,000 ft., Macbride & Featherstone 14-08. — Sin. Dept.:
Pawn s.n. (coll. 1827, specimen at P).
The most variable of all the new world species of this genus.
More or less frequently encountered in low-lying meadows to
medium altitude oak and pine woods, and rocky hillsides from Mex-
ico through Central America to South America including Venezuela,
Brazil, British Guiana, Colombia, Ecuador, and Peru.
This species and B. longifolia HBK. are very closely allied and
perhaps even conspecific. The distribution of the latter taxon is
essentially that of B. pusila in South America.
Buchnera rosea HBK., Nov. Gen et Sp. 2: 342. 1818. B. rosea
var. congesta Schmidt, in Mart., Fl. Bras. 8: 328. 1862. B. litho-
spermifolia sensu Stand!., in Contr. U. S. Nat. Herb. 27: 334. 1928,
non HBK.
Perennial herb (30-)45-60(-115) cm. tall. Stem simple or
branched, upright, glabrescent above to hispid-pilose below; leaves
opposite to subopposite (the lowermost sometimes alternate), linear-
lanceolate, 4.5-8.5 mm. wide, entire or serrulate, rigid, subcoriaceous,
3-5-nerved, nerves prominent beneath, glabrous to scabrous above,
subglabrous to hispid-scabrous beneath, especially on the nerves and
margins; inflorescence spicate, variable, elongate and lax (20 cm.
long) to tightly compact and many-flowered (3.5 cm. long); bracts
ovate-lanceolate, about 2.5-4.0 mm. long, hispid-scabrid, ciliate;
514 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
bracteoles linear-subulate, hispid; calyx erect, up to about 7.5 mm.
long in fruit, densely reticulate between the main nerves, spreading
hispid, pubescent with short, stiff hairs or glabrescent on the tube,
teeth subequal, short-deltoid, densely to finely hispid-scabrid; co-
rolla blue, violet to purple, tube slender, almost twice the length of
the calyx, externally softly pubescent, lobes obovate, emarginate or
entire; capsule ovoid, compressed, slightly longer than the mature
calyx, style base partly persistent.
San Martin: low mountain pastures near Tarapoto, Spruce 4862;
Sin. Loc. : Jussieu, s.n.
This species is found in low-lying savannas and riversides to
medium altitude grassy hillsides from Panama and Colombia, east
through Venezuela to British Guiana, south through Brazil to Bolivia
and westward to Peru.
Buchnera weberbaueri Diels in Engler, Bot. Jahrb. 37: 430.
1906. B. leiantha Standl., in Field Mus. Publ. Bot. 22: 105. 1940.
Annual herb (15-) 25-50 (-78) cm. tall. Stem slender, simple or
very rarely branched, glabrous to subglabrous above with scant, soft
pubescence near and just above the nodes, sparsely pubescent at
base; leaves opposite to subopposite, erect, linear, 0.5-2.0 mm. wide,
entire, trinerved, a few adpressed hairs on revolute margins at base,
upper leaves becoming ovate-lanceolate with hispid margins similar
to bracts; inflorescence spicate, elongate, 1.0-3.5 (-7.0) cm. long,
laxly flowered; bracts broadly ovate, acuminate, about 3.0 mm. long
and 1.5 mm. wide at base, ciliate; bracteoles narrowly ovate-lanceo-
late, ciliate, little shorter than bracts; calyx about 5.0-7.0 mm. long,
glabrous, finely reticulate between the nerves, teeth triangular-
acuminate, subequal; corolla tube slender, up to about 5.0-8.5 mm.
long, longer than the calyx, externally glabrous, lobes 2.5 mm. long,
broadly ovate; capsule cylindric, slightly compressed and shorter
than the calyx with a few hairs at the suture, style-base deciduous.
Cuzco: Prov. Urubamba, grass slope, petals blue, alt. 2,900 m.,
Vargas 2702.- — Junin: Petals pink-lilac, appendages fringed, alt.
about 9,000 ft., Macbride & Featherstone 1408; Congos, Vidal-Senege
4825.— Loreto: Moyobamba, Mt. Morro, alt. 900-1,100 m., Weber-
bauer 4595 (photo of type). — San Martin: high mountain pastures
near Tarapoto, Spruce 4397 p.p.
Lowlands to high mountain passes from British Honduras and
Panama to Colombia and Peru and east to Venezuela and Trinidad.
FLORA OF PERU 515
Buchnera ternifolia HBK., collected north, east, and south of Peru
could well be encountered in lowland areas (up to 1,600 m.) within
its borders.
CALCEOLARIA L.
Calceolaria L., Kongl. Vetensk. Akad. Handl. 31:288.1770,
non (Heister) Fabricius, Enum. Meth. PL Hort. Helmstad. ed 2, 37.
1763 (Orchidaceae) nor Loefl., Reise Span. Land. Europa u. Amer.
(Violaceae) -1 Fagelia Schwenke, Verh. Bataafsch. Genootsch. Rot-
terdam 1: 474. 1774. Jovellana R. & P., Fl. Peruv. 1: 12, 1. 18. 1798.
Annual small herbs, perennial herbs, vines, lianas or shrubs, pros-
trate, repent, twining or erect, stems usually terete, occasionally
squarrose, simple to much branched, vines occasionally more than
2 m. long, shrubs up to about 1.7 m. tall, plants glabrous to pubes-
cent or glandular-pubescent, foliage and calyces sometimes punctate,
internodes longer or shorter than the leaves. Leaves opposite or
sometimes ternately verticellate, rarely fasciculate, very rarely alter-
nate-fasciculate, petiolate or sessile, linear or aciculate to ovate, obo-
vate or hemispherical, entire, toothed or lobed to pinnate or twice
pinnate, minute to about 20 cm. long, usually 1-8 cm. long, thinly
chartaceous to coriaceous, costa usually impressed above, elevated
beneath, sometimes leaves bizarre, the blade broadly winging the
petiole and the opposite leaves connate-perf oliate ; petiole sometimes
thickened, woody at base and thinly connate; flowers solitary, axil-
lary or forming cymes, dichasia or other regular or irregular inflores-
cences, these usually terminal, less often lateral; calyx of four sepa^,
fused only at or near the base, the lobes usually a little unequal ; co-
rolla most often yellow, occasionally yellow and white or white, rarely
purple, frequently blotched, banded or marked, very strongly modi-
fied and the structure difficult to verbalize, upper lip annulate or
hooded, 1-12(-14) mm. long, when hooded the genitalia often in-
cluded, composed of 2 wholly fused lobes, lower lip of 3 fused lobes,
6-50 mm. long, calceolate, saccate, the sac almost lacking (C. aperta)
to sometimes almost closing the throat, the upper part of the sac
proximally either underfolded and forming a glandular nectary (sub-
genus Cheiloncos) or forming a hollow external cavity and glandular
or not (subgenus Calceolaria) ; stamens 2 in number and of 2 distinct
1 Due to nomenclatural technicalities Calceolaria L. takes precedence and need
not be conserved over Calceolaria (Heister) Fabr. nor Calceolaria Loefl., data to
be published by Dr. W. Stern (Brit. Mus.) fide Mr. Hunt (Kew. Herb.) in personal
correspondence, 1966.
PLATE I. Anther types, Calceolaria: A, herzogiana Kranzl.; B, lobata Cav.;
C, phaceliaefolia Edwin; D, simulans Edwin; E, engleriana Kranzl.; F, utricu-
larioides Benth.; G, reichlinii Edwin; H, pavonii Benth.; I, hyssopifolia HBK.
516
PLATE II. Anther types, Calceolaria: A, endopogon Kranzl.; B, cypripedtiflora
Kranzl.; C, scabra R. & P.; D, brachyantha Edwin; E, solanifolia Edwin; F, tri-
partita R. & P.; G, salicifolia R. & P.; H, triloba Edwin; I, weberbaueriana Kranzl.;
J, annua Edwin; K, delicatula Kranzl.
517
518 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
kinds, either anther cells contiguous, both fertile (subgenus Cheilon-
cos) or cells separated on a connective and both fertile or one cell
(highly) modified, fertile or sterile or wholly lacking (subgenus Cal-
ceolaria) ; style short, often recurved, stigma lobes united all or part
way, ovary ellipsoid to ovate; fruit a septicidal capsule, secondarily
barely loculicidal apically, placenta rupturing, shedding the numer-
ous, minute, loosely tubercular- testiculate seed.
The largest genus in terms of number of species of the Schrophu-
lariaceae in the new world, with 550-600 species. Predominantly of
the Andes mountains of South America but representatives found
through Central America, as far north as Mexico. There are three
species disjunct in New Zealand.
Taxonomically difficult, due to both the number of taxa and the
lack of reliable characters below the level of subgenus. The arrange-
ments of Bentham (1846) and Kranzlin (1907) based on habit and
inflorescence types have become untenable since a number of taxa
include individuals that are both herbs and vines, shrubs and lianas,
erect and reclining, etc. Inflorescence types, while useful, appear to
have their best application at the subsectional level. Other means of
arranging the species in the genus must be found. Perhaps some
of the more useful characteristics in this regard in the subgenus Chei-
loncos are the anther cells (equal or unequal and length in relation
to width), the sac of the corolla (more or less than half the length of
the lower lip), and the size relationships between the corolla lips.
The subgenus Calceolaria presents a quite distinct set of problems.
My knowledge of this subgenus is far too limited to offer useful
suggestions.
KEY TO THE SUBGENERA AND SPECIES
Anther cells contiguous, sessile on the filament, connective lacking,
both cells fertile, very rarely one cell rudimentary, cells equal or
unequal in size and shape; sac of the lower corolla lip with the
upper, proximal end recurved into a glandular, tongue-like nec-
tary, leaves entire to grossly toothed, rarely small-lobate.
Subgenus Cheiloncos (p. 519).
Anther cells separated by the usually well-developed connective,
when connective reduced anther cells often thinly blackened on
lower edge, usually (very) unequal in size, shape and position, the
upper fertile, unmodified and larger than the usually modified,
FLORA OF PERU 519
fertile or sterile lower cell; sac of the lower corolla lip with the
proximal end usually hollowed into an external cavity, glandular
nectary present or lacking; leaves usually deeply lobed, pinnate or
pinnatifid, occasionally merely coarsely toothed, rarely subentire.
Subgenus Calceolaria (p. 552).
KEY TO THE SPECIES OF THE SUBGENUS Cheiloncos
Filaments at least 3 times longer than the anthers, thin or very
little thickened ; erect or repent, usually annual herbs, rarely ex-
ceeding 20 cm. tall, very rarely a small, caespitose shrublet.
2. Lower lip of the corolla 4-5 times longer than the upper lip;
filaments little thickened, usually 1-nerved ; anther locules con-
fluent or distinct.
3. Plants caespitose or subcaespitose; leaves dentate; shrublet.
C. plectranthifolia.
3. Plants with soft, branched, leafy stems; leaves entire or al-
most so; herb C. dichotoma.
2. Lower lip of the corolla not more than twice as long as the
upper lip ; filaments very elongate and thin or shorter and little
thickened, 1- or 2-nerved ; anthers all distinct.
4. Leaf margins entire to very shallowly denticulate; filaments
thin, 4 to 10 or more times longer than the anthers; flowers
usually pedicellate and solitary in axils of the upper leaves,
less often in simple dichasia, rarely in once- or twice-branched
dichasia C. utricularioides.
4. Leaf margins dentate, filaments shorter and thicker, occa-
sionally reaching 5 times longer than the anthers; flowers
pedunculate, in compound, branched dichasia, in upper leaf-
axils or terminal.
4a. Leaves mostly 8 cm. long or longer C. fiebrigiana.
4a. Leaves up to 4.1 cm. long C. lopezii.
Filaments thickened, usually shorter than the anthers, when
longer, less than 3 times as long; usually perennial herbs or shrubs
more than 20 cm. tall, sometimes scapose or acaulescent, or vines
or lianas.
5. One cell of the anther most often very reduced or rudimentary
or lacking, occasionally the 2 cells developed, when both cells
apparent the larger at least twice the size of the smaller.
C. phaceliaefolia.
520 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
5. Both cells of the anther fully developed and functional, cells
equal or unequal in size, when unequal the larger not twice
the smaller.
6. At least the upper leaves dissected to the mid-rib, small
herbs, usually not more than 30 cm. tall C. simulans.
6. Leaves entire, toothed, cut or lobed, but not cut to the mid-
rib, when deeply cut or lobed, woody and more than 30 cm.
tall.
7. Lower lip of the corolla not twice as long as the upper lip,
lips sometimes subequal, rarely upper longer than the
lower (see p. 523).
8. Plants scapose or acaulescent C. scapiflora.
8. Plants with well-developed, leafy stems.
9. Leaf blades or the winged or expanded petioles con-
nate-perfoliate.
10. Leaf blades sessile and widest at the stem.
C. connatifolia.
10. Leaf blades petiolate, widest at the apex of the
petioles; petioles (sometimes the uppermost ex-
cepted) expanded or winged.
11. Lips of the corolla subequal in length and width,
each ca. 7-8 mm. long C. sonchensis.
11. Lower lip of the corolla almost twice as long as
the upper lip, both lips often 12 mm. long or
longer C. calycina.
9. Neither leaves nor petioles connate-perfoliate; peti-
oles only rarely broadly winged or expanded; blades
sessile or petiolate.
12. At least most of the leaf blades less than 2 cm.
long and 0.5 cm. wide, usually lanate to densely
pubescent, margins frequently obviously to ob-
scurely 3-lobed.
13. Anther cells equal in size.
14. Some leaf blades sessile, margins essentially
entire or obscurely 3-lobed.
15. Leaves globose, trilobate C. inaudita.
15. Leaves linear, aciculiform or filiform.
C. callunoides.
FLORA OF PERU 521
14. Leaf blades petiolate, margins clearly 3-lobed.
C. triloba.
13. Anther cells unequal in size C. pallascensis.
12. At least most of the leaf blades more than 2 cm.
long or 0.5 cm. wide or both, never 3-lobed, usually
glabrous to puberulent, rarely lanate.
16. Inner surface of the calyx lobes with a distinct
row of hairs near the margin, other pubescence
present or lacking.
17. Blades sessile or on petioles not more than
2 mm. long.
18. Blades linear, linear-lanceolate or linearly-
ligulate C. hyssopifolia.
18. Blades ovate or oblong.
19. Blades sparsely puberulent above, essen-
tially glabrous beneath . . . . C. pearceana.
19. Blades canescent on both surfaces.
C. pulverulenta.
17. Blades on petioles more than 2 mm. long.
20. Anther cells subhemispherical, or broadly
ovoid, almost sessile, usually bigger than
the filaments.
21. Leaf blades up to 8 mm. wide.
C. hyssopifolia.
21. Leaf blades most often more than 8 mm.
wide, up to 3 cm. in width.
22. Corolla lips subequal C. padifolia.
22. Upper lip only little more than half as
long as the lower C. nivalis.
20. Anther cells oblong, elliptic-lanceolate, or
ovoid, clearly stalked, usually smaller than
the filament C. cerasifolia.
16. Inner surface of the calyx lobes without a defi-
nite row of hairs near the margin, glabrous to
densely pubescent or glandular.
23. Leaf blades sessile.
24. Leaves ovate or oblong. . . .C. pulverulenta.
522 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
24. Leaves linear, linear-lanceolate or linear-
ligulate.
25. Leaves glabrous C. hyssopifolia.
25. Leaves lanate beneath.
C. rosmariniifolia.
23. Leaf blades petiolate.
26. Leaves linear, lance-linear or ligulate,
mostly less than 9 mm. wide.
27. Leaf blades lanate beneath.
C. rosmariniifolia.
27. Leaf blades glabrous . . . . C. hyssopifolia.
26. Leaf blades ovate, lance-ovate, oblong ellip-
tical or rarely obovate, rarely less than 1.3
cm. wide.
28. Anther cells unequal, 2-3 times longer
than wide, large for the flower.
29. Leaf blade margins irregularly and
sharply appressed-serrulate; anther
cells a little unequal C. involuta.
29. Leaf blade margins usually entire, oc-
casionally; regularly, minutely serru-
late or crenulate; anther cells very
unequal C. tetragona.
28. Anther cells equal.
30. Anther cells at least twice as long as
wide, sometimes with an acumin at the
apex of the filament.
31. Stems and foliage often glabrous;
leaf blade margins sharp-toothed.
C. involuta.
31. At least the younger stems and foli-
age lanate; teeth of the leaf blade
margins rounded. C. croceopunctata.
30. Anther cells less than twice as long as
wide, occasionally wider than long,
acumin lacking at the apex of the fila-
ment.
FLORA OF PERU 523
32. Leaf blades usually 3 or more times
longer than wide.
C. deflexa var. deflexa.
32. Leaf blades up to twice as long as
wide.
33. Calyx and the veins on the lower
surfaces of the leaves with uni-
cellular hairs.
C. deflexa var. aurantiaca.
33. Calyx and the veins on the lower
surfaces of the leaves with multi-
cellular hairs C. oblonga.
7. Lower lip of the corolla at least twice as long as the upper
lip.
34. Anther cells unequal in size or shape or both (see
p. 529).
35. At least most of the leaf blades not exceeding 8 mm.
wide, linear or almost so to sometimes narrowly
ovate.
36. Corolla purple, rarely yellow with prominent pur-
ple markings; anther cells little longer than wide.
C. weberbaueriana.
36. Corolla yellow or yellow and white; anther cells
much longer than wide, frequently at least twice
as long as wide.
37. Leaf blades up to 6 cm. long or longer, mostly
more than 1.8 cm. long C. procera.
37. Leaf blades mostly not exceeding 2.5 cm. long,
often less than 1.8 cm. long.
38. Teeth of the leaf blade margin many and ap-
parent, blades narrowly ovate, rarely lanceo-
late or lance-linear, very rarely aciculiform,
pubescence long or short, sometimes mixed
with a few glandular hairs, on both surfaces,
rarely pubescence sparse, tissue usually visible.
C. scabra.
38. Teeth of the leaf blade margin usually few
and obscure, when apparent or many the
linear, lance-linear, aculiform or rarely nar-
524 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
rowly ovate blades with short, erect pubes-
cence, tissue sometimes hidden.
39. Calyces and lower surfaces of the leaf blades
densely covered with long, white, soft hairs,
at least partly obscuring the tissue; anther
cells very unequal C. reichlinii.
39. Calyces and lower surfaces of the leaf blades
variously pubescent or glandular-pubescent
to occasionally glabrous, vesture not of
long, soft white hairs; anther cells almost
equal to less often very unequal.
40. Anther cells sagittate, black or dark
brown C. revoluta.
40. Anther cells divergent or divaricate, yel-
low or light brown.
41. Corolla pilose, lower lip with the sac
half or little more than half its length ;
calyx lobes up to 7 mm. long, fre-
quently exceeding 5 mm. long, usu-
ally 5 mm. wide at base.
C. cajabambae.
41. Corolla either glabrous or pubescent,
vesture when present papillose (lower
lip) or pilose on upper lip only, lower
lip saccate either almost its entire
length or less than half its length;
calyx lobes not exceeding 5 mm. long
and 4 mm. wide, often smaller.
42. Anthers about 2-3 mm. long; calyx
lobes lance-ovate or narrowly ovate.
C. brachyantha.
42. Anthers about 4-5 mm. long; calyx
lobes ovate to broadly ovate.
43. Lateral veins clearly visible on
both surfaces of the leaf blade,
impressed above, elevated be-
neath, blades often exceeding 4
mm. wide; inflorescences racemose.
C. linearioides.
FLORA OF PERU 525
43. Lateral veins of the leaf blade
wanting or only barely visible
above, blades rarely up to 4 mm.
wide; inflorescences dichasiate.
C. Unearis.
35. Leaf blades mostly more than 8 mm. wide, wider
than linear.
44. Leaf blades perfoliate or petioles broadly winged
and connate-perfoliate.
45. The larger cell and often the smaller of each
anther usually less than twice as long as wide.
46. Ovary covered with long, tangled hair; petiole
margins entire or with rounded teeth or lobes.
C. calycina.
46. Ovary glabrous or sparsely glandular-puberu-
lent, hair descrete; petiole margins with small,
sharp teeth C. herzogiana.
45. The larger cell and often the smaller of each
anther more than twice as long as wide.
47. Cauline leaf blades, uppermost excepted, most
often wider than long, sometimes as long as
wide C. aequilateralis.
47. Cauline leaf blades mostly longer than wide.
48. Leaf blades puberulent with the vesture
primarily on the veins C. pavonii.
48. Leaf blades tomentose or lanate at least
beneath, the long, often matted hair on
both veins and tissue C. tomentosa.
44. Leaf blades sessile or petiolate, blades and petioles
neither connate nor perfoliate.
49. Anther cells sagittate and subparallel, little di-
vergent.
50. Style 2-3 mm. long, shorter than to occasion-
ally as long as the calyx; anther cells at least
twice as long as wide; corolla yellow.
51. Leaf blades cordate at base C. lobata.
51. Leaf blades broadly obtuse at base.
C. phaceliaefolia.
526 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
50. Style 4-6 mm. long, little shorter to longer
than the calyx; anther cells less than twice as
long as wide; corolla white or yellow.
52. Leaf blades pubescent beneath epunc-
tate C. baliotifolia.
52. Leaf blades glabrous beneath, densely
red-punctate C. leucantha.
49. Anther cells widely divergent or divaricate,
sometimes slightly downturned distally.
53. Anther cells less than twice as long as wide.
54. Most of the leaf blades not exceeding 2.5
cm. long, usually less than 1.5 cm. long,
bullate or plane; stem pubescence erect.
C. scabra.
54. Leaf blades 2.5-6.0 cm. long, usually more
than 3 cm. long, plane; stem puberulence
flattened or erect.
55. Apex of leaf blade acuminate, margins
regularly, shallowly toothed.
C. variefolia.
55. Apex of leaf blade obtuse; margins sub-
regularly more deeply toothed.
C. pseudoscabra.
53. Anther cells, occasionally only one of the pair,
at least twice as long as wide.
56. Foliage glandular-pubescent on tissue.
57. Leaf blades from 4-10 cm. long, usually
6 cm. or longer C. dentifolia.
57. Leaf blades up to 3.5 cm. long, mostly
less than 3 cm. long.
58. Leaf blades 2.0-3.5 cm. long.
C. pseudoscabra.
58. Leaf blades up to 2.0 cm. long, usually
not exceeding 1.5 cm. long. .C. scabra.
56. Foliage glabrous to pubescent, occasionally
only the blade margin glandular.
59. Foliage glabrous or occasionally very
sparsely pubescent on tissue and costa
above.
FLORA OF PERU 527
60. Peduncles and pedicels pubescent or
papillose; calyx glabrous or pubescent.
61. Inflorescences usually 20- or more
flowered, rarely less than 10; sec-
ondary and tertiary veins clearly
visible beneath on the subcoriaceous,
viscid leaf blade. . . .C. arborescens.
61. Inflorescences usually not more than
6-8 flowered.
62. Stem and calyx lobes externally
glabrous C. myrtiiloides.
62. Stem and calyx lobes externally
puberulent or pubescent.
C. viscosa.
60. Entire plant glabrous, occasionally ca-
lyces glaucous or pustulate.
63. Leaf-blades ovate, broadly ovate
and less often elliptically ovate,
rarely less than 2 cm. wide, up to
about 4 cm., margins sinuate or dis-
tantly toothed, teeth broadly serru-
late; ovary pubescent and glandu-
lar-pubescent; calyx lobes usually
acute C. endopogon.
63. Leaf -blades lanceolate, lance-elliptic
or narrowly lance-ovate, rarely ex-
ceeding 2 cm. wide, margins closely
and sharply serrate or doubly ser-
rate; ovary usually glabrous; calyx
lobes usually abruptly acuminate.
C. salicifolia.
59. Foliage pubescent.
64. Foliage and calyx densely lanate, pu-
bescence masking the tissue; leaf blade
margins entire or almost so.
C. reichlinii.
64. Foliage and calyx variously pubescent,
often with short curly hair, tissue vis-
ible or hidden by the pubescence; blade
528 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
margins variously toothed, less often
entire.
65. All or most of the leaf blades at least
twice as long as wide (occasionally
1.7-1.9 times longer than wide) usu-
ally acute to obtuse at base.
66. Petioles gibbous and sometimes
woody at base and laterally con-
nate part way around the stem.
67. Leaf blades entire to very finely,
minutely toothed.
C. engleriana.
67. Leaf blades coarsely toothed.
68. Leaf blades mostly ovate,
margins regularly or subreg-
ularly crenate-dentate.
C. bicrenata.
68. Leaf blades mostly lanceolate
to narrowly lance-ovate, mar-
gins irregularly serrate.
C. longiinternodia.
66. Petioles neither gibbose nor woody
at base and not decurrent.
C. scabra.
65. All or most of the leaf blades not
more than 1.6 times longer than
wide, sometimes almost as wide as
long, often obcordate at base.
69. Most of the mature leaf
blades less than 3.5 cm.
long, margins crenate-den-
tate or crenate-serrate.
C. scabra.
69. Most of the mature leaf
blades more than 3.5 cm.
long, up to 6.0 cm. long or
longer.
70. Leaf blade margins
grossly and irregularly
toothed or lobed.
C. rhacodes.
FLORA OF PERU 529
70. Leaf blade margins
subentire to finely ser-
rate or serrulate.
C. solanifolia.
34. Anther cells equal in size and shape, rarely very slightly
unequal in shape only.
71. All or most of the leaves more than 1 cm. wide (see
p. 542).
72. All or most of the leaf blades less than 2 cm. long,
occasionally blades 1.5-2.8 cm. long, with most
2.0-2.8 cm. long (see p. 532).
73. Petioles mostly 1-3 cm. long or longer, mostly
more than half the length of the blade, some-
times longer than the blade C. lobata.
73. Petioles mostly less than 1 cm. long, only rarely
(C. hedera) up to half the length of the blade.
74. Leaf blade densely lanate, tomentose or pul-
verulent beneath on both veins and tissue.
75. Leaf blade lanate beneath, of various shapes
but not obovate, margins deeply toothed or
lobed.
76. At least most of the leaf blades twice as
long as wide or longer C. aperta.
76. At least most of the blades less than
twice as long as wide.
77. Blade margins crenate-dentate; peti-
oles ca. 1-2 mm. long or lacking.
C. cordiformis.
77. Blade margins lobed, mostly with two
lateral pairs and one terminal lobe; pet-
ioles often 5-7 mm. long.
C. hedera.
75. Leaf blades tomentose or pulverulent be-
neath, obovate, entire to deeply toothed.
78. At least some of the leaf blades wider
than long C. hutchisonii.
78. Leaf blades longer than wide.
530 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
79. Blades pulverulent beneath, margins
entire or crenulate-serrulate. C. inflexa.
79. Blades tomentose beneath, margins
regularly or irregularly, grossly or nar-
rowly dentate, crenate or crenate-ser-
rate.
80. Leaves ovate or obovate, margins
grossly, irregularly, distantly den-
tate C. sparsiflora.
80. Leaves elliptical or less often ellip-
tically ovate or blades lance-elliptic,
margins crenate or crenate-serrate.
C. atahualpae.
74. Leaves glabrous, pubescent or glandular-pu-
bescent, vesture when present on venation or
on venation and tissue, but tissue neither
densely lanate, tomentose nor pulverulent
beneath.
81. Only the venation densely lanate, hair
curled and matted.
82. Style ca. 6 mm. long ; anther cells apically
acuminate C. croceopunctata.
82. Style 2-3 mm. long; anther cells without
an apical acumin C. velutinoides.
81. Venation glabrous or pubescent with short,
straight, descrete hair.
83. Corolla mostly (20-) 25-37 mm. long and
about half as wide; style 6-8 mm. long.
C. boliviano,.
83. Corolla mostly less than 20 mm. long,
when longer almost as wide as or wider
than long, style 1.5-5.0 (-6.0) mm. long.
84. Style 4-6 mm. long C. pearceana.
84. Style 1.5-3.0 mm. long, rarely to 3.5
mm. long.
85. Anthers 3.2-4.2 mm. long, more than
twice as long as wide.
FLORA OF PERU 531
86. Leaves usually elliptic; corolla
about 25 mm. to occasionally 30
(-40) mm. long. . . .C. atahualpae.
86. Leaves mostly ovate; corolla usu-
ally not more than 20 mm. long.
C. pisacomensis var. arequipensis.
85. Anthers not more than 3 mm. long,
mostly 1.5-2.8 mm., sometimes less
than twice as long as wide.
87. Leaf blades often entire, teeth
when present distant and shal-
low C. ramosissima.
87. Leaf blade margins distinctly
closely toothed or blades few-
lobed.
88. Leaf blades with few-lobed
margins, lobes toothed.
C. hedera.
88. Leaf blade margins toothed,
lobes not developed.
89. Ovary glabrous to glandu-
lar-pubescent.
90. Blade margins dentate or
doubly so, leaves often
ternate . . . C. verticillata.
90. Blade margins serrate,
serrulate or serrate-ser-
rulate, leaves usually op-
posite.
91. Calyx lobes 4-6 mm.
long C. virgata.
91. Calyx lobes 2.0-3.5
mm. long.
92. Leaves ovate or
broadly ovate.
C. ramosissima.
92. Leaves lanceolate to
narrowly lance-ovate
C. angustiflora.
532 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
89. Ovary densely pubescent
with short, stiff, white hair.
C. sibthorpioides.
72. At least most of the leaf blades more than 2 cm.
long.
93. Leaves mostly or entirely radical or basal;
plants scapose, or subscapose with stem leaves
few and very reduced, occasionally caulescent
with few branches C. corymbosa.
93. Leaves disposed over leafy, well-developed, usu-
ally branching stems.
94. All or only the upper leaves perfoliate or con-
nate.
95. Only the upper leaves perfoliate or connate,
the others petiolate or sessile.
C. pisacomensis.
95. All the leaves perfoliate or connate.
C. aequilaterahs.
94. Leaves not perfoliate nor connate, petiolate or
sessile; petioles sometimes only thinly, partly
surrounding the stem.
96. Leaf blades up to twice as long as wide (see
p. 536).
97. Corolla 25 mm. long or longer, up to
about 45 mm.
98. Bases of the petioles of the lower
(longer) leaves only very little thick-
ened and not at all woody and some-
times not thinly surrounding the stems
or branches.
99. Anther cells at least 1.5 times longer
than wide.
100. Leaf blade margins crenate-den-
tate, lobes lacking.
C. cypripediiflora.
100. Leaf blade margins 5-7-lobed.
C. rhacodes.
99. Anther cells less than 1.5 times
longer than wide C. boliviano,.
FLORA OF PERU 533
98. Bases of the petioles of the lower leaves
thickened, woody and usually sur-
rounding the stems and branches.
101. Calyx lobes up to 10 mm. long, in
life and when dry, light green to
dark brown-green.
102. Leaf blade margins coarsely
toothed C. bicrenata.
102. Leaf blade margins entire or fine-
toothed C. inflexa.
101. Calyx lobes 11-15 mm. long, in life
and when dry, bright yellow some-
times also red distally.
C. chrysocalyx.
97. Corolla up to 25 mm. long.
103. Leaves pubescent beneath with multi-
septate hair.
104. At least most of the petioles more
than 4 mm. long, up to 2 cm. long
or longer C. heterophylla.
104. All or most of the petioles less than
4 mm. long C. oblonga.
103. Leaves variously pubescent, hair non-
septate (occasionally uni- or bi-sep-
tate) or glabrous beneath.
105. Leaf blades densely lanate, canes-
cent or tomentose beneath.
106. Calyx lobes bright yellow or pale
yellow-green, mostly 8 mm. long
or longer.
107. Calyx lobes 11-15 mm. long.
C. chrysocalyx.
107. Calyx lobes up to 10 mm. long.
108. Calyx lobes bright yellow;
anther cells less than twice
as long as wide.
C. luteocalyx.
534 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
108. Calyx lobes pale green-yel-
low; anther cells twice as
long as wide. .C. bicrenata.
106. Calyx lobes dark brownish-green
to green, usually not more than
7 mm. long.
109. Leaves ferrugineous-tomen-
tose beneath C. maculata.
109. Leaves otherwise pubescent.
110. Corolla 10-20 mm. long.
111. Leaf blade margins ser-
rate, unlobed ; corolla (8-)
10-14 mm. long.
C. serrata.
111. Leaf blade margins 5-7-
lobed, lobes dentate; co-
rolla 16-18 (-20) mm.
long C. rhacodes.
110. Corolla 20-35 mm. long.
C. bicrenata.
105. Leaf blades glabrous to variously
pubescent beneath but neither la-
nate, canescent nor tomentose.
112. Leaf blade margins few (5-9)-
lobed, lobes large.
113. Petioles (1-) 2-5 cm. long or
longer, frequently at least half
as long as the blade, vesture
often of red, stalked glands;
leaf blades densely pilose on
both surfaces C. lobata.
113. Petioles up to 2 cm. long, usu-
ally shorter, not reaching half
the length of the blade; only
the calyx and flower stalks
glandular-pubescent; blades
when densely pubescent only
beneath C. rhacodes.
112. Leaf blade margins variously
toothed, occasionally subentire.
FLORA OF PERU 535
114. Calyx lobes yellow or pale
green-yellow; usually 8 mm.
long or longer.
115. Calyx lobes 11-15 mm.
long C. chrysocalyx.
115. Calyx lobes mostly (5-)
8-10 mm. long.
C. bicrenata.
114. Calyx lobes dark brown-
green to green, usually not
more than 8 mm. long.
116. Leaf blade margins mostly
subentire or (sometimes)
inconspicuosly toothed,
rarely (the largest) with
obvious teeth.
117. Leaves usually more
than 2 cm. wide.
C. variefolia.
117. Leaves usually less than
than 2 cm. wide.
C. inflexa.
116. Leaf blade margins con-
spicuously toothed.
(See choice 118.)
118. Calyx lobes glabrous or almost so, margins frequently ciliolate
or fimbriate.
119. Lower lip of the corolla 17-25 (-28) mm. long; calyx lobes
5-9 mm. long, ciliolate.
120. Lower lip of the corolla 20-28 mm. long; calyx lobes 5-
7 mm. long, ovate, acute C. leptantha.
120. Lower lip of the corolla 17-18 mm. long; calyx lobes 8-9
mm. long, lanceolate, attenuate-acuminate. . .C. scandens.
119. Lower lip of the corolla 9-15 mm. long; calyx lobes 3-4 mm.
long, sparsely setose, margins sometimes fimbriate.
121. Leaf blade teeth sharply dentate to doubly dentate.
C. verticillata.
121. Leaf blade teeth rounded, crenate C. leptantha.
536 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
118. Calyx lobes variously pubescent.
122. Calyx lobes of the mature flower less than 4 mm. long.
123. Leaf blades very short-puberulent on the upper surface;
anthers less than 3 mm. long.
124. Style up to 2 mm. long; leaf blade teeth sharp.
C. ramosissima.
124 . Style 3-4 mm. long; leaf blade teeth usually rounded.
C. corymbosa var. perennis.
123. Leaf blade glabrous on the upper surface; anthers more
than 3 mm. long C. divaricata.
122. Calyx lobes of the mature flowers 4-7 mm. long or longer.
125. Corolla 1.5 cm. long or longer, rarely 1.5-3.5 cm. long co-
rollas on one plant.
126. Capsule ovoid or lance-ovoid; leaf blade teeth mostly
large and extrorse C. bicrenata.
126. Capsule oblong, conic-ovoid or elliptic; leaf blade teeth
small and antrorse.
127. Corolla 20-35 mm. long, anther cells only little longer
than wide C. boliviana.
127. Corolla mostly 15-20 mm. long; anther cells about
twice as long as wide C. virgata.
125. Corolla only rarely exceeding 1.5 cm. long, usually 1.0-
1.5 cm. long; anthers 1.3-1.8 times longer than wide.
128. Leaf blades rugulose or rugose; corolla up to 1.0 cm.
long C. rugulosa.
128. Leaf blades plane; corollas 1.2-1.5 cm. long.
129. Calyx lobes 4-5 mm. long. . C. corymbosa var. perennis.
129. Calyx lobes 5-7 mm. long, occasionally longer.
C. virgata.
96. Leaf blades 2 or more times longer than wide
(see choice number 130) .
130. Leaf blades densely lanate, canescent, tomentose or pulveru-
lent beneath.
131. Blade pulverulent beneath, marginal teeth lacking.
C. inflexa.
131. Blades lanate or canescent beneath, marginal teeth present
or absent.
FLORA OF PERU 537
132. Anther cells up to 1.3 times longer than wide, usually less,
and varying to wider than long.
133. Style 7-9 mm. long; stigma much larger in diameter
than the style C. aperta.
133. Style 2-4 mm. long; stigma only little larger in diameter
than the style.
134. Upper lip of the corolla ca. 1 cm. long, lower lip ca.
2.5-3.5 cm. long C. atahualpae.
134. Upper lip of the corolla 2-3 mm. long, lower lip ca.
10-12 mm. long C. lasiocalyx.
132. Anther cells at least 1.4 times longer than wide, most often
1.7 times or more longer than wide.
135. Leaf blade lanceolate or narrowly lance-ovate, usually
more than 3 times longer than wide.
136. Blade margins subentire to very finely serrulate ; stems
hirsute C. hirsutula.
136. Blade margins very coarsely dentate; stems glabrous
or almost so C. longiinternodia.
135. Leaf blade ovate to broadly lance-ovate, most often less
than three times longer than wide.
137. Corollas 35-45 mm. long; leaf blades ferruginious-
pubescent beneath C. cypripediiflora.
137. Corolla often not more than 25 mm. long, rarely
reaching 35 mm. ; leaf blades white- or yellow-pubes-
cent beneath.
138. Corolla 6-12 mm. long; anther ceils distally plane.
139. Calyx lobes 5-7 mm. long; corolla sac much
shorter than the open, proximal part of the lip.
140. Style 1.0-1.5 mm. long; anther cells divaricate.
C. rugulosa.
140. Style ca. 6 mm. long; anther cells sagittate,
apex acuminate C. croceopunctata.
139. Calyx lobes 4 mm. long; corolla sac almost equal
to the proximal, non-saccate part of the lip in
length C. serrata.
138. Corolla ca. 13-25(-35) mm. long; anther cells often
distally downturned C. bicrenata.
538 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
130. Leaf blades varying from glabrous to more or less densely
pubescent or glandular-pubescent beneath.
141. Lower lip of the corolla 25 mm. long or longer, the upper lip
often well developed and longer than the calyx-lobes, some-
times only equaling or little shorter than the calyx lobes
(C. bicrenata).
142. Anther cells only little longer than wide to little wider than
long, frequently drying black or dark brown, occasionally
white to tan, parallel to divergent; upper lip of the corolla
longer than the calyx lobes (very rarely shorter) .
143. Blades regularly serrulate, occasionally crenulate or
denticulate to entire.
144. Style 6-8 mm. long; corolla externally puberulent.
C. boliviano,.
144. Style 2-4 mm. long; corolla externally glabrous.
145. Calyx lobes 3-4 mm. long, internally glabrous; co-
rolla orange C. deflexa.
145. Calyx lobes ca. 6 mm. long, internally with a row
of marginal yellow hair; corolla yellow . . . C. nivalis.
143. Leaf blades irregularly crenate-dentate, densely pubes-
cent beneath or on both sides; anthers dark brown to
black when dry C. atahualpae.
142. Anther cells from 1.5 to 3 times or more longer than wide,
drying light brown to white, divergent or divaricate; upper
lip of the corolla usually equal to or shorter than the calyx
lobes.
146. Corolla (25-) 35-45 mm. long, externally pilose; petioles
doubly pubescent (hair of 2 distinct sizes).
C. cypripediiflora.
146. Corolla 25-30 mm. long; petioles glabrous to pubescent.
147. Upper leaf blades amplexicaul or sessile.
C. pisacomensis.
147. Upper leaf blades usually short-petiolate or subsessile,
occasionally sessile.
148. At least most of the leaves not more than 2.6 cm.
wide.
149. Plants glabrous; leaves usually less than 3 cm.
long C. myrtittoides.
FLORA OF PERU 539
149. Plants pubescent; leaves up to 6 cm. long.
C. engleriana.
148. At least most of the leaves more than 2.6 cm. wide,
up to 5 cm. wide or wider.
150. Pubescence unicellular, non-septate.
C. bicrenata.
150. Pubescence septate C. vulpina.
141. Lower lip of the corolla less than 25 cm. long, upper lip most
often shorter than the calyx lobes, sometimes only ca. 1 mm.
long (except C. boliviano, and C. crenata).
151. Corolla white or yellow and white C. bicrenata.
151. Corolla most often yellow, occasionally orange.
152. Wood and foliage glabrous or almost so, only the mid-
vein on the upper surface of the leaf blade sparsely
pubescent.
153. Corolla usually orange, occasionally yellow, anthers
less than twice as long as wide C. deflexa.
153. Corolla yellow; anthers at least twice as long as wide,
usually longer C. myrtilloides.
152. Wood or foliage or both pubescent or puberulent.
154. Hair on stems and branches septate.
155. Calyx lobes ca. 10 mm. long; leaf blades coarsely,
irregularly toothed C. vulpina.
155. Calyx lobes ca. 3-5 mm. long; leaf blades with fine,
regular teeth.
156. Corolla ca. 20 mm. long C. crenulata.
156. Corolla ca. 10-12 mm. long C. lasiocalyx.
154. Hair on stems and branches lacking obvious septa.
157. Anther cells at least twice as long as wide, often
distally downturned.
158. Leaf blades lanceolate, lance-elliptic or narrowly
lance-ovate, when narrowly lance-ovate fre-
quently more than 2.5 times longer than wide,
margins sometimes entire or almost so.
159. Leaves mostly entire or subentire, occasionally
shallowly serrulate.
160. Upper lip of the corolla 3-5 (-6) mm. long,
sometimes equaling, rarely exceeding the
length of the calyx lobes C. engleriana.
540 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
160. Upper lip of the corolla ca. 1.5-2.5 mm. long,
much shorter than the calyx lobes.
C. rhizomatosa.
159. Leaves variously serrate or serrulate, occasion-
ally crenate, only rarely the young blades en-
tire or subentire . . C. angustiflora.
158. Leaf blades ovate, broadly elliptic or broadly
lance-ovate, when broadly lance-ovate blades less
than 2.5 times longer than wide, margins deeply
and irregularly cut (except C. viscosa).
161. Leaves glabrous C. viscosa.
161. Leaves pubescent.
162. Leaves and calyx appressed-pubescent.
C. colcamarensis.
162. Pubescence of leaves and calyx otherwise.
C. bicrenata.
157. Anther cells less than twice as long as wide, distally
usually plane.
163. Leaves lanceolate, entire or subentire, occasion-
ally (C. hirsutula) a few leaves serrulate.
164. Calyx lobes 10-12 mm. long . . C. chrysosphaera.
164. Calyx lobes mostly 5-6 mm. long, occasionally
reaching 7 mm C. hirsutula.
163. Leaves usually ovate, broadly elliptical or
lance-ovate, sometimes lanceolate, when lanceo-
late margins usually crenate, dentate or serrate
and frequently the teeth irregular.
165. Leaves lanceolate, irregularly toothed, rarely
teeth regular, or leaves ovate or elliptic with
small or obsolete teeth.
166. Blades lanceolate, clearly irregularly toothed.
167. Lower corolla lip saccate less than half its
length.
168. Upper lip of the corolla longer than the
calyx lobes, ca. 4 mm. long. .C. crenata.
168. Upper lip of the corolla shorter than the
calyx lobes, ca. 1-2 mm. long.
C. angustiflora.
FLORA OF PERU 541
167. Lower corolla lip saccate at least three-
fourths of its length, throat almost closed.
C. glauca.
166. Blades ovate, broadly elliptical or elliptically
ovate, teeth small to almost obsolete.
169. Calyx lobes mostly 5-6 mm. long or longer.
170. Calyx lobes lanceolate; style 6-8 mm.
long C. boliviano,.
170. Calyx lobes mostly ovate; style 2 mm.
long C. variefolia.
169. Calyx lobes mostly 3 mm. long.
C. crenata
165. Leaves ovate, broadly elliptic or lance-ovate,
marginal teeth or lobes obvious, large, regular
or irregular.
171. Calyx lobes lanceolate.
172. Calyx lobes 8-9 mm. long, acuminate.
C. scandens.
172. Calyx lobes 3-4 mm. long, acute.
C. verticiUata.
171. Calyx lobes ovate or elliptic-ovate or occa-
sionally lance-ovate, usually acute.
173. Calyx lobes 7-10 mm. long.
174. Style 3-4 mm. long.
175. Leaf blades sharply serrate or serrate-
serrulate . . . C. virgata var. llamaensis.
175. Leaf blades variously doubly and ir-
regularly dentate or crenate, only
rarely serrate C. bicrenata.
174. Style 1.5-2.0 mm. long.
176. Bases of the petioles of the larger
leaves swollen and woody ; corolla sac
less than half the length of the lower
lip C. rugulosa.
176. Bases of the petioles of the larger
blades little if at all swollen or woody;
corolla sac at least half the length of
the lower lip C. colcamarensis.
542 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
173. Calyx lobes 3-7 mm. long.
177. Calyx glabrous C. verticillata.
177. Calyx pubescent or puberulent.
178. Corolla 1 cm. wide and 1.5 cm. long
or longer C. bicrenata.
178. Corolla up to 7 mm. wide and only
rarely more than 1.3 cm. long, then
up to 1.5 cm. long.
179. Corolla sac at least half the length
of the lip.
180. Inflorescences compound and
well branched, paniculate or pan-
icles of dichasia; lower lip of the
corolla saccate about three-
fourths of its length . . . C. glauca.
180. Inflorescences simple or once-
branched ; lower lip of the corolla
saccate about half its length.
C. lasiocalyx.
179. Corolla sac less than half as long as
the lower lip.
181. Calyx lobes of the mature flowers
less than 4 mm. long.
C. ramosissima.
181. Calyx lobes of the mature flowers
4-7 mm. long.
182. Leaf blades rugulose or rugose;
corolla up to 1 cm. long.
C. rugulosa.
182. Leaf blades plane; corolla 1.2-
1.5 cm. long C. virgata.
71. At least most of the leaf blades less than 1 cm. wide.
(Please return to the margin; entry 183.)
183. At least most of the leaves linear, filiform or aciculiform, less
than 3 mm. wide, margins entire or with 2-3-pairs of teeth.
184. Anther cells sagittate, each ca. 1.5-3.0 mm. long; style 4-
5 mm. long; upper lip of the corolla ca. 3-4 mm. long; leaf
blades tomentose or pulverulent C. revoluta.
184. Anther cells divaricate or divergent, each ca. 0.7-1.5 mm.
long; style 1.5-3.0 mm. long; upper lip up to 2 cm. long;
FLORA OF PERU 543
leaf blades glabrous, puberulent or pulverulent, sometimes
glandular.
185. Leaf blades, stems and branches glabrous or almost so.
C. alba.
185. Leaf blades and the newer wood variously pubescent or
puberulent and often pulverulent, sometimes glandular.
186. At least most of the leaf blades more than 1.5 cm. long.
C. linearis.
186. At least most of the leaf blades less than 1.5 cm. long.
187. Anther cells mostly 2 or more times longer than wide.
188. Leaves pulverulent beneath, mostly about 1 mm.
wide, margins entire, leaf -fascicles and leaves alter-
nate C. alternifolia.
188. Leaves glandular-puberulent, either 2-3 mm. wide
with entire margins, or about 1 or 2 mm. wide with
2-3 pairs of apical teeth, leaf-fascicles and leaves
opposite C. brachyantha.
187. Anther cells about as long as wide to 1.6 times longer
than wide C. callunoides.
183. At least most of the leaf blades 4-10 mm. wide or when up to
3 mm. wide, broader than linear; margins toothed, lobed or
occasionally entire.
189. Leaf blades up to 3 mm. wide, broader than linear, margin
toothed or lobed, occasionally entire.
190. Leaf blade margins distinctly revolute, sometimes rolled
or folded to the midrib, toothed or with 3-9 pairs of lobes,
rarely more than 9 pairs.
191. Calyx ca. 4 mm. long; corolla ca. 8-10 mm. long; pedi-
cels sometimes 1-2 cm. long, usually englandular; south
Peru C. santalinoides.
191. Calyx ca. 7-8 mm. long; corolla ca. 13-17 mm. long;
pedicels usually less than 1 cm. long, glandular; north
Peru C. incarum.
190. Leaf blade margins plane or weakly revolute, toothed or
with 1-2 pairs of lobes or sometimes entire.
192. Walls of anther cells and septa very thick, polleniferous
part of cells only little more than half of the locule.
C. scabra.
544 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
192. Walls of the anther cells and septa thin, polleniferous
area at least three-fourths of the locules.
193. Petioles gibbose, woody at base and surrounding the
node as a thin collar; internodes long, leaves distant.
C. ajugoides.
193. Petioles neither swollen at base nor woody, descrete;
internodes short, leaves crowded.
194. Leaves, stems and inflorescences lanate.
195. Calyx lobes pubescent within . . C. decus montium.
195. Calyx lobes glabrous within C. argentea.
194. Occasionally leaves or stems or inflorescences la-
nate, never the entire plant.
196. Style 4-5 (-6) mm. long C. pearceana.
196. Style 1.5-4.0 mm. long.
197. Leaf blade with 1 pair of broad, divergent lobes
at the base, apex obtuse or rounded, tissue
lanate beneath C. triloba.
197. Leaves not as above, lanceolate to ovate or
cuneiform, glabrous to lanate, entire to vari-
ously lobed.
198. Leaf blade margin entire or few-crenulate.
C. myriophylla.
198. Leaf blade margins variously toothed or
lobed.
199. Leaves triangular or broadly ovate or
ovate, usually less than twice as long as
wide.
200. Leaf blades ovate, minute, up to 5 mm.
long and 4 mm. wide, margin regularly
with 5-7 pairs of small lobes.
C. millefoliata.
200. Leaf blades triangular to ovate, mostly
more than 5 mm. long, regularly or ir-
regularly toothed or lobed.
201. Calyx lobes 3-4 mm. long; corolla up
to 10 mm. long C. inamoena.
201. Calyx lobes 5-8 mm. long; corolla
ca. 15 mm. long C. cuneiformis.
FLORA OF PERU 545
199. Leaf blades lanceolate to lance-ovate, ob-
long-ovate or cuneately obovate, usually
at least twice as long as wide.
202. Lower lip of the corolla saccate less than
half its length C. sparsiflora.
202. Lower lip of the corolla saccate at least
half its length.
203. Leaves lanceolate or lance-subovate.
C. extensa.
203. Leaves oblong ovate.
204. Leaves glabrous above.
C. bartsiaefolia.
204. Leaves pubescent above.
C. cuneiformis.
189. Leaf blades 4-10 mm. wide.
205. Plants annual, less than 10 cm. tall; flower less than 8 mm.
long C. pumila.
205. Plants perennial, more than 10 cm. tall, up to 50 cm. or
taller; flowers most often 10 mm. long or longer.
206. Leaves entire or obscurely few-toothed, more than 3
times longer than wide.
207. Plants silvery lanate.
208. Calyx lobes 5-8 mm. long, acute.
209. Calyx lobes ca. 8 mm. long and 3 mm. wide at
base, lance-ovate, internally with a row of hair
along the margins C. decus montium.
209. Calyx lobes 5-6 mm. long and 4-5 mm. wide at
base, ovate, internally glabrous. . . .C. argentea.
208. Calyx lobes 4 mm. long, obtuse . . C. rhododendroides.
207. Plants glabrous or variously pubescent, not lanate.
210. Style (3.5-) 4.0-5.5 mm. long.
211. Subshrubs; corolla sac much shorter than the
non-saccate part of the lower lip.
C. rhododendroides.
211. Lianas; corolla sac equal to or longer than the
non-saccate, proximal part of the lip ... C. inflexa.
210. Style 1.5-3.0 mm. long.
212. Calyx lobes 10-12 mm. long. . .C. chrysosphaera.
546 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
212. Calyx lobes 3.5-7.0 (-8) mm. long.
213. Anther cells at least twice as long as wide.
214. Leaf blades less than 2 cm. long.
C. brachyantha.
214. Leaf blades more than 2 cm. long.
215. Lower lip of the corolla saccate more
than half its length C. engleriana.
215. Lower lip of the corolla saccate less than
half its length C. rhizomatosa.
213. Anther cells less than twice as long as wide.
216. Calyx lobes internally pubescent or puberu-
lent over most of the tissue. . C. angustiflora
216. Calyx lobes internally glabrous or with a row
of hairs near the margins and at the apex.
217. Upper corolla lip longer than the calyx
lobes C. rosmariniifolia.
217. Upper corolla lip shorter than the calyx
lobes.
218. Leaf blades covered with a short, dense
scurfy, brownish indument beneath
(pulverulent) C. inflexa.
218. Leaf blades most often hirsute, varying
to glabrous C. hirsutula.
206. Leaf blade margins obviously toothed or lobed, or if
entire or subentire less than three times longer than
wide.
219. Leaf blades obviously lobed and sometimes secondar-
ily toothed.
220. Blades lanate on tissue beneath.
221. Blades with 1 pair of lobes near the base.
C. triloba.
221. Blades with at least 2 pairs of lobes, one or more
pairs at or above the middle.
222. Blades less than 1 cm. long . . . . C. mittefoliata.
222. At least some of the blades 1 cm. long or longer.
C. hedera.
FLORA OF PERU 547
220. Blades usually sparingly pubescent on veins and
tissue beneath or densely glandular, or canescent,
tomentose or pulverulent.
223. Blades lanceolate in outline, occasionally lance-
ovate, 2 or more times longer than wide.
224. Blades usually tomentose beneath, margins
little revolute or plane C. cuneiformis,
224. Blades glandular beneath, margins revolute.
often folded to the midrib C. incarum.
223. Blades ovate to very broadly ovate in outline,
usually less than twice as long as wide.
225. Calyx lobes 6-8 mm. long, anthers divergent.
226. Calyx lobes 7-8 mm. long, glandular-pilose.
C. cuneiformis.
226. Calyx lobes ca. 6 mm. long, puberulent to
glabrous C. collanensis.
225. Calyx lobes 4-5 mm. long.
227. Anthers sagittate, blades mostly more than
1.5 cm. long C. lobata.
227. Anthers divaricate, blades mostly or entirely
less than 1.5 cm. long C. inamoena.
219. Leaf blades entire to toothed, lobes lacking.
228. Blades densely lanate beneath, at least on the veins
and veinlets.
229. Only the venation lanate C. croceopunctata.
229. Venation and tissue lanate.
230. Anther cells less than twice as long as wide.
231. Blade margins toothed C. sparsiflora.
231. Blades entire C. rhododendroides.
230. Anther cells at least twice as long as wide.
232. Teeth of the blade margins small, sharp,
blade occasionally subentire . . C. velutinoides-
232. Teeth of the blade margin large, rounded.
C. cordiformis.
228. Blades glabrous to variously pubescent but indu-
ment not long and lax.
233. At least some of the blades wider than long.
C. hutchisonii.
548 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
233. All the blades longer than wide.
234. At least most of the blades less than 1.2 cm.
long.
235. Style 3-5 (-6) mm. long, mostly 3.5-5.0 mm.
long.
236. Calyx lobes ca. 3 mm. long.
C. angustiflora.
236. Calyx lobes ca. 6 mm. long. .C. pearceana.
235. Style 1-3 mm. long, mostly up to 2.5 mm.
237. Ovary densely beset with erect, short, stiff,
white hair C. sibthorpioides.
237. Ovary glabrous to glandular-pubescent.
238. Polleniferous part of the anther-cells
only little more than half their volume,
walls and septa thick C. scabra.
238. Polleniferous part of the anther-cells at
least three-fourths of their volume, walls
and septa thin.
239. Leaves broadest at or near the bases.
240. Calyx lobes up to 4 mm. long.
241. Blades brown-pulverulent be-
neath C. bartsiaefolia.
241. Blades with white pubescence.
C. inamoena.
240. Calyx lobes 5-6 mm. long, mostly
more than 5.0 mm.
242. Blade margins serrulate with
very shallow, sharp, appressed
serrulations C. virgata.
242. Blade margins crenate, the often
rounded teeth extrorse.
243. Corolla 17-22 mm. long, occa-
sionally longer C. aurea.
243. Corolla up to 12 mm. long.
C. collanensis.
239. Leaves broadest at or near the middle.
244. Calyx lobes 3.0-3.5 mm. long.
FLORA OF PERU 549
245. Lower lip of the corolla saccate
more than half its length.
C. humilis.
245. Lower lip of the corolla saccate
less than half its length.
C. ramosissima.
244. Calyx lobes 4.5-8.0 mm. long,
mostly 5-7 mm. long.
246. Leaf blades less than twice as
long as wide.
247. Corolla sac often oblique,
lower lip of the corolla 12-15
mm. long C. virgata.
247. Corolla sac hemispherical,
lower lip of the corolla ca. 17
mm. long C. aurea.
246. Leaf blades at least twice as long
as wide.
248. Teeth when present on the leaf
blade margin few, 2-4 pairs
and shallow, less than one-fifth
the width of the blade.
C. myriophylla.
248. Leaf blade margins always
toothed or lobed, teeth few to
more often more than 4 pairs,
often at least one-fifth the
width of the blade.
249. Calyx lobes and capsules
glandless.
250. Leaves sometimes oblong
or cuneate-obovate, nar-
rowed to base and apex,
dentate on upper one-half
to two-thirds.
C. sparsiflora.
250. Leaves mostly subovate
to ovate, sometimes nar-
550 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
rowed to apex only,
toothed the entire length.
251. Calyx lobes 4 mm. long
and 3 mm. wide at base.
C. bartsiaefolia.
251. Calyx lobes (5-) 6 mm.
long and 4 mm. wide at
base C. extensa.
249. Calyx lobes and capsules
glandular . . . C. cuneiformis.
234. At least most of the leaf blades more than 1.2
cm. long.
252. Calyx lobes 3-4 mm. long.
253. Style 4-5 (-6) mm. long.
254. Blade margins coarsely doubly dentate.
C. verticillata.
254. Blade margins entire or crenulate.
255. Leaf blades ovate . . C. rhododendroides.
255. Leaf blades lanceolate, elliptic or nar-
rowly lance-ovate.
C. inflexa var. altissima.
253. Style 1.5-4.0 mm. long.
256. Leaf blades up to twice as long as wide,
usually less, lance-ovate or ovate, cre-
nate to serrate.
257. Calyx and pedicels glandular-pubes-
cent C. inamoena.
257. Calyx and pedicels eglandular.
C. ramosissima.
256. Leaf blades mostly at least twice as long
as wide, occasionally less than twice as
long as wide, then broadly serrate.
258. Upper lip of the corolla ca. 4-5 mm.
long, usually equaling to longer than
the calyx lobes C. glauca.
258. Upper lip of the corolla ca. 1-3 mm.
long, shorter than the calyx lobes.
259. Leaf blades usually not exceeding
2 cm. long C. ajugoides.
FLORA OF PERU 551
259. Leaf blades usually longer than 2
cm., up to 6 cm.
260. Uppermost leaf blades narrowly
lanceolate with fewer and smaller
marginal teeth than lower blades,
occasionally entire; calyx lobes
elliptical, obtuse or acute, pubes-
cent or puberulent on both sides.
C. angustiflora.
260. Uppermost leaf blades narrowly
ovate, marginal teeth as the lower
blades; calyx lobes lance-ovate or
lanceolate, internally glabrous,
acute or short-acuminate.
C. verticillata.
252. Calyx lobes 5-8 mm. long, rarely longer.
261. Style 4-6 mm. long or longer.
262. Flowers 20-37 mm. long; calyx lobes 7-
8 mm. long C. boliviano,.
262. Flowers 12-20 mm. long; calyx lobes
ca. 5-6 mm. long C. pearceana.
261. Style (1-) 2.0-3.0 (-3.5) mm. long.
263. Leaf blade margins entire or subentire.
264. Anther cells at least twice as long as
wide; calyx glandular C. viscosa.
264. Anther cells less than twice as long as
wide; calyx pubescent. . . .C. inflexa.
263. Leaf blade margins obviously toothed.
265. Upper lip of the corolla 8-10 mm.
long, often exceeding the calyx.
C. atahualpae.
265. Upper lip of the corolla up to 6 mm.
long, most often little to much shorter
than the calyx lobes.
266. Blade margins with rounded teeth,
crenate.
267. Corolla mostly 15-22 mm. long,
occasionally longer.
552 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
268. Leaf blades less than twice as
as long as wide C. aurea.
268. Leaf blades more than twice as
long as wide C. extensa.
267. Corolla up to 12 mm. long.
C. collanensis.
266. Blade margins with sharp teeth,
serrate, serrulate or dentate.
269. Lower lip of the corolla saccate
more than half its length.
C. inflexa.
269. Lower lip of the corolla saccate
less than half its length.
270. Leaf blades glabrous.
C. viscosa.
270. Leaf blades puberulent, pustu-
lar or glandular, at least above.
271. Calyx lobes 7-8 mm. long,
scabrous; capsule conic.
C. macrocarpa.
271. Calyx lobes 5-7 mm. long,
pilose or glandular-pilose;
capsule ovoid.
272. Calyx lobes 6-7 mm. long,
densely pilose externally.
C. virgata.
272. Calyx lobes 5 mm. long,
glandular-pilose on both
surfaces. . . .C. sparsiflora.
KEY TO THE SPECIES OF THE SUBGENUS Calceolaria
1. Both cells of the anther clearly fertile, the lower cell sometimes a
little modified.
2. Plants subscapose C. subscaposa.
2. Plants with well-developed leafy stems.
3. Leaf blades more than 3 cm. long.
4. Connective shorter than the anther cells that sometimes
appear to be contiguous C. delicatula.
FLORA OF PERU 553
4. Connective longer than at least one of the anther cells.
5. Capsule usually equal to or longer than the calyx, upper
leaf blades pinnate, some twice pinnate; flowers borne
in terminal or less often lateral dichasia . . . . C. pinnata.
5. Capsule shorter than the calyx lobes; upper leaf blades
pinnate or not; flowers solitary, axillary. . . .C. palustris.
3. Leaf blades less than 3 cm. long.
6. Lower lip of the corolla less than twice the length of the
upper C. flosparva.
6. Lower lip at least twice as long as the upper.
7. Connective longer than the anther cells . . . . C. palustris.
7. Connective shorter than the anther cells.
8. Lower lip of the corolla less than 12 mm. long.
C. sarmentosa.
8. Lower lip of the corolla more than 12 mm. long.
9. Leaf blades, at least the upper dissected to the mid-
rib, the pinnae lanceolate C. delicatula.
9. Leaf blades only rarely, if at all, dissected to the
midrib, when so dissected, pinnae usually ovate.
C. annua.
The lower cell of each anther sterile or so modified as to appear
sterile, or lacking.
10. Connective longer than at least one of the anther cells.
11. At least the larger leaf blades with bases thinly decurrent
over the length of the connate petioles and frequently with
more than 3 pairs of pinnae.
12. Pinnae most often decreasing in size base to apex; blades,
especially the lower, more than 12 cm. long and mostly
with 7 or more pairs of pinnae; nodes frequently beset
with dark exudate-filled pubescence. . .C. grandepinnata.
12. Pinnae increasing in size from base to apex; at least most
of the blades less than 12 cm. long and usually with not
more than 4 pairs of pinnae (rarely to 6 pairs); nodal
pubescence as on the stem C. tripartite!,.
11. Leaves not at all or only short-decurrent, rarely very thinly
decurrent the length of the petiole; petioles descrete; blades
most frequently merely dentate, sometimes varying to with
3 pairs of pinnae C. ranunculoides.
554 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
10. Connective shorter than the anther cells.
13. Leaf blades more than 3 cm. long.
14. Both anther cells developed, one sterile; lower lip of the
corolla longer than wide C. tripartita.
14. One anther cell frequently wholly lacking; lower lip of
the corolla wider than long.
15. Leaf blades often merely dentate C. chaetostemon.
15. Leaf blades mostly deeply incised or pinnate.
C. micrantha.
13. Leaf blades less than 3 cm. long.
16. Connective not auriculate.
17. Capsule wider than long C. tripartita.
17. Capsule longer than wide C. tennis.
16. Connective auriculate near the septum apex.
C. micrantha.
Calceolaria aequilateralis Edwin, Phytologia 19: 369. 1970.
Perennial herb, often scrambling, stem soft, 4-angled, frequently
pubescent or puberulent, especially below. Leaves bizarre, petioles
broadly winged, connate, broadest at the stem, usually serrulate, at
least on the lower half, occasionally irregularly dentate, usually a
little shorter than the blade, midvein flattened, elevated beneath, up-
permost blades sessile, broadly ovate, usually the smallest, remaining
blades (petiolate) very broadly deltoid or triangular, less often ovate-
triangular, frequently wider than long or as wide as long, very big
for the genus, 8-12 (-14) cm. long and 9-14 (-15) cm. wide, irregu-
larly coarsely crenate-dentate, narrowing to apex, cordate or truncate
at base, occasionally broadly ovate, puberulent; inflorescences usu-
ally few-flowered, ca. 7-15, dichasiate; flowers up to ca. 40 mm. long,
sepals up to 23 mm. long, broadly ovate, acute or acuminate, gla-
brous or puberulent on lower half, upper corolla lip ca. 8-15 mm.
long, cucullate, lower lip to 30-35 mm. long, orifice almost closed,
nectiferous flap much lighter colored, sac much inflated in life, gla-
brous, except for a tuft of hair inside at base; anthers large, cells
usually ca. 2 times as long as wide, mostly unequal, occasionally sub-
equal or equal, filaments short; style 8-10 mm. long; ovary pubes-
cent; capsule broadly ovoid, narrowed to apex, densely pubescent,
much smaller than and overtopped by the closely investing calyx.
FLORA OF PERU 555
Sporadic, from Prov. Amazonas in the north to Cuzco in the south.
Amazonas: Rocky cliff, alt. 3,300-3,400 m., Pennell 15694', Cerro
Puma-urco southeast of Chachapoyas. Rocky bank near summit,
alt. 3,300-3,400 m., Pennell 15905.— Cuzco: Macchu Picchu, Uru-
bamba Valley, alt. ca. 5,000 ft., Balls 6812; 2,000 m. Rauh-Hirsch
P 854; 809. — Lima: Hualgayoc: Monte Seco, alt. 1,800 m., Soukup
8834. — Pasco: Huariaca, wet place, alt. ca. 9,500 ft., Macbride 3124.
Calceolaria ajugoides Kranzl. in Engler, Bot. Jahrb. 50: Beibl.
110, 75. 1914.
Shrub (ex desc. up to 50 cm. tall, dark red branches curving-
ascending) with wood pilose or only sparsely so, foliage usually
densely tomentulose and internodes 2.5-5.0 cm. long, usually longer
than the leaves. Leaves of the branches short-petiolate, usually ter-
nate, lanceolate, few-dentate, upper often linear and entire, up to
2.51 cm. long and 4.0 mm. wide, usually less than 2.0 mm. wide,
sparsely pilose above, usually densely tomentulose beneath, the tissue
obscured by the vesture; petioles dilated and woody at base, thinly
coalescing and bearing in their axils abbreviated shoots whose leaves
are linear, entire and subsessile or sessile; inflorescences long and lax
dichasia or modified dichasia, few-flowered, the basal stalks only
once branched; bracts linear; peduncles and pedicels pilose; calyx
split to the base, sepals ovate, 3-4 mm. long and ca. 1 mm. wide at
base, densely pilose; corolla up to 20 mm. long, upper lip reduced to
a narrow ring; sac of the 18 mm. long lower lip ample, usually a little
more than half its length and ca. 1.2 cm. wide, minutely pilose; sta-
mens small, filaments ca. 1.0-1.5 mm. long, anthers a little longer
than wide; style 2.0-3. 5 (-4.0) mm. long, ovary glabrous or sparsely
puberulent; capsule unknown.
Ayacucho: Confluence of Lomas and Yanca Rivers at alt. 3,000
m., Weberbauer 5765 (Isotype F) ; Ocros2 alt. 3,200 m., Weberbauer
5801.
Calceolaria alba R. & P., FL, Peruv. 1: 19, t. 27. fig. a. 1798.
Subshrub, glabrous or almost so, stems branched and stems and
bracts both with abbreviated shoots clustered in leaf-axils; leaves
sessile, filiform, up to 2.5 cm. long, often less than 2.0 cm. long and
1 Ex Kranzlin, no leaves approaching this length seen, even on the isotype.
2 Erroneously attributed to Dept. Ancash by the collector; probably collected
at Parinacochas, fide Pennell annotation.
556 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
rarely reaching 2 mm. wide, entire, costa elevated beneath, abbrevi-
ated shoots in many axils densely foliose, therefore axils of leaves of
the stem and branches appearing very densely foliate; inflorescences
few- to many-flowered, lax, distant, paniculate or cymose clustered ;
pedicels, calyces and sometimes the corollas minutely papillose-
glutinous or -tomentose; peduncles 2-3 cm. long; pedicels ca. 3-5 mm.
long; calyx lobes ovate to elliptical, obtuse, 2.0-2.5 mm. long and
ca. 1.0-1.2 mm. wide; corolla white, drying yellow or both corolla
and calyx yellow (ex Kranzl.) upper lip annulate, ca. 1 mm. wide,
lower lip narrowed at base, up to 1.2 cm. long and 7-8 mm. wide;
filaments minute, the divaricate anthers about 1.5 mm. end to end;
style about 1.5 mm. long, ovary sparsely papillose; fruit unknown.
Sin. Loc. : Dombey s.n.
Also known from Chile and Argentina.
Calceolaria alternifolia Cav., Icon. Rar. 5: 28, t. 445, f. 2. 1799.
Differing from C. linearis R. & P. to which it is closely allied in
the following ways: the branches all very short and strictly appressed
to the stem, occasionally elongate; anthers much smaller and about
2 times as long as wide, the ovoid to almost round capsules lacking
a suddenly narrowed, long upper half; some of the leaves suboppo-
site; the upper lip of the deep-yellow corolla only a little shorter than
the 4-5 mm. long calyx lobes, the lower lip reaching up to 20 mm.
long, abruptly narrowed throughout the non-saccate proximal half.
Amazonas: Llui, alt. 11,000 ft., Carricker, s.n.', Cerro Puma-urco,
summit, 3,400-3,500 m., Pennell 15909. — Ancash: Catuc, 15 mi. east
of Huaraz, dry river bluffs, ca. 10,500 ft., Macbride & Featherstone
2511. — Piura: Huancabamba (Porculla?) Piura Divide, alt. 8,500 ft.
Sandeman 4.290. — Sin. Dept.: Buenaventura, Nee s.n.; Chasqui,
stream banks, alt. ca. 10,500 ft., Macbride & Featherstone 1772,
("Okush Romaro").
Calceolaria angustiflora R. & P., Fl. Peruv. 1: 17, t. 28a. 1798,
non Hook. C. terniflora Cav., Icon. Rar. 5: 30, t. 450, f. 2. 1799.
Upright, erect, puberulent subshrub with usually verticillate
leaves, mostly with ascending, elongate branches, abbreviated
branches sometimes present in leaf-axils. Leaves lanceolate to more
often lance-elliptic or narrowly lance-ovate, 1.5-4.6 cm. long and
0.7-2.1 cm. wide, serrate or serrate-serrulate, rarely uppermost nar-
rowly linear and subentire, apex acute, base obtusish, usually puber-
FLORA OF PERU 557
ulent on both surfaces, more densely beneath, especially on the veins,
occasionally puberulent only beneath, rarely glabrous, often with
raised, red-black, glandular punctations, short-petiolate, occasion-
ally sessile; flowers in elongate panicles or dichasia; peduncles and
pedicels pilose, up to 3 cm. long; calyx pilose, segments ovate or ellip-
tical, ca. 2.5-3.5 mm. long, pubescent on both surfaces, often white
margined ; upper lip of the corolla shorter than the sepals, lower lip
narrowed proximally, elongate, distally flaring, saccate to about half
its 10-18 mm. length, the sac only little narrower than long, slightly
upturned, purple marked internally on lower surface; anther cells
equal, less than to more than 2 times longer than wide, divaricate or
almost so, not exceeding 2 mm. long, each cell usually less than 1 mm.
long, when exceeding 2 times longer than wide sometimes downturned
at the free ends; style erect 2-4 (-5) mm. long, ovary glabrous to
densely beset with black-red glands; capsule shorter than and in-
vested by the calyx, glabrous or glandular as the ovary, ovoid, nar-
rowed to the blunt apex.
Ancash: Monterrey, 3 km. below Huaraz, stony slope, alt. 3,000-
3,100 m., Pennell 15303; slope along stream Rio Santa, opposite
Huaraz, alt. 3,100-3,150 m., Pennell 15327; Banks, alt. 2,700-2,900
m., Colca, Pennell 15383; toward El Cerro Shaurema, southeast of
Huaraz, alt. 3,500 m., Pennell 15427; Que"brada de Huaqui, Huaraz,
alt. 3,000-3,050 m., Pennell 15433; Hillside north of Huaraz, alt.
3,250 m., Pennell 15443.— Ay acucho: Coracora, alt. 2,800-2,900 m.,
Weberbauer 5791. — Lima: Prov. Huarochiri, Surco, shore of rivulet,
alt. ca. 2,000 m., Asplund 11060; Huaquisha, above Surco, alt. 2,600-
2,700 m., Ferreyra 6073; Surco, rocky banks, alt. 2,300-2,700 m.,
Pennell & Ferreyra 15240; South of Surco, alt. 2,400-2,500 m., Pen-
nell & Ferreyra 15244; Matucana, Rose & Rose 18638; Sin Loc.:
Ruiz & Pavon s.n.
The collections from Ancash have the glandular ovaries and cap-
sules. The Ayacucho collection has the least pubescence. Although
most collections are from Lima Department, this taxon is probably
widespread from the central to southern Andes in Peru.
Calceolaria annua Edwin, Phytologia 19: 370. 1970.
Annual, branched herb, upper part of the stems pilose with 2-
celled, usually gland-tipped hairs, glands minute, becoming glabrous
near the base, internodes almost always longer than the leaves; leaves
petiolate, very variable, especially as to margins, usually ovate in out-
line, mostly 1-2 cm. long and 6-9 mm. wide, rarely reaching 3 cm.
558 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
long, margins from grossly dentate especially below to occasionally
pinnate or almost so above, when pinnate, the pinnae variable in size
and outline, tissue usually glabrous above, occasionally pubescent,
most often pulverulent beneath, base of the blade only very short
decurrent on the densely pubescent, descrete petioles; inflorescences
most often 3-flowered cymes, rarely reduced to 1 flower, borne in
pairs from the apices of the uppermost leaves or occasionally from
the apices of the upper 2-3 pairs of leaves; peduncles up to 3.5 cm.
long, pilose as the upper part of the stems; flower size variable; calyx
tube pubescent, lobes externally subglabrous, veins parallel, inter-
nally glabrous, ovate or elliptical, acute, sometimes 2 different sizes
on one flower, 6-8 mm. long and 2-5 mm. wide; corolla yellow, upper
lip 2-4 mm. long, sparsely glandular, glands red, sessile, internally
glabrous, nectary almost marginal on the sac, 2-parted, very reduced;
filaments less than 1 mm. long, anther cell connective shorter than
both of the fertile locules the larger of which is about 2 mm. long and
the smaller about 1.3-1.4 mm. long; style glabrous, little curved,
1.9-2.4 mm. long; capsule about as long as to little longer than the
calyx, glabrous, ovoid, the upper part abruptly narrowed into a short,
lanceolate neck, style soon fugacious.
Distributed in Dept. of Lima and possibly one collection from
Junin (Iltis, Iltis, Ugent & Ugent 226, WIS).
Lima: Rio Blanco, between San Mateo and Casapalca, alt. 3,500-
3,550 m., Ferreyra 6974-', Moist rock ledges and granitic soil, S. of
Surco, alt. 2,400-3,000 m., Pennell & Ferreyra 152^5 (US); Rio
Blanco, alt. ca. 15,000 ft., Macbride 3038; alt. ca. 12,000 ft., Mac-
bride & Featherstone 683; Canta, alt. 2,700-3,200 m., Pennell 1^333.
The combination of usually dentate leaves and relatively large
corolla with copious sac and very reduced nectary distinguishes this
taxon from C. delicatula Kranzl., to which it is morphologically
closely allied.
Calceolaria aperta Edwin, Phytologia 19: 371. 1970.
Shrub to about 1 m. tall, foliage densely lanate or tomentose,
stem branched, drying red-brown, appressed-puberulent or -pilose
when young, glabrate in age. Leaves narrowly ovate or ovate or
oblong-ovate, those of the stem up to 4.2 cm. long and 2.0 mm. wide,
usually a little smaller, those of the branches smaller, all leaves
densely lanate when young, becoming tomentose with age, margins
crenate-dentate or crenulate-denticulate, apices acute, bases broadly
FLORA OF PERU 559
obtuse to (especially the larger) cordate, short-petiolate; petioles
lanate; flowers in compound dichasia with long basal peduncles (up
to 5.5 cm.) and short pedicels, ca. 0.5-2.0 cm. long, flower-stalks pu-
berulent to tomentose; calyx lobes ovate, ca. 6 mm. long and almost
as wide at base, apex obtuse, tomentulose on both surfaces; corolla
13-17 mm. long, essentially glabrous, upper lip erect, up to 5 mm.
long, almost as long as the sepals, hooded, lower lip saccate merely
along the margin, about 1-3 mm., nectary thin, linear; filaments
short, ca. 1 mm. long, thickest at base; anthers large for the corolla,
each cell about 2 mm. long and almost as wide medianly, broadly
ovoid, papillose-glandular, drying brown with buff-colored lower
margins; style thickened, 7-8 mm. long, stigma very thickened and
gibbose much larger than the style, ovary short, broadly ovoid,
densely beset with sessile glands; capsule unknown.
Known only from the type collection.
Lima: Prov. Canta: near Antaicocha, Cerro Colorado, east of
Canta, rocky wall of canyon, alt. 3,600-3,800 m., Pennell 14642.
A very clearly marked taxon with no morphologically similar species
in Peru.
Calceolaria arborescens Edwin, Phytologia 19: 372. 1970.
Erect, much-branched, subglabrous shrub with rigid stems and
subcoriaceous, bicolorous, viscid leaves, pubescence restricted to the
inflorescences and sometimes the younger wood. Leaves glabrous,
ovate to less often elliptical, variable in size from 1-6 cm. long and
0.5-2.5 mm. wide, usually ca. 3-5 cm. and 1-2 cm., entire to serru-
late, narrowed to apex and base, often dimidiate, drying dark green
above and pale yellow-green beneath, veins faintly visible above,
prominent beneath, costa a little elevated, other venation reticulate,
petiolate; petioles up to 1.4 cm. long, mostly 5-8 mm. long; inflores-
cences many-flowered, paniculate dichasia, terminal on the branches;
the flower-stalks puberulent or pubescent, basal peduncles up to 4.5
cm. long, terminated by a pair of reduced leaves, decreasing to ulti-
mate pedicels a few mm. long; calyces puberulent-pubescent, unequal
in width, ovate to elliptical 6-8 mm. long and 3-5 mm. wide at base,
margins minutely ciliolate, apices acute, internally usually glabrous;
corolla yellow, 25-27 (-30) mm. long, both lips externally more or
less densely beset with minute stalked glands; upper lip hooded, up
to about 4 mm. long, lower lip up to 25 mm. long, saccate to three-
fifths its length, only little narrowed proximally, glabrous internally
or with a tuft of white hairs at base; filaments very short, anther cells
560 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
divaricate, slightly to very unequal in size, each pair of cells 5 mm.
long, each cell most often at least twice as long as wide, anthers de-
hiscing through the septum; style 3-4 mm. long, a little curved at
apex, stigma punctiform, ovary glandular as the corolla; capsule not
known.
Amazonas: Cerro Calla Calla above Leimebamba (19 km.), alt.
3,000 m., Hutchison & Wright 6916. — Ancash: Bolognesi, between
Tallenga and Pachapaque, alt. 3,500-3,600 m., Ferreyra 7^79; West
slope of Cordillera Negra below Paso de Collan, rocky slopes and
banks, alt. 3,650 m., Pennell 154.76; Moist bank above stream, south
slope of Paso de Fortale'za, alt. 3,550 m., Pennell 15398. — Junin: alt.
3,500m., Ochoa721.
Morphologically closest to C. deflexa var. aurantiaca and also C.
deflexa var. typica differing from both in having the lower lip of the
corolla more than twice as long as the upper, in unequal anther cells
and in the upper lip of the corolla shorter than the sepals as well as
different vesture in the inflorescence. C. arborescens differs from the
typical deflexa also in leaf shape. The superficial resemblance be-
tween arborescens and deflexa var. aurantiaca is striking but the taxa
differ profoundly on a technical basis.
Calceolaria argentea HBK., Nov. Gen. et Sp. 2: 382, t. 170.
1817.
Erect, branched, usually white-lanate shrub, older wood excepted.
Stems and branches smooth; leaves oblong or oblong-lanceolate, up
to 2 cm. long and 3-4 mm. wide, entire, narrowed to both ends,
sparsely lanate above, densely so and obscuring the tissue beneath,
short-petiolate; petiole thickened and woody at base; inflorescences
at the apex of stem and branches, terminal and axillary, branched
dichasia; flower stalks lanate, 1.0-1.5 cm. long; calyx segments ovate,
acute, 5-6 mm. long and 4-5 mm. wide at base, internally glabrous;
upper lip of the corolla shorter than the sepals, lower lip much larger,
up to 2 cm. long, obovoid, minutely puberulent externally, saccate
over half of its length, the sac suddenly flaring from near the base;
anther cells, equal, divaricate, filaments short. (Ex original descrip-
tion of Kranzlin and type photograph.)
Close to C. decus-montium from which it differs in size and shape
and internal vesture of the calyx lobes.
Northern Peru in Cajamarca and Paramo de Guamani.
FLORA OF PERU 561
Calceolaria atahualpae KranzL, Pflanzenr. IV (257C): 107.
1907. C. excelsior KranzL, Bot. Jahrb. 50: Beibl. Ill, 74. 1913.
Tall, scandent vine, branches square, diverging, glabrous below
and short-dense- tomentose above, interned es varying in length.
Leaves elliptical, less often oblong, occasionally ovate or ovate-
elliptical, 1-4 cm. long and up to 1.3 (-2.0) cm. wide, vesture very
variable, glabrous to minutely pilose above, sparsely to densely to-
mentose or pilose beneath, crenulate, apically narrowed, basally acute
to subobtuse, rarely obtuse, short-petiolate; petiole usually not at all
thickened nor woody at base; inflorescences frequently 3-flowered
dichasia at the apices of the branches; flowers most often on thin,
pilose, up to 4 cm. long stalks or sometimes stalks with a few glandu-
lar hairs intermixed; calyx lobes elliptical to ovate, 6-8 (-9) mm. long
and 3-4 mm. wide, externally pubescent, internally with a distinct
line of very short hairs along the margins, otherwise glabrous to
pilose; corollas among the largest in the genus, upper lip longer than
the calyx lobes, ca. 1.0-1.2 cm. long, wider than long, lower lip up
to 3.2 cm. long and almost as wide, saccate more than half its length
with transverse or rectangular orifice, externally glabrous or minutely
pilose, internally glabrous to densely pilose, especially at the base of
the lower lip; filaments very short, anthers small, each locule ca. 1
mm. long, drying (dark) brown; stigma 2-3 mm. long, ovary gla-
brous; glutinous; capsule unknown.
Ayachuco: Pampalca between Huanta and Rio Apurimac, alt.
3,200 m., Killip & Smith 23237; below Tambo and Rio Apurimac,
alt. 3,500-3,600 m., Weberbauer 5576, the type collection of C. excel-
sior; between Tambo and Rio Apurimac below Yanamonte, Weber-
bauer 5596; Prov. Abancay, alt. 3,200 m., Vargas 77 b. — Cuzco: Prov.
Urubamba, alt. 2,100-2,400 m., Vargas 797; alt. 2,800 m., Vargas
2655; Prov. Calca, below Lares, alt. 2,800-2,900 m., Weberbauer 7898.
— Huancavelica: Prov. Tayacaja, Que"brada 1 km. N. of Salcabamba,
alt. 3,500 m., Stork & Horton 10333. — Junin: Carpapata above Hua-
capistana, alt. 2,700-3,200 m., Killip & Smith 24377.
Specimens originally determined as Calceolaria excelsior Kr. differ
from C. atahualpae in a few details, especially as to vesture (as de-
scribed by Kranzlin) but the differences do not appear to me to be
worthy of specific distinction.
Calceolaria aurea Pennell, Proc. Acad. Sci. Phila. 97: 159. 1945.
Shrub with much branched, finely pubescent stem, older wood
puberulent. Leaf blades oblong-ovate, 1.0-2.5 cm. long and 8-12
562 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
mm. wide, dentate, the 6-8 pairs of teeth rounded, plane, pilose
above, veins pilose beneath, intervening tissue reticulate, glabrescent,
rounded at both ends; petioles pubescent, 2-5 mm. long; inflores-
cences of solitary flowers or few-flowered, paired cymes; pedicels
finely pubescent, glandless; bracts and bracteoles leaf -like but re-
duced; sepals ovate, 5-6 mm. long, entire, apically rounded, greenish
yellow, pubescent on both surfaces, finely so internally; corolla
golden-yellow, externally finely pubescent, upper lip less than 2 mm.
long, arching over but not covering the stamens, lower lip 16-22
mm. long, sac shorter than the non-saccate part, internally red-
purple blotched and with a tuft of white hairs at base; anthers 3.0-
3.5 mm. long, brown, cells cylindric, glabrous, divaricate, about
three-fifths as wide as long, dehiscing through the septum; style
2 mm. long, glabrous; capsule 7-9 mm. long, ovoid-conic, densely
pubescent, glandless. (Desc. ex Pennell.)
Cuzco: Paucartambo Valley 3,500 m., Jan. 1927, Herrera 1416;
same data except Jan. 1926, Herrera s.n.; Colquipata, rocky banks,
alt. 3,200-3,500 m., Pennell 13772, Isotype (F; US); Quispican-
chis, alt. 3,700-3,900 m., Vargas 922; 9816; alt. 3,500 m., Vargas
1366; Prov. Calca, Amaru, Pisac, alt. 3,500 m., Vargas 5026; Prov.
Canas, Asuncion Bridge, Apurimac River, alt. 3,760 m., Vargas
11036.
Calceolaria ballotifolia Kranzl. in Fedde, Rep. Sp. Nov. 27:
18. 1929.
Tall shrubby climber, stem profusely branching, wood sparsely to
densely pubescent, the older sometimes glabrate. Leaf blades ovate,
1.5-6.0(-8.0) cm. long and 0.8-3.5 cm. wide, more or less densely
pubescent on both surfaces, especially on the veins beneath, little
asymmetrical, coarsely crenate-dentate, apically acute, broadly ob-
tuse and often cordate and dimidiate at base, upper surface usually
much darker green than the lower, petiolate, uppermost occasionally
subsessile; petioles pubescent, very short to about 1 cm. long, those
of the main stem thickened at the base and each pair surrounding the
stem as a thin ring; inflorescences terminal or axillary in uppermost
leaves, few-flowered, unbranched to twice branched dichasia; flower-
stalks pilose; flowers small up to about 16 mm. long; calyx lobes ex-
ternally pilose, ovate, 6-7 mm. long, acute, margin fimbriate or en-
tire, internally glabrous to pilose; corolla white to pale yellow, upper
lip of the corolla well developed, two-thirds to almost as long as the
calyx lobes, lower lip ca. 10-12 mm. long, broadly oblong, saccate
FLORA OF PERU 563
about half-way ; stamens short, anther cells subparallel to sagittate,
each pair joined about half their length apically, usually drying
brown, about two-thirds as wide as long, slightly unequal in length
and width; style 5-6 mm. long, ovary shining black; capsule conic-
ovoid, a little longer than the calyx.
Amazonas: Cerro Puma-urco, southeast of Chachapoyas, alt.
3,400 m., Pennell 15706; Chachapoyas, alt. 2,700 m., LI. Williams
7575.— Libertad : Prov. Pataz, Weberbauer 7036, Isotype (F).
Calceolaria bartsiaefolia Wedd., Chi. And. Nov. 2: 134. 1860.
Differing from C. aurea Pennell in having densely tomentose
stems with the vesture black, cinereous near the apex; leaf blade
margins grossly and sharply few-dentate, the teeth lightly recurved,
glabrous above, canescent-pilose beneath, ovate and oblong, 5-10
mm. long and 3-4 mm. wide; flowers in few-flowered corymbs (usu-
ally 3-flowered); pedicels glandular-pilose; calyx segments ovate,
acute, 4 mm. long, basally 3 mm. wide.
Reported by Kranzlin from Chicla at 4,300 m., Ball s.n. If these
obviously closely related taxa were considered conspecific bartsiae-
folia, the older epithet, would take precedence.
Calceolaria bicrenata R. & P., Fl. Peruv. 1: 15, t. 23 b. 1798.
C. bicolor R. & P., Fl. Peruv. 1: 16, t. 25b. 1798. C. trifida R. & P.,
I.e. 17, t. 26b. C. flexuosa R. & P., I.e. 17, t. 26a. C. saxatilis HBK.,
Nov. Gen. et Sp. 2: 382. 1817. C. lamiifolia HBK., Nov. Gen. et
Sp. 2: 383. 1817. C. chimborazensis Willd. ex Link, in Spreng.,
Jahrb. 1(3) : 52. 1820. C. lancifolia Willd. ex Link, in Spreng., Jahrb.
1(3): 52. 1820. C. diffusa Lindl., Bot. Reg. 16: t. 137 b. 1830. C.
macrocalyx Kranzl. in Fedde, Rep. Sp. Nov. 1: 99. 1905.
Shrub or subshrub, occasionally a climber, from densely pubes-
cent to glabrous or almost so, stems and branches erect to semi-
prostrate. Leaves all petiolate or the upper sometimes almost sessile,
opposite or ternately verticillate above, ovate, sometimes cordate,
occasionally oblong, rarely very broadly ovate, 3-10 cm. long and
1-6 cm. wide, mostly 5-8 cm. and 1.5-4.0 cm., acute, occasionally
subobtuse, margins usually deeply doubly toothed, the teeth sharp
or dull, rarely regularly dentate, base when not cordate, obtuse,
tissue chartaceous to firm and thickened ; inflorescences few- to many-
flowered, dichasiate, corymbose or paniculate, terminal or in upper
leaf-axles of the branches; interfloral leaves reduced; flower-stalks
decreasing base to apex; flowers large for the genus; calyx lobes
564 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
ovate, 5-10 mm. long, mostly 6-9 mm., light yellow-green to dark
brown-green, acute, entire or few-toothed, sometimes reflexed after
anthesis, rarely 2 sizes on one flower; corolla yellow and white or all
yellow, often red-glandular-dotted and pustulate externally, upper
lip of the corolla minute, ca. 1 mm. long to about half as long as the
sepals, rarely to two-thirds as long, usually arched over the genitalia,
lower lip 15-30 mm. long, saccate about half way or less, sac some-
times furrowed, erect, ascending or patent, up to about two-thirds
as wide as long, usually half as wide, internally with a tuft of multi-
cellular, white hairs at base; stamens usually short, shorter than the
style, anther cells at least 2 times longer than wide, divergent or
divaricate, distally downturned or plane, sometimes when divergent,
proximally joined apically and appearing subsagittate; style 1.5-
2.5(-3.3) mm. long or rarely 4-5 mm. long; ovary usually glandular,
glands red; capsule ovoid-conic, usually glandular and a little longer
than the calyx, frequently blunt at apex.
Generally distributed in the Andes from Cajamarca in the north
to Arequipa in the south. Citations are selected from the quite
copious material.
Ancash: Prov. Bolognesi, Arriba de Rasrachaca, alt. 3,300-3,400
m., Ferreyra 9508. — Arequipa: Condesuyas, Chuquibamba, alt.
10,000 ft., Stafford 1158. — Ayacucho: Pampalca between Huanta and
Rio Apurimac, alt. 3,200 m., Killip & Smith 22251 ; north of Huanta,
near Huaillay, alt. 3,500-3,600 m., Weberbauer 7590. — Cajamarca:
Cajabamba, Carricker s.n.; Osgood & Anderson 55; Along streams on
jalca, Cord, de Cumulloa to Celendin, alt. 3,300 m., Pennell 15169;
Along stream E. of Cajamarca, alt. 2,600-2,700 m., Pennell 15226.—
Huancavelica: Larmes al E. of Conaica, alt. 3,400-3,450 m., Tovar
174; Salcabamba, Prov. Tayacaja, shrubland, alt. 3,250 m., Stork &
Horton 10275; Prov. Huancavelica, 4 km. south of Mejorada, alt.
2,500 m., Stork & Horton 10919. — Huanuco: Huacachi station near
Muna, alt. ca. 6,500 ft., Macbride 3883. — Junin: Between Viques and
Ingahuasi, Mantaro Canyon, S. of Huancayo, alt. 3,150 m., Killip &
Smith 22144; Tarma, along stream, alt. 13,000 ft., Macbride & Feath-
erstone 1071 ; Pueblo Chinchapalca, 5 miles above Mito, alt. ca. 9,500
ft., Macbride & Featherstone 1588. — Libertad: Prov. Otuzco, Agall-
pampa, alt. 3,200 m., Lopez 0307; Dry bank, above Motil, alt. 3,000
m., Pennell & Ferreyra 14840. — Lima: Along Rio Chillon near Obra-
jillo, bushy banks, alt. 2,200-2,500 m., Pennell 14328; Canta, rocky
slopes and along irrigation ditch, alt. 2,700-2,900 m., Pennell 14353;
FLORA OF PERU 565
San Buenaventura, open rocky bank, alt. 2,800-2,900 m., Pennell
14543; Moist thicket, hills south of Surco, alt. 2,000-3,000 m., Pen-
nell & Ferreyra 15251; Canta, Ruiz & Pawn s.n.; Prov. Cajatambo,
Banos de Churin, alt. 8,000 ft., Sandeman 5388. — Pasco: Vicinity of
Cerro de Pasco, ledge of limestone cliff, alt. ca. 12,500 ft., Jenks 40;
Yarush, alturas de Huariaca, alt. 4,000 m., Ochoa 1116. — Sin. Loc.:
Dombey s.n.; Mathews 1681.
Calceolaria boliviana (Britton ex Rusby) Pennell, Proc. Acad.
Nat. Sci. Phila. 97: 170. 1945.
Perennial herb or climbing shrub, young wood 4-angled and pu-
berulent, older glabrate, often with peeling bark. Leaf blades lanceo-
late to usually lance-ovate or ovate, 1.5-6.0 (-7.0) cm. long and
0.5-3.0 (-3. 5) cm. wide, crenate or crenate-dentate, long-acute or
acute apically, usually subobtuse or obtusely rounded basally, usu-
ally puberulent on both surfaces, sometimes only pilose on veins
beneath, most often red-punctate on both surfaces; petioles 0.4-
1.2 cm. long, usually thickened and pubescent and narrowly connate
across the node, not at all woody; inflorescences terminal, dichasiate
or corymbose, not exceeding 10-flowered, usually less; pedicels and
peduncles pubescent or puberulent, peduncles 2-8 cm. long, pedicels
decreasing base to apex, the ultimate pedicels smallest; bracts and
bracteoles reduced, lanceolate, crenate or crenulate; flowers very
variable in size; calyx lobes often reflexed in anthesis, 6-9 mm. long,
and 5-6 mm. wide at base, ovate, obtuse or acute, entire or few-
dentate; corolla yellow or orange, drying orange, usually large for
the genus, 20-35 (-37) mm. long and about half as wide distally,
most often externally puberulent and internally with a tuft of white
hairs at the base of the lips; upper lip longer than the sepals, up to
1.2 cm. long, usually erect, lower lip 20-28 mm. long, saccate over
half its length; stamens small, anthers divergent or almost parallel,
only little longer than wide, reniform or ovoid, or as wide as long
and almost spherical, brown in life and when dry; ovary red-glandu-
lar or pubescent, style 6-8 mm. long; capsule about as long as wide
at base and about equal to the calyx in length, glabrous, glandular
or pubescent.
Two varieties both from southern Peru (Cuzco and Puno) are
easily separable.
Leaves mostly 3.5 cm. long or longer, crenate; ovary and capsule
glandular or glabrous C. boliviana var. boliviana.
566 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Leaves all or mostly shorter than 3.5 cm., dentate; ovary and capsule
pubescent C. boliviano, var. speciosa.
Calceolaria boliviana (Britton ex Rusby) Pennell var. bolivi-
ano,. Fagelia boliviana Britton ex Rusby, in Bull. Torr. Bot. Club
27: 24. 1900. Calceolaria zanatilla Kranzl. in Fedde, Rep. Sp. Nov.
1:107. 1905.
Cuzco: Prov. Quispicanchis: Near Marcapata, alt. ca. 3,000 m.,
Metcalf 30733; Near Rio Yanamayo, below Pillahuata, alt. 2,100-
2,300 m., Pennell 13931+', Between Mistiana and Kerous, Valle Cos-
fiipata, near Paucartambo, alt. 600-950 m., Scolnik 888; Prov. Pau-
cartambo, between Tanamayo and Tambamayo, alt. 1,500-2,200 m.,
Vargas 67; Prov. Calca, near Vilcabamba, alt. 2,800 m., Vargas 3937;
Paucartambo, Rio Tambomayo, West 7095 (cult, at U. Calif.).—
Puno: Prov. Sandia, near Limbani, alt. 3,200-3,450 m., Metcaif
30530.
Calceolaria boliviana var. speciosa (Pennell) Edwin, stat. nov.
C. speciosa Pennell, Proc. Acad. Nat. Sci. Phila. 97: 167. 1945.
Cuzco: Prov. Paucartambo, alt. 12,000-13,000 ft., Balls 6701;
6776; Paucartambo to Tre's Cruces, Cerro de Cusilloyoc, alt. 3,400-
3,500 m., Pennell 13813; Alt. 3,400-3,600 m., Pennell 13816, type
collection of C. speciosa; Alt. 3,700 m., Vargas 1547; Between Acja-
macu and Tr6s Cruces, alt. 3,500-3,700 m., Vargas 2249; Prov. Uru-
bamba, Puyupatamarca, alt. 3,400 m., Vargas 2779a; Prov. Paucar-
tambo, Dist. Marcachea, vicinity of Marcachea, alt. 3,400-3,500 m.,
Vargas 1175.
Calceolaria brachyantha Edwin, Phytologia 19: 373. 1970.
Glandular, erect shrub, much branched with ascending branches.
Leaves minute to very small, sometimes with abbreviated shoots in
the axils, the leaves then appearing clustered, linear or occasionally
lanceolate-linear or aciculiform, mostly 1-3 mm. wide but infre-
quently up to 6-7 mm. and up to about 1.2 cm. long, mostly shorter,
margin entire or with 2-3 pairs of teeth near the apex, usually revo-
lute, apically acute, either sessile or short-petiolate, when present the
minute petiole thickened and the leaf-base short-sagittate, blades
glandular-puberulent or -pubescent on both surfaces, especially on
the costa beneath, main vein impressed above, elevated beneath,
other venation obsolete, tissue dark green above, pale green beneath,
coriaceous and usually thickened; inflorescences terminal on the
FLORA OF PERU 567
branches, few- to about 15-flowered, in regularly or irregularly
branching dichasia; peduncles and pedicels glandular and erect, pe-
duncles usually 2-3 times longer than the pedicels; flowers very
variable in size; sepals lance-ovate to ovate on the same flower,
usually about 4 mm. long and 3 mm. wide at base or 5 mm. long and
2 mm. wide at base (those of the smallest flowered collections about
two-thirds the others), acute, glandular-puberulent on both surfaces;
corolla yellow, 13-26 mm. long, externally glabrous or the upper lip
puberulent; upper lip about a half to two- thirds the length of the
sepals, covering the stamens, lower lip oblong, saccate about three-
fourths to four-fifths its length, internally glabrous; stamens with
very short (subobsolete) filaments, anthers among the smallest in the
genus, cells a little unequal or almost equal, divaricate, each pair ca.
2.5-3.0 mm. long and about one-third to a half as wide; style thick-
ened, ca. 2.0-2.5 mm. long, about the size of the densely glandular
ovary; capsule glandular, exceeding the calyx, 5-7 (-8) mm. long, and
2-3 mm. wide at base, ovoid-conic, long-acute, style persistent.
Distributed in the Andes from north central to northern Peru,
departments Ancash, Libertad and Cajamarca.
Ancash: Shincush above Chiquian in Prov. Bolognesi, alt. 3,980
m., Ferreyra 5820. — Cajamarca: Between Cajabamba and Huama-
chuco, Prov. Cajabamba, alt. 3,000-3,200 m., Ferreyra 3053; Cumbre
above Cajamarca, alt. 3,500 m., Olsson 2; Cajamarca: Culentin
(prob. Celendin) Road, km. 15.5, Olsson s.n., Feb. 11, 1948; Caja-
marca: Rocky limestone, alt. 2,750-2,850 m., ridge, 11 km. S. of
Cajamarca, Pennell & Anderson 15076; Rocky limestone, alt. 2,750-
2,850 m., ridge 11 km. S. of Cajamarca, Pennell & Anderson 15077;
Banos del Ynca, just outside Cajamarca, Soukup 4627 and 4630.—
Libertad: Toyabamba, in timber-line scrub, alt. 11,000 ft., Carricker
s.n.; Otuzco, Ferreyra 3043; Above Huamacucho, alt. 3,400 m., Pen-
nell & Ferreyra 14872.
This species is close to Calceolaria linearis R. & P., C. cajabambae
Kr., C. ludens Kr., and C. engleriana Kr., differing from these by
various character combinations, especially anther size, calyx lobe
shape, and vesture and leaf blade shape and size.
Calceolaria cajabambae Kranzl. in Fedde., Rep. Sp. Nov. 1:
102. 1905.
Shrub with branched stems, wood pilose, sometimes drying black
or black-dotted. Leaves numerous, small, mostly oblanceolate or
568 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
lanceolate, up to 2-3 mm. wide often less and up to 1-2 cm. long,
margin revolute, entire or with small rounded-obtuse teeth, espe-
cially on upper half, acute at both ends, sometimes long-acute to
base, coriaceous, pilose above and below, tissue clearly visible espe-
cially when mature; the pubescence of wood and foliage erect or al-
most so and short; inflorescences terminal or in upper leaf-axils,
2-3-flowered or flowers solitary; pedicels 2-3 cm. long, pilose or vil-
lous; bracts merely reduced leaves; flowers 20-25 mm. long; calyx
densely pilose externally, pubescent internally, lobes broadly ovate,
ca. 7 mm. long and 5 mm. wide at base, acute; upper lip of the yellow
corolla a little shorter than the calyx, ca. 4-6 mm. long, lower lip
obovate-orbicular, 16-19 mm. long, saccate less than half its length,
sac erect, entire corolla densely to sparsely pilosulous externally, the
throat (ex desc.) pulvinate-papillose or glabrous; stamens on short
filaments; anther cells 2 or more times longer than wide, unequal,
divaricate, each pair of cells ca. 5 mm. long, septum thickened; style
3.0-3.5 mm. long; ovary glutinous, sometimes also glandular, capsule
not seen.
Sin. Dept. : Steep grassy slope, alt. about 12,000 ft. Rio Blanco,
Macbride & Feather stone 662.
Close to C. reichlinii and C. brachyantha Edwin and C. linearis
R. & P. The type from Dept. Ancash; photograph seen.
Calceolaria callunoides Kranzl. in Fedde, Rep. Sp. Nov. 1:
103. 1905.
Erect, branched shrub, wood glabrous below, white or dark pilose
or puberulent above. Leaves very numerous, aciculiform or linear,
less than 1 cm. long and 1 mm. wide, margin revolute, apex hammate
or plane, slightly wider at base, puberulent beneath, sessile; inflores-
cences of crowded flowers at the apex of stems and branches or axil-
lary, irregularly dichasiate, spicate occasionally (ex. desc.) corym-
bose; pedicels and peduncles puberulent; flowers small, ca. 10 mm.
long; calyces puberulent, 2 -3 (-4) mm. long, narrowly to broadly
ovate, acute or obtuse; upper lip of the corolla longer than the calyx;
lower lip ca. 7 mm. long, saccate about half way, narrowly obovate;
stamens on short filaments, anthers about as wide as long to 1.5
times longer than wide, globose or subglobose, cells equal; style long
for the corolla, ca. 2.5-3.5 mm. long, exserted; capsule shorter than to
equaling the calyx, globose to broadly ovoid, glabrous or pubescent.
FLORA OF PERU 569
Amazonas: Grassy open, Cerro Puma-urcu, alt. 3,250-3,400 m.,
Pennell 15688.— Libertad: Stony soil, above Motil, alt. 3,200 m.,
Pennell & Ferreyra 1^882.
This species and C. alternifolia Cav., C. brachyantha Edwin, and
C. hypericifolia Edwin form a close complex.
Calceolaria calycina Benth. in DC., Prodr. 10: 211. 1846.
C. kraenzlinii Kranzl. in EngL, Bot. Jahrb. 54: Beibl. 119, 19. 1916.
Tall, erect, usually glutinous-hispid, perennial herbs. Leaves,
except the upper, bizarre; petioles broadly winged, connate-perfoli-
ate, usually shorter than the blades, varying from narrower than to
almost as wide as the blades, widest at the stem, entire or with a few
small, rounded teeth; blades deltoid-triangular to broadly ovate,
very variable in size, up to over 10 cm. long, irregularly doubly cre-
nate, acutish at apex, rugose-hispid above, usually canescent be-
neath; flowers forming large, ebracteolate panicles, often big for the
genus, the yellow calyx lobes reaching 30 mm. long and the bright
yellow corolla up to 55 mm. long; calyx lobes broadly ovate, most
often long-acuminate, occasionally acute, mostly 17-25 mm. long;
upper lip of the corolla a little shorter than the calyx lobes, lower lip
little less than to more than twice as long as the upper, orifice usually
less than half the length of the sac; filaments thickened, shorter than
the 5-7 mm. long, apically sigmoid style, anthers divaricate, thecae
1.3 to 1.8 times longer than wide; ovary densely beset with long,
tangled hair; capsule much smaller than the loosely enclosing calyx
lobes, almost spherical to very broadly ovoid, densely pubescent.
Amazonas: Cerro Puma-urcu Southeast of Chachapoyas, alt.
2,800-3,200 m., Pennell 15535; Alt. 2,900-3,500 m., Wurdock 788.—
Junin: Carpapata above Huacapistana, edge of forest, alt. 2,700-
3,200 m., Killip & Smith 2U33.— Libertad: Prov. Otuzco, El Gra-
nero (Hda. Llaguen), alt. 2,900 m., Lopez 599.
Calceolaria cerasifolia Benth. in DC., Prodr. 10: 218. 1846.
Erect, branched, glutinous, often viscose shrub, branches fre-
quently 4-angled, especially above, young wood sometimes sparsely
brown-pilose, more often glabrous, tufts of white or brown hairs fre-
quently present at the younger nodes. Leaves petiolate, mostly
lance-ovate, occasionally lanceolate or broadly ovate, 2-3 cm. long
and 1 cm. wide or up to 10-12 cm. long and 4 cm. wide, serrulate to
serrate, glabrous, subcoriaceous, acute at the apex, obtuse at base,
dark brown-green above, much paler beneath; petioles 1-2 cm. long;
570 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
inflorescences paniculate or corymbose, ultimate pedicels ca. 1 cm.
long, shining; bracts and bracteoles minute; calyx lobes ovate, 5-
7 mm. long, acute, externally glabrous, internally with a distinct row
of oriented hairs along the margins, otherwise glabrous or puberulent;
corolla yellow, upper lip longer than the calyx, lower lip usually less
than twice the length of the upper lip, occasionally reaching twice as
long, saccate about half way, entire corolla up to 22 mm. long, some-
times externally puberulent, usually with a tuft of hairs internally
at the base; stamens short, anther cells equal, divaricate, a little
longer than wide, dehiscing through the septum; style thin, ca. 1.5-
2.5 mm. long; ovary and capsule usually glabrous, sometimes both
puberulent, capsule broadly ovoid, acute, most often a little shorter
than the calyx.
Amazonas: N. of Chachapoyas, along stream, alt. 2,350-2,400 m.,
Pennell 15501; Southeast of Chachapoyas, Cerro Puma-urcu, alt.
2,400-2,900 m., Pennell 15503; alt. 3,000-3,200 m., Pennell 15548;
Ridge, SE. of Calcomar, alt. 2,100-2,200 m., Pennell 15596; Banks,
alt. 2,100 m.; Pennell 15599; roadsides, alt. 2,400 m., Pennell 15605;
Rocky, mountain slopes, alt. 2,700-2,800 m., above Calcomar, Pen-
nell 15614; Thickets on mountains, alt. 3,200-3,300 m., Pennell
15628; Grassy jalca, alt. 3,500 m., Pennell 15651; On rocks, Cerro
Puma-urcu, alt. 3,400 m., Pennell 15713; Lower slopes of Puma-
urcu, abundant, 2,400-2,500 m. alt., Wurdack 429. — Anchash : Yanar,
weed in cult, field and on roadsides, etc., Carricker s.n.; Bushy
banks, N. of Chancos, alt. 3,250 m., Pennell 15326; Bushy bank,
mountain, E. of Copa Grande, N. of Chancos, alt. 3,300-3,500 m.,
Pennell 15366; Ravine on jalca, alt. 3,500-3,550 m., Cerro San Chris-
tobal, N. of Huaraz, Pennell 15453; Puerto Domingo, alt. 4,200 m.,
Rauh-Hirsch P2047. — Cajamarca: Vicinity of Cajamarca, alt. 2,800-
2,850 m., Ferreyra 3271 ; Along stream, E. of Cordillera de Cumulloa,
alt. 3,400 m., Pennell 15163; Mountain E. of Celendin, alt. 2,700-
2,900 m., Pennell 15192; 15208; Hills midway between Cerro de
Cumulloa and Celendin, alt. 2,700-3,000 m., Pennell 15222; Below
spring, Paso de Gavilan, alt. 3,200 m., S. of Cajamarca, Pennell &
Anderson 15089; 2,850 m. alt., Pennell & Anderson 15299; 3,000-
3,300 m. alt., Pennell & Ferreyra 14945; Prov. Contumaza, .Tuque",
Trinidad, alt. 3,230 m., Sagastegui 3841; Roadside, Hualgayoc to
Bombamarca, alt. 2,650 to 2,625 m., Scolnik 1350; Prov. Chota,
Huambos, alt. ca. 2,000 m., Soukup 4558. — Libertad: Prov. Otuzco,
between Motil and Shorey, km. 105, alt. 3,500 m., Lopez 1131 ; 2367;
FLORA OF PERU 571
Yanac, ca. 15 km. SE. of Huamacucho, alt. 3,080 m., Straw 2510. —
Piura: El Tambo, alt. 9,300 ft., Carricker s.n. ; Banks at edge of forest,
alt. 1,700-1,900 m., above Chanqui, Pennell & Ferreyra 14-914.; Aya-
baca, alt. 2,743 m., Soukup 1+353; no location in Piura, Weber-
bauer 6039.
Calceolaria chaetostemon Pennell, Proc. Acad. Nat. Sci. Phila.
97: 171, /.i 5o. 1945.
Herbaceous, probably perennial, 3 dm. tall or taller, stem hirsute,
internodes longer than the leaves. Leaves petiolate, ovate, mostly
5-9 cm. long and 3.5-8.0 cm. wide, pubescent on both surfaces, mar-
gins dentate, usually acute at apex, base subtruncate-obtuse to cor-
date; petioles about 1 cm. long, lax-pubescent; inflorescences typi-
cally a pair of twice branched, ebracteate cymes, sometimes one cyme
suppressed; flower stalks minutely glandular, multicellular-pubes-
cent, the glands apically placed, minute; basal peduncle 6-9 cm.
long or longer; secondary and ultimate stalks (pedicels) abruptly
much shorter; calyx lobes lance-ovate, 2-3 mm. long, subobtuse, ex-
ternally hirsute, hairs gland-tipped, multicellular, internally gla-
brous; corolla yellow, upper lip ca. 3 mm. long, externally pubescent
as the sepals, lower lip 7-9 mm. long, saccate more than half its
length, lobes crenate, nectary downturned, narrowed, pubescent with
minute gland-tipped hairs; lower anther cell subobsolete or lacking,
connective as long as or longer than the upper cell, thickened and
auriculate above the insertion of the filament; style 2.0-2.5 mm. long,
strongly curving inward, basally puberulent; the young capsule
glandular-pubescent .
Apurimac: Abancay Prov., Curahuasi, alt. 2,800 m., Marin 1953.
— Cuzco: Anta, alt. 2,700 m., Vargas 231 (type F) ; Ayusbamba, alt.
3,000 m., Vargas 852; Limatambo, alt. 3,400 m., Vargas 7079.—
Huanco: Hills N. of Acomayo, alt. 2,400 m., Woytkowski 257.
Calceolaria chrysocalyx Pennell, Proc. Acad. Nat. Sci. Phila.
97: 166. 1945.
Perennial herb, erect and glandless, stems branched, distally pi-
lose and becoming glabrous and subshining, internodes longer than
the leaves, drying light brown. Leaves ovate, 4-10 cm. long and
3-8 cm. wide, dentate, teeth rounded, apically acute, basally rounded-
truncate to slightly cordate, dark green and pilose above, tomentose
beneath, rarely tomentum wanting; petioles 5 mm. long, pilose or
glabrate, tumid, woody at base and narrowly connate at the node;
inflorescences in terminal groups of 2 pairs of 3-compound cymes;
572 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
basal peduncles 7-15 cm. long, flower-stalks decreasing apically to
1 cm. long, hirsute; primary bractlets ovate-lanceolate, dentate, acu-
minate, mostly 2-5 cm. long; sepals ovate, 11-15 mm. long, acute,
reticulate veined, externally pilose, bright yellow; corolla yellow,
upper lip ca. 5-6 mm. long, minutely pilose externally, lower lip
18-23 mm. long, sac globular, less than half the length of the lip,
glabrous internally and externally; anther cells equal, gray-brown,
dehiscing through the septum, subparallel to somewhat divergent, ca.
1.5 times longer than wide; style ca. 2.0-2.5 mm. long, little if at all
thickened; capsule ovoid-conic, 5-8 mm. long, pilose, hair short,
spreading.
Apurimac: Prov. Abancay, Ampai, alt. 3,000 m., Vargas 775;
Prov. Grau, Heights of Oropega Valley, alt. 3,400-3,800 m., Vargas
9810—Cuzco: Paucartambo, 11,500 ft., Balls 6724; Prov. Uru-
bamba, Cerro Machu-picchu, alt. 3,200 m., Mexia 8077; Near Rio
Yanamayo, below Pillahuata, alt. 2,900-3,200 m., Pennell 13792,
type coll; Macchu-picchu, near Puno, alt. 4,000 m., Soukup 144',
Prov. Convencion, Macchu-picchu, alt. 2,400 m., Vargas 797; Prov.
Paramo, near Hda. Ayusbamba, alt. 3,300-3,860 m., Vargas 857;
Macchu-picchu, alt. 2,100 m., Vargas 2090; Prov. Calca, below Lares,
alt. 2,900-3,000 m., Weberbauer 7899; Cultivated in U. of Calif. Bot.
Gard. from a collection from Macchu-picchu, alt. 2,400 m., West
6428.— Sin. Dept.: Valle de San Miguel, alt. 2,060-2,260 m., Her-
rera 1998.
This species is close to C. bicrenata R. & P. which differs in having
smaller calyces, light yellow-green or darker glandular-pubescent
flower stalks and in deeper doubly-toothed leaf-margins.
Calceolaria chrysosphaera Pennell, Proc. Acad. Nat. Sci.
Phila. 97: 156. 1945.
Perennial herb, hirtous-pubescent with spreading, short, brown,
glandless hairs, densest on young growth, stems branched, pilose
beneath the new growth. Leaves lanceolate, the larger 6-11 cm. long
and 1.0-2.5 cm. wide, sometimes shorter and less than 1.0 cm. wide,
obscurely crenate-dentate to subentire, pilose and dark green above,
paler and pubescent on the veins beneath, cuneate at base; petiole
less than 1 cm. long, narrowly connate at the node, little if at all
thickened or woody; inflorescences few-flowered; basal peduncles 8-
9 cm. long; pedicels ca. 2 cm. long; primary bracts lanceolate, as long
as or longer than the pedicels; sepals ovate, acuminate, 10-12 mm.
FLORA OF PERU 573
long, few-dentate, finely pubescent; corolla yellow, externally gla-
brous, upper lip ca. 2 mm. long, lower lip ca. 15 mm. long, hirsute
within at the base, sac globose, ca. half the length of the lip; anther
cells equal, each pair ca. 3 mm. long and two-thirds as wide, divari-
cate, dehiscing through the septum; style 2 mm. long; capsule and
ovary (immature) globose, glandular-pilose. Hair sometimes septate
on new wood.
Cuzco: Upper valley of Rio Sembray, W. affluent of Vilcamota,
alt. ca. 1,700 m., Mexia 8058; near Rio Yanamayo, below "Pilla-
huata," bank in forest, alt. 2,000-2,300 m., Pennett 14045, type
collection.
Calceolaria colcamarensis Edwin, Phytologia 19: 374. 1970.
Erect, appressed-pubescent herb, stems simple or few-branched,
branches, when present, frequently abbreviated in leaf-axils, nodes
mostly much longer than the leaves, pubescence soft, white or light-
gray, densest on younger growth. Leaves short-petiolate, ovate or
less often narrowly ovate, all but the largest at least twice as long as
wide, (1. 5-) 2.0-4. 8 cm. long and 0.6-2.0 (-3.0) cm. wide, margins
serrate-serrulate, apex broadly acute, base cuneate, densely ap-
pressed-short-pubescent and red-punctate above, appressed-pubes-
cent on the elevated, almost white, thickened veins and densely
red-punctate beneath; petioles 2-10 mm. long, sparsely to densely
pubescent, thickened, discrete; inflorescences terminal or in upper
axils, dichasiate-corymbose, few- (ca. 3-5-) to many-flowered (ca.
15-20) ; basal peduncles up to 3 cm. long, flower-stalks decreasing to
ultimate pedicels a few mm. long, all stalks densely appressed-
pubescent; calyx lobes when young with pubescence and venation
as the lower surface of the leaves, veins with age less apparent and
vesture less dense, mostly ovate, acute, 7-10 mm. long, internally
glabrous; corolla yellow, wholly glabrous, 13-17 mm. long; upper lip
2^4 mm. long; lower lip distally almost round, saccate less than half-
way, orifice oblique; stamens small, filaments short, anther cells
equal, divaricate, a little less to a little more than twice as long as
wide, each cell ca. 1.2-1.4 mm. long, red-brown when dry; style 1.5-
2.0 mm. long, glabrous, ovary resinous, dark red gland-dotted; cap-
sule broadly ovoid, abruptly long-acute, 5-6 mm. long and wide,
red-gland-dotted .
Amazonas: Roadsides on dry loam, alt. 2,300-2,800 m., Colca-
mar, Pennell 15604-
574 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
This species differs from all other Calceolarias by the combination
of the following characters: flattened, elevated, white venation on the
leaf blades beneath, dense-red-gland-dotted fruit, calyx and blade,
sepals often about half as long as the lower lip of the corolla and ap-
pressed pubescence on the stems and branches.
Calceolaria collanensis Edwin, Phytologia 19: 375. 1970.
Erect, much-branched shrub, the wood when young densely
white-puberulent-pubescent, becoming glabrous with age, branches
and branchlets twisted, drying brown to dark brown or black, inter-
nodes much longer than the leaves, sometimes 5-7 times as long.
Leaves petiolate, ovate to very broadly ovate, 0.6-1.5 cm. long and
4-10 mm. wide, margins narrowly revolute, thickened, with 3-7 pairs
of obtuse lobes or teeth, pubescent-puberulent on both surfaces,
hairs curly beneath, apically acutish, basally broadly obtuse to sub-
truncate; petioles short, up to 5 mm. long, pubescent-puberulent,
thickened, swollen and woody at base, the petioles of the main stems
and branches with abbreviated branchlets in the axils; inflorescences
simple or branched dichasia, borne in upper leaf-axils on stems and
branches, occasionally the dichasium suppressed and axils 1-flowered;
flower-stalks short, pubescent-puberulent; bractlets when present,
minute; calyx lobes elliptically ovate or elliptical, 5-6 mm. long and
1.5-2.5 mm. wide, pubescent on both surfaces; corolla 11-12 mm.
long, drying dark brown (orange-red in life?), lower lip ca. 2 mm.
long, upper lip saccate distally only 1-2 mm., sac undulate, nectary
only partly reflexed-hidden, almost as long as the sac, entire lip gla-
brous or sparsely papillose-puberulent; filaments 1.0-1.5 mm. long,
anther cells equal, divaricate, little longer than wide; style ca. 1.0-
1.5 mm. long, ovary red-shining-glandular; capsule exceeding the
calyx, 7-8 mm. longer, narrowly ovate to ovate, upper half narrowly
elongate, beaklike, red-shining-glandular, glands short-stalked.
Known only from the holotype.
Ancash: Western slope of Cordillera Negra, below Paso de Col-
Ian, bank along stream, alt. 3,500 m., Pennell 15470.
This taxon is quite distinct from all other taxa due to the com-
bination of open corolla with undulate sac and prominent nectary as
well as the swollen petioles with abbreviated axillary shoots and
finally the long internodes.
FLORA OF PERU 575
Calceolaria connatifolia Pennell, Not. Nat. 256. 1951.
Lax scrambling herb up to 1 m. tall, glabrous or subglabrous ex-
cept for the inflorescence. Stems drying purple or brown; leaves
connate-perfoliate, tapering to apices, oblong-lanceolate, 5-8 cm.
long and 2.0-3.5 cm. wide, serrate, dark green above, pubescent on
the sunken mid-vein, paler beneath with sharply marked reticula-
tions; flowers in simple or compound cymes, in axils of upper leaves;
stalks of the inflorescences short-pilose to subglabrous; basal pedun-
cles up to 4.0 cm. long, the intermediate peduncles shorter; the ped-
icels shortest, 1.5-2.0 cm. long; sepals 4-5 mm. long, broadly ovate,
mucronulate, glutinous and sometimes externally pilose, internally
with a dense row of short hairs near the margin; corolla lemon-yellow,
globose, upper lip 2-3 mm. long, projecting over the anthers, lower
lip up to 8-9 mm. long, saccate about one-third its length, basally
hirsute within; anthers pale brown, cells rounded, dehiscing through
the septum; style much exceeding the stamens, ca. 4.0 mm. long,
sigmoid at apex, ovary glabrous, glutinous; capsules very broadly
ovoid or deltoid-ovoid, narrowed to apex, subglutinous, much smaller
and included in the loosely investing sepals.
Amazonas: Cerro Puma-urco, near summit, seepage forest, in
thicket, alt. 3,300-3,400 m., Pennell 15906; alt. 2,700-3,100 m.,
Wurdack 1631. — Cajamarca: Carricker s.n.
Calceolaria cordiformis Edwin, Phytologia 19: 376. 1970.
Climbing perennial herb or subshrub, stem branched, when young
densely tomentose-lanate, becoming almost glabrous with age, nodes
longer or shorter than the leaf blades. Leaves opposite or verticil-
late, broadly ovate-cordiform, mostly up to 1.5 cm. long, rarely up
to 2.5 cm. and up to 9 mm. wide, occasionally up to 2 cm. wide,
tomentose above, very densely tomentose-lanate beneath, the pubes-
cence completely obscuring the tissue, margins revolute, dentate,
teeth large, round, apex subacute or acute, bases cordate, occasion-
ally truncate or broadly obtuse, venation impressed above, sessile or
subsessile; petioles when present up to 4 mm. long, very densely
lanate; inflorescences dichasiate, dense, terminal or in upper leaf-
axils; flower-stalks not exceeding 2 cm. long, often less than 1 cm.
long, densely lanate; calyx externally lanate, internally glabrous or
sparsely puberulent at the apex, especially when young, ovate, ca.
5 mm. long and 4 mm. wide near the base, entire, acute; corolla
yellow, wholly glabrous, 7-9 mm. long, lower lip 1-2 cm. long, upper
lip saccate less than half its length, ca. 3-4 mm. wide; anther cells
576 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
equal, each pair divaricate, ca. 1.5-2.2 mm. long, about twice as long
as wide, filaments ca. 2 mm. long; ovary glandular, style short, ca.
1-2 mm. long; capsule glandular, ovoid-conic, acute, shorter than
and loosely invested by the calyx lobes.
This taxon is keyed as leaves more or less than 1 cm. wide, since
the largest leaves indicate the strong possibility that future collec-
tions may well have all or most leaf blades wider than 1 cm.
Cajamarca: Along streamlet, above Llama, alt. 2,450 m., Pennell
15924; Prov. Chota, Huambos, alt. ca. 2,000 m., Soukup 4549.—
Piura: Road from Huancabamba to the three acequias, alt. ca.
2,500 m., Scolnik 1422.
This taxon is closest to C. hedera, C. aperta, and C. velutinoides
Edwin, but quite readily separable from these taxa.
Calceolaria corymbosa R. & P., Fl. Peruv. 1: 14, t. 20b. 1798.
Herb, nodding and scapose to erect and caulescent, the stems pu-
bescent, usually unbranched and few-leaved, up to 60 cm. tall, foliage
and inflorescences usually variously pubescent; internodes gibbose or
not. Leaves mostly radicle, long- to short-petiolate, the stem leaves
when present petiolate to sessile or almost so, ovate to elliptically
ovate, 4-15 cm. long and 2-8 cm. wide, usually irregularly crenate to
obtusely dentate, rarely teeth reduced, apex obtuse, base sometimes
attenuate to acute, usually obtuse, subglabrous to densely pubescent ;
inflorescences usually few-flowered, regularly, bifurcately branching
or corymbose, either terminating a scape or in axils of upper leaves ;
uppermost leaves bracteate; bracteoles lacking; flower-stalks usually
pubescent, from much longer to shorter than the flowers, the pedicels
shortest; calyx lobes deltoid-ovate or broadly ovate, acute or short-
acuminate, usually densely pubescent and soft-ciliate, pubescence
occasionally sparse, dark red-green externally, light yelow-green in-
ternally, 7-9 mm. long; calyx tube 1-2 mm. long, pubescent; corolla
10-18 (-22) mm. long, lemon-yellow, marked with brown lines, gla-
brous or sometimes viscid-puberuelnt, upper lip much shorter than
the calyx lobes, with a tuft of long, white pubescence internally at
base, lower lip 4 to 5 times longer than the upper, about two-thirds as
wide as long, distally strongly upcurved, the upper fold one-half to
almost completely closing the orifice; filaments short, dilated, anthers
divaricate, about twice as long as wide; style usually shorter but
varying to longer than the filaments, ovary glabrous or glandular-
pubescent.
FLORA OF PERU 577
The northern collections usually more densely pubescent than the
southern. The species as now constituted shows a disjunct distri-
bution.
Plants scapose C. corymbosa var. corymbosa.
Plants caulescent C. corymbosa var. perennis.
Calceolaria corymbosa R. & P. var. corymbosa 19: 377. 1970.
C. paralia Hook., Exot. Fl. t. 75. 1823, non Cav. C. hians Penny
ex Don and C. hopeana Morris ex Don, Gen. Syst. 4: 603 and 606.
1838. C. elliptica Wedd., Chi. And. Nov. 2: 137. 1860. C. heterophyl-
ioides Kranzl. in Fedde, Rep. Sp. Nov. 1: 100. 1905.
Cajamarca: East of Celendin, alt. 2,700-2,900 m., Pennell 15209;
Paso de Gavilan South of Cajamarca, alt. 3,200 m., Pennell & An-
derson 15094 and Pennell & Ferreyra 14954; Michiquillo, alt. 3,400-
3,500 m., Pennell & Reichlin 15036. — Sin. Loc.: Dombey s.n. — Sin.
Depto. : Huasohuasi, Weddell s.n. and reported by Kranzlin, Tarma,
Junin : Weberbauer 2423.
Calceolaria corymbosa var. perennis Edwin, Phytologia 19:
377. 1970.
Stems leafy, erect and elongate; inflorescences in axils of upper
leaves.
Lambayeque: Hacienda Taulis, vicinity of the Casa Hacienda,
Prov. Chiclayo, alt. 1,900 m., Hutchison & von Bismarck 6339.—
Libertad: Samne, Prov. Otusco, along Rio Moche, below junction,
cliff, alt. 2,500 m., Pennell & Ferreyra 14887; Road banks, Samne,
alt. 1,900 m., Hutchison & Wright 6130 (type).
Calceolaria crenata Lam., Encycl. 1: 556. 1783. C. amplexi-
caulis HBK., Nov. Gen. et Sp. 2: 384. 1817. C. floribunda HBK.,
Nov. Gen. et Sp. 2: 385. 1817.
Erect subshrub, stems little to much branched, fistulose, pilose or
glandular-pilose. Leaf blades sessile, lance-oblong to narrowly lance-
ovate, up to 12 cm. long and 2.0-3.5 cm. wide, hirtellus or pilose
above, hirsute or glandular-pilose beneath, crenate- or dentate-ser-
rate, ovate or cordate at base, amplexicaul or almost so, often gluti-
inous; flowers in corymbs or umbellate corymbs; peduncles glandular-
pilose, up to 12 cm. long; calyx lobes externally and often internally
glandular-pilose, ovate, 2-3 mm. long and 2.0-2.5 mm. wide, veins
often obvious ; corolla yellow, glabrous, upper lip a little longer than
578 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
the calyx, lower lip up to 2 cm. long and 1.5 cm. wide, saccate less
than half its length, the orifice closed by the upturning of the sac to
the upper lip; stamens very short, anther locules equal, less than
twice as long as wide. (Description after Kranzlin.)
Only photographs and fragments seen. This taxon is included
provisionally. The material attributed by Kranzlin to Peru may
pertain rather to Ecuador.
Calceolaria crenulata Pennell, Proc. Acad. Nat. Sci. Phila. 97:
169, pi 14 C. 1945.
Erect, lax, branched herb, stems appressed-hirsute, with multi-
cellular, brown-jointed hairs. Leaves oblong-ovate, blades 5-8 cm.
long and 2.5-3.0 cm. wide, crenulate, glandular-puberulent above,
hirsute with brown, appressed hairs, on mid-rib and main veins be-
neath, apex acute, rounded to base; petioles frequently more than
1 cm. long, thickened, densely hirsute with appressed, brown hairs;
inflorescences of a few pairs of 1-compound cymes; basal peduncles
4-5 cm. long; pedicels decreasing apically, all stalks hirsute; primary
bractlets small, leaf -like; sepals ca. 3 mm. long, lanceolate, obtuse,
entire, densely, minutely hirsute on both sides; corolla yellow, exter-
nally puberulent, internally hirsute in a circle around the orifice,
upper lip ca. 5 mm. long, slightly wider than the base of the 30 mm.
long lower lip that is upturned and forms a sac oneh-alf to two-thirds
its length; anthers brown, rounded, the almost globose cells dehiscing
through the septum; style 2-3 mm. long, glandular-puberulent bas-
ally. Known only from the type collection. (After original de-
scription.)
Cuzco: Prov. Paucartambo, E. slope of Cord, de Caravaya along
Rio Yanamayo, rocky stream-bank in forest, below "Pillahuata"
SE. of Paso de Tre"s Cruces, alt. 2,100-2,300 m., Pennell 14036.
Calceolaria croceopunctata Edwin, Phytologia 19: 378. 1970.
Shrub, erect, branched, internodes distant, up to ca. 13 cm. long,
about 4-6 times longer than the leaves near the base to only 2-3
times longer than the leaves below the inflorescences, current wood
sparsely pubescent with recumbent white hairs, soon glabrate, bark
drying smooth. Leaf blades (very) short-petiolate, ovate, mostly
2-3 cm. long and 1.0-1.3 cm. wide, crenate, acute at apex, obtuse to
subtruncate at base, rugulose, sparsely puberulent above, densely
tomentose-lanose on the veins and very densely punctate with sessile,
shining, orange punctations beneath; petiole densely lanate, up to
FLORA OF PERU 579
3 mm. long, reduced leaves often present in the axils; inflorescences
paired, many-flowered, dichasiate-panicles, in the upper leaf axils of
stems and branches; peduncles 2-3 cm. long; pedicels 1-2 cm. long,
all flower-stalks densely lanate; flowers 13-17 mm. long; calyx lobes
dimorphic on each flower, lance-elliptic or ovate, 7 mm. long and
3 mm. wide or 6-7 mm. long and 5 mm. wide, pubescent-puberulent
on both surfaces; corolla yellow, minutely papillose, a tuft of white
hair within at the base, upper lip 4-6 mm. long, lower lip 9-12 mm.
long, saccate less than half its length, sac only little wider than prox-
imal part of lip; filaments 1.0-1.5 mm. long, anther cells equal, sagit-
tate, each pair 3.0-3.4 mm. long, each cell ca. 1.2 mm. wide, an acu-
min developed at the apex of the septum; style ca. 6 mm. long, very
long for the flower, exceeding the upper lip of the corolla; ovary
glandular-pubescent, glands stalked, red; capsule broadly deltoid-
ovoid, acute vesture as the ovary, much shorter than the calyx, 3.5
mm. long and about as wide at the base.
Known only from the type.
Chachapoyas: Cerro Tinaja, alt. 3,200 m., Ochoa 1686 (US).
Close to C. rugulosa Edwin, from which it differs in leaf -size, ves-
ture (orange punctations beneath), style length, and anther size and
shape.
Calceolaria cuneiformis R. & P., Fl. Peruv. 1: 19, t. 27, fig. b.
1798. C. rotundifolia HBK., Nov. Gen. et Sp. 2: 382. 1817. C. foli-
osa Meyen. ex. Walp. et Schau., in Nov. Acta. Acad. Nat. Cur. 19:
Suppl. 1. 295. 1843.
Shrubby, erect and branched, pubescence on wood and foliage
variable, occasionally almost glabrous, wood usually hirsute or pilose.
Leaves variable in shape, ovate to more often triangular-ovate, occa-
sionally very broadly ovate, not infrequently triangular, rarely sub-
rotund, 0.4-2.0(-3.2) cm. long and 0.2-1.0(-1.5) cm. wide, mostly
0.7-1.5 cm. long and 0.5-0.9 cm. wide, margin crenate or dentate or
few-lobed, rarely many-lobed, cuneate at base, obtuse to subacute
apically, hirsute above, pilose beneath, rarely vesture wanting, often
obscuring the tissue beneath, at least in part; inflorescences terminal,
axillary, irregularly branched, few- to usually many-flowered ; flower
stalks up to 2 cm. long, glandular-pilose; bracts minute, scattered;
calyx segments shining, ovate, ca. 8 mm. long and 5 mm. wide near
the base, acute, densely glandular-pilose; corolla yellow, ca. 1.5 cm.
long and distally 1 cm. wide; upper lip shorter than the calyx, lower
580 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
lip saccate less than half its length; filaments short, anther cells con-
tiguous, almost twice as long as wide to only little longer than wide;
style 2-3 mm. long; ovary and capsule glandular-pilose.
The distribution of this taxon is quite broad. It has been col-
lected in concentration in Dept. Lima and then scattered collections
to almost the southernmost department (Moquegua).
Apurimac: Prov. Abancay, Curahuasi, alt. 2,800 m., Marin 1926;
Prov. Condehuaylos, Moyobamba, alt. 3,650 m., Vargas 8730', 2 km.
S. of Chincheras, shrubland, alt. 2,800 m., Stork & Horton 10734.—
Arequipa: Canyon above Arequipa, alt. 2,700-3,300 m., Pennell
14271 ; Lower slope of Chachami 3,170 m., Hutchison & Wright 7227;
9,000 ft. alt., Sandeman 3809. — Huancavelica, Prov. Castro virreina,
near Cordova, alt. 3,050-3,300 m., Ferreyra 6976; Valley of Rio
Rimac, near Lima-Aroya, 70 km. E. of Lima, alt. 1,850 m., Goodspeed
33122; Rio Blanco, alt. 3,000-3,500 m., open hillside, Killip & Smith
21584; 21585; 21703; 21736; Matucana, rocky hilltop, alt. ca. 8,000
ft., Macbride & Featherstone 682; Huaros, alt. 3,400-3,500 m., Pen-
nell 14713; Rocky bank above Rio Blanco, alt. 3,500-3,700 m.,
Pennell & Maldonado 15939. — Moquequa: Carumas, rainy-green
formation, alt. 3,200 m., Weberbauer 7292. — Puno: Prov. Puno, 10
km. from Chucuito, on road to Puno, alt. ca. 4,000 m., Metcalf 30690.
The wide altitudinal range, from 1,850 to 4,000 m., as well as its
latitudinal dispersion (see above) may account for the morphological
variability expressed by this species.
Calceolaria cypripediiflora Kranzl. in Fedde, Rep. Spec. Nov.
1: 101. 1905. C. furcata Pennell, Proc. Acad. Nat. Sci. Phila. 97:
152. 1945. C. drepanantha Pennell, I.e. 97: 154. 1945.
Herb, erect and most often pubescent, hair occasionally white to
usually brownish, or doubly pubescent with long brown and short
white hairs, mostly much branched. Leaves long-petioled, ovate or
oblong-ovate, occasionally elliptic-ovate, rarely subglabrous, usually
hirsute or pilose, sometimes densely so beneath, the vesture obscuring
the tissue; blades 2.5-7.5 cm. long and 2.5-5.0 cm. wide, margins
grossly crenate-dentate or doubly dentate, apex acute to obtuse, base
rounded to cordate, tissue most often paler beneath; petiole mostly
0.8-2.5 cm. long; inflorescences branched, often corymbose, few- to
several-flowered; flower-stalks with vesture as the stems or denser;
flowers 30-45 mm. long; calyx lobes equal or unequal, often reflexed
at anthesis, 5-10 mm. long, lanceolate to ovate, acute, entire, exter-
nally hirsute or pilose, internally glabrous; corolla yellow, externally
FLORA OF PERU 581
pubescent, upper lip 4-10 mm. long, lower lip 25-40 mm. long, sac-
cate up to half its length and the sac often canaliculate, the non-
saccate, proximal part very narrow, ca. 3-4 mm. wide; filaments
short, anther cells equal, 1.5 times or more longer than wide, each
pair of cells 3-4 mm. long; style 4-6 mm. long, little thickened, api-
cally incurved, ovary densely glandular-pubescent; capsule little
longer to little shorter than the calyx, vesture as the ovary.
Ayacucho: Moist rocks near a brook, Choimacota Valley, Prov.
Huanta, alt. 3,100-3,200 m., Weberbauer 7577.— Cuzco: Macchu-
picchu, alt. 2,100 m., Cook & Gilbert 847; alt. 2,000 m., Rauh-Hirsch
P855; Macchu-picchu, Prov. Convencion, alt. 2,400 m., Vargas 8^0,
type of C.furcata Penn. ; Puyupatamarca, Prov. Urubamba, on rocks,
alt. 3,400 m., Vargas 2779; Shrub on open rocky places, Lares Valley,
Huallhuayoj, Prov. Calca, alt. 2,200-2,300 m., Weberbauer 7901, the
type of C. drepanantha Pennell. — Puno: Near Limbani, Prov. Sandia,
wet mountainside, alt. 3,200-3,450 m., Metcalf 305^1.
Calceolaria decus-montium Kranzl. in Engler., Bot. Jahrb.
54: Beibl. 119. 20. 1917.
Small, caespitose, densely lanate, little-branched shrub, sterile
shoots not exceeding 5 cm. tall, fertile shoots up to 18 cm., wood,
foliage and inflorescences densely beset with long, soft, white hairs.
Leaves lanceolate, with the hair silvery, the largest 8 mm. long and
3 mm. wide; flowers apical in 1 or 2 pairs on the fertile stems (due to
the abortion of 1 of the flowers of a simple dichasium) ; pedicels up
to 2 cm. long, usually shorter; calyx lobes ovate-lanceolate, 8 mm.
long and 3 mm. wide, acute, internally glabrous except for the pilose
margins; corolla yellow, up to 27 mm. long, usually at least 23 mm.
long and 2 cm. wide, densely white villose internally at base, upper
lip ca. 5 mm. long, cap-like, short-pilose, lower lip obovate to almost
orbicular, externally glabrous; filaments minute, anthers 2-3 mm.
long, 1.5 times longer than wide, cells divaricate; style 1-2 mm. long,
ovary ovate, short-setose; capsule unknown. (Description after
Kranzlin.)
Known only from the type material.
Piura: Prov. Huancabamba, between Huancabamba and Jicate,
alt. 2,500-2,600 m., Weberbauer 6329 (F).
Calceolaria deflexa R. & P., Fl. Peruv. 1: 18, t. 30, f.b. 1798.
C. fuchasiaefolia Hemsl., The Garden, 258. 1879.
582 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Essentially glabrous, erect or climbing shrub, stems branching.
Leaves either glabrous, lanceolate, lance-ovate or narrowly oblong-
ovate, usually 4-8 cm., long and ca. 1-3 cm. wide, minutely serrate
to entire, mostly acute at both ends, sometimes subobtuse at base,
bicolorous, paler beneath, veins subobscurely reticulate to subobso-
lete, petiolate, or leaves ovate, 2.5-4.0 (-4.5) cm. long and 0.7-
2.0 cm. wide, with veins reticulate beneath ; petioles up to 1 cm. long;
inflorescences few-flowered, not exceeding 8 (-12) -flowered, terminal
or axillary, subumbellate or corymbose or occasionally dichasiate;
flower-stalks minutely puberulent or glabrous; basal peduncles 2-
4 cm. long; calyx glabrous, sepals ovate, acute, 3-4 (-6) mm. long,
2-3 (-5) mm. wide; corolla yellow or orange, upper lip of corolla
about twice as long as the calyx (ca. 0.8-1.0 cm. long), hooded, paler
than the lower lip that is from little longer to almost twice as
long as the upper, occasionally a little more than twice as long
(up to 2.2 cm. long), suborbicular, usually saccate over three-
fourths of its length, glabrous except for a dense tuft of white hairs
at the base internally; filaments very short, often pilulose, thecae
equally divergent, each pair ca. 2.5-3.0 mm. long, less than twice
as long as wide; style 3.5-4.5 mm. long, stigma punctiform, ovary
glabrous; capsule ovoid, acute, glabrous, much longer than the
calyx. Separable into 2 distinct varieties.
Leaves more than 2.5 times longer than wide, lanceolate, lance-ovate
or narrowly oblong-ovate C. deflexa var. deflexa.
Leaves up to 2.2 times longer than wide, usually less, ovate or ellip-
tically ovate C. deflexa var. aurantiaca.
Calceolaria deflexa R. & P. var. deflexa
Ancash: Prov. Bolognesi: Huayalpampa, alt. 3,600 m., Ferreyra
381 ; Cerro Chiquian, alt. 3,360 m., Ferreyra 400', Arriba de Chiquian,
alt. 3,500-3,800 m., Ferreyra 6220; Prov. Huaraz, alt. 3,000 m.,
Nunez 3194.', Copa Grande, north of Chancos, along stream, alt. 3,350
m., Pennell 15362; Along stream Colcas, alt. 2,800-2,900 m., Pen-
nell 14382; Paso de Fortale"za, above stream, alt. 3,350 m., Pennell
15397; Cerro San Cristobal, west of Huaraz, alt. 3,300-3,350 m.,
Pennell 15444; Cajamarca: Along Rio Lulichuco, alt. 2,600-2,700 m.,
above (sw.) Cajabamba, Pennell & Ferreyra 14860. — Libertad: Bom-
bamarca, city of Bombamarca, slope of E. Cordillera, alt. 10,000-
12,000 ft., Carricker s.n.; Huamachuco: wet cliff, alt. 3,250 m., Pen-
nell & Ferreyra 14870. — Lima: near Infiernillo, between Matucana
and Tarma, alt. 3,350-3,400 m., Ferreyra 6255; Open hillside, Rio
FLORA OF PERU 583
Blanco, alt. 3,000-3,300 m., Kittip & Smith 21674; 21676; along Rio
Chillon, above Obrajillo alt. 3,100-3,300 m., Pennell 14408; Cerro
Colorado east of Canta, alt. 3,800-4,000 m., Pennell 14659; Moist
stream bank, above Rio Blanco, alt. 3,500-3,700 m., Pennell & Mol-
denado 15932; Alt. 3,500-3,600 m., Pennell & Moldenado 15941 —
Sin. Dept.: Rio Blanco (Lima?), alt. ca. 12,000 ft., Macbride & Feath-
er stone 713; In Rubus thicket below Chavanillo, Rio Maranon, alt.
7,500 ft., Macbride & Featherstone 1983; Obragillo, Nee s.n.; Ceuchin
or Cheuchin, Ruiz and Pavon s.n. — Sin. Loc.: Dombey s.n., Ruiz s.n.
Calceolaria deflexa R. & P. var. aurantiaca Edwin, Phytologia
19: 378. 1970.
Climbing shrub with orange corolla, differing from the typical
variety in having ovate leaves usually less than twice as long as
wide, rarely reaching twice as long as wide.
Ancash: Western slope of Cordillera Negra below Paso de Collan.
Rocky slopes and banks, alt. 3,650 m., Pennell 15478, type collection.
Calceolaria delicatula Kranzl. in Fedde, Rep. Sp. Nov. 1: 102.
1905. C. conocarpa Pennell, Proc. Acad. Nat. Sci. Phila. 97: 172,
fig. 15b. 1945.
Erect or lax, branched herb with glandless or gland-tipped hair,
internodes usually longer than the leaves, stems and inflorescences
soft-pubescent. Leaves petiolate, ovate in outline, very variable in
size with most either pinnatifid proximally with 2 pairs of pinnae cut
almost to the midrib and distally with further pinnae that unite into
a wide mid-blade, the distal segments irregularly doubly, sharply
dentate, pilose above and beneath or blades irregularly dentate-
laciniate, lobed near the base, occasionally lowest blades basically
dentate with 2 pairs of denticulate lobes, 1.3-7.0 cm. long and up to
5 cm. wide, acute apically; petioles 1.0-2.5 cm. long; inflorescences
glandular-hirsute, simple to 3-4 times compound cymes, in terminal
pairs or from the upper leaf-axils, rarely flowers solitary, basal pe-
duncles 4-10 cm. long, remaining stalks proportionately shorter;
bracts and bracteoles reduced, leaf -like, pinnate, smallest bracteoles
lanceolate; sepals ovate or triangular-ovate, 6-8 mm. long, margins
ciliate; corolla yellow, externally glabrous or subglabrous, internally
pubescent at base beneath; upper lip ca. 2.0-2.5 mm. long, lower lip
10-14 mm. long, saccate over most of its length, sac brighter yellow;
filaments pubescent, both anther cells fertile, upper only little shorter
584 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
than the combined length of the connective and lower, smaller cell;
style 1-2 mm. long, slightly distally curved; capsule conic, glandular-
pubescent, longer than the calyx, 7-8 mm. wide at base.
This taxon is distributed from Cajamarca in the north to Puno
in the south with its greatest concentration from Lima to Cuzco.
Citations selected from the copious material.
Ancash: Lomas de Casma, Prov. Santa, alt. 400-500 m., Ferreyra
8054.. — Ayacucho: Pampalca, between Huanta and Rio Apurimac,
alt. ca. 3,200 m., Kittip & Smith 22210; 22257.— Cajamarca: Monte
seco, Prov. Hualgayoc, alt. 1,800 m., Soukup 2835. — Cuzco: Above
Paucartambo, alt. 11,500 ft., Balls 6722; Near Rio Quencomayo, be-
low Colquipata, alt. 3,200-3,300 m., Pennell 13774; Near Rio Yana-
mayo, below "Pillahuata," alt. 2,900-3,200 m., Pennell 13794 (type
collection of C. conocarpa); Macchu-picchu, alt. 2,000 m., Rauh-
Hirsch P856; Hda. Ayusbamba, alt. 3,400 m., Vargas 859; Tio, Mar-
capata, alt. 1,900 m., Vargas 3074; Que"brada de Llulluchayoc, alt.
3,100 m., Vargas 4292; "Pillahuata," Paucartambo, alt. 2,400 m.,
Vargas 4899; Near Asuncion Bridge, Rio Apurimac, alt. 3,730 m.,
Vargas 11028. — Huancavelica: near Cordova, alt. 3,050-3,300 m.,
alt. Metcalf 30289. — Huanuco: Carpish, 60 km. between Huanuco
and Tingo Maria, Swingle 72. — Junin: At bridge, valley bottom,
Canyon of Rio Huasahuasi, below Huasahuasi, alt. 2,400 m., Hutchi-
son 1139; Carpapata, above Huacapistana, alt. 2,700-3,200 m., Killip
& Smith 24368.— Libertad: Trujillo, alt. 500 m., Angulo 2004; Cerro
Campana, Prov. Trujillo, alt. 675 m., Lopez 1688. — Lima: Between
Huaral and Huacho, lomas de Lachay, alt. 400-600 m., Ferreyra
3862; Various locations alt. 2,700-3,600 m., Pennell 14363; 14364;
14399; 14403; 14534; 14703; Matucana, alt. ca. 8,000 ft., Macbride &
Feather stone 103; Hacienda Balconcillo, Soukup 2147; Chosica, alt.
800 m., Soukup 2149. — Puno: Near rocks and in water, near Lim-
bani, Prov. Sandia, alt. 3,200-3,450 m., Metcalf 30486.
Calceolaria dentifolia Edwin, Phytologia 19: 379. 1970.
Erect shrub, little branching on the upper half, wood, foliage,
flower-stalks, calyx tube and calyx lobe margins densely beset with
gland-tipped hair and with sessile or subsessile glands, the hair fre-
quently 2-celled. Leaves subcoriaceous, short-petiolate, ovate,
mostly 6-10 cm. long and 2-4 cm. wide, long-acute to acuminate at
apex, subobtuse and dimidiate at base, margins irregularly and very
sharply, doubly serrate, midvein often conspicuously elevated be-
FLORA OF PERU 585
neath; petioles thickened, up to 1.3 cm. long; inflorescences simple or
branching, corymbose, subtended by a pair of reduced leaves, termi-
nal on the main stem and on branches from the upper leaf -axils;
basal and intermediate peduncles 2-3 cm. long; pedicels much shorter
than the peduncles; calyx lobes lance-elliptic to lance-ovate, long-
acute to acuminate, the anterior lobe narrower than the others, ca.
10 cm. long and 4-6 mm. wide, sparsely puberulent externally, oscil-
late internally, oscillae sunken; corolla large for the genus, yellow,
upper lip sparsely puberulent, hooded, up to 10 mm. long, much
overtopping the genitalia, lower lip 30-35 mm. long and up to ca.
25 mm. wide, saccate most of its length, externally sparsely puberu-
lent or glandular with sessile glands, the vesture in patches, occasion-
ally glabrous, internally with a tuft of hair at the base; filaments
about 3 mm. long, very thickened, flattened, about 2 mm. wide, an-
ther cells unequal, large, each pair 6.5-7.5 mm. long; style ca. 7 mm.
long, thickened, apically incurved, stigma enlarged or not, puncti-
form-capitate, ovary glutinous, sparsely glandular, glands golden,
sessile; capsule not seen.
Cajamarca: Shrub, along stream E. of Cordillera de Cumulloa
on road to Celendin, 3,400-3,500 m. alt., Pennell 15162, type col-
lection; Cajamarca-Celendin road, km. 90, Olsson s.n., February 11,
1948. Leaves sticky.
Close to Calceolaria endopogon Kranzl. and C. salicifolia R. & P.
differing from both in vesture. In addition, this taxon differs from
endopogon in leaf-margins and in the structure of the inflorescence.
Dentifolia differs from salicifolia in leaf-shape and petiole length.
Calceolaria dichotoma Lam., Encyc. Meth. Bot. 1: 555. 1783.
C. ovata Smith, Icones 1 : 3, tab. 3. 1789. C. biflora R. & P., Fl. Peruv.
1: 16, tab. 20. 1798. C. anagalloides Kranzl. in Fedde, Rep. Sp. Nov.
1: 84. 1905. C. lysimachioides Kranzl., I.e., 84. C. meistantha Pen-
nell, Proc. Acad. Nat. Sci. Phila. 103: 96, fig. 1. 1951.
Annual herb, usually up to about 15 cm. tall, erect or ascending,
branching dichotomously over most of its length, foliage, branches,
flower-stalks and calyces variously soft, white pubescent. Leaves
ovate, short- to moderately petiolate, up to about 5 cm. long and
4 cm. wide, usually not exceeding 3.5 cm. long and 2.8 cm. wide,
acute to subacute at apex, rounded at base, sometimes subobtuse,
pubescence most often scattered on both surfaces, margins entire to
irregularly serrulate; petioles up to about 1 cm. long, little if at all
thickened; flowers usually axillary, solitary and in pairs, sometimes
586 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
also in few-flowered, terminal dichasia on the branches; pedicels and
peduncles usually short, up to ca. 2.5 cm. long, most often not ex-
ceeding 1.5 cm.; flowers small, 6-9(-10) mm. long; calyx lobes 2.5-
5.0 mm. long, elliptical to ovate; corolla light yellow, upper lip ca.
0.5-2.0 mm. long, lower lip 5-8 mm. long, saccate up to about one-
half its length; filaments long, up to 5 mm., thin, anther cells very
small, usually each pair not exceeding 1 mm. long, sometimes con-
fluent, the anther seeming unilocular; style ca. 2 mm. long, ovary
glabrous or almost so; capsule broadly ovoid, a little longer than the
calyx, apically obtuse, often notched, the usually persisting style
within.
Ayacucho: Aina between Huanta and Rio Apurimac, alt. 750-
1,000 m., open woods, Killip & Smith 22813; Evergreen bush wood
near Quillamito, Choimacota Valley, Prov. Huanta, alt. 1,300-1,400
m., Weberbauer 754.2. — Cajamarca: In full exposure, Llama, Cutervo,
alt. 7,500 ft., Sandeman 4200; Plowed ground alt. 3,000 m., SW slope,
Paso de Gavilan, S. of Cajamarca, Pennell & Ferreyra 14965.—
Cuzco: San Miguel, Urubamba Valley, alt. ca. 1,800 m., Cook &
Gilbert 924; Amaibamba, Prov. Convencion, in cultivation, Marin
1575; alt. 1,650 m., Scolnik 813; Vargas 3423. — Lima: Lomas de
Quilmana, between Lurin and Cafiete, alt. 480-500 m., Ferreyra
3989; Lomas de Atocongo, Prov. Lima, alt. 300-400 m., Ferryra
4017; Lomas de Caracolas, between Lurin and Cafiete, alt. 490-
500 m., Ferreyra 6281; Sandy lomas along the sea, Lurin, alt. ca.
200 ft., Macbride 5969.—Piura,: Herb in wet maize field, Canchaqui,
alt. 1,200-1,600 m., Pennell & Ferreyra 14893.— Sin. Dept.: Lucu-
mayo Valley, alt. 1,800-3,600 m., Cook & Gilbert 1397.
Calceolaria divaricata HBK., Nov. Gen. et Sp. 2: 381. 1817.
Herb or subshrub, stem glabrous, floriferous, branchlets sparsely
puberulent at the apex and very divergent. Leaves short-petiolate
except the sessile uppermost, ovate, up to 4 cm. long and 3 cm. wide,
grossly crenate-serrate, acute at apex, subcordate at base, bright
green and glabrous above, manifestly reticulate-venose and pilose,
especially on the margins, beneath; petioles (the lowermost) up to
1.2 cm. long, basally connate and thickened; flowers in corymbose
inflorescences, ultimate branchlets divaricate, dichasiate; the thin
pedicels 1.0-1.5 cm. long and with the calyces glandular-pilose;
calyx lobes broadly ovate, acute, 3 mm. long and 2 mm. wide at
base; corolla yellow, 1.5-1.8 cm. long, lower lip saccate less than half
its length; stamens short, anthers large.
FLORA OF PERU 587
No specimens seen. Reported from Piura: near Succhubamba
and Ayavaca (Ayabaca), alt. 2,800 m., Humboldt & Bonpland.
Calceolaria endopogon Kranzl. in Engler, Bot. Jahrb. 54:
Beibl. 119, 18. 1917.
Subglabrous to often glabrous vine, stem and branches sometimes
glutinous, often purple-dotted in drying. Leaves petiolate, elliptic-
ovate to ovate, large for the genus, up to 10 cm. long or even longer
and about 5 cm. wide, entire or almost so, apically acute, frequently
dimidiate near the narrowed base, veins conspicuous or not beneath,
darker above, paler beneath; inflorescences few-flowered, subdicha-
siate to racemose; pedicels and peduncles thin, glabrous, peduncles
up to 10 cm. long, pedicels to 5 cm. long; floral leaves only a little
smaller than the stem leaves; flowers large for the genus; calyx lobes
9-12 mm. long, glabrous, oblong to ovate-oblong, acute, margins
fimbriate; corolla yellow, externally glabrous, upper lip well devel-
oped, sometimes almost as long as the calyx lobes, more often only
about half as long, internally beset with long, white, stiff hairs on the
lower edge; lower lip about 30 mm. long, saccate over three-fourths
its length, internally pubescent as the upper lip, sac very inflated in
life and upcurved distally; anthers on short filaments, cells very un-
equal, each anther large for the genus, 7-9 mm. long and about a
third to a fourth as wide, cells divaricate; style 7-8 mm. long, glan-
dular at the base, apically abruptly curved, ovary densely glandular-
puberulent.
This taxon has been sporadically collected in the Andes of north-
ern Peru.
Cajamarca: Road from Cajamarca to Celendin, km. 95, Olsson
s.n.; Thicket on mountain E. of Celendin, alt. 2,700-2,900 m., Pen-
nell 15210; Hills W. of Celendin, alt. 2,700-3,000 m., Pennell 15218.
—Piura: Huancabamba, West side of Cordillera E. of Huancabamba,
alt. 3,000 m., Weberbauer 6136 (Isotype F) ; Above Huancabamba, alt.
3,000 m., Weberbauer 6147.
Calceolaria engleriana Kranzl. in Fedde, Rep. Sp. Nov. 1: 106.
1905. C. epilobioides Kranzl., Ann. K. K. Naturh. Hoefm. Wein. 22:
194. 1907. C. urubamba Kranzl., Pflanzenr. IV(257C) : 109. 1907.
C. incachacensis Kranzl. in Fedde, Rep. Sp. Nov. 27: 20. 1929.
C. viminalis Kranzl. in Fedde, I.e. 21. 1929. C. endotrachys Kranzl.
in Fedde, I.e. 22. 1929. C. engleriana subsp. acuminata Pennell,
Proc. Acad. Nat. Sci. Phila. 97: 158, t. 10A. 1945.
588 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Erect branched shrub, wood usually tomentulose above becoming
glabrous below. Leaves short-petiolate or the upper blades sessile,
narrowly ovate-lanceolate to lanceolate, occasionally ovate, up to
about 6.0 cm. long and 2.6 cm. wide, usually narrower, not infre-
quently 5-9 mm. wide, entire to serrulate, apex usually acute, less
often short-acuminate, variously pubescent to occasionally glabrous;
petioles thickened and laterally connate, sometimes woody; inflores-
cences few-flowered corymbs; flower-stalks puberulent; calyx lobes
green to yellow-green, ovate or narrowly ovate, acute, 5-7 (-8) mm.
long, externally sparsely puberulent and often red-gland-dotted,
sometimes reflexed at anthesis, internally more densely hairy, some
of the hairs arranged in a row along the margins; corolla yellow,
upper lip ca. 4-6 mm. long, shorter to occasionally little longer than
the calyx lobes, lower lip 18-28 mm. long, saccate more than half its
length, distally upturned; anther cells more than twice as long as
wide, filaments shorter than the 2 mm. long, apically curved style;
ovary glandular; capsule a little shorter to equaling the calyx, ovate,
gradually narrowed to apex but lacking an abruptly constricted neck,
densely glandular.
Found throughout the Andes of southern Peru, especially in
Cuzco.
Apurimac: Quishuala, road between Abancay and Andahualas,
alt. 11,500 ft., Balls 6893. — Ayacucho: Between Huanta and Haci-
enda Pagora, alt. 3,700-4,200 m., Killip & Smith 22189; 23297;
23300.— Cuzco: Alt. 3,300-3,500 m., Herrera 526; "Ppucuchu-ppu-
cuchu," Urubamba Valley, alt. 3,000 m., Herrera 1525; Colinas de
Saxaihuaman, alt. 3,600 m., Herrera 2367; Que"brada de Quispicanchi,
alt. 2,350 m., Herrera 2621; Valle de Paucartambo, alt. 3,400 m.,
Herrera 2989; near Wenner Gren ruins, Prov. Urubamba, alt. 3,400-
3,600 m., wet places, Metcalf 30763; Sacsahuaman, above Cuzco,
stream bank, alt. 3,500-3,600 m., Pennell 13545; Cuzco, alt. 3,500-
3,600 m., Pennell 13702; Huancalli, open rocky banks, alt. 3,400-
3,500 m., Pennell 13712; Pisac, rocky canyon E. of Pisac, alt.
3,300-3,500 m., Pennell 13722; Cerro de Colquipata, puna, alt. 3,900-
4,000 m., Pennell 13746; near Rio Quencomayo, below Colquipata,
alt. 3,200-3,300 m., Pennell 13778 (type of C. engleriana acuminata
Pennell); Cerro de Colquipata, trail from Paucartambo to Caicai,
alt. 3,400-3,700 m., Pennell 14178; Vilcanota below Caicai, alt. 3,300-
3,500 m., Pennell 14193; Cordillera Veronica, alt. 2,900 m., Rauh-
Hirsch P969; 3,700 m. alt., Reitz 5986; Sacsahuaman, Soukup 21;
outskirts of Cuzco, Soukup 127; Chocta, near roadside, alt. 3,400 m.,
FLORA OF PERU 589
Vargas 228; Anta, alt. 3,650 m., Vargas 298; Huanoquita, Prov.
Paruro, alt. 3,380 m., Vargas 2359; Juncaipata-Sta Rita, Prov. Uru-
bamba, alt. 2,800 m., Vargas 2670; Hacienda Cachupata, Prov.
Paucartambo, alt. 3,500 m., Vargas 2851; Colinas del Rodadero,
Vargas 3132; Prov. Calca, alt. 3,200 m., Vargas 3620; Chenchamayo,
Prov. Paucartambo, Vargas 4303. — Huancavelica: Pumalucana to
Conaica, alt. 3,500-3,550 m., Tovar 79. — Puno: 1 km. from Cuyo-
cuyo, on road to Sandia, alt. 3,600 m., Metcalf 30715. — Sin Depto:
Lucamayo Valley, alt. 1,800-3,600 m., Cook & Gilbert 1327; Ollan-
taytambo, 3,000 m. alt., Cook & Gilbert 597; 658; Pinasniocj, Panti-
calla Pass, alt. 3,600 m., Cook & Gilbert 1815. — Sin. Loc.: Weberbauer
5478.
One of the commonest "slipper-flowers" in the southern Andes of
Peru. This taxon is quite easily recognized due to the long, narrow
leaves, anther cells, and corollas.
Calceolaria extensa Benth. in DC., Prodr. 10: 214. 1846.
Shrub, usually much branched, the branches ascending, some-
times square on the upper part, older wood tomentose, youngest
puberulent. Leaves short-petiolate, oblong-lanceolate, up to 2 cm.
long, usually not exceeding 1.5 cm. and 5 mm. wide, crenate or grossly
crenate, acute at apex, cuneate to long-cuneate at base, subglabrous
above, paler and densely pubescent on the venation beneath; inflores-
cences few-flowered corymbs, only little exceeding the leaves; branch-
lets and flower-stalks papillose-pilose; bracts linear or almost so, ob-
tuse; calyx dimorphic, on some plants either densely pilose externally
with the segments ovate, subacute to acuminate, 5-6 mm. long and
4-5 mm. wide, or oblong-lanceolate, 7-10 mm. long and 3 mm. wide
and glabrous; upper lip of the yellow, externally sparsely papillose
or glabrous corolla, minute, reduced to a narrow ring, lower lip ca.
1.5 cm. long, saccate about half way, about 8-10 mm. wide distally;
filament ca. 3 mm. long, anthers divergent, ca. 1.5 times longer than
wide; style ca. 1.5-3.0 mm. long, ovary glabrous to sparsely glandu-
lar; capsule not seen.
Ayacucho: Prov. Lucanas, between Nazca and Puquio, alt. 3,200-
3,300 m., Ferreyra 51+62; above Puquio, alt. 3,400-3,500 m., Ferreyra
5525; Allpaja, ca. 6 km. from Puquio, alt. 3,300-3,400 m., Ferreyra
7191.— Sin. loc. alt. 2,407 m., Soukup 4003.
Also reported from the departments of Ancash and Lima as well
as from southern Bolivia.
590 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria fiebrigiana Kranzl. in Fedde, Rep. Sp. Nov. 1: 84.
1905.
Herb, frequently scapose or subscapose, scape or stems white-
pilose. Leaves broadly ovate, up to 15 cm. long and 8 cm. wide,
variously pubescent on one or both surfaces, veins sometimes prom-
inent beneath, margins grossly sinuate-dentate, apex acute, base
often cordate, sessile or short-petiolate; petiole when present up to
5 mm. long, winged by the attenuate leaf-base; inflorescences usually
much branched and many-flowered, glandular-pilose and often also
glutinous; calyx lobes ca. 5 mm. long and 3 mm. wide, usually glan-
dular-pilose, occasionally merely pilose, acute; upper lip of the yellow
corolla from about half as long as the lower lip to almost as long,
lower lip 6-9 mm. long, saccate more than half its length; fila-
ments linear, long, exceeding the flower, anthers small, cells equal;
style about as long as the stamens, ovary glandular; capsule broadly
ovoid, little longer than the calyx.
Arequipa: Isern 2016. — Cuzco: Paucartambo Valley, Hacienda
Ccapana, alt. 3,600 m., Herrera 1083; La Raya, rock-ledges in cas-
cade, alt. 4,400-4,500 m., Pennell 13509; Sin. Loc.: Alt. 4,710 m.,
Rauh-Hirsch Pi 160; Alturas de Laguna de Sial, alt. 4,300 m., Var-
gas 374.1 ; Environs of San Andres de Checca, alt. 3,860 m., Vargas
9845. — Moquegua; Humid rocks, Saylapa near Carumas, alt. 3,300
m., Weberbauer 7343. — Puno: Granja Salcedo, scattered among rocks,
alt. 3,835 m., Mexia 4253; Santa Rosa 50 mi. SSW. of Llave, alt.
4,500 m., Pearson & Pearson 31 ; Near Puno, alt. 4,000 m., Soukup
255; 370. — Tacna: In Chuvire, 1 km. from Challaquaya, Prov. Ta-
rata, alt. 2,500 m., Metcalf 30408. — Sin. Loc.: Volcanic Ash, alt.
13,500 ft., Stafford 670.— Sin Dept. : Uyumiri, Sicuani, alt. 12,500 ft.,
Stafford 962.
Calceolaria flosparva Edwin, Phytologia 19: 380. 1970.
Prostrate herb, rooting at the nodes, stems and branches glabrous
or almost so. Leaves mostly long-petiolate, blades ovate in outline,
mostly 1.5-2.8 cm. long and 1.3-2.2 cm. wide, densely to thinly pul-
verulent beneath, irregularly coarsely dentate, laciniate-dentate and
varying to with 1 or 2 pairs of lobes, the lobes irregularly laciniate-
dentate, the narrowed base decurrent the length of the petioles that
are at least half as long as the blade to almost as long; flowers axillary
in upper leaves usually a pair in each axil, pedicellate, only rarely
pedunculate; pedicels glabrous, 1.5-2.0 cm. long; calyx glabrous,
lobes ovate, subobtuse, about 5 mm. long and 3 mm. wide, margins
FLORA OF PERU 591
very sparsely crenulate; corolla yellow, glabrous, 6-7 mm. long, lower
lip only little shorter than the almost wholly saccate upper lip; both
anther cells fertile and much smaller than the thickened connective;
style about 1.5 mm. long; capsule broadly ovoid, little shorter than
the calyx, pubescent, neck lacking.
Thus far found only in Dept. Amazonas: Rocky stream in moun-
tain forest, alt. 3,300 m., above Colcamar, Pennell 15633 (PH); in
seepages, infrequent, uppermost slopes and summit of Cerros de Calla
Calla, near Kms. 403-407 of Balsas-Leimebamba road, alt. 3,400-
3,550 m., Wurdack 1711.
The combination of very small corollas with upper lip only little
smaller than lower lip, almost wholly pedicellate flowers, generally
glabrous inflorescences, and long-petiolate leaves is not found any-
where else in this subgenus.
Calceolaria glauca R. & P., Fl. Peruv. 1: 17, tab. 27, fig. e. 1798.
Stiff, erect, branching shrub; wood in age glabrous, younger mi-
nutely puberulent, youngest densely so. Leaves ternate or less often
opposite, youngest densely, minutely puberulent on both surfaces,
older less so, oldest becoming glabrous above and only sparsely pu-
berulent beneath, both surfaces punctate with shining, golden punc-
tations, lanceolate to lance-ovate, margins entire to irregularly
serrulate, denticulate or occasionally ovate with doubly dentate-
denticulate margins, mostly up to about 5.0 cm. long and 1.5 cm.
wide, usually not exceeding 3.5 cm. long and 1.0 cm. wide or occa-
sionally when ovate and doubly toothed just 2 times longer than
wide, acute at apex, usually subobtuse at base; inflorescences irreg-
ular, often paniculate or of panicles of dichasia, many-flowered;
flowers congested on short, puberulent stalks, mostly 10-13 (-15) mm.
long; calyx puberulent, ovate, 3-4 mm. long and almost as wide at
base, apically acute; corolla yellow, glabrous to sparsely puberulent
externally, a tuft of short, white hairs often present internally at the
base, upper lip about as long as the calyx and as wide, overtopping
the genitalia, lower often saccate three-fourths of its length, the prox-
imal part very narrow and stipe-like; filaments short, anther cells
equal, up to almost twice as long as wide, each pair ca. 2-3 mm. long,
wholly divaricate; style 1.5-2.5 mm. long, pustulate at base, ovary
glandular-pustulate; capsule about as long as wide, broadly ovoid,
glandular, apex blunt.
Ancash: Marcara, Prov. Huaras, alt. 3,000 m., Nunez s.n. — Caja-
marca: Cajabamba, near the bridge of Lulichucu, alt. 2,600-2,700
592 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
m., Ferreyra 3047. — Lima: Rio Blanco between Matucana and
Aroya, alt. 3,500-3,600 m., Ferreyra 5271; Rocky ground near San
Mateo, alt. 2,600-2,700 m., Ferreyra 5316; open hillside Rio Blanco,
alt. 300-3,500 m., Kittip & Smith 21654; 21686; Creek bank, Matu-
cana, alt. ca. 8,000 ft., Macbride & Feather stone 97; Surco, Prov.
Huarochiri, alt. 1,750-1,900 m., Nunez 1708; Along Rio Chillon near
Obrajillo, bush banks, alt. 2,200-2,500 m., Pennell 14327; alt. 2,600-
2,800 m., Pennell 14358; Open rocky slopes, alt. 2,800-3,200 m., Pen-
nell 14395; San Buenaventura, alt. 2,700-2,800 m., Pennell 14554;
Canta, alt. 3,000-3,200 m., Pennell 14600; Ruiz & Pavon s.n.; Poca-
tulpe-Hulpa, Matucana, Prov. Huarochiri, alt. 3,000-3,400 m. Var-
gas 5250. — Sin. Loc: Weberbauer 5397.
The single collections from Ancash and Cajamarca indicate a
significantly wider range for this taxon than is presently known.
Calceolaria grandepinnata Edwin, Phytologia 19: 380. 1970.
Soft herb, branching above the middle, stems pubescent with
multicellular hair and these frequently thickened and filled with a
dark-brown exudate above and at the nodes. Leaves up to 23 cm.
long, narrowly ovate to elliptically ovate in outline, very variable,
multi-pinnate or -pinnatifid over all or most of their length, pinna
sessile or less often stalked, irregularly dentate-laciniate, sometimes
the upper part of the leaf merely dentate, pinnae most often reduced
in size toward the apex, alternate or opposite, apically acute or long-
acute, pubescent on both surfaces with multicellular hair, these often
appressed on the upper surface, long- or short-petiolate; petioles thinly
connate; inflorescences simple or once-branched cymes, borne in
pairs on the branches from the axils of the uppermost leaves; basal
peduncle up to 3-4 cm. long, mostly varying in length, densely and
laxly gland-tipped-pubescent; pedicels pubescent as the peduncles,
abruptly much shorter than the peduncles; bracts and bractioles re-
duced, lanceolate, otherwise leaf -like; calyx lobes broadly ovate or
deltoid, ca. 3-4 mm. long and ca. 3 mm. wide in flower, becoming
9 mm. long and 6 mm. wide in fruit, acute, densely pubescent exter-
nally, less so internally, acute, margins entire with the terminals of
the veins little protruding; corolla yellow, more or less densely beset
with sessile, red glands externally, 13-15 mm. long, upper lip ca.
4 mm. long, hooded, margins rounded, lower lip a little narrower than
long, saccate almost its entire length, nectary small, entire, hemi-
spherical, terminal on the internally glabrous sac; one anther cell
sterile, the other fertile, almost globular, the connective between the
FLORA OF PERU 593
two longer than either cell and thickened, filament knob-like; style
ca. 2.5 mm. long, pubescent near the base, ovary pubescent; capsule
as long as or longer than the calyx, broadly triangular-ovoid, pubes-
cent, style persisting at the acute apex, neck lacking.
Distributed in Piura near Canchaqui: Stream ditch, alt. 1,000 m.,
below Canchaqui, Pennell & Ferreyra U891 (US); Alt. 1,000-1,400
m., in wet places Pennell & Ferreyra H892; Ferreyra 3091.
Easily separable from all other Calceolarias by the combination
of pluripinnate, very long leaves, and the peculiar nodal pubescence.
Calceolaria hedera Edwin, Phytologia 19: 381. 1970.
Climbing shrub with the lax branches and the nodes distant,
young wood tomentose, tomentum decreasing with age until older
wood almost glabrous, foliage beneath and flower stalks densely la-
nate. Leaves mostly very broadly ovate in outline, less often almost
subrotund, mostly up to 9 mm. wide, sometimes reaching 2 cm. wide,
only very rarely exceeding 2 cm. long, mostly up to 15 mm. long,
margin revolute, with usually 2 pairs of lateral lobes and a single
apical lobe, varying to 1 or 3 pairs of laterals, lobes rounded, usually
minutely cordate at base, occasionally truncate or broadly obtuse,
pilose above, densely lanate beneath, the indument matted, obscuring
the tissue; flower stalks up to ca. 2 cm. long, usually less, not infre-
quently less than 1 cm. long, occasionally the flowers subsessile;
flowers solitary and axillary or in 3-flowered, simple dichasia;
branches floriferous over half their length; floral leaves similar to
cauline, only little reduced; the short calyx tube and lower parts of
the calyx lobes externally densely tomentose, vesture decreasing to
the apices of the lobes, internally sparsely pubescent, lobes ellipti-
cally, broadly ovate, ca. 6 mm. long and 5 mm. wide, acute, entire;
corolla yellow, wholly glabrous or with a very few hairs internally
at the base of the lower lip, 14-17 mm. long, upper lip 1-2 mm. long,
lower lip saccate more than half its length, sac broadly, abruptly
expanded, much wider than the non-saccate part; anther cells equal,
divaricate, 2 or more times longer than wide, the upturned distal
ends farthest from the floor of the corolla each pair of cells 2-3 mm.
long, filaments about 1.0-1.2 mm. long, about half the length of the
thickened 2.0-2.5 mm. long style; ovary glandular- or granular-
pubescent; capsule unknown.
Known only from the type collection.
Cajamarca: springs in hills, midway between Cordillera de Cum-
ulloa and Celendin, alt. 2,700-3,000 m., Pennell 15221.
594 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
The leaves are much like the common "ivy" in shape. This taxon
is related to C. cordi/ormis and C. triloba Edwin but is most easily
separable at least at present. Although most of the leaves of this
taxon are less than 1 cm. wide, the largest blades indicate the strong
possibility that future collections may uncover specimens whose
leaves are more than 1 cm. wide. The taxon is keyed therefore under
both key entries.
Calceolaria herzogiana Kranzl. in Engler, Bot. Jahrb. 50:
Beibl. 111. 69. 1914.
Erect but flaccid, mixed-pubescent, perennial herb with the square
stems drying 4-sulcate, densely pubescent with glandular hairs inter-
mixed. Leaves bizarre; petioles winged up to 10 cm. long or occa-
sionally longer, basally dilated, connate, auriculate, margins usually
grossly dentate, especially on the lower half; blades (except the ses-
sile, ovate-cordate, smaller, sometimes interfloral uppermost) usually
triangular, often at least as wide as long, very variable in size 6-16
cm. long and 5.0-16.5 cm. wide,1 coarsely, irregularly doubly dentate,
acute, bases truncate, retuse or subcordate, pubescent or puberulent
beneath on the venation, above sparsely to subdensely puberulent;
inflorescences corymbose or dichasiate, few- to many-flowered; pe-
duncles and pedicels pubescent as the stem, basal peduncles 6-10 cm.
long, flower stalks decreasing in size apically; calyx glabrous or pu-
berulent, the free sepals entire, eciliate, ovate, 13-23 mm. long, bas-
ally up to 13 mm. wide, acute or acuminate; corolla up to 40 mm.
long, mostly over 30 mm., glabrous except for a tuft of white hairs
within at base of the lower lip, sac usually about two-thirds the length
of the corolla, more than half the length of the 30-35 mm. lower lip,
upper lip short, ca. 3-5 (-7) mm. long, only partly obscuring the geni-
talia; filaments short, 2-3 mm. long, anther cells divaricate, walls
thin, dehiscing through the septum; style 3-5 mm. long, ovary glan-
dular; capsule broadly ovoid, 8-9 mm. long and about as wide at
base, glandular at least along the sutures, apically obtuse.
Basically found in southern Peru but extending to central Peru
in Junin and Ancash. The southern counterpart of C. tomentosa and
C. pavonii.
Ancash: Prov. Bolognesi, Chiquan, alt. 3,560 m., Ferreyra 5703;
Encima de Chiquian, alt. 3,350-3,400 m., Ferreyra 7463. — Ayacucho:
Ccarrapa, between Huanta and Rio Apurimac, alt. 2,200 m., Killip
1 According to original description up to 19 cm. wide and 13 cm. long.
FLORA OF PERU 595
& Smith 22314-—Cuzco: Vicinity of Macchu-picchu and Urubamba
Valley, alt. 1,800-2,500 m., Balls 6812; Cardenas 4200; Cook & Gil-
bert 1153; Herrera 1555; Mexia 8074; Rick s.n.; Soukup 142; Vargas
798; 9818; Valley of San Miguel, between Cedrabamba and La Ma-
quina, alt. 2,060-2,260 m., Herrera 1997; Cerro de Cusilluyoc, thicket,
alt. 3,400-3,700 m., Pennell 13805; Near Rio Yanamayo, below Pilla-
huata, alt. 2,200-2,400 m., Pennell 14042; Pillahuata, Cerro de
Cusilluyoc, alt. 3,000-3,300 m., Pennell 14135.
Calceolaria heterophylla R. & P., Fl. Peruv. 1: 16, t. 21, fig. b.
1798, non Willd. C. hirsuta Benth. in DC., Prodr. 10: 212. 1846.
Herb decumbent at base, stems and branches densely beset with
long, multicellular, descrete hair, plant drying brown. Leaves dis-
tant, long-petiolate, petioles and the veins at least beneath with
vesture as the stems and branches, blades ovate, 4-9 mm. long and
3-5 mm. wide, margins crenate-dentate or dentate, acute at apex,
subobtuse to obtuse at base; inflorescences corymbose, few-flowered;
peduncles and pedicels with vesture as the stems, basal peduncles
up to 20 cm. long (fide Kranzlin) ; bracts lacking; flowers 20-25 mm.
long; calyx vesture as the stem, lobes ovate, acute, mostly 7-8 mm.
long; corolla yellow, wholly glabrous, ca. 15-18 mm. long, about one-
half to two-thirds as wide, upper lip 2-3 mm. long, lower lip saccate
less than half its length; filaments ca. 2 mm. long, black when dry,
anther cells equal, small, ca. 1.5 mm. long, divergent to divaricate;
style 3.0-3.5 mm. long, little if at all thickened; ovary glandular,
glutinous; capsule shorter than the calyx, broadly ovoid, only little
narrowed to the obtuse apex, the locular suture forming a deep
sulcation.
Amazonas: Mathews 152, the type of C. hirsuta Benth. — Apuri-
mac: Pincos, in sunny sites, Prov. Andahuaylas, alt. 2,600 m., Stork
& Horton 10704- — Huanuco: Huacachi, station near Muna, alt. ca.
6,500 ft., Macbride 4154; Ruiz & Pawn s.n.
Collections are from in or near Muna except the Apurimac and
Amazonas cited above. These latter seem well within the morpho-
logical variation limits for the taxon. The disjunction is quite un-
usual for the subgenus.
Calceolaria hirsutula Pennell, Proc. Acad. Nat. Sci. Phila. 97:
158. 1846.
Shrubby at the base, erect, hirsutulous with short, spreading, up-
curved hairs, stems several from a rhizomatous base, simple or
596 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
branched. Differing from C. chrysosphaera Pennell in pubescence
and also in having smaller calyx lobes, ca. 6-7 mm. long and these
green instead of yellow; smaller corolla, ca. 10-12 mm. long; shorter
leaves, ca. 3-4 cm. long and also smaller capsules; differing from
C. engleriana Kranzl. in many of the same ways as well as in having
smaller anthers, ca. 2 mm. long and capsules shorter than the calyx
lobes; the terminal pedicels in hirsutula are shorter than in both
chrysosphaera and engleriana.
Cuzco: Cult, at U. Calif. Bot. Gard. from Goodspeed 4513, col-
lected Macchu-picchu; Valle de San Miguel, Macchu-picchu, alt.
2,400 m., Herrera 1999 (type); Valle del Urubamba, Macchu-picchu,
alt. 2,200 m., Herrera 3223; Hda. Anaibamba, alt. 1,650 m., Scolnik
791 ; Hda. Potrero, alt. 1,300 m., Vargas 1181; Alt. 2,100 m., Vargas
2507; Macchu-picchu, alt. 2,000 m., Vargas 2173; Hda. Amaibamba,
alt. 1,500 m., Vargas 3426; Prov. Quispicanchis, Tio to Myrayaca,
Marcapata, alt. 1,200-2,000 m., Vargas 5198.
Calceolaria humilis Edwin, Phytologia 19: 382. 1970.
Small, branched shrub from a thickened rootstock, several stems
rising closely grouped, lower nodes longer, upper sometimes shorter
than the small leaves, wood drying wine-red, puberulent as the ped-
icels and calyx lobes. Leaves ovate, up to 1.5 cm. long and 9 mm.
wide, short-petiolate, pulverulent on both surfaces and often puber-
ulent above, yellow-green, glandular beneath, crenulate-serrulate,
plane, acute at apex, subacute to subobtuse at base, widest near but
not at the base, venation subobsolete and impressed above, promi-
nent and elevated beneath, often with flattened, white puberulence;
flowers solitary, crowded, axillary to upper leaves and terminal on
the branches and stems; pedicels reddish under the white puberu-
lence, ca. 1 cm. long; bracts leaf-like, reduced; calyx lobes ovate, 3-4
mm. long and almost as wide at base, internally puberulent on the
red margins, yellow-green-glandular on the light green tissue, acute;
corolla yellow, 9-11 mm. long, upper lip reduced to a ring less than
1 mm. long, lower lip papillose on both surfaces, saccate less than
half its length, the proximal part only little narrower than the distal,
saccate part; filament ca. 1 mm. long, thickened, red; anther cells
equal, divergent or divaricate, each pair 2 mm. long and 1 mm. wide;
style ca. 1.0-1.5 mm. long, thin, ovary papillose-glandular; fruit
not seen.
FLORA OF PERU 597
Cajamarca: Open jalca, southwest slope, Paso de Gavilan, S. of
Cajamarca, alt. 3,100-3,300 m., Pennell 11+953, type collection; Bushy
ravine on jalca, southwest slope, Paso de Gavilan, S. of Cajamarca,
alt. 3,100-3,200 m., Pennell 11+951.— Ancash: Matahuanca, ca. 20
km. S. of Huaraz, alt. 4,000 m., Rick s.n.
Close to C. inamoena Kranzl. which differs in having the flowers
in dichasia, much more deeply toothed or lobed leaves and very dif-
ferent indument.
Calceolaria hutchisonii Edwin, Phytologia 19: 383. 1970.
Erect, branched herb to 1.5 m. tall, stem drying red-brown, stri-
ate on upper part, closely puberulent above to sparsely so or almost
glabrous below. Leaf blades very broadly ovate, sometimes to often
wider than long, the longest up to 2.5 cm. long, mostly 0.8-2.0 cm.
long, and 1.0-2.2 cm. wide, narrowed apically, very broadly obtuse
basally, margins coarsely, irregularly crenate-dentate, teeth usually
few, distant, densely and closely puberulent above, densely tomentu-
lose-pulverulent beneath, drying dark-green above, much paler be-
neath, subcoriaceous, short-petiolate; petioles densely pubescent, up
to 6.0 mm. long, thinly connate across the nodes; inflorescences
paired, from the uppermost leaf-axils, usually in irregular, l-2x
branched, subcymose clusters; flower stalks short, often less than
1 cm. long, densely puberulent; bracts and bracteoles lacking; calyx
lobes ovate, 4.0-4.5 mm. long, 3.0-3.5 mm. wide, acute to obtuse,
densely closely puberulent on both surfaces, calyx tube puberulent;
corolla yellow, densely papillose-glandular externally, 13-17 mm.
long, upper lip annulate, 2-3 mm. long, lower lip saccate about half
its length, internally proximally beset with long, white hair; stamens
very small, filaments hardly 0.5 mm. long, anthers ca. 2.5 mm. long,
cells equal, divergent, a little longer than wide; style 1.5-2.0 mm.
long, very thickened on lower half, incurving to the upper lip from its
base; ovary glandular; capsule not seen.
Piura: Huancabamba Prov., 18.5 km. above Huancabamba, on
road W. to Piura, alt. 3,200 m., Hutchison & Wright 6630; El Tambo,
road between Piura and Huancabamba, alt. ca. 3,000 m., Scolnik
1407; Above Huancabamba alt. 3,200-3,300 m., Weberbauer 6048.
The very short filaments, very broad leaves, compound inflores-
cences and very thickened style form a combination of characters
not seen in any other species in the genus.
598 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Calceolaria hyssopifolia HBK., Nov. Gen. et Sp. 2: 386. 1817.
C. linifolia Willd. ex Link., in Spreng., Jahrb. 3: 54. 1820.
Erect, much branched, sometimes glutinous shrub, wood 4-angled,
drying brown, foliage drying green-brown, externally glabrous except
for puberulence on the branchlets and flower stalks. Leaves sub-
sessile or almost so, lanceolate or linear-lanceolate (in ours), up to
8 (-9) mm. wide, usually not exceeding 5 mm. long, margin entire,
sometimes a little revolute, apex long-acute to acuminate, base usu-
ally acute, much darker above than beneath; petioles reaching 5 mm.
long; inflorescences lax panicles; bracts leaf -like, reduced; calyx lobes
narrowly ovate, acute, 2.0-3.5 mm. long and 1-2 mm. wide; corolla
yellow, glabrous except for a tuft of hair internally at the base, upper
lip much exceeding the calyx, 7-9 mm. long, lower lip 10-14 mm.
long, saccate less than half its length; anther cells equal, broadly
ovoid to subhemispherical, on very short filaments, up to ca. 1.2 mm.
long; style 3 mm. long, ovary glabrous.
Amazonas: Approaches to Cerro Campanaria, NNE. of Diosan,
Prov. Chachapoyas, alt. 3,200-3,500 m., Wurdack 1591.— Libertad:
Moist bank, above Motil, shrub, alt. 3,000 m., Pennell & Ferreyra
14838.
This taxon has its main area of distribution in Ecuador. The
Peruvian material tends to have slightly wider leaves and smaller
flowers.
Calceolaria inamoena Kranzl. in Engler, Pflanzenr. 4(257c):
89. 1907.
Differing from C. cuneiformis R. & P. as in the key and in having
smaller leaves that are pilose both above and beneath and finally in
developing few-flowered dichasia at the apices of the branches.
Found from Arequipa, area of greatest concentration, N. to Junin
and Lima.
Arequipa: S. slope of Chachani, north of Arequipa, Hinkley and
Hinkley 24', Que"brada de San Lazaro, near Arequipa, alt. 9,100 ft.,
Munz. 15502; Slopes of Chachani, alt. 3,600-3,700 m., Pennell 13272;
Pampo de Arrieros, open rock-ledges, alt. 3,700-3,750 m., Pennell
13334; Rose 18952; Pichu-Pichu, among rocks on high banks, alt.
9,000 ft., Stafford 817; Prov. Arequipa, Jesus-Chiquata, alt. 2,800-
3,000 m., Vargas 8078. — Junin: Between Viques and Ingahuasi,
Mantaro Canyon S. of Huancayo, alt. 3,150 m., Killip & Smith
22153; Huancayo, Soukup 3148. — Lima: Prov. Cajatambo, stony
ground near Oyon, alt. 3,400-3,500 m., Ferreyra 3526.
FLORA OF PERU 599
Calceolaria inaudita Kranzl. in Fedde, Rep. Sp. Nov. 1: 104.
1905.
Small, black-dotted shrub up to 1 m. tall, black glandular-pilose.
Leaves very numerous, petiolate and sessile intermixed; petioles
when present broader and attenuate at base, black-setose; blades ob-
scurely trilobate, appearing globose due to the revolute margin, 1.0-
1.5 mm. in diameter equally thickened; flowers terminal on the
branches, forming few flowered corymbs; pedicels 2-3 cm. long,
glandular-papillose, gemmif orm ; minute bracts present on the pedi-
cels; calyx lobes densely glandular- villose on both surfaces, ovate,
obtuse, 2-3 mm. long and 2 mm. wide; corolla yellow, lips subequal,
about 1 cm. long and 6-7 mm. wide, upper lip margin little involute,
lower saccate over most of its length; stamins short, anthers reni-
form. (After Kranzlin, 1907.)
No material seen. Reported from Ancash: Cordillera Negra
above Caraz, alt. 4,200 m., Weberbauer 3085.
The text almost indicates a monstrous form of some other species.
Calceolaria incarum Kranzl. in Fedde., Rep. Sp. Nov. 1: 103.
1905.
Branched, glandular, erect shrub, stem and branches very densely
brown-glandular, especially above. Leaves sometimes bullate or
bullulate, oblong, with the short petiole up to 2.5 cm. long and up
to 8 mm. wide, deeply round lobed and strongly revolute, often
folded to the mid-vein, obtuse at apex and base, densely short-
velvety tomentose above, glandular-pilose beneath, glands often red-
brown when dry; petioles pubescent as the lower leaf surface; in-
florescences many-flowered, irregularly corymbose, flowers densely
disposed; flower stalks densely glandular, the pedicels usually less
than 1 cm. long; calyx lobes oblong to ovate, externally and inter-
nally densely villose and sometimes glandular, 6-7 mm. long and
5 mm. wide at base; upper lip of the corolla a little shorter than the
calyx, hooded, lower lip of the corolla 13-17 mm. long and about as
wide, the suborbicular sac almost its entire length to about half its
length, yellow, usually both internally and externally, sparsely glan-
dular-pilose or -pilosulus, varying to almost glabrous, long, white
glandular or eglandular hairs either present or absent around the
orifice internally; filaments short, anther cells equal, contiguous,
about 1.5 times longer than wide, each pair 1.5-3.5 cm. long, de-
hiscing through the septum; style thickened, a little longer than the
600 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
stamens; ovary glandular, capsule about as long as the calyx, nar-
rowly oblong-subovoid, glandular, except at the base, topped by the
persisting style.
Common in Dept. Ancash and thence north through Libertad to
Cajamarca. This type of distributions is not unusual in the genus.
As collecting continues other stations in Libertad and Cajamarca
should be uncovered.
Ancash (selected from the wealth of collections) : Fragments and
photograph of type, Prov. Cajatambo, alt. 3,000-3,500 m., Weber-
bauer 2762; Prov. Bolognesi, Shincush above Chiquian, alt. 3,890 m.,
Ferreyra 5827; Between Tallenga and Pachapaque, 3,400-3,600 m.,
Ferreyra 7470; Chavanillo, alt. ca. 8,000 ft., Macbride & Featherstone
1969; Catuc, ca. 15 mi. E. of Huaraz, alt. ca. 10,500 ft., Macbride &
Featherstone 2509; Jalca, Copa Grande, N. of Chancos, alt. 3,200-
3,500 m., Pennell 15363; Stony Puna, S. slope of Paso de la Fortale"za,
alt. 3,700 m., Pennell 15394; Open rocky limestone, above Chiquian,
alt. 3,600-3,700 m., Pennell 15418; Dry rocky and sandy bed, Que"-
brada de Huaqui, Huaraz, alt. 3,000-3,050 m., Pennell 15431 ; Jalca,
slopes and meadows, Cerro San Cristobal, N. of Huaraz, alt. 3,450-
3,600 m., Pennell 15448; Western slope of Cord. Negra, below Paso
de Collan, alt. 3,650-3,800 m., Pennell 15475; Matahuanca, ca. 20
km. S. of Huaraz, alt. ca. 4,000 m., Rick s.n. — Cajamarca: Cumbre
El Gavilan, road between Cajamarca and Chilete, alt. 3,200 m.,
Ferreyra 3273; Pennell & Ferreyra 14961; Limestone rocks, Llaca-
nora, alt. 2,550-2,600 m., Pennell 15487. — Libertad: Lake Sausa-
cocha, 10 km. N. of Huamachuco, alt. 3,000-3,200 m., Ferreyra 3031 ;
Pennell 14849.
Calceolaria inflexa R. & P., PL Peruv. 1: 16, t. 25a. 1798.
Vine or lax shrub, well branched, stem and branches sparsely
pilose, vesture occasionally very dense in the inflorescence, branches
opposite. Leaves variable in shape and size, lance-ovate to ovate,
up to 4 cm. long, usually not exceeding 3 cm. and up to 2 cm. wide,
mostly 5-13 mm., sparsely pubescent or glabrous above, densely
pulverulent beneath, both on the tissue and the thickened, promi-
nent, elevated veins, entire to rarely serrulate or crenulate, often
somewhat sinuate, acute at apex, narrowed to an usually acute or
acutish base, short-petiolate; petioles up to 2-4 mm. long, pubescent,
deeply grooved, thickened, thinly connate at least part way; flowers
in panicles; the basal peduncle usually once branched, up to 2 cm.
long, usually shorter; pedicels ca. half the length of peduncles, flower
FLORA OF PERU 601
stalks subglabrous to pubescent to occasionally densely glandular-
pubescent; bracts leaf -like, little reduced in size; calyx lobes ovate,
4-6 (-7) mm. long, acute, pubescent on both surfaces, margins thick-
ened, a little revolute; corolla yellow, both lips pustulate-pubescent,
from 13-30 (-33) mm. long, upper lip little shorter to often longer
than the calyx, ca. 5-7 mm. long; lower lip 20-28 mm. long, saccate
most of its length, sac much inflated, abruptly upturned, with a tuft
of white hairs at base internally; filaments short, anthers small, cells
equal, notched at the apex of the septum; style short ca. 1-2 mm.
long, densely beset with stalked glands at base; ovary densely covered
with sessile glands.
This emended description combines the information presented in
the text and picture of Ruiz and Pavon with the general run of speci-
mens most nearly matching these data.
Leaves entire or sinuate; flower stalks eglandular, sparsely puberu-
lent C. inflexa f . inflexa.
Leaves serrulate-crenulate; flower stalks densely glandular-puberu-
lent C. inflexa f . altissima.
Calceolaria inflexa R. & P., f. inflexa. C. laxa Benth. in DC.,
Prodr. 10:215. 1848.
Huanuco: Tambillo, 7 mi. SW of Panao, alt. ca. 8,000 ft., Mac-
bride 3580; Huacachi near Mufia, alt. ca. 6,500 ft., Macbride 3888;
between Acamayo and Carpish, beside stream in semi-shade, alt.
8,500 ft., Sandeman 5224; Sariapampa, alt. 3,600 m., Woytkowski
34322.— Junin: 6 mi. S. of Mito, alt. ca. 10,000 ft., Macbride & Feath-
erstone 1838; Pichis Trail, Dos de Mayo, dense forest, alt. 1,700-
1,900 m., Killip & Smith 25851; Oxapampa Soukup 2414; 1,768 m.
alt., Soukup 3345.
Also reported from Canta (Lima Dept.) and Sandia (Puno Dept.) .
Calceolaria inflexa f. altissima (Kranzl.) Edwin, stat. nov.
C. altissima Kranzl. in Engler, Pflanzenr. IV(257c): 107. 1907.
Sin. Dept.: Tambo de Vaca, alt. ca. 13,000 ft., Macbride 4437.
Originally described from Bolivia.
Calceolaria involuta R. & P., Fl. Peruv. 1: 15, t. 23a. 1798.
Tall, glutinous, branched shrub, branchlets drying shining brown-
ish-purple, older wood brown, plant almost entirely glabrous on new
wood, flower-stalks and calyces sparsely puberulent. Leaves petio-
602 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
late, blades lance-ovate, occasionally ovate, rarely lanceolate, mostly
4-8 cm. long and 1.5-3.0 cm. wide, dentate or serrate usually on
upper three-fourths of the blade only, entire toward the acute to
subobtuse base, acute or long-acute apically; petioles thickened, ca-
naliculate, up to almost 2 cm. long, usually 1.0-1.5 cm. long; inflores-
cences terminal on the branches, paired, few-flowered dichasia or
corymbs; calyx glabrous or the narrowly ovate lobes acute and
sparsely puberulent externally, 6-8 mm. long, ca. 3.0-3.5 mm. wide
at base; corolla yellow, glabrous externally or lower lip sparsely pu-
berulent, upper lip about as long as the calyx lobes or little longer,
lower ca. 12-15 mm. long, not 2 times longer than the upper lip,
internally a tuft of hairs developed at the base or more often lacking,
sac a little shorter than the non-saccate part; filaments short, anther
cells unequal or less often equal, more than 2 times longer than wide,
walls and septa thickened, each anther ca. 4.5 cm. long; style ca.
7 mm. long, apically curved, ovary puberulent; capsule not seen.
Libertad: Bombamarca, W. slope of eastern Cordillera, alt. 10,000
ft., Carricker s.n., Sin. Loc: Ruiz & Pavon s.n.
Calceolaria lasiocalyx Pennell, Proc. Acad. Nat. Sci. Phila. 97:
168, fig. 14b. 1945.
Herbaceous at least on upper part of the plant, stems lax,
branched, stem and branches pubescent with septate hair, pubes-
cence glandless and sometimes reflexed. Leaves petiolate, lance-
ovate, obtuse to subacute, upper 3-5 cm. long and 1.5-2.5 cm. wide,
glandular-punctate above, densely to very densely pubescent above
and beneath, crenate-dentate ; petioles 6-7 mm. long, densely pubes-
cent; inflorescences a pair of simple or once-branched cymes from the
axils of the uppermost pair of leaves; peduncles up to 2.5-4.0 cm.
long, pedicels abruptly much shorter, flower stalks septate-pubescent ;
bracts and bractlets usually lacking; calyx lobes ca. 5 mm. long,
ovate, obtuse, pubescent externally; corolla yellow, externally, prox-
imally or over its entire 10-13 mm. length finely glandular-pubescent
(fide Pennell), upper lip 2-3 mm. long, lower lip 10-12 mm. long,
saccate about half its length, with a tuft of white hairs internally at
base; anthers 2.5-3.0 mm. long, cells equal, a little longer than wide,
widely divergent; style ca. 1.5 mm. long, basally frequently glandu-
lar-puberulent; capsule ovoid-acute, glandular-puberulent, longer
than the calyx.
Cuzco: Tio to Murayaca, Marcapata Valley, Prov. Quispicanchis,
alt. 1,200 m., Vargas 5199.
FLORA OF PERU 603
The type Vargas 3760 also collected in the Marcapata Valley.
Calceolaria leptantha Pennell, Proc. Acad. Nat. Sci. Phila. 97:
163. 1945.
Climbing or sprawling shrub, stem finely glandular-pubescent.
Leaves on glandular-pubescent petioles that are mostly 12-20 mm.
long, thickened at base and connate across the nodes; blades ovate,
mostly 3-7 mm. long and 2.2-4.8 cm. wide, margins dentate, apex
acute, base lightly cordate, tissue above pilose, beneath reticulate-
veined, finely pubescent, especially on the veins and sparsely gland-
dotted; inflorescences compound cymes; basal peduncles 5-8 cm.
long; bracts petioled, reduced, leaf -like, but hardly cordate; inter-
mediate peduncles and bractlets very reduced, all flower stalks glan-
dular-pubescent; pedicels 15-35 mm. long; sepals glabrous, ovate,
acute, entire to few-toothed, ciliolate, 3-7 mm. long; corolla yellow,
externally essentially glabrous, upper lip ca. 2-3 mm. long, lower lip
15-28 mm. long, internally pubescent; anthers glabrous, cells equal,
divaricate, from little longer than wide to almost twice as long; style
ca. 1.5 mm. long, curved; capsule ovoid-conic, 6-7 mm. long, glan-
dular-puberulent.
Cuzco: Near Rio Paucartambo, below Paucartambo, alt. 2,900-
3,200 m., Pennell 13973 (type); Que"brada de Llulluchayocc, alt.
3,100 m., Vargas 4293', 5 km. NE. of Paucartambo on road to Pilla-
huata, alt. 2,900 m., Vargas and Ugent 4409.
Calceolaria leucantha Edwin, Phytologia 19: 384. 1970.
Differing from C. ballotifolia Kranzl. to which it is very closely
related in the following ways: wood and foliage glabrous, leaves
0.8-5.5 cm. long and 0.5-1.6 cm. wide, densely red-punctate beneath;
lower lip of the corolla saccate only about a third its length.
Tall shrubby climber, glabrous except for the peduncles, pedicels
and calyces, older wood unisulcate. Leaves ovate, very coarsely
dentate, asymmetrical, acute at apex, obtuse to sometimes cordate
at base, short-petiolate; inflorescences dichasiate or corymbose, few-
flowered, terminal on the branches or in upper leaf axils; basal pedun-
cles 3-5 times longer than the remaining flower stalks; calyx tube and
the lower part of the lobes pubescent; corolla white, drying yellow.
Amazonas: above Colcamar, rocky stream in mountain forest,
alt. 3,300 m., Pennell 15634-
604 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria linearioides Edwin, Phytologia 19: 385. 1970.
Shrub, erect or apically nodding, stems and branches densely
puberulent or sometimes also glandular-puberulent. Leaves puberu-
lent on both surfaces, more so beneath, blades short-petiolate, broadly
linear or narrowly elliptically ovate, 1.0-2.5 cm. long and 4-8 mm.
wide, margins few-serrulate on the upper half, often thinly revolute,
apex acute or subacute, base acute or long-acute, venation visible,
impressed above, elevated beneath; petioles ca. 1 mm. long, vesture
as the blades, those of the stem and principal branches swollen and
often with reduced leaves or abbreviated shoots in the axils; inflores-
cences racemose or true racemes, borne in upper leaf-axils; pedicels
puberulent, ca. 1-2 cm. long; interfloral leaves reduced, otherwise as
the cauline; calyx lobes sparsely puberulent on both surfaces or in-
ternally glabrous, broadly ovate, acute 4-5 mm. long, almost as wide
near the base; corolla yellow, sometimes red marked within near the
base, upper lip 2-3 mm. long, puberulent or papillose, lower lip glan-
dular-puberulent beneath on the non-saccate proximal part or papil-
lose beneath along its length, sometimes with a tuft of long white
hairs at the base within, ca. 11-15 mm. long, saccate a little less than
half its length; anthers large for the corolla, 4-5 mm. long, cells un-
equal, 2-3 times longer than wide, divaricate; style 2-3 mm. long,
apically curved, ovary puberulent or sparsely glandular with fugitive
glands; capsule ovoid, a little longer than the calyx, puberulent, nar-
rowed to a long neck, apex obtuse.
Cajamarca: Dry banks (volcanic soil), alt. 2,900 m., 15-20 km.
above Cajabamba, Pennell & Ferreyra 14855. — Pasco: Yanahuanca,
northwestern slope, alt. ca. 10,000 ft., Macbride & Featherstone 1184
(type).
Very close to Calceolaria linearis R. & P., differing only as in the
key. The new species is proposed also because of the distance be-
tween the two collections, possibly indicating its wider occurrence in
North Central Peru.
Calceolaria linearis R. & P., Fl. Peruv. 1: 19. 1798. C. tenui-
folia Pennell, Proc. Phila. Acad. Nat. Sci. 97: 155. 1945.
Stems up to 10 dm. tall, branched and densely foliaceous, pubes-
cence absent or when present variable. Leaves linear or narrowly
lanceolate, minute to about 3 cm. long, from less than 1 mm. to
about 3 mm. wide, margin revolute, entire, costa prominent beneath
apex usually acute, often hirtellus or pulverulent or tomentose on
one or both surfaces, occasionally glabrous; flowers in usually elon-
FLORA OF PERU 605
gate, dichasiate inflorescences, on the apices of the branches or axil-
lary, occasionally solitary or in pairs; peduncles usually glutinous-
pilose, apically bracteate; pedicels pilose; bractioles at apex of inter-
mediate flower stalks; calyx segments broadly ovate, 4-5 mm. long
and 3-4 mm. wide, obtuse, entire, pilose on both surfaces, often
yellowish, even when dry; upper lip of the corolla minute, much
shorter than the sepals, lower lip minutely pubescent, 10-15 mm.
long, the sac more than half the length of the lip; anthers on very
short, ca. 1.0-1.5 mm. filaments, cells usually unequal, less often sub-
equal or equal, about 3 times longer than wide, divaricate; style very
short ca. 1.0-2.0 mm. long, ovary glabrous or more often glandular-
puberulent; capsule 4-7 mm. long and 2.0-3.5 mm. wide at base,
usually glandular-puberulent, narrowly ovoid below with a long,
abruptly narrowed upper part.
Citations selected from the copious material at most major
herbaria.
Ancash: Above Chiquian, alt. 3,500-3,600 m., Pennell 15417;
north of Chancos, alt. 3,250 m., Pennell 15322; toward El Huascaran,
north of Yungay, alt. 2,850-2,900 m., Pennell 15423; Jalca slopes
and meadow, Cerro San Cristobal, north of Huaraz, alt. 3,450-
3,550 m., Pennell 15449. — Ayacucho: between Huanta and Hacienda
Pagora, alt. 4,200 m., Killip & Smith 23298; Prov. Huamanga, alt.
3,500 m., Weberbauer 5476. — Cajamarca: Cerro Tinaya, alt. 3,150 m.,
Ochoa 1688; Along stream east of Cord, de Cumulloa, to Celendin,
alt. 3,400 m., Pennell 15160; Thickets, mtn. north of Celendin, alt.
2,700-2,900 m., Pennell 15206; Ridge, rocky limestone, 11 km. south
of Cajamarca, alt. 2,750-2,850 m., Pennell & Anderson 15076; 15-
20 km. above Cajabamba, alt. 2,900 m., Pennell & Ferreyra 14855;
10 km. above Cajamarca, alt. 3,300 m., Pennell & Ferreyra 14866;
2-3 km. above Huamachucho, alt. 3,200 m., Pennell & Ferreyra 14860;
West of Llacamora, chert hill, alt. 2,500-2,600 m., Pennell & Ferreyra
14937; Open jalca, Paso de Gavilan south of Cajamarca, alt. 3,100
m., Pennell & Reichlin 15032. — Cuzco: Jucay, Soukup 854. — Huan-
cavelica; Rauh-Hirsch 3473; Huancavelica, alt. 3,798 m., Soukup
2781 ; Prov. Huancavelica, north of Huancavelica, calcareous rocks,
alt. 3,700 m., Stork & Horton 10827; Bunbunya, cerro west of Co-
naica, alt. 3,680-3,700 m., Tovar 210. — Huancayo: Incahuasi, alt.
4,000 m., Soukup 3158. — Junin: Ocopa, alt. ca. 3,300 m., rocky
stream bed, Killip & Smith 21979; Concepcion, alt. 3,300 m., Ochoa
733; Huancayo, alt. 4,000 m., Ledig 21; Concepcion, Huancayo, alt.
606 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
10,800 ft., Sandeman 4918; Soukup 1981. — La Libertad: Between
Trujillo and Huamachuco, alt. 2,900-3,300 m., Ferreyra 3013; Otuz-
co, Agallpampa, alt. 3,270 m., Lopez 0328; S. of Otuzco, alt. 2,500 m.,
Pennell & Ferreyra 14820; km. 82, west of and below Agallpampa,
alt. 2,900 m., Pennell & Ferreyra 14831; 14832; above Motil, alt.
3,200 m., Pennell & Ferreyra 14881 ; Santiago de Chuco: above Cachi-
cadon, alt. 2,800 m., Stork & Horton 9974- — Sin. Dept.: Concepcion:
Ingenio, Soukup 3670. — Sin. Loc. Ruiz & Pavon s.n.
Calceolaria lobata Cav., Icon. 5: 26, t. 443, f. 1. 1799.
Subshrub, stems ascending, woody at base with vesture of stalked,
red glands, internodes from much longer to shorter than the leaves,
stems usually few-branched, branches thin. Leaves long-petiolate,
very broadly ovate in outline, base cordate to less often truncate,
usually 7-lobed, occasionally 5- or 9-lobed, lobes irregularly dentate,
apical lobe acute, size very variable from about 1 to 10 cm. long and
almost as wide, usually densely pilose on both surfaces; petioles
mostly 1-5 cm. long, on the largest leaves even longer, frequently
from half to as long as the blade; inflorescences on long basal pedun-
cles, these often up to 10 cm. or more and frequently aphyllous except
at their summit where a pair of leaf -like bracts are produced, corym-
bose, few- to many-flowered; peduncles, pedicels, calyces and ovaries
densely glandular-pilose; calyx lobes ovate, 4-6 mm. long and 3.5-
4.5 mm. wide; corolla yellow, usually purple marked internally, upper
lip of the corolla much shorter than the calyx lobes, lower lip ca. 14-
18 mm. long, saccate less than half its length, distally upturned; fila-
ments exceeding the lower lip, anther cells equal, apically fused and
diverging to the base, each pair of cells sagittate or almost so, large
for the corolla orifice; style ca. 2 mm. long; capsule ovate to broadly
ovate, abruptly narrowed to a long neck, glandular-puberulent, a
little longer than the calyx, drying violet-red.
Ancash: Rocky wall, Paso de Fortale'za, alt. 3,700 m., Pennell
15262; Alt. 3,800-3,850 m., Pennell 15390; 15391; above Chiquian,
alt. 3,500-3,600 m., Pennell 15416; W. slope of Cerro Negra, below
Paso de Collan, Pennell 15480; Ocros, alt. 3,200 m., Weberbauer 5804-
— Arequipa: Que"brada de San Lazaro, above Arequipa, alt. 9,800 ft.,
Munz 15516; Volcan de Misti, base of volcano, rocky wall of canyon,
canyon, alt. 3,300-3,400 m., Pennell 13226; 14273; Picchu-picchu,
near moisture, alt. 10,500 ft., Sandeman 4015; Alt. 12,000 ft., Staf-
ford 669; Chachani, amongst rocks in deep ravine, alt. 11,000 ft.,
FLORA OF PERU 607
Stafford 596; Chivay, alt. 3,900 m., Weberbauer 6890.—Ayacucho:
Puquio- Valley, alt. 3,400 m., Rauh-Hirsch P439. — Junin: Prov.
Tarma, rocky ground between Tarma and La Oroya, alt. 3,600-
3,700 m., Ferreyra 3830. — Lima: Prov. Huarochiri, Chicla, steep
stony slope, alt. 3,700 m., Asplund 11249; Stony ground near Casa-
palca, between San Mateo and Matucana, Ferreyra 3947; Rio Blanco,
alt. 3,000-3,500 m., open hillside, Killip & Smith 21651; 21711;
21731 ; Rio Blanco, bunch grass, stony slopes, about 15,000 ft., Mac-
bride 2976; Alt. 12,000 ft., leaning against rock faces, Macbride &
Feather stone 708; near Antaicocha, Cerro Colorado, E. of Canta, alt.
3,500-4,000 m., Pennell 14625; Huaros, rock ledges on open slope,
alt. 3,500-3,600 m., Pennell 14738; Rocky bank above Rio Blanco,
alt. 3,500-3,700 m., Pennell & Maldonado 15937; Rio Blanco-Casa-
palca, alt. 3,600-4,000 m., Vargas 5265. — Moquegua: Saylapa near
Carumas, humid rocks, alt. 3,500 m., Weberbauer 7342.
Calceolaria longiinternodia Edwin, Phytologia 19: 386. 1970.
Tall herb, branching near the apex, branches swollen at base,
nodes, especially the lower, several times (ca. 4-5) longer than the
largest leaves, stems drying dark- to purple-brown, striatulate,
mostly glabrous except hirsute-puberulent in the inflorescence.
Leaves opposite or ternate, blades narrowly ovate to lance-ovate,
densely tomentose and pale-green beneath, glabrous, darker and dull-
green above, very variable in size, the major leaves 4-6 cm. long and
usually 1.5-3.0 cm. wide, margins very coarsely dentate, apex long-
acute, variable at base; petioles up to 6 mm. long, densely tomentose,
often with abbreviated shoots or reduced leaves at base in the axils;
peduncles and pedicels densely tomentose, the longest peduncles ca.
4.5 cm. long, once or twice branching, forming dichasia, pedicels
about half as long or less than half as long as the peduncles; inflores-
cences on upper part of the stem, terminal on the branches; calyx
lobes broadly ovate, 7-9 (-10) mm. long, acute, entire, externally
densely puberulent, internally puberulent with a row of hairs near
the margin; corolla yellow, 25-30 mm. long, lower lip externally
pustulate, internally with a tuft of hair at the base, upper lip ca.
4-6 mm. long, lower lip 21-24 mm. long, saccate less than half its
length; anthers on short filaments, cells equal or a little unequal,
sometimes dimorphic in a flower, mostly at least 2 times longer than
wide, less often only little longer than wide to almost as wide as long,
each pair of cells 4.5-5.5 mm. long; style a little thickened, erect,
ca. 4 mm. long, ovary glandular.
608 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Ancash: Herb, banks near stream Tinco, alt. 3,400-3,650 m.,
E. slope of Cordillera Negra opposite Huaraz, Pennell 151+68 (type).
Close to C. impressifolia Edwin, from which it differs in leaf and
anther shape and C. hirsutula Pennell and C. bicrenata R. & P. The
former has leaf margins with very fine teeth and the latter differs
in leaf-shape. C. longiinternodia differs from all other Calceolarias in
the combination of dimorphic anthers, leaves about three times as
long as wide with coarsely dentate margins and calyx lobes usually
7-9 mm. long with a distinct row of hairs internally near the margins.
Calceolaria lopezii Edwin, Phytologia 19: 387. 1970.
Soft erect herb, branches and leaves opposite, stems, branches
and flower-stalks more or less densely pubescent with gland-tipped
hairs, glands small and frequently fugitive in age, hairs white of in-
termixed, varying sizes, upper nodes bearing densest vesture. Upper-
most leaves sessile, others short-petiolate, blades ovate, mostly 2.8-
4.1 cm. long, sometimes shorter, and 2.0-3.8(-4.4) cm. wide, some-
times narrower, dentate-denticulate, apex acute, base obtuse to trun-
cate, occasionally cordate, subglabrous to sparsely puberulent with
2-celled hairs, mostly concentrated on the margins and the veins
above, chartaceous, a little paler beneath than above, both surfaces
dull green; petioles only 1-2 mm. long, rarely a little longer, vesture
denser than the blades; inflorescences terminal on stem and axillary
or terminal on the branches with a pair of reduced bract-like leaves
at the base of the peduncle, simple or lax branched dichasia; calyx
lobes externally sparsely pubescent, margins ciliate, internally gla-
brous, main veins parallel, ovate to elliptical, 2.3-3.0 mm. long and
1.2-2.0 mm. wide, acute; corolla yellow, 12-22 mm. long, almost
completely glabrous, upper lip more than half the length of the lower,
7-8 mm. long, lower lip 11-13 mm. long and 7-9 mm. wide, saccate
about half its length; filaments about 4 mm. long, flattened, with one
main vein, anthers bilocular, locules very small, each anther about
1 mm. long, usually smaller, a little longer than wide; style glabrous,
elongate, 4.5-5.0 mm. long, glabrous, thin, stigma very small, scarcely
wider than the style, ovary drying dark red; capsule not seen.
Libertad: Cerro Campana, Prov. Trujillo, alt. 500 m., Aug. 9,
1948, Lopez 0221 (US); same location August 21, 1949, alt. 420 m.,
Angulo 0698.
Calceolaria luteocalyx Edwin, Phytologia 19: 388. 1970.
Climbing shrub, main branches and stem canaliculate at least
part way, young wood densely, older wood sparsely puberulent.
FLORA OF PERU 609
Leaves petiolate, blades ovate or elliptic-ovate, mostly 2.0-4.0 cm.
long and 1.5-2.2 cm. wide, usually not twice as long as wide, nar-
rowed at both ends, margins coarsely, irregularly serrate-dentate;
petioles densely pubescent, mostly 0.8-1.8 cm. long, basally little
thickened and thinly connate across the nodes that are frequently
much longer than the leaves; inflorescences arising from the upper
leaf-axils, basally dichotomously branched, ultimately 3-flowered
cymes, basal branching sometimes wanting, then the inflorescences
simple or once-branched cymes; bracts and bractlets foliose, reduced;
basal peduncles 5-9 cm. long, secondary peduncles ca. a half to two-
thirds as long as the basal; pedicels little shorter than secondary
peduncles; flower stalks densely puberulent; calyx yellow, lobes sub-
glabrous, sometimes broader than long, broadly ovate, obtuse, 7-
8 mm. long and 7-9 mm. wide, margins sinuate or irregularly broadly
crenulate and sparsely ciliolate, calyx tube sparsely to densely pu-
berulent; corolla yellow, wholly glabrous, upper lip 3 mm. long with
hood 1 mm. long; lower lip 18-22 mm. long, saccate less than half its
length, proximal, non-saccate part only little narrower than distally;
anther cells equal, ca. 2 mm. long, divaricate, subsessile, filaments
ca. 0.3-0.5 mm. long and about as wide; style ca. 2.5-3.0 mm. long,
thickened, ovary pubescent, flattened; capsule not seen.
Known only from the type collection.
Amazonas: Forest, alt. 3,000 m., Cerro Puma-urco, SE. of Cha-
chapoyas, Pennell 1551+9 (PH).
This species differs from all other taxa in the genus by virtue of
the following combination of characteristics: calyx yellow, with lobes
often wider than long, non-saccate part of the lower lip of the corolla
only little narrower than the saccate part and the anthers subsessile.
Calceolaria macrocarpa Pennell, Proc. Acad. Nat. Sci. Phila.
97: 161. 1945.
Branched shrub forming clumps, stems scabrous. Leaves short-
petiolate, elliptical, 1.5-2.0 cm. long and 0.8-1.0 cm. wide, acute,
dentate, with about 5 pairs of sharp teeth, pilose above, puberulent
and gland-dotted beneath, base cuneate; petioles 3-4 mm. long,
basally thickened; inflorescences in pairs of 1-3-flowered cymes; pe-
duncles 2.5-4.5 cm. long; bracts reduced foliage leaves; pedicels 9-
13 mm. long; flower stalks finely scabrous; sepals ovate to lance-
ovate, acute, 7-8 mm. long, minutely scabrous; corolla yellow, ex-
ternally proximally puberulent, upper lip 1.5-2.5 mm. long, lower lip
610 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
ca. 20 mm. long, internally glabrous, sac less than half its length;
anther cells equally divaricate, ca. 2 mm. long, about two-thirds as
wide as long; style 1.2 mm. long, curved apically; capsule narrowly
conic, 10 mm. long, obsoletely glandular-pubescent.
Cuzco: Checacupe, open rocky bank, alt. 3,400-3,450 m., Pennell
13535 (type); Cook & Gilbert 93 pp., from Sicuani reported by Pen-
nell but unknown to me.
Calceolaria maculata Edwin, Phytologia 19: 389. 1970.
Climbing herb with opposite branches, young stems and branches
densely pubescent, older densely puberulent, internodes longer than
the leaves. Leaves short-petiolate, densely eglandular, ferrugineous-
tomentose beneath, densely pubescent above with shorter, frequently
glandular hairs, ovate, grossly dentate, apex acute, base broadly
obtuse to subtruncate, mostly 1.8-3.0 cm. long and 1.0-1.9 cm. wide;
petioles up to ca. 8 mm. long, densely pubescent, those of the stem
leaves thick, indurate, woody, persistent, with the branches forming
swollen nodes; flowers clustered at the apices of the upper branches
in irregular cymose or corymbose inflorescences; bracts and flower
stalks densely pubescent; basal peduncles up to 3 cm. long, interme-
diate peduncles and pedicels much shorter, frequently less than half
as long; calyx lobes ovate, acute, 5.0-6.5 mm. long, almost as wide
at base, externally densely pubescent, internally with a thick row of
short hairs along the margins and a few, scattered hairs on the lower
third; corolla yellow, the internal surface of the lower lip red-brown
marked or spotted and with a tuft of long white hairs at the base,
mostly 19-23 mm. long, less than half as wide, upper lip up to 2.5 mm.
long, lower lip beneath, externally sparsely puberulent, 16.5-22.0
mm. long, saccate only about a third to a fourth its length, the sac
at the distal end with 2 short sulcations therefore appearing 3-lobate;
stamens glabrous, filaments ca. 1.5 mm. long, anthers ca. 3 mm. long,
cells equal, almost twice as long as wide; style 1.5-2.0 mm. long, erect
or little curved at apex, glabrous, most often thickened, ovary glan-
dular with sessile, golden glands; capsule ovoid, obtuse, glandular,
shorter than the calyx.
Known only from the type collection.
Cajamarca: Meadow, E. of Celendin, alt. 2,500-2,700 m., Pen-
nell 15215 (US).
This very distinct taxon is probably close to C. aperta Edwin,
but easily distinguished from it. The corolla is very unusual.
FLORA OF PERU 611
Calceolaria micrantha Pennell, Proc. Acad. Nat. Sci. Phila.
97: 176-177. 1945.
Annual, mostly glandular-pubescent herb, up to 30-40 cm. tall,
stem glabrescent near the base, internodes usually longer than the
leaves. Lower leaves ovate in outline, 1.5-2.7 cm. long and almost
as wide, usually pinnatifid or with 2 pairs of pinnae, on petioles 1.5-
2.0 cm. long, upper leaves and bracts lanceolate, about 1.5 cm. long
and 3-6 mm. wide, on petioles about 5 mm. long; petioles connate
across the nodes; inflorescences compound, one-sided cymes; pedicels
reduced apically, 2-20 mm. long; basal peduncles 35-50 mm. long;
sepals ovate, 4-5 mm. long, acute to acuminate, irregularly sparsely
denticulate, ciliate, externally with gland-tipped pubescence; corolla
yellow, externally glabrous, upper lip 2-3 mm. long, lower 5-8 mm.
long, saccate most of its length; connective thickened, longer than
the sterile lower anther, shorter than the fertile upper anther; style
glabrous 1.0-1.5 mm. long; capsule about as long as the calyx, ovoid,
constricted to the conic neck, glandular-puberulent.
Ayacucho: Ccarrapa, between Huanta and Rio Apurimac, alt.
ca. 1,500 m., Killip & Smith 22381. — Cuzco: Near Rio Yanamayo,
below "Pillahuata," alt. 2,200-2,400 m., Pennell (type).
This is a forest species of southern Peru which probably has
greater distribution than the two collections indicate. It is incon-
spicuous and easily overlooked.
Calceolaria millefoliata Kranzl. in Engler, Bot. Jahrb. 50:
Beibl. Ill, 72. 1914.
Much branched, erect shrub, bark glabrescent below and glandu-
lar-pilose above. Leaves very numerous, petiolate, blades up to
5 mm. long and 4 mm. wide, margin with 5-7 obtuse lobes and a
little revolute, densely white-pilose above and beneath, the hairs
curly and loosely matted; petioles up to almost 1.5 mm. long, red-
dish; flowers solitary or in few-flowered dichasia at the apices of
stems and principal branches; flower stalks thin, densely pilose, 1.0-
2.5 cm. long; calyx lobes ovate, 4.0-5.0 mm. long and 2 mm. wide at
base, acute; corolla orange-red, entirely glabrous, 1.5 mm. long and
1.0 cm. wide distally, upper lip 3 mm. long, hooded, lower lip 1.2 cm.
long, saccate more than half its length, the transverse orifice 2 times
as wide as long; filaments very short, anthers ca. 2 mm. long, up to
almost 2 times longer than wide; style about as long as the stamens;
ovary papillose. (Description after Kranzl.)
612 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Only two collections seen, the type material from the department
of Ayacucho and the other from Arequipa. The distance between
collections, about 350-400 km., indicates strongly that future speci-
mens will be found in the area now separating them.
Arequipa: Rocky canyon above Arequipa, alt. 3,300 m., Pennell
13212. — Ayacucho: between Rio de Lomas and Rio de Yanca in
Prov. Parinacochas, alt. 2,900-3,000 m., Weberbauer 5770.
Calceolaria myriophylla Kranzl. in Engler, Pflanzenr. 4(257c):
88. 1907.
Shrub, much branched, branches thin often curved, older wood
glabrous, light or dark grey, younger densely white-tomentose.
Leaves minute, very numerous, short-petiolate, ovate and oblong,
mostly up to 5 mm. long and 3 mm. wide, occasionally reaching al-
most 10 mm. long and 6 mm. wide, margin plane or little revolute,
entire or sometimes crenulate-crenate, minutely white-pilose with
impressed nervation above, long-white-pilose with prominent ner-
vation beneath, apically obtuse; inflorescences of 3-flowered dichasia
or flowers solitary in upper leaf-axils, dichasia sometimes crowded;
pedicels thin, ca. 1.0 cm. long, sometimes shorter or longer and with
the calyces minutely papillose; bracts minute; calyx 4.5-6.0 mm.
long, pubescent, lobes ovate, 3.5-4.5 (5.0) mm. long, 3.5-4.0 mm.
wide, puberulent on both sides, obtusish; corolla yellow, minutely
pilose or glabrous, mostly 2.0-2.8 cm. long, upper lip shorter than
the calyx, lower saccate less than half its length, orifice rectangular;
filaments 1.5-2.5 mm. long, anther cells equal, up to 2 times longer
than wide, variable in size, usually large for the corolla, divaricate;
style short, ovary glabrous or usually sparsely glandular, drying shin-
ing, red-black; capsule glandular or glabrous, longer than the calyx
and filling it tightly, spherical or very broadly ovoid with a long-
acuminate beak.
Well represented in Dept. Cuzco, unknown elsewhere.
Cuzco: Rocky slope, alt. 3,300 m., Cardenas 4186; Ollantaitambo,
alt. ca. 3,000 m., Cook & Gilbert 377, 496, 627; "Puru-puru," Rio
Japfi, alt. 3,500-3,600 m., Herrera 776; "Pulu-pulu," Huasco, alt.
3,200 m., Herrera 3032; Cusipata, alt. 3,300 m., Pennell 13537; Sacsa-
huaman, alt. 3,500-3,600 m., Pennell 13569, 13588; San Sebastian,
alt. 3,300-3,400 m., Pennell 13597; Ollantaitambo, alt. 3,100-3,300
m., Pennell 13676: Pisac, rocky canyon, alt. 3,400-3,500 m., Pennell
13720; Qurquijana, alt. 3,200 m., Pennell 14208; Sajsaihuaman, alt.
3,600 m., Vargas 229; Sin Loc., Watkins s.n.
FLORA OF PERU 613
Calceolaria myrtilloides Kranzl. in Fedde, Rep. Spec. Nov. 1:
99. 1905.
Erect essentially glabrous shrub, wood frequently drying black
or dark brown. Leaves short-petiolate, ovate or elliptic-ovate,
mostly 1.5-3.0 cm. long and 1.2-2.2 cm. wide, dark green with ob-
scure venation above, light yellow-green with obvious, reticulate
venation beneath, obscurely dentate-serrate, narrowed to both ends;
petioles ca. 3-6 mm. long; inflorescences few-flowered, corymbose;
flower-stalks minutely papillose; bracteoles lanceolate or lance-ovate;
calyx lobes ovate, acute, 5.0-7.5 mm. long, externally glabrous, in-
ternally with a row of hairs along the margins and with scattered
puberulence; corolla yellow, variable in size, glandular-puberulent
or -pilose or occasionally only very sparsely so, upper lip from 3 mm.
long to only little shorter than the calyx lobes, lower lip from 20-
30 mm. long, sac round, more than half the length of the lip; anther
cells equal or unequal, at least twice as long as wide, each pair of
cells divaricate, 3.5-4.5 (-5.0) mm. long, filaments short, thickened,
especially at the base; styles abruptly incurved, ca. 2.3 mm. long,
ovary densely glandular-pubescent.
Ancash: Near Shincash above Chiquian, Prov. Bolognesi, alt.
3,885 m., Ferreyra 5848; Between Tallenga and Pachapaque, Bo-
lognesi, alt. 3,500-3,600 m., Ferreyra 7477; Near Ocros, Prov. Caja-
tambo, alt. 3,500-3,700 m., Weberbauer 2689 (fragment of type).—
Libertad: Toyabamba, alt. 10,000-12,000 ft., Carricker s.n.
Calceolaria nivalis HBK., Nov. Gen. et Sp. 2: 381. 1817.
Shrub, essentially glabrous, stems, branches and upper surface
of the leaves shining, even when dry, plants frequently glutinous.
Leaves short-petiolate, blades ovate or lance-ovate, rarely almost
lanceolate, subcoriaceous, largest up to 6 cm. long and 2 cm. wide,
margin dentate-serrate, apex acute, narrowed to base, obviously re-
ticulate-venose beneath; petioles 4-6 mm. long; inflorescences cymose-
subumbellate, up to 10-flowered, usually less; pedicels black-hirtellus,
rarely elongate; calyx lobes ovate, acute, 5-6 mm. long and 3-4 mm.
wide, sparsely glandular-pubescent externally, most often with a row
of yellow hair internally along the margin; corolla yellow, glabrous,
upper lip longer than the calyx, ca. 7-8 mm. long, lower lip up to
12 mm. long, saccate less than half its length; filaments very short,
anther cells equal, little longer than wide, longer than the filament.
Reported (by Kranzlin) in Peru. No material seen. The report
may be erroneous. This species is well collected in Ecuador.
614 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria oblonga R. & P., Fl. Peruv. 1: 15, t. 24, fig. b. 1798.
C. punicea R. & P. Ic. t. 24, fig- «•
Herb, sometimes woody at base, leaves, especially on the veins
beneath, calyx lobe margins and often also the stems and branches
beset with multicellular hair. Leaf blades very short-petiolate, peti-
oles up to 4 mm. long, densely pubescent, blades ovate, acute at apex,
obtuse or subobtuse at base, irregularly serrate-serrulate; inflores-
cences usually paired, from the axils of the uppermost leaves, few-
flowered, corymbose or dichasiate (ex desc.); flower stalks densely
pubescent with multicellular hair, basal peduncles 3-4 times longer
than the remaining peduncles and pedicels; interfloral leaves much
reduced; flowers small, 12-18 mm. long; calyx lobes, margins ex-
cepted, externally, sparsely pubescent, internally glabrous, ovate,
acute, 3.5-4.0 mm. long and 2.5-3.0 mm. wide, veins obvious within;
corolla yellow to pale yellow, upper lip sometimes almost white, 4.5-
6.5 mm. long, hood very short, externally glandular-pubescent, ori-
fice internally surrounded by a tuft of long, white hairs, lower lip
more or less densely pubescent externally with short, gland-tipped
hairs, 7-12 mm. long, from little more to little less than twice as long
as the upper lip, saccate over most of its length; anthers very short-
stalked, locules equal, divaricate, each pair 2.0-2.5 mm. long, only
little longer than wide; style very short, ca. 1.0-1.3 mm. long, thick-
ened, erect, ovary pustulate-glandular.
Cuzco: Valley of Cosnipata, between Mistiana and Deros, alt.
ca. 800 m., Scolnik 889. — Junin: Pichis Trail, Porvenir, alt. 1,500-
1,900 m., dense forest, Kittip & Smith 25952.
The distance between collections of this distinct species indicates
its occurrence in the area between and possibly even beyond them.
Calceolaria padifolia HBK., Nov. Gen. et Sp. 2: 380. 1817.
C. deflexa Willd. ex Link in Spreng., Bot. Jahrb. 1(3): 52. 1820.
Slightly glutinous, branchlets excepted, erect, much branched
shrub. Leaves on petioles 5-10 mm. long, subconnate (ex Kunth,
pilose in the canal) ; blades ovate-lanceolate, base rounded or short-
acute, apex acute and acuminate, dentate, reticulate-venose and
paler beneath than above, up to 6 cm. long and 2 cm. wide; inflores-
cences subdichasiate, basally equally 3-branched, the ultimate branch-
ing dichasial; pedicels about 2 cm. long, all flower stalks thin; calyx
segments ovate, obtuse, 6 mm. long and 3 mm. wide at base; corolla
yellow, glabrous, upper lip longer than the calyx, 1 cm. long; lower
FLORA OF PERU 615
lip 1.2 cm. long; stamens very short, anthers much larger than the
filaments. (Description after Kranzlin.)
Reported from Piura: Ayavaca (Ayabaca) Humboldt & Bonpland
3586; photo seen.
Calceolaria pallascensis Edwin, Phytologia 19: 390. 1970.
Erect, strictly branching from about the middle, mostly densely
white-lanate, nodose shrub, stems very densely lanate above, vesture
decreasing basally, thickened at the closely spaced nodes. Leaves a
little shorter to a little longer than the internodes, with the branches
and flower stalks strictly pointing upward, almost parallel to the
main stem, sessile, linear or aciculiform, mostly 0.5-1.5 cm. long up
to 2 mm. wide, densely lanate in the groove of the costa and varying
to densely puberulent-pustulate on the tissue, the puberulence mi-
nutely gland-tipped, the bases of the outermost leaves at each node
thickened, mostly with very abbreviated shoots in the axils, the
leaves appearing clustered, occasionally the shoots a little elongated,
margins revolute, entire, apex hammate, tissue thickened, venation
obsolete; inflorescences terminal on the main stem and branches and
in the uppermost leaf-axils, simply cymose; pedicels 5-10 mm. long,
densely lanate; calyx tissue thickened, calyx lobes externally densely
lanate, internally glabrous or minutely glandular-puberulent, or with
a line of hairs near the margin, 3 segments broadly ovate, the fourth
segment ellliptically ovate, 4.0-4.5 mm. long and ca. 3.0 mm. wide
or 5.0-5.5 mm. long and 2.0 mm. wide, all acute; corolla yellow, gla-
brous or often minutely puberulent, 7-11 mm. long, upper lip 3-4
mm. long, hooded but not obscuring the genitalia, lower lip saccate
about half its length, sac distally crenulate, internally glabrous; an-
ther cells large for the corolla on very short filaments, divaricate,
unequal and almost oribicular; style 2.5-4.5 mm. long, curved, ovary
granular; fruit not seen.
Known only from the type.
Ancash: Laguna de Pelagatos, Prov. Pallasca, alt. 4,300 m., Lopez
2400 (LA).
Unequal almost orbicular anther cells are rare in the genus. This
characteristic along with the strict branching, strict flower stalks,
very small corolla, simple inflorescence, dense woolly vesture and
very narrow leaves aggregated in nodose masses renders the taxon
quite distinct.
616 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Calceolaria palustris Sodiro ex Kranzl. Pflanzenr. 4(257c) : 27.
1907.
Repent to ascending annual herb, upper half of the stems usually
glandular-pubescent, becoming glabrous below, pubescence several
celled. Leaves variable, uppermost lanceolate, acute, few-dentate,
dentations deep, large acute, becoming pinnatifid to pinnate below
and ovate in inclusive outline, subglabrous to sparsely pubescent
above, puberulent beneath, sometimes only on the costa, lowest
leaves largest, up to 4 cm. long and 2.5 cm. wide, decreasing in size
to the apex of the stems; petioles flattened, mostly 1-2 cm. long,
those of the largest leaves very thinly connate across the stem;
flowers borne singly in the axils of the upper leaves; the appressed-
glandular-pubescent pedicels shorter to longer than the subtending
leaves; calyx lobes lance-ovate to occasionally ovate, acute, denticu-
late, 6-8 mm. long and 2.5-4.0 mm. wide, glandular-puberulent on
the margins, venation externally prominent; corolla pale yellow, up-
per lip 3-5 mm. long, lower lip occasionally drying creamy white,
externally glabrous, 8-15 mm. long, saccate most of its length; calyx
and corolla often purple-maculate; connective longer than the anther
cells, upper fertile, almost globular, lower cell sterile, rudimentary;
style ca. 1.0 mm. long; capsule as long as wide or more often wider
than long, 4 mm. long and 4-6 mm. wide, broadly ovoid, shorter than
the calyx, glabrous.
Cuzco: Near Rio Yanamayo, below "Pillahuata," alt. 2,000-
2,300 m., Pennell 14044', Prov. Paucartambo, Tambomayo, alt.
1,300 m., Vargas 7288. — Huanuco: Prov. Huanuco, Carpish, alt. ca.
2,750 m., Asplund 128^7; Cordillera Azul, alt. 1,600 m., Scolnik 1113.
— Puno: Prov. Sandia, Limbani, alt. 3,300 m., Vargas 9636.
Distributed from Colombia to Bolivia.
Calceolaria pavonii Benth., in DC., Prodr. 10: 211. 1846. C. per-
foliata R. & P., Fl. Peruv. 1: 15, t. 210. 1798, non L.f.
Erect, perennial, viscose-villose or viscose-puberulent herb. Peti-
oles winged, dilated at base, connate, subdentate; leaf tissue not ob-
scured by the vesture. Otherwise much like C. tomentosa R. & P.
with which it may be conspecific.
Amazonas: Prov. Chachapoyas: near Leimebamba, alt. 3,000 m.,
moist open clay soil (Cerro Calla-Calla?), Metcalf 30797; Ridge south-
east of Colcamar, alt. 2,100-2,200 m., Pennell 15595; 15600; Above
Colcamar, alt. 2,500 m., Pennell 15607; Mathews 3042. — Huanuco:
FLORA OF PERU 617
El Tambillo, camino a Panao, alt. 2,650 m., Scolnik 1053; Several
km. west of summit of Carpish, alt. 2,800 m., Stork & Horton 9899.
— Junin: Huancapistana, thickets and open woods, alt. 1,800-2,400
m., Killip & Smith 21+179; Prov. Tarma: Aqua Dulce, alt. 2,000 m.,
Woytkowski 37008.— Piura: Porculla, alt. 6,000-7,000 ft., Carricker
s.n. — Sin. Dept.: Yanano, alt. 6,000 ft., Macbride 1+923. — Sin. Loc.:
Ruiz & Pavon s.n.
Calceolaria pearceana Kranzl. Pflanzenr. 4(257C) : 107. 1907.
Vine or climbing shrub, occasionally a climbing perennial herb,
stem, branches and flower stalks usually brown-puberulent, puberu-
lence decreasing with age, main stem ultimately frequently glabrate
near the base. Leaves small for the plant, sessile or almost so, ovate,
reaching 2.5 cm. long and 1 cm. wide, mostly smaller, especially in
width, crenate, acute at apex, acute to cuneate at base, puberulent
or pilose on both surfaces or glabrous above, sometimes few-gold-
punctate beneath; inflorescences of few-flowered corymbs, terminal
on the branches; flower stalks thin, up to 2 cm. long; calyx lobes
ovate, 6 mm. long, acute, puberulent externally, few-gold-punctate
internally and often with a row of golden hairs along the margins;
corolla 14-20 mm. long, sparsely muricate with red puncticulations
and golden puncta, upper lip well developed, sometimes exceeding
the calyx, usually more than half as long as the calyx, yellow or
whitish, lower lip yellow, saccate more or less than half its length;
filaments short, anther cells drying black or brown, equal, walls
thickened, divergent to subsagittate, large for the corolla, less than
twice as long as wide to little longer than wide; style 4-6 mm. long,
little or not at all thickened, with the ovary, glandular punctate at
base; capsule ovate, when immature shorter than the explanate calyx.
Cuzco: Paucartambo to Tre's Cruces, Cerro de Cusilluyoc, thick-
ets, alt. 3,400-3,700 m., Pennell 13803; "Pillahuata," Cerro de Cusil-
luyoc, thickets above town, alt. 3,000-3,500 m., Pennell 13928;
forests above "Pillahuata," alt. 2,200-2,600 m., Pennell 13930;
among shrubs near Marcachea, Prov. Paucartambo, alt. 2,450 m.,
Vargas 11174- — Puno: Sandia, near Limbani, alt. 3,200-3,450 m.,
Metcalf 301+81; Sin. Loc.: Soukup 1228, no data.
The small corolla enables rapid separation of this taxon from its
near relatives, such as C. boliviana and C. atahualpae. Explanate
fruiting calyces are rare in the genus.
618 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Calceolaria phaceliaefolia Edwin, Phytologia 19: 390. 1970.
Subshrub dying back to near the ground or occasionally an herb
or small shrub, usually branching over much of its length, younger
stems puberulent, often with mixed glandular and eglandular hair,
older at length glabrate. Leaves petiolate, ovate in outline, very
variable in size, mostly 1-3 cm. long and 0.8-2.5 cm. wide, in herba-
ceous material up to 6 cm. long and 5 cm. wide, margins with 2-4
pairs of irregular lobes, these sometimes secondarily toothed, long-
acute at apex, obtuse to subtruncate at base, puberulent above and
beneath with gland-tipped hairs; petioles sometimes almost as long
as the blades, most often a third to a fourth the length of the blades,
glandular-puberal ent to -pubescent; inflorescences variable, corym-
bose to paniculate, when paniculate, the ultimate branchings cymose
or dichasiate, few- to many-flowered, usually terminal on the branches;
bractlets very reduced, leaf-like, ca. 0.5 mm. long, lobes short, 1-2
pairs; basal peduncles 3-10 times longer than the remaining flower
stalks, peduncles and pedicels gland-tipped-puberulent, sometimes
densely so; flowers variable in size, 22-40 mm. long when dry, up
to only half as wide; calyx lobes ovate, acute, 7-9 mm. long and 4-
6 mm. wide, internally and externally gland-tipped puberulent; co-
rolla yellow, 20-35 (-38) mm. long, externally sparsely to densely
very short-puberulent, the hairs gland-tipped, upper lip very short,
ca. 2-4 mm. long, not hiding the stamens and stigma; lower lip 20-
35 mm. long, distally saccate less than half its length, the proximal
part abruptly much narrower than the upcurving sac, internally long-
pubescent with white hairs at the base; stamens with filaments ca.
1.5-2.5 mm. long, anther cells unequal or very rarely equal, sagittate,
the larger of each pair about 3 mm. long and 1.0-1.5 mm. wide, the
other cell from subobsolete to about three-fourths as large, often
varying from flower to flower on a single plant, when almost lacking
reduced to a sterile flap at the apex of the filament, the stamen ap-
pearing uniloculate unless magnified (ca. 30 X); style erect, 2.5-
3.0 mm. long, ovary glandular; capsule ovoid, little shorter than the
calyx, apically narrowed, style persisting.
Ancash: Tranca, near Chiquian, alt. 3,220 m., Ferreyra 5691;
Capillapunta, mountain S. of Chiquian, alt. 3,540 m., Ferreyra 5732;
Mountain E. of Chiquian, alt. 3,500-3,550 m., Ferreyra 7334; 7342
(herbaceous); Marcara, Prov. Huaraz, alt. 3,000 m., Nunez 3234;
Stream bed, S. of Huaraz, alt. 3,100 m., Pennell 15263; Monterrey,
3 km. below Huaraz, alt. 3,000-3,100 m., Pennell 15309; Shrub,
bushy banks, alt. 3,200-3,500 m., N. of Chancos, Pennell 15320;
FLORA OF PERU 619
Bushy, rocky slope along stream, S. of Rio Santa, alt. 3,100-3,150 m.,
opposite Huaraz, Pennell 15328; Sandy banks, alt. 3,400-3,450 m.,
Cerro Shaurema, SE. of Huaraz, Pennell 15352; Low mountain
toward El Huascaron, N. of Yungay, alt. 2,850-2,900 m., Pennell
15^.22 (anther cells equal); Summit of Cerro Shaurema, SE. of Hu-
araz, alt. ca. 3,500 m., Pennell 154-28; Bed of Que"brada de Huaqui,
alt. 3,000-3,050 m., N. of Huaraz, Pennell 15436; Hills N. of Huaraz,
alt. 3,100-3,150 m., Pennell 15438; Jalca, E. of slope of Cordillera
Negra, opposite Huaraz, alt. 3,600 m., Pennell 15464- — Lima: Chicla
between San Mateo and Casapalca, alt. 3,700 m., Ferreyra 6515;
Rimac Valley, alt. 4,000 m., Rauh-Hirsch P1794-
This species is easily separated from the other members of the
genus by the combination of very unequal anther cells, deeply lobed
leaves, long petioles, and the irregularly varying inflorescence.
Calceolaria pinnata L., Mant. II. 171. 1771. C. pulchella
Salisb., Prodr. 89. 1796. C. bipinnatifida Phil., Fl. Atac. 36. 1860.
Annual, branching herb, sparsely to densely puberulent to occa-
sionally almost glabrous, stems sometimes repent. Leaves petiolate,
blades up to 7-10 cm. long, all pinnate, with 4-5 pairs of subopposite
pinnae, sometimes bipinnate or pinnae pinnatifid and often variable
in outline, usually irregularly coarsely dentate, uppermost pinnae
frequently alternate; petioles up to about 3 cm. long, mostly a little
shorter; inflorescences usually 3-flowered, terminal dichasia, rarely
in branched dichasia; pedicels 6-12 mm. long, glandular-pubescent;
calyx lobes ovate to broadly ovate, 4.5-7.0 mm. long and up to 5 mm.
wide, sparsely, minutely, gland-tipped puberulent, venation parallel,
conspicuous; corolla yellow, variable in size, 8-20 mm. long or even
longer, externally glabrous, upper lip shorter than the calyx, hooded,
3-6 mm. long; lower lip saccate two-thirds to three-fourths of its
length, sac expanded, little longer than wide; both cells of each an-
ther fertile, the upper larger, all cells exceeding the connective arm
in length; style ca. 1.5 mm. long, persisting on the capsule; ovary
sparsely glandular-puberulent; capsule broadly ovoid, pubescent as
the ovary, apically narrowed, little longer than the calyx and about
as long as wide.
Libertad: Prov. Trujillo, Cerro Campana — "Farolillo," alt. 420
m., Angulo 0699. — Lima: Amancaes, vicinity of Lima, loma vegeta-
tion, Asplund 13724 and alt. 380-420 m., Ferreyra 3969; Valley of
Amancaes, alt. 245-275 m., Mexia 4018; Cerro de Amancaes, alt.
200-600 m., Pennell 15925; Rock ledges on hills S. of Atocongo, alt.
620 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
200-600 m., Pennell & Anderson 159^2; Chancai, Lomas de "Lachai,"
among rocks, alt. 500 m., Vargas 4688.
This is one of the very few wholly low elevation species of Cal-
ceolaria in Peru.
Calceolaria pisacomensis Meyen, Reise 1: 469. 1830.
Herb or subshrub up to 50 cm. tall or even taller, newer branches,
foliage, flower stalks, calyx margins externally and calyx tissue in-
ternally more or less densely, glandular-puberulent. Leaves ovate
or deltoid, 1-5 cm. long and 0.6-4.0 cm. wide, uppermost perfoliate-
amplexicaul, middle sessile, lower short-petiolate, acute or upper
acuminate at apex, broadly obtuse, subtruncate or subcordate at
base, grossly and irregularly dentate-serrate; inflorescences sub-
umbellate-corymbose, often many-flowered but varying to few-flow-
ered; basal peduncle 2-3 times longer than intermediate peduncles,
these latter branching into pedicels only little shorter; intern1 oral
leaves apparently lacking; calyx lobes ovate, acute, 4-6 mm. long
and 3-4 mm. wide; corolla essentially glabrous within and without,
upper lip and the lower distally red or red-brown, proximally yellow,
upper lip 2-3 mm. long, lower 20-28 (-30) mm. long, saccate less
than half its length; stamens longer than the upper lip, anther cells
equal, divaricate, each pair of cells 3.0-3.5 mm. long, each cell at
least 2-3 times longer than wide; style ca. 2 mm. long, curved at
apex, ovary glandular; capsule ovoid, glandular, little shorter than
the calyx, the abrupt narrowing to a neck absent, the short style
persisting.
Arequipa: Plateau between Arequipa and Puno, near Arequipa,
alt. 2,600-4,000 m., Ferreyra 2598; Pichu-pichu, among rocks, in dry
steam bed and rocky slopes, alt. 10,000-12,000 ft., Stafford 668;
Je"sus-Chiquata, alt. 3,100 m., Vargas 8091.
This endemic is unusual in having essentially red flowers.
Calceolaria plectranthifolia Walp., Nova Acta Acad. Nat.
Cur. 19 (Suppl. 1): 395. 1843.
Small, much-branched, caespitose or subcaespitose shrublet, with
the leaves densely congested. Leaf-blades on basally expanded,
ochraceous petioles up to 5 mm. long, oblong or obovate, ca. 2 cm.
long and 1 cm. wide, margin doubly dentate, rugose above, reticulate
venose and brown-puberulent beneath; floral leaves usually entire;
inflorescences dichotomously branching, many-flowered; pedicels and
FLORA OF PERU 621
calyx lobes very sparsely pilose or glabrous; base of the calyx densely
pilose, lobes oblong, obtuse, the posterior larger than the others;
corolla yellow, 8-10 mm. in diameter, upper lip minute, lower lip
globose, inflated; filaments much longer than the equal-celled anthers.
Description from Kranzlin and type photograph. Originally col-
lected by Meyen in Arequipa.
Calceolaria procera Pennell, Proc. Acad. Nat. Sci. Phila. 97:
154. 1945.
Tall shrub, at least the lower nodes longer than the sessile or very
short-petiolate leaves, young growth puberulent, becoming glabrate
with age. Leaf blades widely linear or oblinear, mostly 3-7 cm. long
and 3-8 mm. wide, subacute to rounded at apex, attenuate at base,
sparsely denticulate to entire, slightly revolute, dark green and gla-
brous above, paler and puberulent beneath; petioles when present
1 cm. long, narrowly connate across the nodes; inflorescences paired
cymes from upper leaf-axils; bracts lanceolate, bractlets shorter, a
little wider, bracts, bractlets and calyx yellow (at least in life) ; pedi-
cels and peduncles glabrous, peduncles ca. 3 cm. long about equaling
the combined length of the secondary peduncles and pedicels, all
flower stalks more or less puberulent; sepals ovate, apex rounded, ca.
10 mm. long, pubescent on both surfaces; corolla darker yellow than
bracts and calyx, minutely puberulent proximally, upper lip ca. 2-3
mm. long, lower lip 10-20 mm. long (mostly 16-20 mm.), saccate less
than half its length, non-saccate part internally puberulent and also
hirsute at the orifice with much longer hairs; anthers 5 mm. long,
cells divergent (sagittately so) or divaricate, unequal, at least two
times longer than wide; style ca. 3 mm. long, thickened on lower
half and also pubescent, apically curved, ovary pubescent; capsule
little shorter than the calyx, ca. 8 mm. long, globose-ovoid, eglandu-
lar, pubescent.
Cuzco: Prov. Urubamba, Lucumayo Valley, alt. 1,800-3,600 m.,
Cook & Gilbert 1315; Prov. Calca, below Lares, Weberbauer 7891.
This endemic in the two provinces of Dept. Cuzco is unusual in
having yellow floral leaves.
Calceolaria pseudoscabra Edwin, Phytologia 19: 392. 1970.
Coarse perennial herb, erect, branched, foliage, stems and branches
drying brown, stems and branches glandular-puberulent. Leaves
mixed glandular and eglandular-puberulent, sessile or less often
short-petiolate, ovate to broadly elliptically ovate, mostly 2.0-3.5
622 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
cm. long and 1.0-2.1 mm. wide, margins revolute, crenate, narrowed
to an obtuse apex, base broadly obtuse, tissue scabrous, costa ele-
vated and prominent beneath; young inflorescences arising in pairs
from the upper leaf -axils, cymose, few-flowered; peduncles glandular-
pubescent, 2-5 times longer than the pedicels; bracts and bracteoles
reduced floral leaves, bracts occasionally with a pair of lobes near
the base; calyx lobes mixed glandular- and eglandular-pubescent on
both surfaces, elliptic-ovate, long-acute, 4-5 mm. long and ca. 3.5
mm. wide; corolla yellow, upper lip glabrous, 1-2 mm. long; lower
lip externally short-glandular-puberulent, saccate over most of its
length, 13-15 mm. long and about 5 mm. wide, a tuft of white hairs
present at base within; stamens short, anthers large for the small
corolla, anther cells divaricate, unequal, usually at least twice as long
as wide, the larger 1.8-2.0 mm. long, the smaller 1.2-1.5 mm.; style
ca. 1.5 mm. long, thickened, ovary glandular; capsule unknown.
Cajamarca: Wet place, alt. 2,700-2,900 m., mountain E. of Celen-
din, Pennell 15213 (PH).
Closest to Calceolaria scabra and C. variefolia, differing from the
former in being a much coarser plant with larger leaves and from the
latter in having the internodes much shorter as well as in growth
form, an herb as opposed to a shrub.
Calceolaria pulverulenta R. & P., Fl. Peruv. 1: 19. 1798.
Shrub, erect or scandent, stems and branches white-tomentose.
Leaves canescent above and more densely beneath, ovate or elliptical,
up to 5 cm. long and 2.3 mm. wide, crenulate or crenate, subacute to
acute apically, broadly acute to subobtuse basally, short-petiolate
or sessile; inflorescences paniculate, sometimes reaching 20 cm. long,
most often shorter, ultimate inflorescence branches corymbose or
dichasiate; calyx lobes externally canescent, internally with a distinct
row of hairs along the margins, ovate, 5-7 mm. long, acute; upper lip
of the yellow corolla, about twice as long as the calyx, lower lip 1.0-
1.2 cm. long and ca. 1.5 cm. wide, saccate about half its length; fila-
ments short, anther cells equal, drying black or dark brown.
No specimens seen. Reported from northern and north-central
Peru and also from Ecuador. Killip & Smith 2^377 from Junin,
alt. 2,700-3,200 ft., may possibly pertain here.
Calceolaria pumila Edwin, Phytologia 19: 393. 1970.
Tiny unbranched herb up to 6 cm. tall, the simple stem densely
glandular-pubescent, bearing 1-3 pairs of oblong or ovate leaves.
FLORA OF PERU 623
Leaf blades sessile, entire, up to 1.2 cm. long and 0.8 cm. wide, usually
reaching 0.9 cm. by 0.7 cm., puberulent-pubescent on both surfaces;
flowers solitary, pedicellate, terminal or in the upper leaf -axils, tiny,
up to 7 mm. long and 2 mm. wide, yellow; pedicels short, up to 7-8
mm. long, glandular-pubescent; calyx lobes elliptical, acute, 3.0-
3.5 mm. long and 1.5-2.2 mm. wide, externally glandular-pubescent;
upper lip of the glabrous corolla ca. 2 mm. long, lower lip 5 mm. long,
sac laterally placed the length of the lip but only about 0.5 mm. thick
medianly from the distal end; stamens very small, filaments thick-
ened, less than 1.0 mm. long, anther cells equal, drying white to light
yellow, each anther ca. 1.0-1.2 mm. long, each cell a little longer than
wide, divaricate; style thickened, ca. 1.0 mm. long, ovary sparsely
glandular-puberulent; capsule elliptical, sparsely glandular-puberu-
lent, 6.0-6.5 mm. long and 2.0-2.5 mm. wide, much exceeding the
calyx.
This taxon has the smallest flowers of any in Peru.
Known only from the type specimen.
Cajamarca: Rocky limestone on ridge 11 km. S. of Cajamarca,
alt. 2,750-2,850 m., Pennell & Anderson 15078 (PH).
Since the sheet is not marked, unicate isotypes are probable.
A unique taxon in the genus, at least in Peru. No close relative
is known at this time.
Calceolaria ramosissima Kranzl. in Fedde, Rep. Sp. Nov. 1:
107. 1905; non 1913.
Vining shrub, newer wood pubescent, older ultimately glabrous,
much branched, branches ascending, drying brown to dark purple-
brown. Leaves ovate, opposite or ternate, little longer than wide to
almost twice as long as wide, 0.8-3.0 cm. long and 0.5-1. 5(-2.0) cm.
wide, pubescent on both surfaces, serrate, occasionally serrate-den-
tate, apically narrowed, basally obtuse to very broadly obtuse; in-
florescences terminal and in the axils of the upper leaves, dichasiate,
few to several-flowered; flower stalks pubescent, up to about 1 cm.
long; calyx lobes dimorphic, ovate, usually obtuse and venose, 2.5-
3.5 mm. long and 2.0-2.5 mm. wide, or lanceolate, usually acute and
veins obscure, 2.5-3.5 mm. long and 1.5-2.0 mm. wide, mostly pu-
bescent on both surfaces, especially at the margins; corolla yellow,
minutely and densely pilose, 10-15 mm. long, upper lip much shorter
than the calyx lobes, lower lip saccate a little less than half way,
nectary on the upper margin of the sac, thickened and very narrow,
lip usually less than half as wide as long, even distally; anther cells
624 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
reniform, approaching twice as long as wide, each pair less than
3 mm. long on thickened, short filaments; style 1-2 mm. long, ovary
mostly few-glandular, glands sessile, golden resinous; capsule ovoid
about as long as the calyx, gradually and evenly tapering to the acute
apex, vesture as the ovary.
Ancash: Chiquian, Prov. Bolognesi, alt. 3,220 m., Ferreyra 5698,
Stony puna, S. slope of Paso de la Fortale"za, alt. 3,550-3,600 m.;
Pennell 15395; Thickety bank below Chiquian, alt. 3,200-3,300 m.,
Pennell 15404- — Cajamarca: Sandy soil on cumbra above Cajamarca,
alt. 3,500 m., Olsson 2a; 6 miles N. of Cajamarca, on road to Chota,
sandy soil, alt. 2,800 m., Olsson 1+a; Road from Cajamarca to Celen-
din, alt. 2,750-4,000 m., Scolnik 1314. — Lima: between Matucana
and Surco, Prov. Huarochiri, alt. 2,100-2,200 m., Ferreyra 7599.
The southernmost collections (Ancash and Lima) possess the
lanceolate sepals.
Calceolaria ranunculoides Kranzl. in Fedde, Rep. Sp. Nov. 1:
97. 1905.
Annual herb in wet to dry places, stems rooting freely, prostrate
or ascending, plant frequently glandular-pubescent, glands often
black. Leaves variably petiolate; petioles descrete; blades long-
petiolate below and petioles decreasing in length to the apex, or
sometimes blades all sessile or subsessile, variable in shape and size,
mostly ovate and dentate or lobed up to the middle, ovate and doubly
dentate to entire above the middle, all blades occasionally varying
to deeply lobate or subpinnatifid, from 1 to 6 cm. long or even longer
(petiole included); flowers most often borne singly on the ends of
short branches, occasionally in few-flowered dichasia; pedicels up to
5 cm. long, glandular-pubescent; calyx lobes ovate, 6.5-8.5 mm. long,
acute, glandular-pubescent; corolla yellow often glandular-pubescent
on both surfaces, sometimes only externally, upper lip 5-7 mm. long,
lower up to 3.0 cm. long, usually 2.0-2.5 cm. long, saccate about
two-thirds its length, sac large, much inflated; filaments short, an-
ther cells very unequal, upper fertile cell much larger than the lower
cell which is sometimes sterile, connective longer than the cells; style
ca. 1.5-2.0 mm. long; capsules ovoid, acute, from as long as to 1^
times longer than the calyx, glandular-pubescent.
Amazonas: Colcamar, valley below jalca, alt. 3,300 m., Pennell
15640; Puente de Pipos, on Rio Sonche, opposite Daguas, E. of
Chachapoyas, alt. 2,000-2,100 m., Pennell 15755; "Hidalgo," below
FLORA OF PERU 625
Chachapoyas, alt. 1,800 m., Tarnawiecki for Pennell 15912. — An-
cash: Consuzo, Prov. Pallasca, alt. 3,700 m., Lopez 2384', Above
Racrachaca, Prov. Bolognesi, alt. 3,300-3,400 m., Ferreyra 7511;
Usger, below Chiquian, alt. 3,250-3,300 m., Ferreyra 7560; Huaraz,
alt. 3,000 m., Pennell 15301; Chancos, alt. 3,000-3,200 m., Pennell
15311; 15317; Copa grande, N. of Chancos, alt. 3,250 m., Pennell
15361 ; in stream and bog, Colcas, alt. 2,500-2,600 m., Pennell 15385;
Paso de la Fortale"za, alt. 3,550 m., Pennell 15396. — Cajamarca,
Hacienda Limon, Carricker s.n.; NE. of Cajamarca, alt. 3,650 m.,
Pennell 15143; Along streams on jalca, CordiUera de Cumulloa, to
Celendin, alt. 3,400-3,500 m., Pennell 15168; Mountain E. of Celen-
din, alt. 2,600-2,700 m., Pennell 15199; Llacanora, alt. 2,550-2,600
m., Pennell 15490; 15-20 km. above Cajabamba, alt. 2,900 m., Pen-
nell & Ferreyra 14582; 10 km. above Cajabamba, 2,900 m., Pennell
& Ferreyra 14662; SW. of Cajamarca, alt. 2,800 m., Pennell & Reich-
lin 15022. — Cuzco: Macchu-picchu, Valley of Rio Urubamba, ruins,
Ugent & Ugent 5340. — Huanuco: Dist. Churubamba, trail Puente
Durand to Exito, Que"brada de Gaza, alt. 1,450 m., Mexia 8244-—
Junin: Huacapistana, alt. 1,800-2,400 m., KilUp & Smith 24095;
Pichis Trail, Enenas, alt. 1,600-1,900 m., Killip & Smith 25768;
Huancayo, alt. 3,317 m., Soukup 3001. — La Libertad: Above Motil,
dry open jalca, alt. 3,200 m., Pennell & Ferreyra 14847; Wet cliff,
3-4 km. above Huamachuco, alt. 3,250 m., Pennell & Ferreyra 14871 ;
W. edge of Huamachuco, alt. 3,050 m., Straw 2527. — Lima: Chosica,
alt. 800 m., Soukup 2051. — Sin. Dept.: Chasqui, Macbride 3294-
The description here is much broader than that of Kranzlin who
saw far less material. The texture as well as the size, shape, and
margins of the leaves is quite variable. The blades are thinly char-
taceous to very firm or almost subcoriaceous. The thin leaves are
most frequently the larger.
This taxon may be conspecific with Calceolaria aquatica Brown
and Bouch6 of Bolivia but is provisionally held distinct on the bases
of vesture and venation differences.
Calceolaria reichlinii Edwin, Phytologia 19: 394. 1970.
Shrub erect and branched, wood, leaves, flower stalks, and calyces
more or less densely beset with long, silvery, soft hairs (lanate), the
leaves beneath and calyx lobes most densely so, the vesture some-
times fugitive with age, usually eglandular except occasionally for
the corolla and frequently the ovary. Leaves most often linear or
626 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
aciculiform, occasionally broadening to lance-ovate, sessile or almost
so, the vesture frequently completely obscuring the tissue beneath,
margins mostly entire, revolute, obtuse apically, blades sessile; in-
florescences 1- to few-flowered; flower stalks up to 2 cm. long, usually
less; calyx-lobes elliptical or elliptically ovate, rarely ovate, 5-8 mm.
long, and up to 4 mm. wide, acute, the tissue frequently completely
obscured by the vesture externally, glabrous or sometimes with a
few hairs paralleling the margins or very rarely a few glands present
internally; corolla yellow, variable in size, 12-25 (-28) mm. long,
externally glabrous or few-glandular, rarely papillose, saccate more
than half its length, internally glabrous; anther cells unequal, 2 or
more times longer than wide, completely divaricate, each pair of cells
up to 4 mm. long, septum thickened; style 2-4 mm. long, glandular
at base; ovary glandular-pubescent or long-pubescent, rarely merely
few-glandular; capsule narrowly conic-ovoid, about as long as the
calyx, vesture as the ovary, blunt at the apex, attenuate, style per-
sisting.
Common in Dept. Cajamarca, in the Andes of northern Peru and
three collections also seen just to the south in Dept. Libertad.
Cajamarca: Dry banks (volcanic soil), 15-20 km. above Caja-
bamba, alt. 2,900 m., Pennell & Ferreyra 14,853', Prov. Cajabamba,
km. 38, N. of Huanachuco, near Cajabamba, alt. 2,900-3,000 m.,
Ferreyra 301+1+; On cumbre above Cajamarca, alt. 3,500 m., Olsson 2;
6 miles N. of Cajamarca on road to Chota, sandy soil, alt. 2,800 m.,
Olsson 1+, pro. parte; Celendin-Balsas road-km. 124, Olsson 1+a;
Mountain E. of Celendin, alt. 2,700-3,000 m., Pennell 15173; Alt.
2,400-2,800 m., Pennell 15177; Rocky calcareous hillside, NE. of
Cajamarca, alt. 2,600-2,700 m., Pennell 15230; Rocky limestone
ridge, 11 km. S. of Cajamarca, alt. 2,750-2,850 m., Pennell & Ander-
son 15082; Cliff about 10 km. above Cajamarca, alt. 3,200 m., Pen-
nell & Ferreyra 11+865; Open jalca, Paso de Gavilan, S. of Cajamarca,
alt. 3,000-3,300 m., Pennell & Ferreyra 11+959; Jalca SW. of Caja-
marca, alt. 2,700 m., Pennell & Reichlin 11+981; Stony limestone,
Michiquillo, alt. 3,100 m., Pennell & Reichlin 15031+; Alt. 2,750-4,000
m., on the Cajamarca-Celendin road, Scolnik 1308. — Libertad: 2-3
km. above Otuzco, Trujillo-Otuzco road, alt. 2,500 m., Ferreyra 2977;
Sandy area, S. of Otuzco, alt. 2,500 m., Pennell & Ferreyra 11+820;
km. 82 W. and below Agallpampa, alt. 2,900 m., Pennell & Ferreyra
11+832.
This species is close to C. linearis R. & P. The latter differs in
having much shorter and sparser indument, apically acute leaves,
FLORA OF PERU 627
usually dichasiate inflorescences, shorter calyx segments that are in-
ternally pilose, minute upper lip of the corolla, and much shorter
style.
Calceolaria revoluta Penn., Proc. Acad. Nat. Sci. Phila. 97:
165, PL ISA. 1945.
Glandless, erect or ascending, branched shrub, ascending-pubes-
cent or puberulent on the wood and in the inflorescence. Leaves
linear or linear-oblanceolate, 2.0-2.5 (-3.0) cm. long and 2-3 (-5) mm.
wide, margin entire, revolute, blade minutely appressed-pilose above,
paler and minutely tomentose beneath; petiole ca. 1 mm. long, white-
villose, basally indurate and narrowly connate; inflorescences ter-
minal, clustered, usually 1-compound cymes; peduncles up to 2-3 cm.
long below, appressed-pubescent as the 7-15 mm. long pedicels;
bracts lanceolate, smaller than the leaves, bractlets minute or want-
ing; sepals yellow in life, yellow-green when dry, ovate, 4-5 mm. long,
entire, acute to subobtuse, finely appressed-puberulent; corolla yel-
low, externally finely puberulent, upper lip ca. 3-4 mm. long, hooded,
lower lip to 19 (-22) mm. long, saccate less than half its length, inter-
nally on the lower surface a tuft of hair at the orifice, crossed with
red, dot-like marking above the tuft and then distally minutely
puberulent; anther cells subparallel to little diverging (1.5)2.0-3.0
mm. long, usually equal, occasionally a little unequal, acute above
the septum which parts at dehiscence; style 4-5 mm. long, basally
pubescent, apically curved, ovary densely pubescent; capsule ovoid-
subconic, 4-5 mm. long, ca. equaling the calyx, densely pubescent
with upcurving indument.
Cuzco: Valley of Paucartambo, alt. 3,400 m., Herrera 2991 ; Pau-
cartambo to Ire's Cruces, Cerro de Cusilluyoc, alt. 3,300-3,500 m.,
Pennell 13802 (Isotypes F; PH; US); Paso de Tre"s Cruces, Cerro
de Cusilluyoc, paramo, alt. 3,800-3,900 m., Pennell 13885; Pampa
Corral (Lares), Prov. Calca, alt. 3,400 m., Vargas 3208; Que"brada
de Llulluchayocc, Prov. Paucartambo, river bank, alt. 3,650 m.,
Vargas J>282; Below Lares, alt. 3,100-3,200 m., Weberbauer 7891.—
Sin. Loc.: Diehl2533.
Calceolaria rhacodes Kranzl. in Engler, Bot. Jahrb. 50: Beibl.
Ill, 71. 1913.
Shrub to occasionally herbaceous, most often erect, older stems
and branches glabrate, younger usually pilose, drying violet, usually
well-branched, internodes increasing in length toward the base.
628 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Leaves broadly ovate in outline, occasionally oblong-ovate, up to
6 cm. long and about 4 cm. wide, vesture variable, from sparsely
puberulent on both surfaces to densely lanose beneath, upper sur-
face usually sparsely pubescent, 5-7-lobed, the lobes frequently den-
tate, blades broadly acute at apex, truncate to shallowly cordate
basally, occasionally broadly obtuse, lowermost subsessile, upper and
median on petioles up to 2 cm. long; inflorescences variable, irregu-
larly corymbose or dichasiate, or simply cymose or flowers merely
singly paired; bracts of the basal branches of the inflorescences foli-
ose; pedicels and peduncles thin, densely glandular-pilose, peduncles
up to about 2 cm. long, rarely longer; calyx externally glandular-
pilose or pilose, with 2-3-celled hairs, lobes ovate, narrowly obtuse,
5.0-6.5 mm. long; corolla yellow, 16-30 (-35) mm. long, varying in
length from plant to plant but closely fixed on a single plant, mi-
nutely glandular-pilose, upper lip shorter than the calyx, lower sac-
cate more than half way, distally almost as wide as long; anther
cells two or more times longer than wide, wholly divaricate, usually
large for the corolla, each pair up to 5-6 mm. long, cells of a pair
equal or unequal; style ca. 3.0-3.5 mm. long, glabrous, apically
curved, ovary glandular, shorter than the style; capsule oblong-
ovoid, shorter than the calyx, apically rounded or obtuse, neck lack-
ing, style persisting.
Ancash: Cerro S. of Chiquian, Capillapunta, alt. 3,540 m., Fer-
reyra 5725', Cerro E. of Chiquian, alt. 3,400-3,500 m., Ferreyra 7338.
— Ayacucho: Above Puquio, Prov. Lucanas, alt. 3,400-3,500 m.,
Ferreyra 5508; Marcahuasi, above Puquio, alt. 3,400-3,450 m., Fer-
reyra 7212; Coracora, alt. 3,200 m., Weberbauer 5809 (Isotypes PH;
US); 6038; 6040; 6400.— Libertad: Usquil, Prov. Otuzco, alt. 3,000-
3,100 m., Ferreyra 7641 ; 7649 ; Agallpampa, alt. 3,200 m., Lopez 0337.
—Lima: Prov. Huarochiri, above Surco, alt. 2,800-3,000 m., Fer-
reyra 699; 3446; Alt. 3,200-3,300 m., 6094; Alt. 3,000 m., Hrdlicka
s.n.; Matucana, alt. ca. 9,000 ft., Macbride & Featherstone 468; S.
of Surco, alt. 2,700-3,000 m., Pennell & Ferreyra 15257.— Piura:
Palambla, alt. 3,500 m., Carricker s.n.; Above Canchaqui, alt. 1,700-
1,900 m., Pennell & Ferreyra 14911. — Sin. Dept.: Al Paso del Indio,
alt. 3,000 m., Ochoa 1625.
Calceolaria rhizomatosa Kranzl. in Engler, Bot. Jahrb. 50:
Beibl. Ill, 70. 1913.
Repent, twining or erect shrub with a woody rhizome, usually
with more branches above than below, sometimes (ex desc.) sub-
FLORA OF PERU 629
caespitose, wood strigulose and puberulent. Leaves sessile or short-
petiolate, oblong or lance-ovate, mostly 2-3 cm. long and up to 1 cm.
wide, occasionally lower leaves on the stem reaching 5 cm. long and
almost 2 cm. wide, entire to distantly, minutely dentate, narrowed
to base and apex, short-pilose above, punctate and glabrous beneath,
except on the veins; inflorescences branched, arising from the axils
of the upper leaves; peduncles up to 4 cm. long; pedicels 1.5 cm. long,
both stalks puberulent; bracts lanceolate, conspicuous; flowers yel-
low, ca. 18-20 mm. long; calyx lobes broadly ovate, acute, ca. 5-6
mm. long, puberulent on both surfaces; upper lip of the corolla ca.
1.3-2.5 mm. long, lower lip ca. 16 mm. long, saccate less than half
its length, proximally white-puberulent externally beneath, sparsely
long-pubescent proximally within and with a tuft of hair at the long
orifice; anther cells equal, ca. 3 times longer than wide, plane or
downturned distally, divaricate or divergent; style ca. 2 mm. long;
semimature ovary ovate. (Ex. desc.)
Ayacucho: Near Coracora, Prov. Parinacochas, alt. 3,200 m.,
Weberbauer 5803, Isotype; Below Totorabamba, Prov. Huamanga,
alt. 3,300-3,400 m., Weberbauer 5478.
Calceolaria rhododendroides Kranzl. Pflanzenr. 4(257c): 88,
fig. 17, 0-6. 1907.
Small, erect, branched shrublet to 50 cm. tall, older wood sparsely
pilose, younger wood densely lanate. Leaves short-petiolate, sub-
dense, younger blades, densely lanate above and beneath, older be-
coming glabrous except for the revolute margins and midvein, ovate,
1.2-1.8 cm. long and 4-6 mm. wide, acute at apex, little dimidiate
at base, rigid, midvein beneath wide and prominent, older leaves
f errugineous below with bright, green venation ; inflorescences in up-
permost leaf-axils of the branches, few-flowered cymes ; flower stalks
densely lanate; calyx segments lanate externally at least on lower
half, glandular internally, elliptic, obtuse, ca. 4 mm. long and 2 mm.
wide; corolla yellow, densely pilose, 13-17 mm. long, upper lip
hooded, 2-3 mm. long, lower lip obovate, dilated, sac shorter than
the open part; filaments short, anther locules equal, only little longer
than wide, large, walls and septa thickened; style 5-7 mm. long,
ovary densely beset with long, gland-tipped, golden-brown hair.
Only fragments of the type seen, description largely after Kranzlin
(1907).
Cajamarca: Between Cajamarca and Celendin, alt. 3,700-3,800
m., Weberbauer
630 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria rosmariniifolia Lam., Encycl. 1: 556. 1783. C. jer-
ruginea Cav., Icon. 5: 27, t. 4.4-5, fig. 1. 1799. C. graminifolia HBK.,
Nov. Gen. et Sp. 2: 386. 1817. C. bifida Willd. ex Link, in Spreng.,
Bot. Jahrb. 1: 52. 1820. C. hartwegii Benth., PI. Hartweg. 147. 1844.
Erect, sparsely or wholly unbranched subshrub, sometimes herba-
ceous or vine-like above, wood glabrous below, sparsely to densely
pilose or glandular-pilose above. Leaves sessile and often narrowly
connate, linear to lance-linear, 5-10 cm. long and 6-10 mm. wide,
entire and often revolute, apex acuminate, glabrous, often shining
and glutinous above, usually tomentose or pilose beneath; inflores-
cences in upper leaf-axils, paniculate, the basal stalks repeatedly
branching; peduncles and pedicels glandular-pilose and often also
glutinous ; calyces broadly ovate to triangular-ovate, glandular-pilose
or occasionally merely glutinous-puberulent, 3.5-4.0 mm. long and
3.0-3.5 mm. wide, sometimes as long as wide; corolla yellow, glandu-
lar-pilose externally, sparsely so internally or varying to wholly gla-
brous, 13-15 mm. long, upper lip little shorter to little longer than
the calyx, lower lip 2-3 times longer than the upper; filaments short,
anther cells equal, almost as wide as long to 1.5 times longer than
wide; style 2-3 mm. long, ovary glutinous, papillose to puberulent.
Only fragments seen, description after Kranzlin.
Sin. Dept.: At the foot of Chimborazo, alt. 3,600 m., Bonpland
3145; near Tarca, alt. 2,800 m., Bonpland 3204- This species is also
known from Ecuador.
Calceolaria rugulosa Edwin, Phytologia 19: 395. 1970.
Erect, branched, pilose-tomentose shrub, stems appressed-pilose
below, tomentose above, and with the leaves drying light brown, the
leaves also sometimes drying brown-green, tomentum densest on
young growth. Leaves opposite or ternate, ovate or elliptic-ovate,
occasionally elliptical (1.5-)3. 0-5.0 cm. long and (1.0)2.0-3.5 cm.
wide, margins plane, serrate to doubly serrate, sometimes grossly so,
narrowed to an acute apex, obtuse to broadly obtuse at base, rugose
or rugulose, pilose above, tomentose beneath, sometimes, especially
when young, the vesture obscuring the leaf tissue, often with golden-
yellow glands intermixed beneath; petioles l-4(-6) mm. long,
thickened, woody, densely tomentose, usually narrowly connate; in-
florescences in upper leaf-axils and also terminal, many-flowered,
compound dichasia or corymbose; flower stalks densely long-soft-
pubescent; bracts merely reduced leaves; calyx tube and lobes densely
pilose externally, glabrous internally, lobes ovate or broadly ovate,
FLORA OF PERU 631
acute, 6-8 mm. long; corolla yellow, up to 1 cm. long, externally very
short-puberulent, internally glabrous; upper lip 1-2 mm. long; lower
lip ca. 6-8 mm. long, saccate only 1-2 mm. ; anther cells equal, each
pair ca. 2.0-2.2 mm. long, almost twice as long as wide; style thin,
glabrous, 1.0-1.5 mm. long, ovary glandular; capsule ovoid-conic, a
little shorter and loosely invested by the calyx, apically long-acute,
sparsely golden glandular.
Cajamarca: Rock cliff below Llama, alt. 1,800-1,850 m., Pennell
15917, (US); Earth bank, Llama, alt. 1,850 m., Pennell 15916.
This taxon is related to C, aperta Edwin, but it is morphologically
quite distinct.
Calceolaria salicifolia R. & P., Fl. Peruv. 1: 18, t. 19, fig. b.
1798, non Morris, nee Griseb.
Erect shrub, or occasionally woody vine, essentially glabrous,
often up to 2 m. tall, glutinous and shining. Leaves petiolate, often
coriaceous, lanceolate, lance-elliptic or lance-ovate, very rarely a few
leaves ovate, often longer than 8 cm., occasionally not exceeding 3-
4 cm. long and usually not more than 2 cm. wide, rarely reaching
4 cm., margins closely and sharply serrate or doubly serrate, rarely
subentire, apex acute or long-acute, base subobtuse, sometimes di-
midiate, much darker above than beneath, often drying brown, veins
mostly apparent beneath, costa usually elevated ; inflorescences cor-
ymbs, dichasiate or dichasia, with two reduced leaf-like bracts at
the base and 2-3 bracteoles at the forking or branching of the pe-
duncles; both pedicels and peduncles elongate; flowers large for the
genus, up to 5 cm. long and 2.5 cm. wide; calyx lobes most often
lance-elliptic and abruptly acuminate, varying to lance-ovate, the
anterior narrower than the others, 10-15 mm. long and 5-8 mm.
wide; upper lip of the yellow corolla 7-10 mm. long, hooded, margin
irregularly cut with rounded teeth; lower lip up to 3.7 cm. long and
2.5 cm. wide, saccate over most of its length, a tuft of white hairs
present internally at the base, the lower surface microscopically
glandular; filaments 3.5-4.5 mm. long, thickened, anther cells un-
equal, most often two or more times longer than wide, each pair
7-9 (-10) mm. long; style thickened, uncurved apically, 6-8 mm.
long, ovary glutinous; capsule only little more than half as long as
the calyx, glabrous, very broadly urceolate but the neck little con-
stricted, median locular suture present as a sulcation.
Amazonas: Cerro Tinaja, alt. 3,200 m., Ochoa 1684', 1687; Moun-
tains E. of Balsas, alt. 10,000 ft., Osgood & Anderson 75; Cold moun-
632 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
tain forest above Colcamar, alt. 3,200-3,300 m., Pennell 15623;
Bushy paramillo, Cerro Puma-urco SE. of Chachapoyas, alt. 3,000-
3,500 m., Pennell 15673; Open cold swamp on summit of Cerros de
Calla-Calla, alt. 3,500-3,750 m., Wurdack 1193; Summit and upper-
most slopes of Cerros de Calla-Calla, near Kms. 403-407 Balsas-
Leimebamba road, alt. 3,400-3,550 m., Wurdack 1710. — Huanuco:
15 m. SE. of Huanuco, 10,500 ft., Macbride & Featherstone 2110;
2122; Between Kms. 46-48 Huanuco-Panao, alt. 3,000 m., Ochoa
1105; Alt. 2,750 m., Skolnik 1062; Pillao, alt. 2,700 m., Woytkowski
47. — Junin: Sunny grass -shrub slope, alt. ca. 9,000 ft., Macbride &
Featherstone 1604- — Libertad: In open pasture above (NE) Cachica-
don, Prov. Santiago de Chuco, alt. 2,700 m., Stork & Horton 9987.
—Sin. Loc.: Dombey 308; Ruiz & Pawn s.n. (two collections).
Calceolaria santalinoides Kranzl. in Engler, Bot. Jahrb. 50:
Beibl. Ill, 73. 1914.
Differs from C. incarum Kranzl. as in the key and also in having
smaller leaf blades and less densely glandular and shorter pubescent
wood ; inflorescences few-flowered and with the ultimate flower stalks
(pedicels) among the largest in the genus, the bizarre leaved species
excepted.
This taxon is known from collections in Southern Peru in Depts.
Ayacucho, Arequipa, and Tacna. It is closely allied to C. incarum
and the two taxa form a vicarious pair, separated by the middle pass
(Morococha-Ticlas) of the Peruvian Andes. The original material
was found in Argentina.
Arequipa: Vulcan Chachain, alt. 3,000 m., Rauh-Hirsch P575.—
Ayacucho: between Nazca and Puquio, near Lucanas, alt. 3,200-
3,300 m., Ferreyra 5474; Puquio Valley, alt. 3,200 m., Rauh-Hirsch
P419; S. of Chavina, alt. 3,200 m., Weberbauer 5781 ; Alto de Tacora,
Isern 2018. — Tacna: Prov. Tarata, dry granite, rock wall, 8 km. on
trail from Candarave to Puno, alt. 3,250 m., Metcalf 30385.
Calceolaria sarmentosa Kranzl. in Fedde, Rep. Sp. Nov. 1: 37.
1905.
Usually repent, simple herb, varying to suberect and branched,
sometimes dwarfing, ca. 10-15 cm. tall, when erect, stems sparsely
pubescent. Leaves short-petiolate, blades ovate or elliptically ovate
in outline, only rarely reaching 3 cm. long, usually 0.6-2.5 cm. long,
most often deeply 5-7-lobed, lower sometimes 2-3-lobed, upper oc-
casionally almost pluri-pinnate and margins coarsely, irregularly,
FLORA OF PERU 633
doubly dentate, sparsely pubescent above, paler and little or not at
all pubescent beneath; petioles up to 2 cm. long, most often densely
pubescent; plants flowering from upper leaf -axils only or over most
of their length, when apically aggregated the inflorescences irregu-
larly branched cymes; calyx lobes ovate or less often lance-ovate,
4-6 mm. long, acute, varying from glabrous to glandular-pubescent
on one or both sides, pubescence frequently lost at time of fruiting;
corolla yellow, glabrous or occasionally puberulent externally, 18-
30 mm. long, upper lip 2-4 mm. long, lower saccate more than half
its length; one cell of each anther sterile and often wholly lacking,
connective shorter than the fertile cell, thickened; style persisting in
fruit, 1.5-2.0 mm. long, dilated at base at least in fruit; capsule occa-
sionally glandular-pubescent or pubescent, most often sparsely pu-
berulent to glabrous, much longer than the calyx, usually abruptly
narrowed to a long neck, sometimes only gradually tapering, either
ovoid-conic or ovoid-triangular.
Amazonas: Cerro Puma-urcu, SE. of Chachapoyas, alt. 3,400-
3,500 m., Pennell 15716. — Ancash: Rio Santa, opposite Huaraz,
along stream, alt. 3,000-3,150 m., Pennell 15331; Sin. Data, Weber-
bauer 3121 (type photograph), ex Kranzlin, lower part of Hacienda
Cajabamba between Samenco and Caraz. — Ayacucho: Pampalca
between Huanta and Rio Apurimac, alt. ca. 3,200 m., open hillside,
Killip & Smith 12257. — Cajamarca: SW. of Cajamarca, moist bank,
alt. 2,800 m., Pennell & Reichlin 15022. — Huanuco: Carpish, Chan-
cai, alt. 2,200 m., Vargas 5410.
Since this species is distinguished by a combination of leaf size,
flower size, and fruit size in relation to calyx size and, in addition,
since the few collections are so widely dispersed and, finally, since
this material is from such diverse habitats it seems not unlikely that
this taxon may be more apparent than real. However, at this time
it is retained because the material cited does not lend itself to place-
ment elsewhere. Further study of the entire subgenus is required.
Calceolaria scabra R. & P., Fl. Peruv. 1: 19, t. 29a. 1798.
C. melissifolia Benth. in DC., Prodr. 10: 214. 1846. C. glechomifolia
Kranzl. in Engler, Bot. Jahrb. 54: Beibl. 119, 20. 1917.
Variable, erect or ascending shrub, plant with dense vesture to
almost glabrous, stems and branches usually pubescent with 2-3-
celled, erect hairs, occasionally a few short, gland-tipped hairs inter-
mixed, sometimes wholly glabrous, the wood of the usually copiously
branched, densely foliaceous stem drying yellow to dark red. Leaves
634 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
mostly ovate to rhomboidal, occasionally obovate, not exceeding
3.5 cm. long, mostly 0.9-2.5 cm. long and from a few millimeters
wide to almost as wide as long, mostly pubescent above as the stem
and with more gland-tipped hairs intermixed beneath, the vesture
sometimes very dense, obscuring the tissue, rarely wholly glabrous,
margin plane or revolute, crenate or dentate, sometimes teeth prom-
inent, apex usually acute, base mostly cuneate to obtuse, tissue
bullate or plane, usually thickened, rarely chartaceous, sessile or al-
most so; flower 10-25 mm. long, solitary or in 2-3-flowered inflores-
cences, very rarely up to 10-12-flowered; peduncles and pedicels up
to 3 or more cm. long, usually glandular-pubescent; bracts reduced
or wanting; calyx lobes mostly glandular-pubescent on both sides,
sometimes only externally, ovate, acute, mostly 6-8 mm. long and
4-5 mm. wide at base; upper corolla lip minute to 3 mm. long, lower
yellow, frequently red-marked within, mostly pubescent with soft
hairs externally, glabrous internally, saccate more than half its
length, about a half to two-thirds as wide as long; filaments short,
anthers large for the corolla, divaricate, almost always at least 2
times longer than wide, cells unequal, septum and often the walls
thickened, mostly 4-6 mm. long, shorter in the smaller flowers; ovary
mostly pubescent or glandular-pubescent; capsule narrowly ovoid,
or lance-ovoid, a little shorter to longer than the calyx; vesture as
the ovary.
Widely distributed in the Andes from Cajamarca in the north to
Arequipa in the south. The taxon has not yet been reported from
the southernmost parts of Peru. The citations below are selected
from the copious available material.
Amazonas: Prov. Chachapoyas near Ubil6n between Chacha-
poyas and Leimebamba, alt. 1,500-2,000 m., Ferreyra 7120; alt.
2,400 m., N. of Chachapoyas, Pennell 15499; Cerro Puma-urcu,
alt. 2,500-2,600 m., Pennell 15519; Bushy slope, Taquia, E. of Cha-
chapoyas, alt. 2,400-2,500 m., Pennell 15584; SE. of Colcamar, alt.
2,050 m., Pennell 15597; lower slopes of Puma-urcu, above Hda. Sta.
Isabel, 1.5-2.0 km. SE. of Chachapoyas, alt. 2,400-2,500 m., Wur-
dack 436. — Ancash: Prov. Bolognesi, Cerro N. of Chiquian, alt.
3,540-3,560 m., Ferreyra 5608; above Aquia, alt. 3,250-3,300 m.,
Ferreyra 7516; below Chiquian, alt. 3,200-3,300 m., Pennell 15490;
toward El Huascaran, N. of Yungay, alt. 2,850-2,900 m., Pennell
15426. — Arequipa: Moquegua Prov., Rainy-green formation, alt.
3,200 m., Weberbauer 7992. — Ayacucho: Pampalca between Huanta
FLORA OF PERU 635
and Rio Apurimac, alt. about 3,200 m., Killip & Smith 22222; 23280.
— Cajamarca: Cajabamba, Carricker s.n.; Limestone rocks, NE. of
Cajamarca, alt. 3,500 m., Pennell 15139; Limestone rocks NE. of
Cordillera de Cumulloa to Celendin, alt. 3,500-3,600 m., Pennell
1515^; mountain E. of Celendin, alt. 2,700-3,000 m., Pennell 15175;
Quinamayo, alt. 3,500-3,600 m., Pennell & Reichlin 1504-8. — Huan-
cavelica: Prov. Huancavelica, 3 km. SE. of Conaica, alt. 3,560-
3,590 m., Tovar 94; Bunbunya, W. of Conaica, alt. 3,700-3,750 m.,
Tovar 319. — Junin: Near Huancayo, open rocky hillside, alt. 3,300-
3,500 m., Killip & Smith 22133; Limestone cliff ledges, La Oroya,
alt. ca. 12,000 ft., Macbride & Featherstone 972; Limestone, Tarma,
alt. ca. 13,000 ft., Macbride & Featherstone 1059; Mito, alt. ca. 9,000
ft., Macbride & Featherstone 1472; Incahuasi, Soukup 3181 ; Mantaro
Valley, near Huachiacna, alt. 3,200-3,300 m., Weberbauer 6552 —
Libertad: Prov. Otusco: Usquil, alt. 3,000-3,100 m., Ferreyra 7637.
— Sin. Loc.: Ruiz & Pavon s.n.
Calceolaria scandens Pennell, Proc. Acad. Nat. Sci. Phila. 97:
164. 1945.
Climbing shrub with elongated, yellowish-brown, glandular-pu-
bescent stems. Leaves petiolate, ovate, 5-6 cm. long and 3-4 cm.
wide, acute, dentate, minutely pilose above, puberulent beneath and
reticulate- veined, cordate at base; petioles 15-25 mm. long, glandu-
lar-pubescent, basally thickened and connate across the nodes; in-
florescences of compound cymes; bracts petioled, bractlets smaller,
both of these foliose but less cordate than cauline leaves; flower stalks
glandular-pubescent; basal peduncles 6-8 cm. long; remaining stalks
decreasing in length to the 20-30 mm. pedicels; sepals lanceolate,
8-9 mm. long, acuminate, sparsely ciliolate; corolla yellow, exter-
nally glabrous; upper lip ca. 1.5 mm. long, lower lip ca. 16-19 mm.
long, saccate less than half its length, few hairs inside near the orifice;
anther cells equal, divaricate, each pair ca. 2.5 mm. long, little longer
than wide, dehiscing through the septum; style 1.5 mm. long, api-
cally curved.
Cuzco: Ollantaitambo, alt. 3,000 m., Cook & Gilbert 750 (US).
Calceolaria scapiflora (R. & P.) Benth. in DC., Prodr. 10: 207.
1846. Jovellana scapiflora R. & P., Fl. Peruv. 1: 13, t. 18, f.6. 1798.
Calceolaria mathewsii Benth. I.e. 207. C. scapiflora subsp. dentifolia
Pennell, Proc. Acad. Nat. Sci. Phila. 97: 152, t. 7a. 1945.
636 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Plants acaulescent, the radical, rosulate leaves ovate, flowers soli-
tary on pedicels mostly 2-10 cm. long. Leaves usually laxly pubes-
cent on one or both surfaces, 1.0-2.5 (-3.5) cm. long, from about half
to almost as wide as long, base attenuate, narrowly winging the short
petiole, margin subentire to dentate; pedicel glutinous, glandular-
pubescent or -puberulent; calyx lobes rarely 5, unequal, the anterior
longest and widest, ovate, 5-7 mm. long and 3.5-5.0 mm. wide, ex-
ternally puberulent, internally glabrous, venose; corolla yellow, red
dotted within, 16-28 mm. long, upper lip from little longer to twice
as long as the calyx, its short hood about as long as the sac of the
lower lip, from a little more than half as long to about as long and
wide as the lower lip that is saccate only up to one-eighth its length;
filaments ca. 2.5-3.5 mm. long, anther cells equal, parallel, about
three-fourths as wide as long; style from about half as long to longer
than the filaments, ovary obconic or obovate, wider than long.
Pasco: Uliachin Ridge, vicinity of Cerro de Pasco, alt. 14,800 ft.,
Jenks 38', Cerro de Pasco; alt. 14,000 ft., Macbride 3062. — Cuzco:
Urcos, alt. 15,000 ft., Stafford S. 43; Paruro, alt. 3,700-3,900 m.,
Vargas 843 (Isotype C. scapiflora subsp. dentifolia Pennell) ; Vargas
9819; Banks of the Checca River, Prov. Canas, Vargas 11009.—
Junin: Between Tarma and Jauja, alt. 4,500 m., Killip & Smith
21967; Puna, Mt. La Juntay, near Huancayo, alt. 4,700 m., Killip &
Smith 22100; Incahuasi, alt. 4,000 m., Soukup 3178. — Lima: Near
Antaicocha, Cerro Colorado E. of Canta, alt. 3,900-4,100 m., Pen-
nell 14662. — Sin. Dept. : Yauli, alt. ca. 13,000 ft., Macbride & Feather-
stone 909. — Sin. Loc.: Ruiz & Pawn, s.n.
Calceolaria serrata Lam., Illus 1: 51. 1783, non G. Don 1838.
Close to C. inflexa R. & P., differing in the following ways:
leaves up to 5 cm. long and 2 cm. wide; calyx lobes obtuse; upper lip
of the corolla minute; entire flower only up to 13 mm. long and 8 mm.
wide; foliage puberulent above and canescent beneath.
Reported from departments Lima, Cajamarca, and Amazonas, as
well as in Bolivia and Ecuador. None seen.
There is general agreement that C. serrata and C. deflexa are
closely allied or possibly conspecific. Their distinct geographic areas
indicates that we are dealing with two taxa.
Calceolaria sibthorpioides HBK., Nov. Gen. et Sp. 2: 382.
1817.
FLORA OF PERU 637
Procumbent herb, branches hispid-pilose. Leaves petiolate,
broadly ovate to subrotund, up to 1.5 cm. long and up to 1.1 cm.
wide, usually a little shorter and narrower, ca. 1.0 cm. long and 6-8
mm. wide, margins crenate, apex acute, base a little cordate, pilose
on both surfaces, especially on the venation beneath, lower surface
drying very pale, dull green; inflorescences corymbose or dichasiate,
terminal few-flowered; flower stalks glandular-pilose; calyx lobes ex-
ternally densely pilose, broadly ovate, 4-5 mm. long, almost as wide
at base, subacute to obtuse, margin ciliate; corolla yellow, sparsely
pilose, ca. 14-17 mm. long, sac of lower lip about 8-10 mm. wide
and deep, upper lip shorter than the calyx-lobes, lower lip 10-14 mm.
long, saccate less than half its length; anthers up to 3 mm. long, cells
equal, each cell about 1.8 times longer than wide; style ca. 2 mm.
long, ovary longer than the style, beset with short, erect hairs; cap-
sule not known.
Reported from Cajamarca (Hualgayoc) and Junin-Lima area
(Palca to Tarma) from 3,600-4,000 m. alt. The only specimen seen
(K) bears no data.
Calceolaria simulans Edwin, Phytologia 19: 395. 1970.
Herb, erect to repent, little- to much-branched, usually up to
25 cm. tall, sometimes only about 10 cm., upper parts of the stems
and branches more or less densely glandular- and eglandular-pubes-
cent, remaining parts of stems and branches glabrous to sparsely
eglandular-pubescent; upper leaves once or twice pinnate, lower very
variable, from dentate to lobed ; pinna dentate, sparsely mixed pubes-
cent beneath as the upper parts of the stems and branches; petioles
1-4 mm. long, pubescent as the lower surfaces of the pinnae; inflores-
cences terminal on the branches and stems, simple or once-branched,
few-flowered; bracts foliose, reduced; flower stalks up to 2.5 cm. long,
usually not exceeding 1.8 cm., vesture as the upper parts of the
branches; flowers yellow, small for the genus; calyx lobes a little
unequal, broadly ovate, 4-5 mm. long and 3-4 mm. wide, acute,
venose on both surfaces, red at base, sparsely glandular-puberulent
on the sinuate margins; corolla yellow, dark brown dotted, glabrous,
ca. 10-13 mm. long, upper lip 1.5-2.5 mm. long, lower lip 9-11 mm.
long, saccate a little more than half its length, proximal, non-
saccate part only little narrower than the distal saccate part;
stamens contiguous, small, anthers ca. 1.3-1.7 mm. long, little
shorter than the filament, cells equal or slightly unequal, divaricate,
little longer than wide, blackened basally near the septum; style very
638 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
short, up to 1 mm. long, apically curved; capsule sparsely glandular-
puberulent, longer than the calyx, lance-ovoid, abruptly narrowed
to a long neck.
Distributed sporadically from Ancash to Ayacucho and lea.
Ancash: Chacchash near Chiquian, 3,220 m. alt., Ferreyra 5687;
Casca, below Chiquian, Prov. Bolognesi, alt. 3,100-3,200 m., Fer-
reyra 7307 (US). — Ayacucho: Pampalca between Huanta and Rio
Apurimac, alt. 3,200 m., open hillside, Killip & Smith 22210. — lea:
Pisco Valley, alt. 2,900 m., Rauh-Hirsch P390. — Lima: Rio Blanco,
wet E. slope, alt. ca. 15,000 ft., Macbride 3037.
This species has the foliage of subgenus Calceolaria and flowers
of subgenus Cheiloncos. It is provisionally placed in the latter group-
ing. In Peru, thus far, this is the only taxon known to bridge the
subgenera.
Calceolaria solanifolia Edwin, Phytologia 19: 397. 1970.
Erect, branched, densely tomentose shrub, up to about 2 m. tall.
Wood, the oldest excepted, leaves, petioles flower stalks and calyx
bases densely beset with short, curled, white to brown, velvety, often
matted hair; leaves ovate, sometimes lanate, up to 8 cm. long and
5 cm. wide, rarely entire, mostly subentire to serrate-serrulate, apex
acute, base obtuse, dark green above, light yellow-green beneath,
veins very apparent to almost concolorous beneath; petioles 1.5-
2.5 cm. long, thickened, very densely pubescent; inflorescences a pair
of compound, cymose dichasia, in the axils of the uppermost leaves of
the stem and branches; basal peduncles up to 8 cm. long, intermedi-
ate flower stalks and pedicels abruptly decreasing in length, ulti-
mately less than 1 cm. long; flowers large for the genus; calyx lobes
ovate to elliptic, acute, 10-14 mm. long, 3-7 (-8) mm. wide at base,
yellow or light green in life, glabrous to sparsely pubescent externally,
the hair on or near the margins; corolla yellow, frequently minutely
papillose, upper lip 4-6 mm. long, lower up to 25 mm. long, fre-
quently 20-22 mm., saccate about half its length, the sac ovoid-
oblong, 12-14 mm. wide, inflated in life; anther cells a little unequal
in length and very unequal in shape, walls and septa thickened, both
cells at least two times longer than wide, each anther 7-8 mm. long,
the smaller cell 3.0-3.3 mm. long; style 5.5-6.5 mm. long, apically
curved; ovary ovoid, pubescent, a little shorter than the style; cap-
sule pubescent, ovoid or elliptic-ovoid, shorter than the calyx, acute
apically.
FLORA OF PERU 639
Amazonas: Prov. Chachapoyas, E. side of Cerro Calla-Calla on
lower and middle slopes. Jalca interfingered with loma and montane
forest, alt. ca. 2,800-3,200 m., Edwin 3628 (F); 3626; 3630; Moun-
tains E. of Balsas, alt. ca. 3,200 m., Osgood & Anderson 76.
This taxon is distinct from all other Calceolarias in having the
combination of unequal anther cells, large corolla, dense tomentose
or lanate pubescence, large thickened leaf blades on long petioles and
paired inflorescences, arising from only the uppermost pair of leaf-
axils of the stems and branches.
Calceolaria sonchensis Pennell ex Edwin, Phytologia 19: 397.
1970.
Pubescent herb up to 50-60 cm. tall, stems square, sulcate-striate.
Leaves bizarre, blades broadly ovate or triangular-ovate, 2.5-4.5 cm.
long and 1.5-2.3 cm. wide, irregularly doubly dentate, apex acute,
base truncate or broadly obtuse, petiolate, the uppermost excepted,
these sessile, pubescent on both sides; petioles broadly winged, con-
nate, widest at or near the stem, entire or minutely serrate, usually
a little shorter than the blades; inflorescences in axils of upper leaves,
cymose-dichasia; the lax pedicels ca. 23-33 mm. long, on peduncles
up to 15 cm. long, usually exceeding 10 cm. ; sepals ovate, 10-12 mm.
long, acute or frequently acuminate, margins distantly or less often
densely serrate, the teeth black, spinose; calyx tube very short, ca.
1 mm. long, calyx externally pubescent; corolla wholly glabrous,
small for the genus, upper and lower lips subequal, ca. 7-8 mm. long
and wide, the hooded upper lip completely covering the brown, di-
varicate, oblate anthers whose filaments do not exceed 1 mm. long;
style ca. 2.5-3.5 mm. long, exceeding the stamens, ovary pubescent
with short, erect hairs, ovoid or hemispherical, shorter than the style;
capsule 2-3 mm. long, subhemispherical or broadly ovoid, tapering
to apex, densely pubescent, loosely invested by the sepals.
Amazonas: Mountain south of Tambo de Ventilla, thin loam over
clay banks, alt. 2,700-2,900 m., Pennell 15791 (PH). The only col-
lection known.
Close to Calceolaria connatifolia Pennell from which it differs in
having mostly petiolate leaves, serrate sepals, and glabrous corolla
with subequal lips, as well as in the general presence of pubescence
and shape of the anthers.
Calceolaria sparsiflora 0. Ktze., Linnaea 24: 228. 1851. C. her-
rerae Kranzl. and C. herreae var. ramosissima Kranzl. in Fedde, Rep.
Sp. Nov. 27: 15. 1929.
640 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Erect, branched shrub, main stem usually aphyllus, branches and
branchlets often rectangular, new wood frequently drying light-yel-
low, puberulent with mixed glandular or eglandular, pustulate hair,
occasionally sparsely eglandular-puberulent, then older wood gla-
brous, older wood usually puberulent as the younger but less densely
so, wood ultimately devoid of hairs, only the red-pustulate bases re-
maining (muricate) . Leaves variable in shape, size, and vesture, from
narrowly cuneate-obovate to ovate, not infrequently lance-ovate or
lance-obovate, 0.6-1.8 (-2.7) cm. long and 2-9 (-18) mm. wide, most
frequently less than 2 cm. long and 3-8 mm. wide, usually densely
beset beneath with gray-white pulverulent tomentum, occasionally
lanate (sericeous) rarely glabrous or almost so, glabrous above, mar-
gin dentate or deeply dentate with usually distant teeth on the upper
half to two-thirds, lower part of margin entire, narrowed to apex and
base, base most often cuneate, usually on petioles 0.5-3.0 mm. long;
petioles frequently villose; inflorescences simple to once-branched
dichasia in the axils of upper leaves; flower stalks mostly puberulent,
10-20 mm. long, frequently 10-15 mm.; calyx lobes puberulent ex-
ternally, pulverulent-puberulent internally, especially near the mar-
gins, ovate to oblong-ovate, 5-6 mm. long, acute; corolla yellow, up
to 25 mm. long, often not exceeding 18 mm., sometimes only 13-
15 mm. long, essentially glabrous, upper lip up to about half the
length of the calyx lobes, lower lip saccate less than half its length,
sac as wide as long or wider; stamens hidden under the lower lip,
filaments short, anther cells equal, divaricate, up to twice as long as
wide, each pair ca. 2.0-3.0 mm. long; style 1.5-2.0 mm. long, ovary
glabrous to occasionally sparsely glandular; capsule with vesture as
the ovary, oblong to narrowly ovoid, neck only little constricted,
apically obtusish, equal to or more often longer than the calyx.
Cuzco: Paucartambo, steep, dry, stony slope, Balls 6723; Cerro
Sape, vicinity of Cuzco, stony ground, alt. 3,400-3,500 m., Ferreyra
2664; Alt. 3,300-3,500 m., Herrera 527; Sin. Loc. Herrera 1704- (frag-
ment of type of C. herrae var. ramosissima Pennell ; Prov. Paucar-
tambo, Hacienda Churu, Herrera s.n.; Sacsahuaman above Cuzco,
alt. 3,500-3,600 m., Pennell 13554; San Sabastian, alt. 3,300-3,400
m., Pennell 13598; Ollantaitambo, alt. 2,900-3,000 m., Pennell 13647;
Huancalli, alt. 3,400-3,500 m., Pennell 13713; E. of Pisac, alt. 3,200-
3,500 m., Pennell 13721; Colquipata, alt. 3,200-3,500 m., Pennell
13773; SW. of Paucartambo, alt. 3,200-3,500 m., Pennell 14169;
1^175; Suburbs of Cuzco, Soukup 48; 128; Alt. 3,400-3,500 m., neigh-
borhood of Kenko or Choko, mixed data for Vargas 230; Sicuani, alt.
FLORA OF PERU 641
slopes 3,600-3,650 m., Vargas 1472; 9840; Slopes of Chicon, alt.
2,800-2,900 m., Vargas 1542; 11066.
Calceolaria subscaposa Edwin, Phytologia 19: 398. 1970.
Erect, annual herb, up to 20 cm. tall, stem with up to 3 pairs of
leaves and unbranched or occasionally once-branched, rarely twice-
branched, apically usually bearing 2 flowers when unbranched, or
each apex 2-flowered when branched, rarely with a 3-flowered cyme
at the apex, stems glabrous, internodes several times longer than the
small leaves. Leaves petiolate, blades ovate in outline, up to about
1.0 cm. long and 0.8 cm. wide, usually sparsely puberulent above,
thinly pulverulent beneath, margins dentate to with 2 pairs of den-
tate-denticulate lobes; petioles up to about half as long as the blade
especially on the lowermost pair of leaves decreasing in size to the
apex of the stem, puberulent; pedicels mixed glandular and eglandu-
lar-pubescent, mostly 3.5-5.5 cm. long; bracteoles occasionally de-
veloped near the base of the flower stalks; sepals glabrous, broadly
ovate, acute, 4.5-5.0 mm. long and 3.8-4.2 mm. wide, margins en-
tire, a little thickened; corolla yellow, 9-11 mm. long, upper lip gla-
brous, ca. 4 mm. long, lower lip not twice as long as the upper,
sparsely red-gland-dotted proximally on the upper part of the sac
that is about three-fourths the length of the ca. 7 mm. long lip; both
anther cells fertile, both larger than the very short, thin connective;
style ca. 1.5 mm. long; capsule from as long as to more often longer
than the calyx, ovoid, basally ca. 4.5 mm. wide, abruptly narrowing
to a neck about a third the length of capsule, sparsely gland-tipped
pubescent, glands soon fugitive.
Known only from the copious type specimen.
Lima: Along Rio Chillon, above Obrajillo, open rocky banks, alt.
2,800-3,200 m., Pennell 14410 (PH).
No other taxon in this subgenus has the following combination
of characters: subscapose stem, almost wholly pedicellate flowers,
lower corolla lip less than twice as long as the upper and anther cells
both fertile and separated by a very short connective.
Calceolaria tenuis Benth. in DC., Prodr. 10: 205. 1846.
Lax annual, stems glabrous below, puberulent to pubescent
above, internodes much longer than the leaves that are most often
small for the plant. Blades variable, usually subsimply lobate be-
low, on longer petioles than the incised, subpinnatifid, occasionally
642 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
sessile upper leaves, lobes usually obtuse and toothed, blades ovate
in outline, often not exceeding 3 cm. long and 2 cm. wide, varying to
5-6 cm. long, sparsely puberulent; flowers borne singly or in pairs in
the upper leaf-axils, most often not exceeding 11 mm. long, frequently
only 8 mm. long; pedicels up to 2 cm. long, glandular-pubescent;
calyx lobes ovate, acute, 5-6 mm. long, glabrous or the entire to
sparsely, finely toothed margins ciliate, veins 5-7, apparent on both
surfaces; corolla yellow, glabrous or sparsely red-glandular, glands
sessile, upper lip ca. 2 mm. long, lower 6-9 mm. long, saccate most
of its length; anther connective enlarged, auriculate, larger, upper
cell about as long as the connective, lower smaller, modified, fertile
or sterile; style ca. 1.0-1.5 mm. long; capsule glabrous, conic-ovoid,
much longer than the calyx, abruptly narrowed to a long neck.
Junin: Oxapampa, Soukup 2J+17. — Lima: Between Huaquicha
and Palacala above Surco, alt. 2,600-2,700 m., Ferreyra 3^8.
This taxon may be merely a depauperate form of Calceolaria
delicatula Kranzl.
Calceolaria tetragona Benth. in DC., Prodr. 10: 218. 1846.
Erect, essentially glabrous shrub, differing from C. cerasifolia
Benth. in the following ways: blade margins almost entire to sparsely
serrulate; petioles usually not exceeding 1.3 cm. long; inflorescences
few-flowered, corymbose; flower stalks longer, pedicels up to 2.5 cm.
long; calyx ca. 7-8 mm. long, externally glabrous or sparsely puber-
ulent, internally glabrous; corolla glabrous; anther cells unequal, 2-
3 times longer than wide, septum and walls thickened, persisting;
ovary and capsule often sparsely to densely glandular-puberulent.
Amazonas: Prov. Chachapoyas, Mathews s.n.; Cerro Puma-urcu,
SE. of Chachapoyas, alt. 3,000-3,300 m., Pennell 15671', 15719;
Bank between Chincha and Tingo, alt. 2,400-2,500 m., along Rio
Sonche, E. of Molinopampa, Pennell 15783; Cerro de Fraijaco
(Huani-Huni) NE. of Tambo de Ventilla, alt. 2,800-2,900 m., Pen-
nell 1584.0; Alt. 3,400 m., Pennell 15863. — Cajamarca: Prov. Cutervo,
trail between Socota and Tambillo, alt. 3,100 m., Stork & Horton
10182. — Piura: Chira, alt. 9,000 ft., Carricker s.n. (northernmost
Peru, near Ecuador).
Calceolaria tomentosa R. & P., Fl. Peruv. 1: 15, t. 22b. 1798.
Tall, erect, perennial herbs, stems, flower stalks and lower sur-
faces of leaves densely lanate, hairs obscuring the tissue, the leaves
FLORA OF PERU 643
rarely lanate only on the venation. Petioles connate, widest at or
near the stems, uppermost blades sessile, entire or minutely toothed;
leaf blades ovate or triangular-ovate, size variable, up to 16 cm. long,
mostly 5-11 cm. long and a little more than half as wide, coarsely
doubly crenate, acute, bases truncate or broadly subcordate; flowers
forming loose panicles or compound dichasia, from medium to very
large; peduncles and pedicels ebracteolate, stalks decreasing in length
toward the apex of the inflorescence; calyx lobes ovate, acute from
12-20 mm. long, densely pubescent on both surfaces, some of the
inner surface hairs very short and arranged in a line near to and
paralleling the margin; corolla glabrous, except for a tuft of long,
white hair at the base of the lower lip, 17-33 mm. long, upper lip
4-6 mm. long, overtopping the genitalia, lower up to 32 (-35) mm.
long, about a half to two-thirds as wide, sac (in life) inflated, about
half the length of the lower lip, longitudinally ridged; anther cells
unequal, usually twice as long as wide or longer, septa and walls
thickened, filaments short, ca. 2-3 mm. long; ovary densely puberu-
lent, style 4-6 mm. long, apically decurved; capsule ovoid, glutinous-
puberulent, much smaller than the loosely investing sepals.
Material from northern Peru more densely lanate than the col-
lections from Central Peru.
Close to C. perfoliata L. f. of Colombia from which it differs in
anther cell and calyx lobe shape and in other details.
Amazonas: Taquia, E. of Chachapoyas, stream bank, alt. 2,300-
2,400 m., Pennell 15568; Daguas on Rio Sonche, east of Chacha-
poyas, alt. 2,400 m., Pennell 15748; Puente de Pipos, opposite
Daguas, alt. 2,000-2,100 m., Pennell 15752; Gullied arroyo-side,
above stream, 1-2 km. NE. of Chachapoyas, alt. 2,300-2,400 m.,
Pennell 15897; Lower slopes of Puma-rucu above Hda. Sta. Isabel,
1.5-2.0 km. SE. of Chachapoyas, alt. 2,400-2,500 m., Wurdack 423.
— Cajamarca: mountain east of Celendin, alt. 2,600-2,700 m., Pen-
nell 15197.— Huanuco: 6 km. north of Pillao, alt. 3,000 m., Woyt-
kowski 76. — Junin: Between Tarma and Palca, alt. 2,600 m., Killip
& Smith 23373; Carpapata, Soukup 3437; Between Palca and Car-
papata, alt. 2,900 m., Stork 10972. — Lima (?): Between Tarma and
San Ramon, growing with small shrubs above Carpapata, alt. 2,300-
2,400 m., Ferreyra 3738; East of Huasahuasi, alt. 2,400-2,500 m.,
Hutchison 1095. — Piura: Chanchaqui, moist rocky banks, alt. 1,700-
1,900 m., Pennell & Ferreyra 14910. — Sin. Loc.: Ruiz & Pavon s.n.
644 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria triloba Edwin, Phytologia 19: 399. 1970.
Climbing or erect shrub, wood densely tomentose when young,
vesture decreasing with age, densely foliaceous on distant or close
branches. Leaves subcoriaceous, broadly ovate in outline, less than
1 cm. long and wide, pilose above, densely lanate beneath, the vesture
obscuring the tissue, margins little to very thickly revolute, with one
pair of broad, basal, lateral lobes, these either divergent or somewhat
antrorse, when divergent base of blade truncate, when antrorse base
broadly ovate; costa and few secondary veins very deeply impressed
above; flowers solitary or in 3-flowered dichasia, axillary in the upper
parts of branches or terminal on the stem; flower stalks up to 2 cm.
long, mostly shorter, less than 1 cm. long, densely tomentose; calyx
externally pilose or puberulent, internally pilose or glandular-puber-
ulent, glands shining red, elliptic to ovate, 4-5 mm. long and 2-4 mm.
wide, acute; corolla yellow, often with 1 or 2 purple bands, externally
glabrous or red-glandular and papillose, internally with a tuft of
white hair at the base of the lower lip, upper lip minute, lower lip
14-18 mm. long, saccate more than half its length, sac abruptly
flaring, globose, upturned; anther cells divaricate, equal, each pair
2-3 mm. long, more or less than twice as long as wide, filaments
shorter than the thickened, 2-3 mm. long, apically curved style;
capsule unknown.
Ancash: Polylepis thicket, alt. 3,500-3,700 m., mountain E. of
Copa Grande, N. of Chancos, Pennell 15371 (PH); Libertad: Open
jalca above Motil, alt. 3,200 m., Pennell & Ferreyra 11^8^8.
Two varieties, the typical and one other, might well be estab-
lished for this taxon based on corolla vesture, amount of recurving
of leaf margins, and calyx lobe shape. In view of the scanty material,
the taxon is provisionally treated as composed of only one variety.
This taxon is close to C. inaudita Kranzl., which differs in having
almost equal corolla lips, only barely lobed leaves, and reniform an-
ther cells.
Calceolaria tripartita R. & P., PI. Peruv. 1: 14, t. 20a. 1798.
C. heterophylla Willd., Enum. Hort. Berol. 29. 1809. C. scabiosae-
folia Roem. & Schult. in L., Syst. Veg. 1 : 187. 1817. C. chelidonioides
HBK., Nov. Gen. et sp. 2: 378. 1817. C. mandoniana Kranzl. in
Pflanzenr. IV (257c): 30. 1907. C. puru-puru Kranzl. in Fedde,
Rep. Sp. Nov. 27: 2. 1929. C. tripartita mandoniana Pennell, Proc.
FLORA OF PERU 645
Acad. Nat. Sci. Phila. 97: 175. 1945. C. obscura Pennell, Proc.
Acad. Nat. Sci. Phila. 97: 176, fig. 16a. 1945. '
Very variable herb or vine, stems from prostrate to erect, when
a vine often large, much-branched, occasionally extending to several
meters long, stems typically glabrous below, increasingly pubescent
toward the apex. Leaves petiolate, blades most often with 1-3 pairs
of pinnae, pinnae larger from the base to apex of the leaf, terminal
leaf-segments ovate, margins usually doubly dentate, teeth coarse,
occasionally blades pinnatifid and pinnate or lobate or merely doubly
dentate, the pinnae usually 1-3 cm. wide, the blade up to 15 cm. or
longer, most often 4-8 cm. long, tissue glabrous to pilose on one or
both surfaces; petioles from up to half as long as and thinly winged
by the blade to almost or quite lacking, when present connate across
the nodes, most often densely pubescent; flowers large for the sub-
genus, the lower 1 or 2 in axils of the branches and the upper typi-
cally in 3-flowered dichasia, occasionally cymose or 5-7-flowered
subumbellate; pedicels mostly 2.0-4.5 cm. long, glandular-pubescent,
often also glutinous; calyx lobes ovate, acute to acuminate, glandu-
lar-pubescent, 5-10 mm. long, margins often distantly minutely
toothed; corolla yellow, pubescent at the orifice internally, upper
lip 4-7 mm. long, lower 15-30 mm. long, saccate most of its length,
sac inflated, about three-fourths as wide as long, non-saccate part
of the lip abruptly narrowed; lower anther cell sterile, connective
sigmoid, auriculate, most often longer than the cells; style 2.0-2.5
mm. long, gradually little tapering to a blunt apex.
The most commonly collected species in the subgenus and one of
the most frequently encountered in the genus in Peru. Very variable
in habit as well as in size of flower parts and leaf shape. It is best
separated from the remaining species of the subgenus by a combi-
nation of large flowers, sterile anther cell, pinnae increasing in size
from base to apex of each leaf and capsule most frequently broader
than long. The leaf blades occasionally are less than 3 cm. long.
Generally distributed in wet sites from Piura to Cuzco, and prob-
ably further south.
The following citations are selected from the copious material.
Amazonas: Cerro Puma-urco, SE. of Chachapoyas, alt. 2,500-
2,600 m., Pennell 15508; Utcubamba, S. of Chachapoyas, alt. 1,700-
1,750 m., Pennell 15588; Along Rio Sanche E. of Molinopampa,
1 This is only a small part of the copious synonymy attached to the name.
The selection is limited to those pertinent in Peru.
646 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Streamlet at Chincha, alt. 2,400 m., Pennell 15781. — Ancash: Cerro
in Rio Fortale'za Valley, Lima-Huaraz road, W. of Comococha, alt.
ca. 2,500 m., Edwin & Schunke V. 3582; Colcas, in stream and bog,
alt. 2,500-2,600 m., Pennell 15384; Small stream above Yungay, alt.
2,650-2,700 m., Pennell 15421.— Apurimac: Prov. Abancay, 12 km.
on road to Cuzco, ca. 5 km. NW. of Abancay, Iltis & Ugent 627.—
Ayacucho: Pampalca, between Huanta and Rio Apurimac, alt. ca.
3,200 m., Killip & Smith 2327k; Mountains NE. of Huanta, alt.
3,100-3,200 m., Weberbauer 7506.— Cajamarca: E. side of Cerro
Calla-Calla, below paramo, Edwin & Schunke V. 3627; Along stream
E. of Cajamarca, alt. 2,600-2,700 m., Pennell 15224; Prov. Cutervo,
in swamp, Rio Sucse Valley, W. of Socota, alt. 2,800 m., Stork &
Horton 10115. — Cuzco: Ollantaitambo, along stream, alt. 2,850-
2,900 m., Pennell 13669; East of Cuzco City, on hills, alt. 3,500-
3,800 m., Pennell 13703; Huancalli, alt. 3,400-3,500 m., Pennell
13715; Near Rio Quencomayo, below Colquipata, alt. 3,200-3,300
m., Pennell 13779; Near Rio Yanamayo, below "Pillahuata," alt.
2,000-2,200 m., Pennell 14043 (type collection of C. obscura Pennell) ;
Limatambo, alt. 2,100 m., Vargas 232; Macchu-picchu, Prov. Uru-
bamba, alt. 2,000 m., Vargas 815; Pueblo Marcapata, Prov. Quispi-
canchis, alt. 3,200 m., Vargas 4828. — Huancavelica: Prov. Castro-
virreina, near Cordova, alt. 3,020-3,300 m., Metcalf 30289; Hills
N. of Pampas, Prov. Tayacaja, alt. 3,250 m., Stork & Horton 10235.
— Huanuco: Carpish Pass, 84 km. from Tingo Maria, on highway to
Lima, alt. ca. 2,750 m., Allard 21112; Quisca, alt. 2,050 m., Scolnik
1042; El Tambillo on road to Panao, alt. 2,650 m., Scolnik 1054;
6 km. S. of Huanuco, in dry stream bed of Huallaga River, alt.
1,920 m., Stork & Horton 9377. — Junin: Alongside ditch, near road,
1-3 km. E. of Palca, alt. ca. 2,000 m., Edwin & Schunke V. 3920;
2-4 km. E. of Huasahuasi, alt. 2,400-2,500 m., Hutchison 1097; At
bridge, Canyon of Rio Huasachuasi, below Huasahuasi, alt. ca. 2,400
m., Hutchison 1139; Prov. Cerro de Pasco, near Ataura, between
Huariaca and Cerro de Pasco, alt. 3,000 m., Hodge 6267; Tarma,
alt. 3,000-3,200 m., Killip & Smith 21865; Huacapistana, alt. 1,800-
2,400 m., Killip & Smith 24095.— Libertad : Otuzco, alt. 2,830 m.,
"zapatos de la reina," Angulo 0935; Above Motil, alt. 3,000 m., Pen-
nell & Ferreyra 14845. — Lima: Matucana, alt. 8,000 ft., sunny brook-
margin, Macbride 2935; Along Rio Chillon, above Obrajillo, alt.
2,700-3,000 m., Pennell 14366; San Buenaventura, alt. 2,800-2,900
m., Pennell 14541; At base of ocean-cliff, wet sandy soil, Barranca,
FLORA OF PERU 647
Pennell 15239; Hills S. of Surco, alt. 2,600-2,800 m., Pennell & Fer-
reyra 15252. — Piura: El Tambo, alt. 9,000 ft., Carricker s.n., July 1,
1933; Cerro Porculla, 3-11 km. W. of continental divide, W. slope,
Edwin & Schunke V. 3739. — Puno: Prov. Sandia, near Limbani, alt.
3,200-3,450 m., Metcalf 30486.
Calceolaria utricularioides Benth. in DC., Prodr. 10: 213.
1846. (ex. nomen. herb. Hooker.)
Soft annual herb, most often with unbranched stem, usually erect,
less often reclining, stems descrete or few-clumped, rarely prostrate
and rooting at the nodes, sparsely pubescent. Leaves few, mostly
much shorter than the internodes, lower short-petiolate, upper ses-
sile, usually ovate, entire, up to 2.5 cm. long, rarely longer and 6-12
(-18) mm. wide, sparsely pubescent on both surfaces; inflorescences
usually few-flowered with 2 basal peduncles either once or twice,
dicotomously branched or once or twice dichasiately branched;
flower stalks pubescent or glandular-pubescent; calyx lobes ovate,
up to about 3 mm. long; corolla yellow, lips from almost equal to the
lower about a third longer than the upper; filaments about 5 times
or more longer than the anthers; anther cells equal, divaricate, each
pair about 1 mm. long and little narrower; style little longer than the
stamens, thin, curving; fruit ovoid, longer than the corolla, sparsely
puberulent or glandular-puberulent.
As seen below, this taxon has been sporadically collected over
most of the length of Andean Peru, Piura to Arequipa. The species
unquestionably occurs in the areas between the presently known
stations and is probably one of the more common "slipper-flowers."
Amazonas: Cerro Puma-urcu, SE. of Chachapoyas, alt. 3,300-
3,400 m., Pennell 15696.— Ancash: Yanac, alt. 11,000-12,000 ft.,
Carricker s.n., March 28, 1932; Bolognesi, a little above Aquia, alt.
3,200-3,360 m., Ferreyra 7517.— Arequipa: Mollendo, Hitchcock
22397. — Cajamarca: SW. slope of Paso de Gavilan, S. of Cajamarca,
alt. 2,700-2,750 m., Pennell & Anderson 15102.— Libertad: Usquil,
Prov. Otuzco, alt. 3,000-3,100 m., Ferreyra 7622. — Lima: Huaquica,
above Surco, alt. 2,600 m., Ferreyra 693; Between Huaquica &
Palacala, alt. 2,600-2,700 m., Ferreyra 3442; Along Rio Chillon,
above Obrajillo, alt. 2,600-2,800 m., Pennell 14359; San Buenaven-
tura, alt. 2,700-2,900 m., Pennell 14530; S. of Surco, alt. 2,400-
3,000 m., Pennell & Ferreyra 15246; Huillacachi by Matucana, alt.
3,300 m., Weberbauer 5726. — Piura: Porculla, Carricker s.n., May 21,
1933.— Sin. Loc.: Weberbauer 6991.
648 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Calceolaria variefolia Edwin, Phytologia 19: 400. 1970.
Divergently branching shrub, wood and flower stalks appressed-
puberulent, branches opposite, distant, main stem aphyllus or almost
so. Leaves ovate, sometimes very broadly ovate, 2.5-6.0 cm. long
and 0.8-4.0 cm. wide, usually 4-5 cm. long, acute at apex, obtuse to
almost acute at base, margin subentire to subdistantly serrulate-
serrate, puberulent above, red-punctate beneath, punctations a little
impressed or on the surface, very short petiolate to subsessile; peti-
oles 0.3-2.0 mm. long, thickened; inflorescences 3-flowered dichasia
in the axils of the upper pairs of leaves; flower stalks up to 1.5 cm.
long, usually a little shorter; calyx lobes pubescent on both surfaces,
especially on and near the margins, ovate, 4-5 mm. long and ca.
3.0-3.5 mm. wide, obtuse; corolla yellow, glabrous, ca. 16-18 mm.
long, upper lip reduced to a ring, ca. 1.0-1.5 mm. long, lower lip
saccate less than half its length; filaments short, equally thickened
base to apex, anther cells less than twice as long as wide, equal or a
little unequal; style ca. 2 mm. long; ovary densely red-glandular;
capsule unknown. Only the type collection thus far seen.
Piura: Rocky bank above Canchaqui, alt. 1,700-1,900 m., Pen-
mil & Ferreyra 1J>916 (US).
Differs from Calceolaria boliviana (Britton ex. Rusby) Pennell,
in having much smaller flowers with ovate calyx lobes as opposed to
lanceolate lobes and short style, ca. 2 mm. long, as opposed to 6-
8 mm. and the vesture and branching habit differ.
Calceolaria velutinoides Edwin, Phytologia 19: 401. 1970.
Erect, branched shrub, wood, except the main stem and flower
stalks, densely lanose-tomentose, main stem glabrate, wood usually
drying light brown. Leaves short-petiolate, densely matted-pilose-
tomentose above and on the veins beneath, orange-red puncticulate
beneath and sometimes sparsely so above, ovate, usually not exceed-
ing 2.0 cm. long and most often 1.0-1.2 cm. wide, a few longer and
occasionally narrower, margins a little thickened, crenulate to entire,
apex acute, base subobtuse; flower stalks mostly not exceeding 1.3
mm. long; inflorescences corymbose-dichasia, usually once branched,
few-flowered, terminal or axillary in upper leaves; calyx lobes exter-
nally pilose and red-puncticulate, internally with a row of white hairs
along the margins and at the acute to short-acuminate apex, ovate,
ca. 6 mm. long and 4 mm. wide; corolla yellow, glabrous, 8-12 mm.
long, upper lip reduced to a narrow ring, lower saccate less than half
its length, almost as wide as long; filaments less than 2 mm. long,
FLORA OF PERU 649
anther cells equal, divergent, usually 2-3 or more times longer than
wide, occasionally only ca. 1.8 times; style ca. 2 mm. long, little if at
all thickened; ovary glabrous to glandular-pustulate; capsule not
seen.
Amazonas: Cerro Puma-urcu, SE. of Chachapoyas, shrub on and
near rocks, alt. 3,100-3,200 m., Pennell 15555 (US); Cerro Puma-
urcu, SE. of Chachapoyas, rocks near base of cliff, alt. 3,300-3,400 m.,
Pennell 15695.
Close to C. croceopunctata and C. rugulosa Edwin, differing from
the former in inflorescence structure, style length, and in lacking an
acumin at the apex of the septum separating the anther locules and
from the latter in having much smaller leaves and shorter petioles,
simpler inflorescences, and also glabrous corolla with longer sac.
Calceolaria verticillata R. & P., Fl. Peruv. 1: 17. 1798. C. mul-
tiflora Cav., Icon. 5: 29, t. 449. 1799. C. gemelliflora Cav., Icon. 5:
30, t. 450. f.l. 1799. C. angustiflora Hook., Bot. Mag. t. 3094. 1831.
Shrubby, much-branched, up to 50 cm. tall and even taller.
Stems sparsely, branches densely setose. Leaves opposite or less
often ternate from the base to the middle of the stem, always ter-
nate on branches and upper stem, ovate, size very variable, from
about 1.5 cm. to 6 cm. long and 0.6-2.8 cm. wide, those of the stem
much bigger than those on the branches, usually sparsely pubescent
above and densely beneath, sometimes glabrous or almost so, apex
acute, base much wider, margins dentate to doubly dentate, blades
short-petiolate; inflorescences paniculate, often interrupted; bract-
lets reduced, foliose; flower stalks more or less densely puberulent;
calyx-lobes lanceolate to lance-acute, 3-4 mm. long, glabrous to
sparsely setose, margins entire, sometimes fimbriate; corolla yellow,
externally papillose, upper lip minute, lower 9-15 mm. long, saccate
less than half its length, proximal part abruptly narrower than the
saccate, distal part; anther cells equal, divaricate, papillose, each
pair ca. 2.5 cm. long, ca. 1.5 times longer than wide; style 1.5-4.0
(-5.0) mm. long; ovary beset with gland-tipped hair; capsule with
vesture as the ovary, ovoid in outline, usually much longer than the
calyx, upper half abruptly constricted, beak-like, style persisting.
A commonly collected, well marked taxon, especially frequent in
and near the Department of Lima.
The citations below have been selected from the copious material
in major herbaria.
650 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Ancash: Rocky jalca, Cerro Shaurema, SE. of Huaraz, alt. 3,200-
3,300 m., Pennell 15293. — Arequipa: Que"quena near Arequon, alt.
8,000 ft., Stafford 295; Characato, alt. 2,400-2,600 m., Vargas 8045.
— Cajamarca: Chaupe, alt. 5,500 ft., Carricker s.n., July 21, 1933.—
Junin: Huancayo, alt. 3,317 m., Soukup 2770; Incahuasi, alt. 4,200
m., Soukup 3202. — Liberyad: Cerro Campana, Prov. Trujillo, alt.
550 m., Lopez 0387. — Lima: Huayopampa, Prov. de Canta, Ferreyra
2000; Valley of Rio Rimac, near Lima-Aroya road at Km. 90, E. of
Lima, alt. 3,000 m., Goodspeed & Weberbauer 33058; Matucana, Mac-
bride 2905; Along Rio Chillon, near Obrajillo, alt. 2,200-2,500 m.,
Pennell 14-320; Canta, open rocky slopes, alt. 2,700-2,900 m., Pennell
14329; San Buenaventura, alt. 2,700-2,800 m., Pennell 14520; S. of
Surco, rocky granitic banks, alt. 2,300-2,600 m., alt. Pennell &
Ferreyra 15241; Cerro Jeronimo, Soukup 1612; Lamboraque, alt.
3,008 m., Soukup 4234- — Moquegua: Rainy-green formation, Caru-
mas, alt. 3,100 m., Weberbauer 7289.
Calceolaria virgata R. & P., Fl. Peruv. 1: 20. t. 31. fig. a. 1798.
Erect shrub with virgate branching, branches, and stem pubes-
cent. Leaf blades subsessile to short-petiolate, opposite or less often
ternate, ovate, 1.2-4.0 cm. long and 0.6-1.8 (-2.2) cm. wide, most
often pubescent on both surfaces, occasionally only on veins beneath,
margins serrate, apex acute to subobtuse, narrowed to base; petioles
on the stems often thickened at base, woody and connate; inflores-
cences irregularly cymose, corymbose or subumbellate, usually aris-
ing from the axils of upper leaves, several on each branch; flower
stalks usually glandular-pubescent; calyx lobes broadly ovate, 5-7
(-10) mm. long, acute, externally more or less, densely pubescent
corolla yellow, mostly 9-15 mm. long, glabrous, upper lip shorter
than the calyx, lower lip saccate less than half its length; filaments
short, anther cells equal, divaricate, more than to less than twice as
long as wide; style 1.5-4.0 mm. long; ovary usually glandular; cap-
sule ovoid, abruptly narrowed to a short neck, about as long as the
calyx or little longer, glandular-punctate, glands golden-yellow.
Leaf blades pubescent beneath on tissue and veins; calyx lobes 5-
7 mm. long; style 1.5-2.5 mm. long C. virgata f. virgata
Leaf blades pubescent beneath on veins only; calyx- lobes 7-10 mm.
long; style 3-4 mm. long C. virgata f. llamaensis.
Calceolaria virgata R. & P. f. virgata. C. extensa Griseb.,
Symb. Ant. 238. 1879; non Benth.
FLORA OF PERU 651
This taxon is the most commonly collected Calceolaria in the
subgenus Cheiloncos, only C. tripartita R. & P. in the subgenus Cal-
ceolaria is better represented in herbaria. Specimens seen from Piura
and Cajamarca in the north to Arequipa and Puno in the south.
The citations below have been selected from the copious material.
Ancash: Yanac, 12,000-13,000 ft., Carricker s.n.; Matarragra,
cerro NW. of Chiquian, alt. 2,560-3,520 m., Ferreyra 5600; S. of
Huaraz, alt. 3,100-3,200 m., Pennell 15264; 15302; N. of Chancos,
alt. 3,200 m., Pennell 15318;Paso de la Fortateza, alt. 3,800-3,850 m.,
Pennell 15389; W. slope of Cordillera Negra, below Paso de Collan,
alt. 3,560 m., Pennell 15474- — Apurimac: Quishuala, alt. 11,900 ft.,
Balls 6902. — Arequipa: Mollendo, Hitchcock 22396. — Ayacucho:
Pampalca, alt. ca. 3,200 m., Killip & Smith 22249. — Cajamarca:
Cajabamba, Carricker s.n.; Bajando, summit of el Gavilan, Contu-
maza Prov., alt. 3,200m., Ferreyra 3332; Mtn. E. of Celendin, alt.
2,700-2,900 m., Pennell 15194; NE. of Cajamarca, alt. 2,600-2,700
m., Pennell 15229; Enconada, alt. 3,000 m., Pennell & Reichlin 15028.
— Quinuamayo, alt. 3,500-3,600 m., Pennell & Reichlin 15058; Vicin-
ity of Celendin, alt. 2,625 m., Woytkowski 3. — Cuzco: Paucartambo,
alt. 11,500 ft., Balls 6825; Sacahuaman, above Cuzco, alt. 3,500-
3,600 m., Pennell 13568; Coraupampa, between Cuzco and Pisac,
alt. 3,400-3,500 m., Pennell 13710; Colquipata, alt. 3,500-3,700 m.,
Pennell 13770; Chincheros, alt. 3,700 m., Vargas 1535; Que"brada de
Llulluchayocc, alt. 3,400 m., Vargas 4291; Marcapata, Quispican-
chis Prov., alt. 3,200 m., Vargas 4829; Amaru, Pisac, alt. 3,500 m.,
Vargas 5025; Calca Prov., alt. 3,100-3,200 m., Weberbauer 7899.—
Huancavelica: Near Cordova, Castrovirreina Prov., alt. 3,050-3,300
m., Metcalf 30253. — Junin: Between Viques and Ingahuasi, S. of
Huancayo, alt. 3,150 m., Killip & Smith 22165; Carpapata, above
Huacapistana, alt. 2,700-3,200 m., Killip & Smith 24369; Mito, ca.
9,000 ft., Macbride & Featherstone 1456; Huancayo, Que*brada Occo-
pilla, Soukup 3629. — Libertad: Above Motil, Km. 96, between Tru-
jillo and Huamacucho, Ferreyra 3016; Agallpampa, alt. 3,050 m.,
Lopez 0874; 2-3 km. above Huamacucho, alt. 3,200 m., Pennell &
Ferreyra 14869; Santiago de Chuco, alt. 3,150 m., Stork & Horton
9946. — Lima: Rio Blanco, Huarochiri Prov., alt. 3,600 m., Asplund
11389; Above Surco, near Llacllighi, alt. 2,500-2,800 m., Ferreyra
687; Below Oyon, Cajatambo Prov., alt. 3,200-3,300 m., Ferreyra
3543; Matucana, alt. ca. 8,000 ft., Macbride & Featherstone 109;
Along Rio Chill6n, above Obrajillo, alt. 2,600-2,800 m., Pennell
652 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
11*357', San Buenaventura, alt. 2,700-2,900 m., Pennell 11*51*0; near
Antaicocha, Cerro Colorado, E. of Canta, alt. 3,200-3,900 m., Pen-
nell 14626; Huaros, alt. 3,300-3,600 m., Pennell 11*717.— Pasco:
Between Salcachupan and Cerro de Pasco, alt. 3,400 m., Ferreyra
6600. — Piura: Canchaque, Soukup 1*680. — Puno: Ollachea a Pte.
Ackopampa, Carabaya Prov., alt. 3,100 m., Vargas 6970.
Calceolaria virgata f. llamaensis Edwin, Phytologia 19: 405.
1970.
Herb or shrub differing from the typical form in the following
ways: leaf blades pubescent beneath on the veins only; calyx lobes
7-10 mm. long; style 3-4 mm. long; petioles very short, hardly
1 mm. long.
Cajamarca: Herb, open bank, below Llama, alt. 1,850 m., Pen-
nell 15911* (PH) ; Shrub, banks and rocks, Llama, alt. 1,900-2,400 m.,
Pennell 15919.
Calceolaria viscosa R. & P., Fl. Peruv. 1: 18, t. 30. fig. a. 1798.
Small branched shrub, upper part sometimes soft and declining,
wood puberulent, internodes usually longer than the leaves, entire
plant red-resinous dotted. Leaves very short-petiolate; petioles 1-
2 mm. long or almost wanting, sometimes with abbreviated shoots in
the axils and the leaves appearing verticillate; blades glabrous, ovate,
mostly 2.1-4.0 (-5.3) cm. long and 1.0-1.8 (-2.2) cm. wide, usually
few- and irregularly serrate, acute at apex, less so at base, veins
sometimes obvious beneath; inflorescences several flowered corymbs
or twice branched dichasia, in axils of upper and uppermost leaves;
calyx lobes ovate, 5-6 mm. long, acute, externally pubescent, corolla
yellow, 17-20 mm. long, externally minutely puberulent, especially
on the margins, upper lip 2-3 mm. long, lower lip 15-17 mm. long,
saccate less than half its length and sac 8-9 mm. wide, wider than
long; filaments short, anthers large for the narrow corolla base, 2.0-
2.5 mm. long, at least twice as long as wide; style 2-3 mm. long, ovary
dark red when dry, hemispherical, shorter than the style; capsule
unknown.
Ancash: Around Cerro de Chiquian, Prov. Bolognesi, alt. 3,100-
3,600 m., Ferreyra 5683; 7296; 731*0.
Fragments of Ruiz and Pavon and Dombey collections from Lima
and Tacna were also studied. The disjunction between the Ancash-
Lima and Tacna material strongly suggests that the species may be
expected to occur between these departments.
FLORA OF PERU 653
Calceolaria vulpina Kranzl. in Engler, Bot. Jahrb. 54: Beibl.
119, 18. 1916.
Plant erect, shrubby and branched or unbranched, stem densely
beset with septate hair and often glandular above, drying dark
brown, 4-angled above, terete below. Leaves narrowly ovate or
rhomboidal, up to 6 cm. long and 2.8 cm. wide, at least twice as long
as wide, pubescent with septate hair on both surfaces, especially
toward the apex, upper sessile, lowest on petioles up to 8 mm. long,
margins dentate on upper two-thirds, entire on lower third, teeth
obtuse, becoming acute on upper leaves, apex acute; inflorescences
in pairs from uppermost leaf -axils, usually bifurcately branching and
often ultimately subcorymbose; flower stalks and calyces densely
glandular-pubescent, hair septate or not; calyx lobes ca. 1 cm. long
and 4 or 5 mm. wide, little dimorphic, the uppermost lobe wider
than the others; corolla yellow, glabrous, velvety near the orifice of
the lower lip, upper lip 4-6 mm. long, lower 18-24 (-28) mm. long
and ca. half as wide, saccate more than three-fifths of its length;
anther cells equal, divaricate, each pair 3-4 mm. long, about twice
as long as wide; ovary densely pubescent and glandular-pubescent;
capsule ovoid, shorter than the calyx, vesture as the ovary.
Junin: Between Viques and Ingahuasi, Mantaro Canyon, S. of
Huachacayo, alt. 3,150 m., Killip & Smith 2211+3; Between Tocas
and Pilcos, near Colcabamba, Prov. Tayacala, alt. 1,900 m., Ochoa
978; Prov. Huancayo, alt. 3,600m., Rauh-Hirsch P1708; Chongos
Bajos, Soukup 3620.
Calceolaria weberbaueriana Kranzl. in Fedde, Rep. Sp. Nov.
1: 101. 1905.
Shrub, erect, with the stem much branched from near the base,
branches usually very numerous, short, strict, sparsely to densely
lanate base to apex. Leaves oblong, obtuse, reaching almost 1 cm.
long and about 2.5 mm. wide, tissue obscured by the silvery white,
lanate pubescence; inflorescences pseudo-paniculate; peduncles up
to 2 cm. long, densely pubescent; calyx lobes ovate, 3-4 mm. long
and 2.5 mm. wide, densely lanate externally; corolla purple or yel-
low and purple, upper lip about as long as the calyx, densely pilose,
lower lip 10-12 mm. long, glabrous, saccate about half its length; an-
ther cells subhemispherically unequal, a little longer than wide; style
longer than the stamens, basally pilose, ca. 5-6 mm. long; ovary
elliptic-ovoid, densely lanate; capsule narrowly ovoid, longer than
the calyx, lanate, style persisting.
654 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Libertad: Toyabamba, among timberline scrub, alt. 11,000 ft.,
Carricker s.n., April 14, 1942; Bombamarca, W. slope of E. cordillera,
11,000 ft., Carricker s.n., July 8, 1932.
Type collected in Ancash (Weberbauer 32^8) which borders
Libertad.
Purple flowers are rare in the genus.
CAPRARIA L.
Xuarezia, R. & P., Syst. Veg. 46. 1798; Fl. Peruv. 2: 13, t. 123a. 1799.
References: Sprague, T. A., Kew Bull. pp. 205-212. Sept. 19, 1921.
Erect shrubs or occasionally subshrubs, rarely herbs (Bought 192),
usually branching, stem terete, glabrous or pubescent with stiff hairs,
especially above. Leaves alternate, usually serrate or dentate, occa-
sionally entire, narrowly elliptic to narrowly obovate or ovate, oc-
casionally lanceolate, sessile or very short-petiolate, gland-dotted;
inflorescences 1 or 2 in each axil; flowers racemose, pedicellate; calyx
of 5, rarely 4, imbricate, free sepals or the narrowly lanceolate sepals
fused only at the base; corolla 4-lobed, yellow-green or white, actino-
morphic or barely zygomorphic, tube short, rotate or almost so;
stamens 4 or 5, when 4, didynamous, inserted on 2 of the anterior
corolla lobes, filaments filiform, anthers subsagittate, locules dis-
tinct, parallel or divergent; ovary bil ocular surrounded by the
nectary, style one, stigma lobes ligulate; capsule septicidal, splitting
dorsally at least part way, ellipsoid, gland-dotted; seed numerous,
testa reticulate.
The genus comprising five species, is found from Texas through
Central America and the West Indies to South America in Peru,
Argentina, and Bolivia, thence East to Brazil. C. biflora L. has been
introduced into and has naturalized in the Bahama Islands and the
Gold Coast, Mauritius, and Cape Verde Islands in Africa. The main
concentration of the genus is in Mexico.
Formerly C. biflora and C. peruviana Benth. were widely em-
ployed as a beverage by the aborigines as well as early European
settlers throughout much of their natural ranges. The introduction
of the former species spread the use of this "tea" to other parts of
the world. C. biflora is used today, but in lesser amounts. Two of
the five species occur in Peru.
Leaf blades mostly lance-elliptic, narrowly elliptically ovate or lan-
ceolate, often 6-8 times longer than broad, margins variously ser-
FLORA OF PERU 655
rate, or subentire, sinuses usually lacking or very small; stamens 5;
corolla actinomorphic; sepals usually less than half as long as the
capsule C. peruviana.
Leaf blades broader than the above, mostly oblanceolate or obovately
oblanceolate or narrowly obovate, 2-43/2 times longer than broad,
margins variously dentate to subincised, sinuses frequently present
and apparent; stamens usually 4, inserted in approximate pairs on
2 of the anterior lobes of the slightly zygomorphic corolla; sepals
free and often as long as or longer than the capsule C. biflora.
Capraria biflora L., Sp. PL 875, 1753; Jacq., Hist. 182, I. 115.
1763; 111. t. 534. fig. 2; HBK., Nov. Gen. et Sp. 2: 354. 1817; Loesener
in Engler, Bot. Jahrb. 23: 120. 1896. Capraria lanceolata Vahl.,
Encyl. 2: 47. 1798; C. semiserrata Willd., Sp. PL 3: 325. 1800; C. Ur-
sula HBK., Nov. Gen. 2: 355. 1817 C. semiserrata var. berterii
(A. DC.) Benth. in DC., Prodr. 10: 429. 1846; Millsp., in Field Mus.
Publ. Bot. 2: 98. 1900; C. mexicana Griseb., Fl. Brit. W. Ind. 427.
1861, non Moric.
Branching shrub, stems hirsute, hairs of various lengths, longi-
tudinally striatulate. Leaves narrowly obovate, up to 5.4 cm. long
and 1.2 cm. wide, mostly shorter and narrower, ca. 3.5 cm. and 0.9
cm., apex acute or subacute, sessile or short-petiolate, the petiole
thinly winged by the tapering blade, margins often pubescent, vari-
ously dentate on the upper third to a half of the blade, rarely sub-
incised, frequently sinuses present between the teeth; pedicels fre-
quently sparsely hirsute, 5-7 mm. long (rarely little shorter), erect
or almost so; calyx tube usually lacking, if present up to 0.8 mm.
long, sepals sometimes sparsely hirsute, 4-6 (-7) mm. long and 1.0-
1.5 mm. wide, narrowly lanceolate; corolla white, subrotate, ca. 12-
15 mm. in diameter, the free part of the ca. 8-9 mm. long petals
elliptical and about 5.0-5.5 mm. long, subacute; stamens 4, didyna-
mous, inserted in pairs anteriorly, the insertion of the longer pair
lower on the corolla tube; filaments about 3 times longer than the
1.2 mm. long, dorsifixed anthers; ovary drying black, shorter than
the 3 mm. long style, stigma ligulate, lobes at least partly free, less
than 1 mm. long, much shorter than the style; capsule broadly ovoid,
sub truncate, 4-8 mm. long; seeds numerous, drying dull brown, less
than 1 mm. long, the brown to black placenta forming a thick column.
Loreto: San Salvador on the Amazon River, LI. Williams 1564'
Leticia on the Amazon River, LL. Williams 3070.
656 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
In the western hemisphere from the Bahama Islands to Argen-
tina, including the West Indies and Central America. Rare in west-
ern South America. Known vernacularly by many names outside of
Peru, none reported within. Cultivated in Colombia as Te" Nacional.
Capraria peruviana Benth. in DC., Prodr. 10: 430. 1846; Robin-
son in Proc. Amer. Acad. 38: 202. 1902; Stewart, in Proc. Calif. Acad.
Sci. 1: 141. 1911. Xuaresia biflora R. & P., Syst. Veg. 46. 1798; Fl.
Peruv. 2: 13, t. 123a. 1799. Witheringia salicifolia Hook., Bot. Misc.
2: 231. 1831.
Erect shrubs or subshrubs, or herbaceous to 1.5 m. tall, stems
drying light to dark brown, glabrous and usually longitudinally stri-
atulate. Leaves glabrous, narrowly elliptic, occasionally lance-sub-
ovate or narrowly lanceolate, tapering to a long, acute apex, base
gradually tapering to a short, narrowly winged petiole, the blade
appearing sessile or almost so, up to about 6 cm. long and 0.8 cm.
wide, usually shorter and a little narrower variously serrate, occa-
sionally serrulate on upper third or subentire or entire, sinuses lack-
ing; pedicels glabrous, mostly 8-12 cm. long; calyx tube very short,
1-2 mm. long, lobes narrowly lanceolate, ca. 3-5 mm. long and 1.0-
1.5 mm. wide; corolla pale green-yellow, rotate, about 12-14 mm. in
diameter, free part of the similar, 6-7 mm. long petals elliptically sub-
ovate, the costa sometimes extending as a very short apiculum at the
little narrowed apex; stamens 5, inserted on the posterior and lateral
corolla lobes, very near the base, filaments 2-3 times longer than the
1.0-1.3 mm. long, dorsifixed anthers; ovary ovoid, drying black,
about twice as long as the style and half the length of the petals,
stigmas ligulate, the lobes at least partly free, almost as long as the
style; capsule ovoid or broadly ovoid, 6-12 mm. long; seeds numer-
ous, light to dark brown, less than 1 mm. long, the black placenta
forming a thick central column.
Lima: Prov. Lima; Banks of Rio Rimac, 20-50 m., Esposito
1^796; Lurin, Moist open soil, alt. 5-20 m., Pennell 12209.— Piura:
Parinas Valley, Haught 192 (near the westernmost point of South
America); Sin. Depart.: Cana Dolce, watercourse, Haught F-109.
From Panama to Peru and Ecuador. Vernacularly known as
T£ del Peru; T<§ de Lima.
Although Hemsley (Biol. Centr.-Amer. Bot. 2: 455. 1882) has
considered this species conspecific with the preceding, it is generally
agreed that they are distinct.
FLORA OF PERU 657
The remaining three species of Capraria, C. mexicana Moric. ex
Benth., C. integrifolia Mart, et Gal., and C. saxifragaefolia C. & S.
(C. frutescens Britt.) have not yet been found south of Panama.
The latter two known only from Mexico and Honduras.
CASTILLEJA Mutis.
Herbs or rarely subshrubs, erect or occasionally repent or pros-
trate and matted, usually branched, glabrous or pubescent. Leaves
alternate, the crowded uppermost occasionally appearing opposite,
narrow, entire, often few-lobed or divided, sessile or almost so; in-
florescences spikes or racemes; flowers sometimes dull, often crowded
near the apex, distant beneath, subtended by brightly colored, usu-
ally strongly modified bracts; calyx tubular, laterally compressed,
deeply cut above, or above and beneath, lobes entire or bifid; corolla
with tube included, bilabiate, upper (posterior) lip galeate of 2 united
lobes, entire, erect, elongated, concave, lower lip (much) smaller,
3-lobed, lobes very short to long; stamens 4, didynamous, with the
style and stigma included in the galea; anther sacs oblong-linear,
unequally placed, upper modifixed, lower attached to the filament
near its apex; style filiform, stigma bilobed or entire; capsule locu-
licidal, ovate or oblong, sometimes oblique; seeds numerous, testa
loose, reticulate.
The genus comprises about 210 species one of which is found in
northern Asia. Commonest in western North America and Mexico
extending north to Canada and south to Chile and Argentina. Nine
species have been found in Peru.
1. Calyx cleft medianly subequally above (posteriorly) and beneath
(anteriorly) .
2. Stigma lobes wholly united, capitate or punctiform; plants
erect, caespitose or dwarfed.
3. Lateral cleavages of the calyx well developed, lobes lance-
attenuate; plants erect, stems few.
4. Lateral cleavages of the calyx 3-5 mm. long, lobes equal,
lanceolate-attenuate, acute; bracts usually with 3 pairs of
lobes C. pseudopallescens.
4. Lateral cleavages of the calyx 1.5-2.5 mm. long, lobes sub-
equal, deltoid or ovate, acute, bracts usually with 2 pairs
of lobes, sometimes only 1 pair C. cerroana.
658 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
3. Lateral cleavages of the calyx minute to lacking, lobes
rounded-obtuse; plants caespitose or dwarfed. . . .C. pumila.
2. Stigma lobes free, plants usually erect, elongate or if clumped,
stems few.
5. Calyx shorter than the exserted corolla C. arevensis.
5. Calyx longer than the included corolla C. communis.
1. Calyx cleft medianly very unequally posteriorly and anteriorly.
6. Bracts broadly ovate in outline, much shorter than the flowers,
3-lobed on the upper half, the lobes redividing into 3 pairs;
matted dwarf C. bracteata.
6. Bracts lanceolate or narrowly ovate in outline, usually as long
as or longer than the flowers, with 1 or 2 pairs of lobes, some
from the lower half of the blade; plants erect, elongate.
7. Lateral cleavages of the calyx a quarter to a half as deep as
the median; flowers sessile or pedicellate (in fruit).
C. fissifolia.
1. Lateral cleavages of the calyx minute or wanting; flowers
short-pedicellate.
8. Calyx 27-30 mm. long, lobes laterally retuse or emargi-
nate; corolla 28-34 mm. long, usually little exceeding the
calyx C. virgata.
8. Calyx 15-17 (-21) mm. long, lateral cleavages wholly lack-
ing; corolla 20-24 mm. long, well exserted. .C. virgatoides.
Castilleja arvensis C. & S., Linnaea 5: 103. 1830.
Erect, hispid-pilose, much branched annual. Leaves sessile,
spathulate or oblong-lanceolate, up to 6 cm. long and usually less
than 1 cm. wide, reaching 1.5 cm. wide on the longest, entire, apex
obtuse, narrowed to base; spikes dense above, interrupted below;
bracts entire, obovate-oblong, wider than the stem leaves, red-
tipped; flowers sessile; calyx 9-13 mm. long, cleft subequally above
and beneath or a little more deeply above, the two lobes entire, sub-
obtuse, truncate or emarginate, the ventricose tube a little shorter
than the lobes; corolla up to 15 mm. long, the tube and throat in-
cluded in the calyx, galea green or yellow-green, concave, externally
puberulent, apex entire, acute, when flattened broadest at the middle
and sharply tapering to both ends, lip subobsolete, weakly saccate,
the 3 lobes short-acuminate; anthers linear, stamens subequal to the
FLORA OF PERU 659
galea in length; stigma bilobed with the style a little longer than
the stamens; capsule black when dry, broadly ovoid-oblong, oblique,
seeds numerous, the testa reticulate, much exceeding the seed body
at both ends.
Ancash: Lachay 90 km. north of Lima, alt. 500 m., Rauh-Hirsch
P61. — Apurimac: Prov. Abancay, Laderas de fruitius, alt. 2,800 m.,
Marin 1862. — Arequipa: Tingo, moist tilled soil, 2,100-2,300 m.,
Pennell 13101. — Cuzco: Ollantaitambo, moist tilled soil, 2,850-
2,900 m., Pennell 13672; Macchu-picchu, alt. 2,000 m., Rauh-Hirsch
P846.— Junin: Mito, cornfield, alt. ca. 9,000 ft., Macbride & Feather-
stone 1609. — Lima: Matucana, southern slide rock slope, about 8,000
ft., Macbride & Featherstone 338; Along Rio Chill6n, near Obrajillo,
open grassy hillside, 2,300-2,500 m., Pennell 14322.— San Martin:
on road to Divisoria, 59 km. from Tingo Maria on highway to Pu-
callpa, about alt. 8,500 ft., Macbride & Featherstone 1316.
The commonest species in Central America and Mexico, extend-
ing southward to Colombia, Venezuela, and Bolivia then eastward
to Brazil (Minas Gerais).
Castilleja bracteata Edwin, Phytologia 19: 402. 1970.
Densely matted, dwarf, perennial herb, up to 5 cm. tall, from a
knob-like root-stock. Stems obscured by the leaves that are longer
than the internodes, puberulent and angled; leaves narrowly lance-
ovate or -oblong, minute to about 1.5 cm. long, usually with 1 or a
pair or 2 pairs of short, linear or narrowly lanceolate lobes, mostly
from the upper part of the blade, occasionally entire, viscid, puberu-
lent-ciliate on the margins, sessile, acute, little or sometimes not at
all narrowed to base; inflorescences short, dense spikes; bracts 11-
14 mm. long, margins ciliolate, broadly ovate in outline, divided from
about the middle into 3 divisions, the lateral divisions usually divid-
ing once again, the median once or twice, lobes lanceolate, acute,
decreasing in length from primary to secondary and tertiary, drying
green (red- tipped?) ; flowers (and fruits) sessile; calyx red-tipped or
green, broadest at the apex, regularly, slightly tapering to the base,
subcylindric or subsalverform, 13-18 mm. long, puberulent on the
main veins and tissue above on both surfaces, ciliolate-puberulent
on the margins, lobes obtuse, median cleavages very unequal, about
a fourth the length of the calyx (above) posteriorly and half way
(below) anteriorly, lateral cleavages lacking or the lobes merely re-
tuse or emarginate; corolla 15-24 (-26) mm. long, exceeding the
calyx, at least part of the galea exserted, usually up to a half its
660 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
length, galea densely puberulent externally, often red-green, 6-10
mm. long, margin thickened, inrolled, lip very short, included in the
calyx, the 3 lobes lanceolate, hardened, infolded, sac moderately
developed; stamens didynamous, pairs almost equal in length, the
longer pair 1-2 mm. shorter than the galea, included; style and
stigma about reaching the longer stamens, stigma lobes capitate-
punctiform, wholly united; capsule narrowly ovoid or oblong, 12-
14 mm. long, oblique near the acuminate apex; seeds numerous,
loosely enveloped by the reticulate testa, reticulations dark.
Cuzco: Prov. Calca, Pisac, prados humedos, alt. 3,400 m., April,
1943, Marin 252.
This well-marked species, especially as to habit and the lobing of
the bracts is also technically unique not fitting into any section of the
genus thus far described.
Castilleja cerroana Edwin, Phytologia 19: 403. 1970.
Erect, pubescent, little branching, perennial herb, 40 cm. tall.
Stem densely pubescent, hair soft and a little curly, neither twisted
nor matted ; leaves pubescent on both surfaces, lanceolate, narrowly
oblong or linear, up to 3 or 4 cm. long, usually with a pair of short,
linear lobes from above the middle, entire, acute, narrowed to the
base, sessile; inflorescences spikes up to almost half the length of the
stem; terminal on the main stem and appressed branchlets; flowers
dense when young, less so at maturity; bracts densely pubescent on
both surfaces, linear or narrowly oblanceolate in outline, sessile, with
usually 2 pairs of linear, acute lobes, the longer pair, from below the
middle always developed, the shorter subapical pair sometimes want-
ing; flowers on very short, ca. 0.5-1.0 mm. pedicels; calyx 17-19 mm.
long, densely long-lax-pubescent without, shorter pubescent within,
tube cleft equally, or subequally medianly to below the middle, lat-
eral cleavages 1.5-2.5 mm. long, the lobes subequal in size and shape,
deltoid or ovate, acute; corolla 20-22 mm. long, lips exserted, lower
lip ca. 6 mm. long, saccate, not indurate, pale yellow, lobes linear,
acute, erect, galea 9-10 mm. long, carmine-red, densely stiff, golden,
pubescent externally, margins inrolled, thickened; stamen pairs very
unequal in length, the upper a little shorter than and included in the
galea; style longer or shorter than the stamens, included, stigma lobes
united, punctiform; capsule narrowly ovoid, 14-16 mm. long, sub-
oblique near the short-acuminate apex; seeds numerous, narrowly
ovoid, the loose testa dark-reticulate.
FLORA OF PERU 661
Ancash: Cordillera Blanca, alt. 4,400 m., Rauh-Hirsch P. 2096.
This taxon is close to Castilleja pseudopallescens Edwin, differing
in the lateral cleavage of the calyx, the calyx lobe size and shape,
general pubescence, and in the lobing of the bracts.
Castilleja communis Benth. in DC., Prodr. 10: 529. 1846.
Erect, usually unbranched, rarely much branched, pilose-hispid
annual or perennial herb. Leaves lanceolate or narrowly oblong-
lanceolate, sometimes only up to 1-2 cm. long, usually 3-6 (-7) cm.
long, entire, narrowed to an acute apex and (usually) an acute base,
sessile; bracts red-tipped, rarely purple- tipped, oblong, broader than
the stem leaves, acute to subobtuse, entire, margins frequently a
little thickened; flowers sessile, green or yellow-green; calyx exceed-
ing the corolla, 14-18 mm. long, cleft about half-way, equally, ante-
riorly and posteriorly, secondary cleavages wanting, lobes oblong,
obtuse, entire, sparsely glandular-pubescent, tube ventricose in fruit,
narrowly cylindric in flower; corolla 12-16 mm. long, included in the
calyx, the cylindric tube about equal to the length of the concave,
externally pubescent, scarious margined galea, lip about 1.5-3.0 mm.
long, the sac subobsolete, the lobes lanceolate, scarious, the median
shorter than the 2 lateral; stamens about as long as the corolla, an-
thers linear; style about as long as the calyx, stigma 2-lobed; capsule
broadly ovoid, oblique, drying black, glabrous, about as long as the
corolla tube; seeds numerous, the loose fitting, reticulate testa nar-
rowed to one end and much exceeding the seed body.
Apurimac: Prov. Abancay, 3,200 m., Vargas 776. — Cuzco: Valle
de Santa Ana, 1,600 m., Herrera 3614-', Macchu-picchu, alt. 8,750 m.,
Saunders 432; alt. 4,000 m., Soukup 276; alt. 8,000 ft., Stafford 789;
Prov. Quispicanchis, Marcapata, humid places, alt. 3,000 m., Vargas
9676 ; Prov. Paucartambo, between Achivani and Medias-Mayu, alt.
2,500-2,800 m., Vargas 11123.— Huanuco: Tingo Maria, Soukup
2267; Prov. Huanuco, in open sun along road, alt. 1,300 m., Stork &
Horton 9596; Pillar, plant purple except base of flowers and leaves,
alt. 2,700 m., Woytkowski 34134; Divisoria, in wet situation, alt.
1,600 m., Woytkowski 34501. — Junin: along road in sun, Chancha-
mayo Valley, alt. ca. 2,000 m., Hutchison 1177; alt. 1,200 m., C.
Schunke 406 and 582; 1,500 m. C. Schunke 1544; Prov. Tarma:
Utcuyacu, upon waste land, alt. 1,900 m., Woytkowski 35413.—
Lima: Valley of Lima, Dombey 6; Mathews 398; Prov. Lima, 6 km.
west of Pachacamoc, alt. 500 ft., Saunders 197; San Bartolo, 8 km.
inland from San Bartolo, alt. 600 ft., Saunders 198. — San Martin:
662 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Zepelacio near Moyobamba, alt. 1,200-1,600 m., Klug 3486; San
Roque alt. 350-1,500 m., LI. Williams 6926 and 7154-
The altitudinal range of this species is 350-4,000 m. and is much
greater than that usually encountered in this family.
Common from Mexico to Chile in dry and wet open sites.
Castilleja fissifolia L. f., Suppl. 293. 1781, non Sesse et Mocino
1894. C. coronopifolia Vent., Choix. 59. 1803. C. fissifolia gamma
longifolia Benth. in DC., Prodr. 10: 534. 1846. C. stricta Benth.,
I.e. 534.
Pubescent or puberulent, rarely subglabrous, often scabrous, erect
annual or perennial. Leaves deeply cut into 3-7 linear lobes (rarely
entire), usually from the upper half of the sessile, oblong, lanceolate
or linear blade, body of the blade up to 4.5 cm. long and not exceed-
ing 1 mm. wide and as the lobes entire, narrowed to a long-acute apex
and sometimes narrowed to base, main veins (3-5) parallel and ele-
vated beneath; flowers pedicellate, pedicels increasing with age, in
fruit reaching 1.5-2.0 cm. long; subtending bracts pubescent on both
surfaces, usually broader than the leaves, mostly with two pairs of
linear lobes, one pair near the apex, the other below the middle of the
blade, lobes acute and with the acute body red- or purple- tipped;
calyx tubular, 17-22 mm. long, densely pubescent on both surfaces,
medianly cleft more deeply anteriorly than posteriorly, laterally cleft
almost a quarter to a half as deeply as medianly, lobes linear or lan-
ceolate, obtuse, shorter than the tube, corolla 19-27 mm. long, lips
exserted, shorter than the narrow tube, lower lip inflated, the pouch
longer than the lanceolate lobes, galea less than to about 2 times the
length of the lip, very densely beset externally with stiff, white pu-
bescence; stamens included, the pairs very unequal in length, anthers
linear; style and stigma included, about equal to the longer stamens
in length, stigma lobes appearing united, separating under pressure;
capsule ovoid, enclosed by the inflated calyx tube, glabrous, acute,
up to 16 mm. long, septum persistent; seeds numerous, the loose
testa reticulate, exceeding the seed body much more apically and
basally than laterally.
Ancash: Cordillera Blanca, Que"brada Cruz Piura, alt. 4,600 m.,
Rauh-Hirsch P. 2090. — Cajamarca: Celendin, Osgood & Anderson
90. — Cuzco: colinas del Laxaihuaman, alt. 3,500-3,700 m., Herrera
832; without data, Herrera 3182; Valle del Urubamba, alt. 2,800 m.,
Herrera 34.37; Prov. Cuzco, alt. 3,500 m., Vargas 254- — Huanca-
velica: Prov. Huancavelica, 1 km. north of city of Huancavelica, alt.
FLORA OF PERU 663
3,700 m., Stork & Horton 10918. — Lima: Rio Blanco, open hillside,
alt, 3,000-3,500 m., Kittip & Smith 21606.
Also collected in Colombia, Venezuela, Ecuador, and Bolivia.
Castilleja pseudopallescens Edwin, Phytologia 19: 404. 1970.
Erect herb up to 22 cm. tall, branches puberulent-pubescent,
foliage puberulent-ciliate, stem alternate branching from near the
base, drying yellow- to purple-brown, subshining, internodes 2-3 cm.
long, at least on lower half of the stem; leaves lanceolate, narrowly
oblong or linear, usually with 1 pair or 2 pairs of long or short, linear
lobes, from minute to about 3 cm. long, most often 1.0-2.5 cm. long,
lobes and blades acute, blades sessile, sometimes clasping; inflores-
cences lax or dense spikes; bracts a little wider than the leaves, with
2 or more often 3 pairs of linear lobes, the pair nearest the base long-
est, the pair at the apex shortest; flowers sessile, red-tinged; calyx
18-21 mm. long, cleft subequally medianly, tube puberulent or sub-
glabrous, veins and lobe margins ciliate-ciliolate, lobes lanceolate,
acute, laterally cleft 3-5 mm. ; corolla well exserted, 25-28 mm. long,
lip ca. 6 mm. long, little more than half as long as the galea, a little
saccate, the 3 lobes linear or narrowly lanceolate, obtuse, galea dor-
sally pubescent, red or purple, 10-11 mm. long; stamen pairs only
little unequal in length, the longer 2-3 mm. shorter than the galea;
stigma lobes wholly united, capitate-punctiform, reaching the longer
pair of stamens; young capsules narrowly ovoid, ca. 15, mm. long
and 6-7 mm. wide, oblique near the acuminate apex; seeds linear to
narrowly ovoid, the loose testa darkly reticulate.
Apurimac: Prov. Grau, cuestra Quellipata, stony slopes, alt.
2,700 m., March 9, 1946, Vargas 5800.
This taxon is related to the species of the section Pallescentes of
western North America and may be placed therein.
Castilleja pumila (Benth.) Wedd., Chi. And. Nov. 2: 119. 1860.
C. nubigena var. pumila Benth. in DC., Prodr. : 10. 543. 1846. C. fis-
sifolia var. pumila (Benth.) Wedd., Chi. And. Nov. 2: 119. 1860.
Caespitose or subcaespitose, sometimes minute, perennial herb,
with usually puberulent stem and foliage, one or both of which may
be glabrous or rarely densely pubescent. Leaves sessile, sometimes
less than 1.0 cm. long, rarely up to 3.5 cm., narrow, rarely exceeding
0.8 cm. wide, usually only up to 4-5 mm., linear to lanceolate, with
or without 1 or 2 pairs of linear lobes, body and lobes entire, acute,
base of the blade obtuse, 3-5 main veins elevated beneath ; inflores-
664 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
cences short spikes, or the sessile or subsessile flowers axillary and
solitary on the dwarfed plants; bracts wider than the leaves, most
often with 2 pairs of lanceolate or linear, red-tipped, acute lobes;
calyx 13-17 mm. long, often red-tinged, tube cylindric, cleft me-
dianly subequally anteriorly and posteriorly, less than half way, the
lateral cleavages minute or occasionally lacking, the lobes narrowly
oblong, blunt or rounded; corolla 15-20 (-22) mm. long, tube cylin-
dric-funnelform, lips exserted from the calyx, shorter than the tube,
galea yellow-green to crimson, only sparsely puberulent externally,
acute, margin entire, scarious, lower lip less than a third the length
of the galea, weakly saccate, the lobe-tips acuminate; stamens in-
cluded, didynamous, the pairs very unequal, anthers linear ; style and
stigma included, equaling or a little shorter than the longer filaments;
stigma lobes wholly united, globose-capitate; capsule ovoid, about as
long as the enclosing perianth tube, shining black; seeds numerous,
ovoid in outline, trigonous in cross-section, the testa loose, reticulate,
exceeding the seed body much more laterally than apically and
basally.
Apurimac: Prov. Andahuaylas, Que"brada in cold grassy slough,
alt. 2,890 m., Stork & Horton 10760.— Cuzco: Colinas de Laxai-
huaman, alt. 3,500 m., Herrera 832; La Raya, alt. 4,300-4,500 m.,
Pennell 13500; Cuzco, alt. 3,500 m.; Pennell 13701; Cerro de Col-
quipata, alt. 3,900-4,000 m., Pennell 13742; Macchu-picchu, alt.
4,000 m., Soukup 300; Prov. Canas, in grassland, alt. 3,800-4,500
m., Vargas 9849; Prov. Quispicanchis, puna vegetation, alt. 3,700 m.,
Vargas 971 4- — Junin: Tarma-Aroya, alt. 4,200 m., Rauh-Hirsch
P. 267; Prov. Huancayo, in puna grass, alt. 4,400 m., Stork 1094.2
and 1091+3. — La Libertad: Prov. Santiago de Chuco, alt. 3,500 m.,
Stork & Horton 9994.. — Lima: near Morococha, moist meadow,
alt. 4,350-4,800 m., V. Grant 7563; near Anticocha, Cerro Colo-
rado, east of Canta, alt. 3,800-4,100 m., Pennell 14658.— Puno:
Araranca, alt. 4,100-4,200 m., Pennell 13428; Soukup 938; Lake
Titicaca, alt. 4,000m., Rauh-Hirsch M 47. — Sin. Depto.: Viso, wet,
rocky, brook banks, ca. 9,000 ft., Macbride & Featherstone 603; Yauli,
"springy" slope, ca. 13,500 ft., Macbride & Featherstone 920; San
Gaban, N. of Camjata Hacienda, Capachica Peninsula, 12,600 ft.
Tutin 1035.
Also found in Ecuador. Occurring at elevations above 2,500 m.,
throughout its range, this species is easily separable from C. fissifolia
in which it is frequently placed as a variety.
FLORA OF PERU 665
Castilleja virgata Dombey ex Wedd., Chi. And. Nov. 2: 119.
1816. C. fissifolia subvar. virgata Wedd., I.e. 119.
Erect perennial herb often forming clumps, unbranched or vari-
ously branching. Stem sparsely puberulent or glabrous, sometimes
subdensely puberulent in the inflorescence; leaves entire or variably
divided, lowermost usually entire, middle with 1 pair or less often
2 pairs of lanceolate or linear, minute or well-developed lobes, upper
leaves with 2 pairs of lobes, narrowly lanceolate or linear, minute to
3.5 cm. long, usually less than 2.5 cm. long, blade body and divisions
entire, acute, usually narrowed to base, sessile, puberulent on main
veins on both surfaces; inflorescences spicate-racemose, dense or lax;
bracts longer and wider than the upper stem leaves and with 2 pairs
of red- or crimson-tipped lobes; flowers on glabrous or puberulent
6-8 mm. long pedicels; calyx tubular, bright green, 27-30 mm. long,
usually puberulent on the veins and at the base externally, on tissue
on the lobes internally, cleft medianly about a third the length pos-
teriorly and two-thirds anteriorly, the lobes laterally retuse or emar-
ginate, obtusely rounded; corolla yellow-green, very little to clearly
exceeding the calyx, 28-34 mm. long, lip usually included, glabrous,
saccate, the lobes indurate, linear, about 2-3 mm. long, galea usually
somewhat nodding, most often at least partly exserted through the
anterior median cleavage, densely golden-puberulent exteriorly, 16-
18 (-19) mm. long, margin thickened, hardened, infolded, drying
black; stamens didynamous, the pairs a little unequal in length, the
longer a little shorter than to equaling the galea, included, anthers
linear; stigma and style equal to or little longer than the galea, stigma
included or exserted, punctiform to subclavate; capsule 15-17 mm.
long, narrowly ovoid, glabrous, drying black, oblique near the short-
acuminate apex; seeds numerous, ovoid, testa loose, reticulate.
Huanuco: Prov. Huanuco, Grassland, alt. about 12,000 ft., Mac-
bride & Featherstone 2190. — Junin Mito, Grass-shrub slope, alt. about
9,000 ft., Macbride & Featherstone 1668. — Lima: near Antaicocha,
Cerro Colorado, east of Canta, rocky wall of canyon, alt. 3,600-
3,800 m., Pennell
Castilleja virgatoides Edwin, Phytologia 19: 405. 1970.
Erect, clumping, perennial herb up to 25 cm. tall. Stems pubes-
cent in the inflorescences and either puberulent below or becoming
glabrate near the base or entirely glabrous and shining brown; leaves
linear or lanceolate, pubescent or puberulent on one or both sides,
up to 3 cm. long, usually with 2 or 3 leaflets in the axil, entire, usually
666 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
with a pair of short, linear lobes, narrowed to an acute apex, sessile;
inflorescence a spike or spicate in flower, racemose in fruit, often in-
terrupted below; bracts with 1 or 2 pairs of lobes, little wider than
the leaves, distally red (in vivo) ; flowers on short, ca. 1-2 mm. long,
pubescent pedicels; calyx tubular, 15-17 mm. long, pubescent on
both surfaces, obtusely rounded apically, medianly cleft very un-
equally, posteriorly only 2-3 mm., anteriorly beyond the middle, ca.
10-12 mm., lateral cleavages lacking; corolla about 20-24 mm. long,
galea yellow-green (in vivo), about 10-11 mm. long, exserted, pubes-
cent externally; lip rudimentary, ca. 1.5-2.5 mm. long, little if at all
inflated, the lobes linear, the median shorter than the lateral ; stamens
included, pairs only a little unequal in length, anthers linear; stigma
and style reaching to about the longer stamens, stigma lobes united,
clavate; capsule drying black, glabrous, oblique near the short-
apiculate apex, borne on pedicels up to 8 mm. long, placenta me-
dianly thickened, septum early deciduous; seeds numerous, ovoid,
the reticulate testa loose.
Cuzco: Prov. Cuzco, open rocky hill on trail to Pisac, alt. 3,500-
3,600 m., April 28-30, 1925, Pennell 13699.— Apurimac: Salcantay,
alt. 4,300 m., Rauh-Hirsch P 1398. — Ayacucho: La Quinau, grassy
banks, alt. 12,000 ft., Macbride & Featherstone 2002. — Cuzco: Cor-
dillera Veronica, alt. 3,300-3,500 m., Rauh-Hirsch P961+. — Huan-
cayo: Huaytapallana, alt. 4,600 m., Rauh-Hirsch P1718. — Junin:
Mount La Juntay, near Huancayo, alt. about 4,700 m., Killip &
Smith 22085. — Lima: Canta, mountains above Canta, Pennell 11+1+29;
East of Canta, alt. 4,000-4,200 m., Pennell 11+686.— Sin. Depto.:
Viso, shrubby slope, alt. ca. 9,000 ft., Macbride & Featherstone 581 ;
Sin. Loc.: Diehl 2563.
This species is closely related to C. virgata Dombey ex Wedd. (see
above) from which it differs in having smaller flowers and fruit,
shorter stigma and style in relation to the stamens and galea, and
in the shorter calyx in relation to the corolla. Further, the calyx
lobes of C. virgata are laterally secondarily retuse and the (lower) lip
of the corolla is clearly saccate. Both C. virgata and virgatoides are
closely allied to species of the western United States that are now
placed in the section Linariaefoliae. However, the Peruvian taxa
differ sufficiently to warrant separate sectional status.
CONOBEA Aubl.
Erect or repent annual herbs, glabrous except the corolla tube
internally, usually branched. Leaves usually petiolate, the petioles
FLORA OF PERU 667
decreasing toward the apex, uppermost leaves sometimes sessile,
toothed or irregularly weakly cut; flowers solitary or 2-3-clustered,
axillary, pedicellate; pedicels bibracteolate at apex; calyx of 5, im-
bricate sepals, free or united at the base, all alike or almost so and
shorter than the corolla tube; corolla bilabiate, lips subequal, lobes
shorter than or equaling the tube, patent, throat open, the 3 lobes of
the anterior lip at least partly free, the two posterior lobes sometimes
wholly united, corolla tube pubescent within anteriorly or some-
times both anteriorly and posteriorly; stamens 4, didynamous, in-
cluded, filaments filiform, anthers dorsifixed, locules proximate,
parallel; style deflexed at apex, stigma lobes 2, distinct; capsule
loculicidal, globose. One (of seven) species, Conobea scoparioides
(C. & S.) Benth., known from Peru. The genus is widely distributed
in the Western Hemisphere from Illinois to Florida, Mexico to Pan-
ama and from Brazil to Venezuela and west to Peru. Two collections
seen from Trinidad. However, no species is commonly found in
great concentration in any small area.1 The greatest concentration
of the taxa in the genus is Eastern lowland South America.
Conobea scoparioides (C. & S.) Benth. in DC., Prodr. 10: 391.
1846. Sphaerotheca scoparioides C. and S., in Linnaea 2: 606. 1827.
Stems and branches 4-angled, the angles ridged, interangular tis-
sue striatulate. Leaves elliptic, lance-elliptic and less often subovate-
elliptic, up to about 6 cm. long and 1.5 cm. wide, usually smaller,
decreasing apically, densely punctate with deeply impressed punc-
tations, margins serrate-serrulate especially on the upper half, acute
to long-acute at apex, attenuate to the base that narrowly wings the
petiole; pedicels erect to spreading, up to 2.4 cm. long, usually about
0.8-1.5 cm. long, filiform, rarely less than 3 times longer than the
calyx, often 4 or more times longer, bibracteolate at apex ; bracteoles
minute, frequently appressed to the calyx, sessile and widest at the
base; sepals imbricate, narrowly lance-ovate, margins frequently
scarious, sometimes also minutely ciliolate, up to 3.5 mm. long, usu-
ally 2.5 to 3.0 mm., usually wholly free, sometimes united only at
the base; corolla about 7-9 mm. long, blue and lavender, bilabiate,
the tube subcylindric, longer than the lobes (occasionally the lobes
equal to the tube), exserted, internally beset with short, sometimes
clavate hairs posteriorly or both posteriorly and anteriorly, the hairs
usually restricted to just below the open throat or continuing to the
tube base; stamens 4, didynamous, glabrous, the filaments inserted
1 Conobea multifida (Michx.) Benth is found scattered in small patches through-
out its range in the United States.
668 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
about halfway on the tube, the longer pair sometimes reflexed, the
anthers almost coherent; ovary globose, shorter than the erect style,
stigma-lobes free and usually spreading; capsule broadly ovoid to
globose, about as long as the sepals; seed minute, testa longitudinally
striate and reticulate, apiculate at both ends, light brown, numerous.
Loreto: Iquitos, 100 m. alt., Killip & Smith 27031; King 1231;
On the Amazon River, Pegas, LI. Williams 17 13 and Caballo-Cocha,
LI. Williams 2432. Also in Brazil and Colombia.
It is to be expected that further collecting in lowland Peru will
uncover C. aquatica Aubl., closely allied to our species and up to now
found in Trinidad, the Guianas, Venezuela, and Brazil.
DIGITALIS L.
Densely golden pubescent herb, erect, simple or branched from
the base. Stems lanate; leaves1 ovate, cauline mostly alternate, up
to 15 cm. long and 8-10 cm. wide, coarsely crenate-dentate, usually
pubescent on both surfaces, the largest leaves radicle, clustered be-
low, decreasing abruptly in size upward, bases of the larger long-
acute, often thinly winging the long petioles, upper petioles very
short, uppermost leaves sessile, apices long-acute to obtuse; flowers
in long racemes, pedicellate; pedicels thickened, densely lanate,
lower up to 2 cm. long, decreasing apically; calyx lobes ovate or ob-
long, acute, long-acute or acuminate, pubescent on both surfaces;
corolla up to 5 cm. long, purple, throat darker purple-dotted, the
dots surrounded by paler tissue, bilabiate, the 3-lobed anterior lip
(the lower) little longer than the 2-lobed posterior median lobe of
anterior lip longer than the lateral lobes, tube anteriorly ventricose,
much longer than the lobes, glabrous without, pubescent or glabrous
within; anthers included, didynamous, filaments flattened; style
much longer than the ovary, stigma lobes distinct, ligulate, ovary lax-
pubescent; capsule ovate (ex desc.), septicidal, secondarily weakly
loculicidal; placenta thickened, septum semi-persistent; seed numer-
ous, minute, oblong-subglobose, testa rugose.
An old world genus of about 30 species, especially abundant in
Europe and Asia.
A single species in Peru D. purpurea.
1 The one collection seen from Peru bears only young leaves. This part of the
description based on extra Peruvian specimens.
FLORA OF PERU 669
Digitalis purpurea L., Sp. PL 621. 1753.
Calyx lobes 5, united only near the base, about 12 mm. long and
7 mm. wide, a little unequal in length and width; anther locules sub-
parallel to divergent, contiguous at apices, ovoid, dehiscing by a
single slit through the septum; style ca. 20-25 mm. long, ovary nar-
rowly ovoid, ca. 8-11 mm. long; capsule little longer than wide,
glandular-hirsute.
Cuzco: Prov. Quispicanchis, along the road at alt. 2,700 m.,
C. Vargas C. 1328.
Probably an escape from cultivation. Naturalized in Canada
and the United States as well as in mountain meadows of Costa
Rica. The taxon is well established in Colombia and becoming so
in Chile.
The species has long been of importance in medicine, yielding
several powerful drugs.
ESCOBEDIA R. & P.
Escobedia R. & P., Prodr. Fl. Peruv. 91, t. 18. 1794, generic
description only; Syst. Veg. Fl. Peruv. 159. 1795, genotype E. scabri-
folia R. & P.; Salvia Veil., Fl. Flumin. 1: 55, t. 149. 1825. Nicalia
Raf., Fl. Tellur. 2:104. 1837.
Ref.: Pennell Proc., Acad. Nat. Sci. Phila. 83: 411-426. 1931.1
Perennial herb dying back to the roots, erect, simple or branched,
puberulent or glabrous. Cauline leaves sessile, opposite; flowers
forming racemes, in the axils of leaf -like bracts; pedicel bibracteo-
late, rarely ebracteolate; calyx tubular or funnellform, lobes shorter
than the costate tube; corolla salverform, white, fragrant, tube ex-
ternally glandular-puberulent, rarely glabrous, internally pilose, lobes
united, internally pilose; genitalia about reaching the corolla throat;
stamens 4, equal, filaments ciliate, anthers narrow, glabrous, locules
equal, base aristate; style elongate, with two stigmatic lines at apex;
capsule ellipsoid, woody, glabrous, ultimately exposed by the break-
ing of the calyx tube; seeds numerous, linear.
A neotropical genus with two easily separable subgenera occur-
ring from Mexico to Chile and Argentina. The extremely showy,
large white corollas simulate Nicotiana spp. of the Solanaceae. The
1 The generic and specific descriptions and the key for Escobedia are essentially
those appearing in this reference. Only two sheets of E. scabrifolia R. & P. were
available to the writer; no other Peruvian material seen.
670 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
genera are sympatric and apparently compete for the same polli-
nators. At present comprising 15 species but conceivably many
more will be described as collecting progresses. Three species occur
in Peru.
1. Peduncles less than 2.0 cm. long; largest leaves 10-16 cm. long.
E. brevipes.
1. Peduncles 2 cm. or more long (2-5 cm.); largest leaves 6-8 cm.
long.
2. Bracteoles on the upper part of the peduncles, 14-20 mm. long;
peduncles 3-5 cm. long E. foliolosa.
2. Bracteoles on about the middle of the peduncles, 2-8 mm. long ;
peduncles 2-3 cm. long E. scabrifolia.
Escobedia brevipes Pennell, Proc. Acad. Nat. Sci. Phila. 83:
418. 1931.
Stem simple or branching near the top, puberulent or glabrous.
Leaves oblong or ovate-oblong, acute or acuminate, serrate or more
or less entire, scabrellus especially above, often glabrate below, the
largest 10-16 cm. long and 4-5 cm. wide; pedicels 1.5-2.0 cm. long;
bracteoles 5-12 mm. long, linear or elliptic-lanceolate, inserted on
the upper half of the peduncle; calyx tube 4 cm. long, teeth 4-6 mm.
long, calyx tissue glabrous or scabrellus with 5 scabrous-puberulent
veins; corolla tube 8-9 cm. long, externally glandular-puberulent,
lobes 5-7 cm. wide, internally pilose; capsule unknown.
Huanuco: Chinchao, Ruiz & Pawn s.n. — San Martin: Tarapoto,
Spruce 4456.
Known also from Bolivia: Beni, Chuquisaca, and La Paz.
Escobedia foliosa Pennell, Proc. Acad. Nat. Sci. Phila. 83: 419.
1931.
Stem up to about 5-7 cm. tall, simple, puberulent, and angled.
Leaves oblong-lanceolate, acute, serrulate or entire, above scabrellus
or glabrate, beneath paler with puberulent veins, the largest 6-8 cm.
long and 3-4 cm. wide; pedicels 3-5 cm. long; bracteoles 14-20 mm.
long, lanceolate or elliptical-ovate, inserted 3-10 mm. below the
calyx; calyx tube 3-5 cm. long, teeth 5-6 mm. long, triangular, acu-
minate, calyx glabrous with 5 prominent, puberulent or glabrous
veins; corolla tube 7-10 cm. long, externally glandular-puberulent
or glabrous, lobes 6-8 cm. wide, internally pilose; capsule not seen.
Chachapoyas: Sesuya, Mathews s.n.
FLORA OF PERU 671
Collected in Ecuador (Tungurhua), as well as northern Peru.
Escobedia scabrifolia R. & P., Syst. Veg. Fl. Peruv. 159. 1798.
Stems simple or branching on the lower half, 5-6 cm. tall, almost
round, puberulent or glabrous. Leaves oblong-lanceolate or ovate,
obtuse or acute or acuminate, serrulate or entire, scabrous above
and with large, prominent, distant, puberulent veins beneath, largest
6-8 cm. long and 1.5-3.5 cm. wide; pedicels 2-3 cm. long; bracteoles
2-8 mm. long, linear, medianly placed on the pedicel; calyx tube
3.0-3.5 cm. long, teeth 5-8 mm. long, triangular, acute or acuminate,
glabrous, beset with 10 veins, 5 larger with 5 smaller medianly be-
tween, puberulent; corolla tube 8-11 cm. long, lobes 5-6 cm. wide,
internally pilose; capsule 2 cm. long, seeds 4.0-4.5 mm. long.
Huanuco: Muna, Macbride 3934. — San Martin, Cerro de Cam-
pana, Spruce s.n.
On the eastern slope of the Andes in central Peru and at higher
elevation than E. brevipes.
EUPHRASIA L.
Annual branched herbs, a few cm. tall. Leaves opposite, sub-
sessile to sessile; flowers in terminal, spike-like inflorescences, ebrac-
teolate, small ; calyx of 4, united sepals, cleft more deeply medianly
than laterally, teeth shorter than the tube; corolla tube exserted,
narrowly cylindric, a little shorter than the lips; upper lip galeate,
reflexed-margined, longer and narrower than the 3-lobed, reflexed
lower lip; stamens 4, included in the galea, anther cells dorsifixed,
parallel, basally short-awned, confluent at apex; style little longer
than the anthers and exceeding the galea, ovary much shorter than
the style; capsule obovoid, loculicidal, little longer than wide,
strongly compressed, seeds few, longitudinally sulcate.
A genus comprising about 90 species mostly cold-temperate to
subarctic in both the northern and southern hemispheres and in the
new and old worlds.
Only one species thus far verified as to its occurrence in Peru,
E. pubescens.1
1 In Anales del Institute Botanico A. J. Cavanilles (Anales del Jardin Botanico
de Madrid) 1 (17): 452-458. 1959, seven species considered to be Euphrasia, six
of which are said to have been collected in Peru, are presented. Of these, possibly
two, E. incana R. & P. and E. dentata R. & P., may be Euphrasias. The remaining
taxa are species of Bartsia or not generically clear to me. Nor can distinctions
between E. dentata and incana as well as pubescens be determined now.
672 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Euphrasia pubescens Benth. in DC., Prodr. 10: 555. 1846.
Stem and foliage densely soft-pubescent. Cauline leaves up to
3 mm. long, deeply cut into 3, almost free lobes, joined only at the
base, lobes linear to narrowly oblong-lanceolate, entire, apices acute;
flowers subtended by bract-like leaves or occasionally these modified
with the median lobe much exceeding the lateral; calyx ca. 5-6 mm.
long, tube little inflated, subcylindric, teeth subequal, acute, poste-
rior median cleavage little deeper than anterior cleavage; corolla
11-13 mm. long, tube well exserted about equaling the lobes, poste-
rior lip (2-lobed galea) longer and narrower than the 3-lobed lacin-
iate, deflexed anterior lip.1
Sin. Loc.: Dombey s.n., ex photograph Paris Herbarium.
GALVESIA Dombey ex Juss.
Galvesia Domb. ex Juss., Gen. PI. 119. 1789, non R. & P.,
Fl. Peruv. Prodr., 56, 1794- Pitavia Molina (Rutaceae).
Branched shrubs or large herbs, new wood sparsely puberulent
so glandular-pubescent or -puberulent, occasionally glabrous, older
wood frequently glabrate. Leaves opposite or ternate-verticillate,
upper sometimes subopposite, rarely uppermost alternate, ovate to
lance-ovate, mostly 1-4 (-5) cm. long, sometimes longer or shorter
and usually less than half as wide, but varying to three-fourths as
wide, glabrate to glandular-pubescent, entire, apices subobtuse to
more often acute, sometimes short-acuminate, bases acute, subobtuse
or obtuse; petioles 2-10 cm. long, usually glandular-pubescent;
flowers pedicellate or forming short racemes, mostly crowded near
the tips of the branches, solitary in the axils of usually reduced
leaves; pedicels glandular-puberulent to glabrous, 5-25 mm. long;
calyces with vesture as the pedicels, when glandular-puberulent
often both internally and externally, 3.0-4.5 mm. long, lobes sub-
equal, lance-ovate or lanceolate; corolla red, 14-23 mm. long, bilabi-
ate, lips subequal, from a quarter to almost a half the length of the
tube, upper lip (posterior) 2-lobed, reflexed, lower 3-lobed, median
lobe little longer than lateral, tube gibbose at base, constricted im-
mediately above gibbosity, then little ampliate to the obstructed
throat, palate of the lower lip composed of 2 swellings and only
partly closing the throat, usually glandular-puberulent; stamens 4,
didynamous, included or exserted, upper, posterior pair shorter than
the lower, anterior, locules parallel and lanceolate before dehiscence,
1 Description drawn largely from a Dombey collection from Chile.
FLORA OF PERU 673
dehiscing longitudinally by slits, locules circular after dehiscence,
contiguous, a fifth very reduced, ca. 2-3 mm. long, staminode usu-
ally present at the base of the corolla posteriorly, filaments of the
fertile stamens pubescent (usually densely) below, glandular or gla-
brous above; ovary about a fourth the length of the style that is
about equal to the posterior stamens, stigma lobes united or barely
free at the tips; capsule depressed-globose, 4-7 mm. long, thin- walled,
dehiscing by two irregular pores; seed numerous, reaching 1 mm.
long, testa irregularly broken-ridged, placenta thickened, persistent.
Two closely allied species (as here interpreted) occur in Peru.
A genus of five to six species of the new world. Distributed from
the southwestern United States to Colombia and Peru. No collec-
tions have been seen from the West Indies. The area of greatest
concentration found in Mexico and secondarily in Peru.
Corolla 12-15 mm. long, the tube about 2-3 times longer than the
lips; leaf -blades mostly less than 2 cm. long; petioles mostly less
than 3.5 mm. long; pedicels mostly as long as or little longer than
the subtending leaves; anthers usually included G. fruticosa.
Corolla 18-23 mm. long, the tube up to 2 times longer than the lips;
leaf -blades mostly 2 cm. long or longer; petioles usually 3.5 mm.
long or longer; pedicels most often shorter than the subtending
leaves; anthers frequently exserted G. ballii.
Galvesia ballii Munz, Proc. Calif. Acad. Sci. ser. 4, 13(12): 379.
1926. G. limensis var. grandiflora Benth. in DC., Prodr. 10: 296
1846.1
A branched herb or shrub. Leaves usually acuminate or acute;
petioles flattened, puberulent, little longer than in G. fruticosa; pedi-
cels 5-10 (-12) mm. long; calyx lobes 3.5-4.5 mm. long, lance-ovate
to lanceolate ; fertile stamens often eglandular above, 12-14 mm. long,
usually exserted; pistil 11-14 mm. long; capsule up to 7 mm. long.
Otherwise as G. fruticosa. Endemic in Departamentos Piura and
Tumbes of northwestern Peru and only with difficulty separable from
fruticosa. Pennell, Not. Nat. 179. 5-6. 1946 considered the two taxa
clearly specifically separable. Munz (I.e.) felt that perhaps they are
only varieties of the same species. Although specific rank is here
1 G. grandiflora (Kell.) Benth. an erroneous citation of Wettstein in Engler &
Prantl, Pflanzenf. IV(3b): 61. 1891, based on Ball's use of G. limensis var. grandi-
flora Benth. in Journ. Linn. Soc. 22: 151. 1887, for a California species caused
Munz to propose G. ballii. The presentation by Wettstein is "too garbled," fide
Pennell.
674 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
maintained for the two taxa, varietal or even synonymic status could
be considered equally correct.
Piura : Negritos, Parinas Valley, Haught F-l 21 ; Paita, l Andr& 4119
(cited by Munz); Abra, 1,350 m., Hutchison 1393; 0-20 m. alt., Pen-
nell 14810; Savatier s.n.; Talara: about the cliffs, "clavela," Johnston
3052; Rocky localities in Qu£brada, 100 m., Rauh-Hirsch PH.—
Sin Depto. : A. H. Olsson s.n. — Tumbes: Prov. Tumbes, alt. 100 m.,
Weberbauer 7752.
Galvesia fruticosa Gmel., Syst. Nat. ed. 13, 2: 927. 1791.
Dodartiafragilis R. & P., Syst. Veg. Fl. Peruv. 1 : 97. 1798. Russeliat
alternifolia Pers., Syn. 2: 164. 1807. Agassizia limensis Domb. ex
Chav., Monogr. Antirrh. 180, t. 11. 1833. Galvesia limensis Domb.
ex Benth. in DC., Prodr. 10: 296. 1846.
Much branched shrub, new wood sparsely puberulent to occa-
sionally glandular-puberulent, vesture decreasing with age. Leaves
broadly ovate to lance-ovate, usually opposite, uppermost suboppo-
site or alternate, usually glandular-puberulent on both surfaces,
more so beneath, occasionally with the wood and calyces essentially
glabrous, from more than to less than twice as long as wide; petioles
grooved above, usually glandular-puberulent; pedicels glandular-
puberulent, slender, little recurved, mostly 8-20 mm. long; calyx
lobes 5, sepals united about a third their length, mostly glandular-
puberulent on both sides, lance-ovate 2-3 mm. long; corolla tube
glandular-pubescent on both surfaces and the palate ; fertile stamens
8-9 mm. long, filaments densely pubescent on lower third and usu-
ally glandular above, inserted at base of the corolla tube, fifth sta-
men bearing sterile anthers on a 1-2 mm. long, filiform filament;
style almost equaling the stamens, about 4 times longer than the
ovoid or subglobose ovary; capsule 4-5 mm. long.
Ancash: Rio Fortale"sa, alt. 2,000 m., Rauh-Hirsch Pi 956; Prov.
Santa, 2-50 m. alt., Las Lomas hacia Huarney, C. Vargas C. 1224-
—lea: Prov. lea, Pampa de Huayuri, 382 km. S. of Lima on Pan
American Highway, alt. 550 m., Hutchison 1223; Prov. Pisco, Pisco
Tal, alt. 7,000 m. (?), Rauh-Hirsch P-402.— Lima: Dombey s.n., Iso-
type2; Canta Valley, alt. 350 m., Hutchison 1001 ; Rio Chillon, near
1 Alternate spelling, Payta.
2 Manuscript citing this collection in Dombey 's protologue, mss. Fl. Peruv.,
in Paris. Seen by Jussieu.
FLORA OF PERU 675
Viscas, alt. 1,800-2,000 m., Pennell 14316; bank of Rio Rimac, W.
of Lima, alt. 10-30 m., Pennell 14288. — Sin. Dept.: Hacienda Hu-
ando, alt. 400 m., Saunders 189; Olmos-Jaen, l alt. 800 m., Rauh-
Hirsch P 2135; Rio de Pariocota, alt. ca. 3,000 ft., Macbride &
Feather stone 2540; Rio de Lomas, Weberbauer 5742.
The wide altitudinal range of these collections may explain in
part at least, the variability of this taxon. Collections confined to
coastal and Andean Peru. The 7,000 m. collection of Rauh-Hirsch
is erroneous as to altitude. There are no mountains of this height
in the area.
GERARDIA Benth.
Gerardia Benth. in DC., Prodr. 10: 514. 1846. Gerardia L.,
Sp. PL 610. 1753 as toG. purpurea only; not as the generic descrip-
tion in L., Gen. PI. ed. 5. 266. 1754, nor as to derivation.2 Virgularia
R. & P., Prodr. Fl. Peruv. 92, 1. 19. 1794. Chytra Gaert., Fr. et Sem.
3: 184, t. 214. 1805. Agalina Raf. New Fl. Amer. 2: 61. 1837.
Herb or less often a shrub, erect and branched. Leaves opposite,
entire, petiolate or sessile, linear to lanceolate; flowers axillary over
most of the branches or terminally clustered in interrupted racemose
or spicate inflorescences; pedicels longer or shorter than the calyx
and the subtending bract-like leaves; calyx campanulate, the 5, al-
most wholly united lobes, valvate or narrowly imbricate and reduced
to spine-like teeth ; corolla pink to dark red to (most often) purple,
tube usually campanulate to campanulate-subfunnelform, externally
pubescent, longer than the flaring lobes, throat open ; stamens 4, di-
dynamous included, filaments pubescent, anther cells equal, usually
appendaged at base ; style elongate, stigmas united ; capsule loculici-
dally dehiscent, secondarily barely septicidal at apex of the valves;
seeds numerous, testa loose.
A temperate zone, new world genus with about 60 species.
One polymorphic species occurs in Peru, G. lanceolata.
Gerardia lanceolata (R. & P.) Benth. in Hook., Comp. Bot.
Mag. 1: 207. 1835; DC., Prodr. 10: 516. 1846. Virgularia lanceolata
& V. revoluta R. & P., Syst. Veg. Fl. Peruv. 161. 1798. Gerardia an-
1 The Jaen-Olmos Road traverses the departments of Cajamarca,Lambayeque,
and Piura.
2 Derivation pertains to a genus in the Acanthaceae.
676 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
gustifolia, G. lanceolata and G. parvifolia Benth. in DC., Prodr. 10:
516. 1846. G. bangii 0. Ktze., Rev. Gen. 3(2) : 233. 1898. G. mega-
lantha andG. stenantha Diels in Engl., Bot. Jahrb. 37: 429. 1906.
Stems and branches striatulate, usually very short-puberulent
with the vesture in rows, occasionally glabrous. Leaves linear to
lanceolate, usually glabrous and gland-dotted, up to 6-7 cm. long,
usually much shorter, sometimes only 1-2 cm. long, longer or shorter
than the nodes, acuminate, entire, upper leaves sessile, bracteolate
lower short-petiolate, costa impressed beneath; petioles indurate,
almost as wide as the base of the blade, usually glabrous; pedicels
from 1-2 mm. to 2 cm. long or longer, glabrous to densely puberu-
lent; calyx tube a little to much longer than the narrowly lanceolate
to lanceolate, long-acute or -acuminate, tooth-like lobes, glabrous or
puberulent as the pedicels, 5-9 mm. long, lobes usually puberulent
on both sides, more densely internally, occasionally one or both sides
glabrous ; corolla pink to purple, often with darker markings and fre-
quently darker within, externally densely villose, corolla tube 2-
3 cm. long, about 4 times longer than the ovate lobes, throat some-
times villose within, lobes frequently puberulent within; stamens 4,
didynamous, posterior pair longer, filaments puberulent, anthers
narrowly oblong, mucronulate at base; style exserted, stigma lobes
united; capsule usually a little longer than the calyx, globose; seeds
shining, irregular in outline, angulate, brown to dark brown-black;
septum deciduous, placenta thin. Common at varying altitudes
throughout the Andes of southern and central Peru.
Apurimac: Prov. Abancay, Upper Marino Valley, alt. 3,000 m.,
Stork, Horton & Vargas 10639; alt. 2,600 m., Weberbauer 5846.—
Chachapoyas: Mathews 3137. — Cuzco: Prov. Quispicanchis, Marca-
pata, alt. 3,600 m., Vargas 1336', Weberbauer 7782; Prov. Urubamba,
Macchu-picchu, alt. 2,200 m., Mexia 8080; alt. 8,750 ft., Saunders
435; alt. 7,000 ft., Stafford 793; Cerro de Cusilluyoc, alt. 3,400-
3,800 m., Pennell 13806; alt. 3,800-3,900 m., Pennell 13827; alt.
2,500-3,000 m., Pennell 13933; Ayusbamba, laderas, alt. 2,700 m.
Vargas 294; Paucartambo, Laderas de Pillahuata, alt. 3,300 m., Var-
gas 6. — Huancavelica: Prov. Huancavelica, Mejorada, alt. 2,400 m.,
Stork & Horton 10905; Prov. Tajacaja, Surcubamba-Salcabamba
Trail, alt. 2,300 m., Stork & Horton 10414; alt. 2,800-2,900 m.,
Weberbauer 7613.— Huanuco: Muna, alt. 7,000-9,000 ft., May 1863,
Pearce, s.n.; alt. 2,700 m., Woytkowski 48. — Junin: Huacapistana,
alt. 1,800-2,400 m., Kittip & Smith 24318; Mito; alt. ca. 9,000 ft.,
FLORA OF PERU 677
Macbride & Feather stone 14-57 p.p.; Prov. Huancayo, Torre- torre,
alt. 3,317 m., Soukup 2939,—Pasco: Huariaca, alt. ca. 9,500 ft.,
Macbride 3095; Yanahuanca, alt. about 10,000 ft., Macbride &
Featherstone 1201. — Puno: Near Puno, alt. 4,000 m., Soukup 233;
393; 4.18; Prov. Sandia, Environs of Limbani, alt. 3,500 m., Vargas
9662. — Sin. Dept.: Uspachaca, alt. about 8,500 ft., Macbride &
Featherstone 1306; Rio Maranon, alt. about 7,500 ft., Macbride
& Featherstone 1981 ; Tabina, Lechler 2084; Poeppig 1373; Pawn, s.n.
HEBE Commers. ex Juss.
Evergreen trees or shrubs with opposite, coriaceous leaves that
leave conspicuous scars upon falling. Inflorescences (in ours), axil-
lary and lateral racemes; flowers dense and several ranked around the
axis; calyx lobes forming a very short tube; corolla more than 2
times longer than the calyx, tube a little longer than the lobes; sta-
mens exserted; ovary ovoid-ellipsoid, much shorter than the exserted
style, stigma lobes punctiform; capsule compressed, septicidally de-
hiscent, the septum subsequently dehiscing medianly to release the
seed, secondarily partly loculicidally dehiscent, ovoid-ellipsoid or
subglobose, much exceeding the calyx; seed few, flattened, testa
smooth.
A genus of about 150 species native to New Zealand. About
three species disjunct in Chile, Argentina, and the Falkland Islands.
A single species, escaped from cultivation, known from Peru,
H. corriganii.
Hebe corriganii Carse, T. N. Z. I. 60: 573. 1930.
Small tree (in ours), essentially glabrous, pedicels, calyces and
leaf-margins excepted. Leaves lanceolate, 7.5-14.5 cm. long and
1.5-2.5 cm. wide, margin entire, minutely pubescent, subhyaline,
narrowed to both ends, the winged petiole 1-3 mm. long; inflores-
cences lateral racemes, most often exceeding the subtending leaves
in length, flowers densely situated on 2-3 mm. long, puberulent ped-
icels, each subtended by a minute bracteole; calyx about 3 mm. long,
lobes 2.0-2.5 mm. long, lanceolate, acute, ciliolate on the pale, mem-
branaceous margin; corolla white, tube cylindric, about 4-5 mm.
long, lobes equal, narrowly ovate, 3-4 mm. long, entire; filaments
about 7.5-9.0 mm. long, anthers parallel or almost so; pistil only
little shorter than the stamens, style apex and stigma exserted; cap-
sule about 4-5 mm. long and 3-4 mm. wide, walls indurate, brown,
smooth, glabrous; septum thickened, persistent, placenta thin.
678 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Cuzco: Granja, Yucay, small tree, Soukup 565.
LAMOUROUXIA HBK1
Erect, usually pubescent, perennial herbs or subshrubs with op-
posite leaves and red or reddish corollas. Leaves narrow, sessile or
short-petiolate, usually deeply toothed, or lobed, the teeth and lobes
sharp; flowers in spike-like inflorescences; pedicels very short or lack-
ing; calyx of 4, united sepals, tube campanulate, teeth acute, equal
in pairs, the median cleavages longer than the lateral; corolla bilabi-
ate, tube elongate, swollen then compressed, posterior lip galeate,
of 2, united petals, entire or emarginate, wider and a little shorter or
longer than the 3-lobed anterior lip, the latter 2-plicate or 2-ridged
and the lobes short; stamens 4, didynamous, included in the galea,
posterior shorter than anterior, anthers bearded, usually mucronate
or short-awned basally, coherent in pairs, locules distinct, parallel,
equal; style entire, stigmatic at apex, obtuse; capsule ovoid, loculici-
dal; seeds many, smooth or striate-sulcate, with or without reticu-
late, hyaline wings.
A tropical American genus of about 30 species, one of which has
been collected in Peru, L. breviflora.
Lamourouxia breviflora Benth. in DC., Prodr. 10: 540. 1846.
L. subincisa Benth., I.e.
Erect perennial herb or sometimes shrubby at base or a subshrub,
profusely branching on upper part of stems. Stems more or less
densely pubescent; leaves lanceolate or narrowly oblong, up to 4 cm.
long and 0.8 cm. wide, usually smaller, subsessile or sessile, often
attenuate to base, irregularly, distantly or closely, coarsely sharp-
toothed or -lobed, most often pubescent beneath, occasionally on
both surfaces, rarely glabrous, veins conspicuously elevated beneath,
obsolete above; inflorescences spikes or spike-like; flowers sessile or
on sparsely pubescent, 0.5-2.5 mm. long pedicels that are thickened,
especially at base; bracts leaf -like, bracteoles lacking; calyx about
half the length of the corolla or shorter, calyx tube shorter than the
lobes, campanulate or broadly campanulate, subglabrous (rarely) to
densely soft-pubescent; calyx teeth pubescent, lanceolate, acute,
margins entire or distantly, irregularly serrulate; corolla up to 20 mm.
long, usually 10-16 mm., externally more or less densely pubescent
or puberulent, internally pubescent in the flaring, elongate, often
Conservandum Proposita, fide L. O. Williams, Nov. 1967.
FLORA OF PERU 679
plicate throat and along one or more of the lines of petal connation,
lobes ovate, entire, anterior little spreading; anthers short-awned
basally; capsule about equaling to little longer than the calyx, nar-
rowed and acute apically, septum papery, soon deciduous.
Amazonas: Prov. Chachapoyas, W. of Chachapoyas, alt. 2,400 m.,
Pennell 15493. — Ancash: Monterrey, 3 km. below Huaraz, alt. 3,000-
3,100 m., Pennell 15307. — Cajamarca: Prov. Cajamarca, "Santa
Polonia" above Cajamarca, alt. 2,700-2,800 m., Pennell & Ferreyra
14.924.; Que*brada de San Vicente, S.W. of Cajamarca, Pennell &
Reichlin 14991. — Junin: Tarma, steep S.W. slope at alt. ca. 7,000 ft.,
Macbride & Feather stone 1007. — Pasco: Huariaca, shrubby canyon,
about alt. 9,500 ft., Macbride 3111. — Sin. Depto.: Viso, alt. about
9,000 ft., Macbride & Featherstone 582. — Sin. Loc.: Mathews 3138.
This species is limited to the Andes of northern and central Peru.
Weberbauer 6025 is probably a new Lamourouxia but the material
is insufficient for adequate description.
Lamourouxia sylvatica HBK. (Nov. Gen. et Sp. 2: 337. 1818) may
be a Bartsia. One collection reported from Peru, not seen.
This genus has a checkered nomenclatural history. Lamourouxia
Agardh, 1817, is not legitimate since it is based on identical material
as Claudia Lamouroux, 1812. However, Claudia Opiz ex Panzer,
1808 in the synonymy of Melica L., 1753 must be considered. The
conservation of Lamourouxia HBK. permits a simple solution to this
tangle. Another generic appellation is available for the algal genus
of Lamouroux and Agardh.
LEUCOCARPUS D. Don
Erect glabrous or puberulent herb, stem 4-angled, the angles usu-
ally with small wings. Leaves cordate-clasping, almost amplexicaul;
flowers congested in axillary, simple or more often compound dicha-
sia, or occasionally cymose, 3-flowered; basal peduncle 1 in each axil,
pedicels from shorter to less often longer than the peduncle; flower-
bearing stalks bracteolate; calyx of 5 united sepals, tubular-campan-
ulate, membraneous, prominently 5-ribbed, lobes short-acuminate;
corolla wholly yellow, tube elongate, throat widened, bilabiate, lips
flattened, upper (posterior) 2-lobed, lower (anterior) 3-lobed, lobes
rounded, plane; stamens 4, didynamous, included, anther locules
distinct, divergent or divaricate; style thickened at apex, stigma 2-
680 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
lobed, interior margins stigmatic; fruit a globose, indehiscent or
loculicidal berry; seed numerous, very small, testa reticulate.
A monotypic genus,1 distributed from Mexico to Chile and Argen-
tina, commonest in Central America. Not yet collected in the West
Indies or the southern United States.
Leucocarpus perfoliatus (HBK.) Benth. in DC., Prodr. 10:
335. 1846. Mimulus perfoliatus HBK., Nov. Gen. et Sp. 2: 371.
1818. Conobea alata Graham, in Edinb. N. Phil. Journ. 3: 1830.
Leucocarpus alatus (Graham) in D. Don. in Sweet, Brit. Fl. Gard. 2:
pi. 124. 1831.
Much branched from the base, perennial. Leaves lanceolate or
elliptic-lanceolate, up to 25 cm. long (occasionally longer) and 5-
6 cm. wide, serrate, acute; peduncles mostly 1.5-3.0 cm. long; pedi-
cels 0.8-2.5 cm. long, all flowering stalks bibracteolate near their
bases, bracteoles decreasing in size to the ultimate pedicels; calyx
oblique, 1.0-1.3 cm. long, tube much longer than the sharply acumi-
nate, subequal, tooth-like lobes; corolla 1.2-1.5 (-1.8) cm. long, tube
internally with 2 pubescent ridges anteriorly, bilabiate, the anterior
lip longer; stamens inserted near the base of the corolla tube; ovary
subglobose shorter than the style; berry exceeding the calyx tube,
chalky white in life, brown when dry; seed dispersed in copious pulp.
(The corolla is very similar to Mimulus L., but the fruit and calyx
differ sufficiently to justify the present arrangement.)
Piura: Prov. Huanacabamba, above Shumaya, alt. 2,200-2,300
m., Weberbauer 6289. — Huanuco: Several km. W. of Carpish, along
stream, alt. 2,800 m., Stork & Horton 9896. — Junin: Latipo, alt.
500 m., Soukup 2846.
LIMOSELLA L.
Reference: Gluck H., Limosella — studien in Engler, Bot. Jahrb.
66:488-566. 1954.
Herbs of brackish or saline mud, occasionally submerged or partly
so, acaulescent or the stems repent or sometimes stoloniferous, rarely
(not in Peru) erect. Leaves aggregated at the nodes or radicle
(rarely opposite on the stem), blades linear or subulate as the peti-
oles or narrowly oval to elliptic, shorter than the long, spreading or
1 Leucocarpus frutescens Benth. in A. DC. Prodr. 10: 336. 1846, pertains to
Hemichaena Benth. "alatus" the only other specific epithet in the genus is in
synonymy here.
FLORA OF PERU 681
erect petioles, blades when differentiated from the petioles entire,
glabrous, up to about 1-3 mm. long, usually not more than half as
wide; flowers solitary, distant or crowded on long or short pedicels,
when pedicels short, flowers appearing almost basal ; calyx tube cam-
panulate, longer than the 5, very short lobes; corolla little exceeding
the calyx, the lips subequal, the 5 lobes shorter than the tube, sub-
equal, the 3 anterior little longer than the 2 posterior, throat open;
stamens 4, 2 anterior, 2 posterior, anther cells wholly confluent,
much shorter than the filaments and little exserted ; ovary globose or
globose-ellipsoid, longer or shorter than the style; stigmas subcapi-
tate, the lobes free or united; capsule globose to ellipsoid, seeds
brown, numerous, minute, testa reticulate.
Placed in the Gratiolinae of the Gratiolae, although having the
calyx of the Mimulinae and sometimes the subcapitate, united stigma
lobes of the other sections of the Scrophularioideae.
A genus of about 11-15 species (requiring much more analytical
study) as interpreted by various authors. Two species are wide-
spread in the new world and both are sporadic in Peru. None of the
few collections displays typical characteristics as described for the
taxa. It seems better at this time to include them in the species to
which they appear closest rather than present them as new taxa.
Cosmopolitan, both generically and (frequently) specifically.
Leaf blades oval or elliptical, clearly distinct from the subulate peti-
oles; style little shorter than the ovary, recurved L. aquatica.
Leaf blades indistinguishable from the petioles; style little longer
than to twice as long as the ovary, erect L. subulata.
Limosella aquatica L., Sp. PI. 641. 1753. Limosella australis
R. Br., Prodr. 44. 1810. L. aquatica var. americana Gliick and
L. aquatica f. terrestris Gliick, in Notizbl. 12: 75. 1934. L. americana
and forma Gluck, I.e. 12: 76. 1934.
Stoloniferous herb, rooting at the nodes, rarely submerged or
partly so, glabrous or almost so. Petioles about 3 times longer than
the elliptical, entire blade that reaches up to 17 mm. long; flowers
solitary, distant (or clustered) ; pedicels about half the length of the
leaves; calyx tube about twice as long as the 0.5-0.7 mm., acute
lobes, campanulate, lobes often reflexed part way; corolla tube about
2 times the length of the ca. 0.5 mm. lobes, little exceeding the calyx
and about 2-3 mm. wide (ex diag.), white with purple tinge; stamens
little exserted, glabrous; ovary globose, much longer than the 0.2-
682 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
0.4 mm. style that is strongly recurved and semi-persistent; capsule
globose or broadly ovoid or obovoid, about 3.0-3.5 mm. long.
Only one collection seen. (Arangaro in glaveosis?) W. Lechler
1877. June, 1854.
Widely distributed, almost world-wide, growing as a "mat" in
muddy places. Possibly conspecific with the following. Most authors
maintain these as distinct species on the bases of numerous small
differences. The only stable characters thus far found for separating
the taxa are in the key. These may be judged insufficient for this
purpose. The Australasian based L. australis R. Br., could be con-
sidered intermediate, but up to now is too poorly known and Brown's
description must be considered scanty, in as much as the entire pro-
tologue is but 4^ lines long.
Limosella subulata Ives, in Trans. Phys. Med. Soc. N. Y. 1:
440. 1817. Ygramela maritima Raf., Atl. Journ. 199. 1833. Limo-
sella lineata Gliick in Engler, Bot. Jahrb. 66: 505. 1934.
Glabrous herb, acaulescent or stems creeping along the ground,
forming linear tufts. Leaf blades and petioles indistinguishable;
leaves up to 4 or more cm. long, linear or linear-subulate or rarely
aciculiform, entire; flowers solitary, crowded or distant, on pedicels
that are longer or shorter than the leaves, up to 4-5 mm. long and
2-3 mm. wide; the weakly reflexed calyx lobes about as wide as the
corolla lobes, calyx campanulate, tube 2.0-2.5 mm. long, almost 3
times longer than the acute lobes; corolla white, tube ca. 1.0-1.5 mm.
longer than the calyx, the short lobes (ca. 1 mm.) usually obtuse and
about half the length of the tube; stamens exserted, anthers one-
celled, about a third to a fourth the length of the filiform filaments;
ovary ellipsoid to globose, (much) shorter than the erect, 1.5-2.5 mm.
style; stigma 2-lobed or lobes united, subcapitate, soon fugacious
after fertilization; capsule ovoid to subglobose, ca. 3.0-3.5 mm. long,
occasionally little shorter; seeds numerous, brown, minute.
Poorly known from Peru.
Arequipa: Prov. Arequipa, Cerrillo Lavandia, near Laspimas,
Eyedam & Beetle 22154; Prov. Chuquibamba (Condesuyos), Huario
below Chuquibamba, 4,000 m. alt., Stafford 1152. — Cuzco: Laura-
marca, Cord. Ausangate, Rauh-Hirsch Pi 132. Also reported fide
Gliick (as L. liveata) from Dept. Junin, Prov. Tarma, Weberbauer.
This species distributed from the eastern United States, where it
is the common representative of the genus, thence sporadically to
FLORA OF PERU 683
Bolivia and Peru with stations in Mexico and Central America. It
is also found in the British Isles.
Limosella acaulis Sesse & Mocino (Flora Mexicana 156. 1895.)
may be another ecological form of L. aquatica L., as suggested by
Pennell in Monog. Phila. Acad. 1: 164, 1935. It is a plant of Mexico
and the southwestern United States. However, Pennell retained
this taxon as a species citing a longer, erect, and slender style in
conjunction with narrower leaf blades. If this interpretation is valid
this taxon stands between L. aquatica and L. subulata in approxi-
mately the same way as does L. australis R. Br. With possible in-
termediates in both the old and new worlds the status as well as the
number of taxa should not be considered as finally resolved. For a
more complete discussion of L. subulata see Fernald, Rhodora 20:
160-164. 1918, and Pennell, Torreya 19: 30-32. 1919.
LINARIA Mill.
Linaria (Bauhin) Mill., Gard. Diet. ed. 4. 1754.
Erect, glabrous herbs, occasionally inflorescences glandular-pubes-
cent, branching basally, with entire, mostly alternate leaves and zygo-
morphic, highly modified corollas. Leaves linear, from 5 mm. to 4-5
cm. long and usually less than 1 mm. wide; flowers in racemose or spi-
cate inflorescences, pedicellate, axillary; pedicels up to about 6-7 mm.
long or more often short, about 1-2 mm. long, usually shorter than
the calyx, always shorter than the corolla with a small leaf -like bract
at base; calyx of 5 linear to lanceolate or narrowly lance-ovate sepals,
united about a fourth their length into a cupular tube; corolla very
zygomorphic, bilabiate, personate, purple, purple-white or white, the
anterior (lower) lip 3-lobed, longer than the 2-lobed, reflexed, arch-
ing posterior, median lobe longest and widest, all lobes ovate or
ellipsoid, obtuse, usually little longer than the cylindric to subfun-
nelform tube, lower lip with 2 raised ridges (a palate) closing the
throat; stamens short, included in the tube, didynamous, inserted
near the base of the corolla anterior filaments longer and anthers
bigger than the posterior pair; ovary ovoid or globose, equal to or a
little shorter than the style, stigma lobes united, punctiform; capsule
globose or ovoid, at maturity a little longer than the calyx, dehiscing
distally (apically) by irregular pores, loculicidal ; seed minute, numer-
ous, testa smooth or tuberculate, angles rounded or sharp, septum
persistent, placenta large.
684 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
A genus of about 100 species found mostly in the old world.
About eight to ten species in the new world of which one occurs in
Peru, L. canadensis (L.) Dumont.
Only one other species seen from South America (Chile), L. vul-
garis Hill, whose occurrence is to be expected in our area. For
L. subandina Diels from Ecuador see under L. canadensis. L. tri-
ornithophora Willd. should be placed in the synonymy of L. vulgaris.
L. elegans Cav. described from Hispaniola and of possible occurrence
in Peru is unknown to me.
Linaria canadensis (L.) Dumont Cours., Bot. Cult. 2: 96. 1802.
Antirrhinum canadense L., Sp. PL 618. 1753. A. canadense assurgens
Raf., Autik. Bot. 155. 1840. Linaria texana Scheele,1 Linnaea 21:
761. 1848. L. subandina Diels in Engler, Bot. Jahrb. 37: 47. 1906.
L. canadensis texana (Scheele) Pennell, Proc. Acad. Nat. Sci. Phila.
73:502. 1921.
Leaves opposite at or near the base, becoming alternate over most
of the length of the plants; calyx lobes 2.5-3.5 mm. long; corolla
8-13 mm. long, varying in color from purple to white and variously
purple or white marked; filaments of larger stamens ca. 3 mm. long,
the shorter ca. 2 mm. long, anthers of longer stamens about 1^
times the diameter of the shorter; ovary 2-3 mm. long, style usually
1.5-2.5 mm. long; capsule about 4-5 mm. long and in diameter;
seeds less than 1 mm. long.
Amazonas: Prov. Chachapoyas, alt. 2,700 m., LI. Williams 7547.
— Arequipa: Prov. Islay, near Mollendo, alt. 40 m., Mexia 7775.—
Huanuco: Prov. Ambo, rock slide, alt. 7,000 ft., Macbride 3157.
— Junin: Prov. Huancayo, Soukup 2947', Mito, stony, grassy slopes,
alt. 9,000 ft., Macbride 3313 and 3380. — Lima: Lurin, cleistogamous,
sandy lomas along the sea, alt. about 200 ft., Macbride 5927; Mtn.
of San Augustin near Lima, alt. 3,000 m., Weberbauer 5706. — Pasco:
Prov. Daniel Carrion, Yanahuanca, alt. about 10,000 ft., abandoned
field, Macbride & Feather stone 1249. — Piura: Prov. Ayavaca, above
Ayavaca, alt. 2,800 m., Weberbauer 6367.
Usually found in open dry sites, generally distributed in the
United States, rare in Mexico and Santo Domingo. Sporadic in
South America with only very few but widely distributed collections
1 L. canadensis is said to differ from L. texana in seed testa markings and seed
angles sharp as opposed to rounded as well as having smaller corolla with shorter
spur. These characters intergrade in the United States (center of concentration)
and appear to do so also in Peru, the "texana" kind little more frequent in our area.
FLORA OF PERU 685
in Colombia, Venezuela, Brazil, Ecuador, Paraguay, Uruguay, Chile,
and Argentina.
LINDERNIA All.
Lindernia All., Misc., Tourin. 3: 178. 1766. Vandellia L., Mart.
1: 12. 1767. Ilysanthes Raf., Ann. Nat. 13. 1820. Bazina Raf.,
Autik. Bot. 44. 1844.
Mostly annual herbs, more often glabrous than pubescent, erect,
procumbent, repent or prostrate. Leaves opposite, sessile or short-
petiolate, usually cut-margined ; flowers axillary, solitary, pedicellate
or almost sessile; calyx of 5, essentially free or partly united sepals;
corolla white, blue or purple and variously marked, zygomorphic,
bilabiate, the lips very unequal, posterior erect, of 2, united lobes
or lobes slightly bifid, much shorter than the spreading 3-lobed an-
terior lip, tube cylindric, longer than the lobes; antheriferous stamens
4 or 2, when 2 the anterior reduced to staminodes or sometimes lack-
ing, anterior filaments each sharply inbending, falsely appearing ap-
pendaged and forked, posterior pair of stamens often connivent,
anther locules proximate, dorsifixed; ovary ellipsoid, shorter than
the style that is surmounted by a 2-lobed, lamellate stigma; capsule
linear to globose or obovoid, septicidal, the placenta remaining as a
plate on the septum after dehiscence, seeds numerous, testa reticulate
or pitted.1
Four of the over 70 species reported in Peru. A genus with
world-wide distribution, mostly old world, especially the Orient.
Area of greatest concentration in the new world is in the southeastern
United States.
1. Anther-bearing stamens 2, the posterior pair; sepals essentially
free L. dubia.
1. Anther-bearing stamens 4; sepals united at least part of their
length or free.
2. Anterior stamens smaller than posterior; leaves often as wide
as long.
3. Pedicels 1-2 mm. long; sepals shorter than the capsule;
leaves sessile or petiolate L. diffusa.
1 The unfortunate choice of the generic name Lendernia by Minod (see under
Stemodia), is now doubly apparent, especially since both Lindernia and Lendernia
are of concern in the same tribe.
686 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
3. Pedicels mostly 10 mm. long or longer; sepals longer than
the capsule; leaves short-petiolate L. Crustacea.
2. Anterior stamens equal to posterior; leaves usually longer than
wide L. thouarsii.
Lindernia Crustacea (L.) F. von Muell., Census 97. 1882.
Capraria Crustacea L., Mant. 87. 1767. Gratiola lucida Vahl., Enum.
1: 95. 1804. Torenia Crustacea (L.) C. & S., Linnaea 2: 570. 1827.
Vandellia Crustacea (L.) Benth., Scroph. Ind. 35. 1835. Pyxidaria
Crustacea Ktze., Rev. Gen. 2: 464. 1891. l
Prostrate, repent or erect, essentially glabrous herb of usually dry
or occasionally wet sites. Leaves usually short-petiolate, broadly
ovate, less than 2 cm. long and almost as wide, obtuse to subacute at
apex, broadly obtuse to subtruncate at base, margins irregularly
coarsely dentate; flowers axillary, solitary, pedicellate; pedicels elon-
gate, much longer than the calyx, up to 20 mm. long, often 12-15
mm. long, usually erect; calyx of 5, more or less united sepals, usu-
ally united at least a third their length, often up to three-fourths,
calyx tube usually oblique, mostly splitting along the sepal margins
with the developing fruit and sepals frequently appearing to be free
in fruit, lanceolate, 3-6 mm. long; corolla about 1^ times longer
than the calyx, blue, purple, or lavender, tube usually shorter than
or equal to the sepals; stamens exserted, posterior anthers connivent,
anterior erect, incurved or connivent, anterior stamens only little
shorter than the posterior; ovary ovoid, shorter than the bilamel-
lately stimatic style; capsule conic or obovoid, apex abruptly fla-
tened, the callosed, slightly thickened style base adherent, seeds few,
testa pale, pitted, pits descrete.
Junin: Puerto Yessup, alt. ca. 400 m., Killip & Smith 26349.—
Loreto: Mishuyacu, near Iquitos, alt. 100 m., Klug 1162 and 1292;
Mouth of Rio Santiago above Pongo de Manseriche, alt. 200 m.,
Mexia 6104; along Rio Itaya, LI. Williams 246; Lower Rio Nanay,
LI. Williams 443; Pebas on the Amazon River, LI. Williams 1713a;
Leticia on the Amazon River, LI. Williams 3037.
An old world species now generally distributed in the warm-
temperate and tropic zones of the world, South Europe excepted.
1 To list in extended form the checkered synonymic history of this taxon will
not serve to better understanding of Peruvian Lindernia species so we add merely
as a selected list the following: Morgania lucida Spreng.; Torenia lucida Ham. in
Wall; T. flaccida Br.; T. varians Roxb.; Vandellia varians G. Don; Gratiola aspera
Roth; Hornemannia aspera Link ex Otto; Tittemannia ovata Reicheb; Antirrhinum
hexandrum Forsk.
FLORA OF PERU 687
Lindernia diffusa (L.) Wettst. in Engler and Prantl, Pflanzenf.
IV(3b): 79. 1891. Vandellia diffusa L., Mant. PI. 1: 89. 1767.
Gratiola origanifolia Vahl., Enum. 1: 99. 1804. Bonnaya origanifolia
Spreng., Syst. 1: 42. 1825. Pyxidaria diffusa (L.) Kuntze, Rev.
Gen. 2:464. 1891.
Repent herb of usually dry places. Puberulent or pubescent on
stems, branches, leaf -margins, veins and calyces; leaves very broadly
ovate, subsessile to less often sessile, petiole when present, thinly
winged by the leaf base, up to 2 cm. long and only little narrower or
sometimes as wide as long, obtuse, margins serrate-crenate; flowers
axillary, solitary, subsessile on pedicels that rarely reach 2 mm. long;
calyx united up to about half its length or almost free, 5-7 mm.
long, lanceolate with long-acute apices; corolla little longer than the
calyx, upper part of the tube and lobes exserted, pinkish white or
blue and white, variously marked; antheriferous stamens 4, included
or exserted, the anterior a little shorter than the posterior; filaments
of the anterior adnate almost all their length to the corolla tube and
the tube appearing 2-ridged, posterior filaments curving proximally,
their anthers connivent; ovary ellipsoid, much shorter than the style,
stigma lobes enlarged, clavate-ovate, pubescent; capsule subobliquely
ellipsoid, up to 12 mm. long, much exceeding the calyx, the callosed
style base when attached only little enlarged; seed numerous, testa
tuberculate or with age pitted, pits distant, few; placenta narrow,
septum often papery, fugitive.
Junin: Puerto Bermudez, dense forest, alt. ca. 375 m., Killip &
Smith 26589.— San Martin: San Roque, alt. 1,350-1,500 m., LI. Wil-
liams 7706; Tarapoto, weed in pasture, alt. 750 m., LI. Williams 55^7.
Widely distributed in the warm-temperate and tropical zones of
the world, southern Europe and East Asia excepted.1
Lindernia dubia (L.) Pennell, Mono. Acad. Nat. Sci. Phila. 1:
141. 1935. Gratiola dubia L., Sp. PI. 17. 1753. Capraria gratioloides
L., Syst. Veg. ed. X. 1117. 1759. Ilysanthes riparia Raf., Ann. Nat.
13. 1820. /. gratioloides (L.) Benth. in DC., Prodr. 10: 419. 1846.2
I. dubia (L.) Barnh., in Bull. Torr. Bot. Club 26: 376. 1899.
1 Throughout the literature, (Benth., 1846; Wettstein, 1891; Pennell, 1935)
the pitting of the seed coat as opposed to tuberculate seed coat has been used to
help distinguish taxa. The study of Peruvian (and North American) material has
led to the belief that this may be a function of age in various species.
2 Complete listing of early synonyms for this taxon can be found here. The
nomenclature of this and several other taxa in the genus is not yet fully understood.
688 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Decumbent or prostrate, glabrous herb growing near water. Leaf
blades sessile, ovate or elliptic-oblong, narrowed to base and apex,
only rarely exceeding 2 cm. long, frequently not more than 1.5 cm.
long, margins denticulate to subentire; flowers axillary, solitary,
pedicellate; pedicels ebracteolate, 5-15 mm. long; calyx of 5 sepals,
joined only at the base, appearing free, very narrowly lanceolate,
3-5 mm. long and ca. 0.5 mm. wide; corolla white to pale blue or
violet, 6-8 mm. long, tube a little longer than the lobes, cylindric
or very narrowly conic; stamens included, only 2, the posterior, an-
theriferous, erect, filaments filiform, 2 anterior staminodes weakly
recurved at apex; ovary shorter than the style, stigma lobes promi-
nent; capsule obliquely ellipsoid, a little longer than the calyx; seed
pale brown to almost cream, testa reticulate.
Loreto: Bank of Rio Huallaga, alt. 400-800 m., King 4379; Hera
near Moyobamba, alt. 890 m., Woytkowski 35309.
Basically a species of the southeastern United States but spread-
ing northward to glaciated areas and west to Texas and Arizona.
It has not yet been reported in Mexico or Central America. How-
ever, along with a collection from Puerto Rico, it has been found spo-
radically across South America from the Guianas to the west coast.
L. dubia has been collected also in Europe.1
Lindernia thouarsii (C. & S.) Edwin, comb, nov., Phytologia
19: 6, 361. 1970. Nortenia thouarsii C. & S., in Linnaea 3: 18. 1828.
Torenia parviflora Ham. in Wall., Cat. 3958. 1828. T. nortenia Stued.,
Nomen. ed. II. 2: 692. 1841. T. thouarsii (C. & S.) Ktze., Rev. Gen.
2: 468. 1891 and Wettst. in Engler and Prantl. Pflanzenf. IV(3b):
79. 1895.
Erect or repent, branched, usually subglabrous annual. Stem
apex and upper parts of the branches sometimes sparsely puberulent;
leaves less than 2 cm. long about half as wide as long, upper reduced
and sometimes narrowly lance-ovate, narrowed to apex, acute to
occasionally obtuse at base,2 subsessile with the leaf blade narrowly
winging the very short petiole, or leaves sessile, margins dentate or
distantly dentate, rarely costa puberulent beneath; petiole when
present often puberulent on the margins and costa, groove obvious;
1 This taxon is very closely related to L. anagallidea (Michx.) Pennell and, in
fact, they may be conspecific. If so, this would have no bearing on the nomencla-
ture as dubia is the older, acceptable epithet.
2 Extra Peruvian material with leaf blades usually obtuse at base and with
short wingless petioles and blades sometimes exceeding 2 cm. long.
FLORA OF PERU 689
flowers axillary,1 solitary, pedicellate, in axils of upper leaves that
are reduced and bract-like; calyx lobes 5, united most of their length
in flower, much less so in fruit, 7-9 mm. long and about 0.5 mm.
wide, the costa depressed extending as a sharp mucron, tissue hya-
line, the suboblique tube when present consequently 5-angled ; corolla
little exceeding the calyx, lips blue or violet-blue with rose markings,
tube white, narrow anterior lip with the lateral lobes little longer
than the median, lobes flattened, corolla throat open; anterior pair
of stamens sometimes connivent, posterior usually so; style exceed-
ing the ovary in length, puberulent at apex as the short, abruptly
bilamellate style and persisting in fruit, enlarged and callosed at
base; capsule narrowly ellipsoid, 4- or 5-angled, usually little exceed-
ing the sepals; seed with testa pitted, pits descrete, seeds round in
outline, pale yellow when young.
Peru: San Martin: Zepelachio, near Moyobamba, alt. 1,200-
1,600 m., mountain forest, Klug 1+301.
An old world species especially of Africa and the Indo-Malaya
area. A few collections seen from Colombia and one from Bolivia.
Torenia thouarsii var. nivea Pennell, I.e. from Colombia, seems con-
specific with our new combination.
Species of Lindernia have been collected from sea-level to over
1,000 m. altitude in Peru and from dry to wet as well as open and
closed sites. In view of this, it would seem that any taxon of the
genus accidentally introduced could establish itself. A likely pos-
sibility appears to be the very widespread L. anagallidea (Michx.)
Pennell.
MAURANDYA Ort.
Maurandya Ort., Nov. PI. Descr. Decad. 21. 1797. Usteria Cav.,
Icon, et Descr. Plant. Hisp. 2: 15, t. 116. 1793.
Ref. Munz, Proc. Calif. Acad. Sci. ser. 4, 15(12): 382-395. 1926.
Woody vines, wood subglabrous or densely soft-pubescent, climb-
ing by means of petioles and pedicels. Leaves alternate, the lower-
most often opposite, petiolate, deeply cut-margined, lobed or grossly
dentate, broadly ovate or deltoid, up to about 3.5 cm. long and al-
most as wide near the base, apices acute, bases cordate, hastate, or
truncate; petioles lax, up to two-thirds as long as the blade; flowers
axillary, solitary, pedicellate; pedicels up to 4 cm. long; calyx lobes
1 Congested in Colombia (according to Pennell, Proc. Amer. Acad. Sci. Phila.
72: 163. 1920).
690 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
5, free most of their length; corolla ample, up to 5.5 cm. long, at least
2.7 cm. long, magenta or bright violet, bilabiate, tube longer than
the lips, campanulate or subfunnelform, a little gibbose at base, in-
ternally anteriorly, on the floor of the throat with 2 lines of hair or
2-plaited, but the throat not closed, posterior lip 2-lobed, erect, a
little shorter than the 3-lobed anterior lip; stamens 4 or 5, the fifth
sterile, reduced or lacking, remaining 4 fertile, didynamous, the pos-
terior pair longer than the anterior; ovary shorter than the style,
stigma barely 2-lobed, internally stigmatic; capsule globose, 1.0-
1.5 cm. long, dehiscing irregularly apically, by 2 pores or slits, thin
or firm walled; seeds numerous, corky- tuberculate, ridged or winged.
A genus comprising about ten species, mainly Mexican but ex-
tending south to Colombia and north to the United States. A few
species are widely cultivated throughout the world. The two species
from Peru are said to have been found in the wild state (fide collector)
but material seen only from the garden.
Calyx lobes lanceolate; leaves glabrous, deltoid, 3-lobed, the lobes
not secondarily toothed; seeds wingless M. barclaiana.
Calyx lobes ovate; leaves softly pubescent, lobes lacking but margins
irregularly grossly dentate; seed winged M. erubescens.
Maurandya barclaiana Lindl., Bot. Reg. 13: t. 1108. 1837.
M. barclayana in Martens & Galeotti, Bull. Acad. Roy. Brux. 12(7) :
3. 1845.
Climbing subshrub with slender, glabrous stems, juvenile fre-
quently reddish. Leaves glabrous, thin, bright green above, paler
beneath, triangular-cordate or obscurely 5-lobed, lobes often mi-
nutely mucronulate, hastate at base, 1.0-3.5 cm. long and 1-3 cm.
wide; petioles 1.0-2.5 cm. long; pedicels glabrous, slender, 3-5 cm.
long; calyx tube very short, the sepals free almost their entire length,
calyx glandular-villose with multicellular hair, segments linear-lan-
ceolate, 3 upper subequal, ca. 1.0-1.3 cm. long, 2 lower 0.9-1.2 cm.
long, curving upward, sometimes reaching 2 cm. long in fruit; corolla
3.0-4.5 m. long, glandular-puberulent externally and pubescent in-
ternally above, bright violet, tube swollen at base, 2 prominent, pale
plaits anteriorly in the throat, lobes 7-9 mm. long, suborbicular;
stamens ca. 1.8 and 2.2 cm. long, filaments yellow, glandular above
and densely glandular-pubescent at the bend, usually glabrous at
insertion, fifth stamen reduced or lacking; style about equaling the
shorter stamens, sparsely glandular-pubescent at base, hair also on
FLORA OF PERU 691
the top of the ovary; capsule globose, ca. 1 cm. long, the 2 valves
projecting slightly beyond the style base; seeds oblong, 1.0-1.5 mm.
long, with broken, tuberculate, corky ridges, wingless, septum soon
deciduous, placenta slender.
Lima: Prov. Lima: in city, in the garden of the British Legation,
Sandeman 4027.
Maurandya erubescens (Don) Gray, Proc. Amer. Acad. 7: 377.
1868. Lophospermum erubescens Don in Sweet, Brit. Fl. Gard. ed. 2.
1: t. 75, note. 1830. L. scandens Don in Sweet, I.e. t. 68. 1830.
Woody vine, slender and densely soft-pubescent on wood, foliage,
and in the inflorescence. Stems soft, glandular-puberulent to pilose;
leaves chartaceous, triangular-hastate to cordate or obscurely 5-
lobed, about 3 cm. long and 2 cm. wide,1 acuminate at apex, subcor-
date at base, petioles almost as long as the blades; flowers solitary,
axillary, pedicellate; pedicels up to ca. 5 cm. long, lax and sometimes
glandular-pubescent, ebracteolate; the 5 calyx lobes 15-20 mm. long,
free most of their length, densely pubescent on both surfaces, broadly
ovate, acute; corolla magenta, up to 5.5 cm. long, as the calyx inter-
nally pubescent with multicellular hairs, tube 2.5-4.0 cm. long, cam-
panulate, gibbose for about 1 cm. of its length at the base, constricted
above gibbosity then broader, lobes about 1 cm. long, upper 2 erect-
reflexed, lower 3 erect and little exceeding upper or occasionally lobes
subequal, throat internally with 2 plaits of golden-yellow hairs ante-
riorly; stamens included, 4 fully developed, didynamous, the fifth
when present about 2-3 mm. long, reaching the bend of the filaments
of the 4 expanded stamens, filaments glandular above, densely pubes-
cent beneath, anther-sacs discrete, circular after dehiscence; pistil
equaling the stamens, ovary glandular-pubescent, stigma bifid ; cap-
sule globose, pubescent, ca. 1.5 cm. long, dehiscing by 2 irregular
slits; seeds oblong, corky, tuberculate with emarginate and lacerated
wings, placenta thin.
Lima: Prov. Lima, in city, garden of British Legation, Sande-
man 4026.
Apparently naturalized in Colombia. This species found through-
out Mexico and Central America; the preceding restricted to Mexico
in the wild state.
The common "snapdragon," Antirrhinum majus L., So. PL, 617.
1753, a native of the Mediterranean area is widely cultivated in gar-
1 Reaching 15 cm. long and wide outside Peru, margins and pubescence also
differing in extra-Peruvian collections.
692 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
dens in Andean Peru along with A. virga Gray, Proc. Amer. Acad. 7:
373. 1868, in numerous varieties and color phases. Permission was
not granted the writer to collect. This genus may be separated from
Maurandya as follows:
Leaves entire, lanceolate to linear; corolla with a definite palate,
closing the throat Antirrhinum.
Leaves triangular-hastate, often lobed or coarsely toothed; corolla
with a palate of 2-plaits or lines of hair,1 not closing the throat.
Maurandya.
MICRANTHEMUM Michx.*
Prostrate herb, forming mats and rooting profusely at the nodes,
glabrous or almost so. Leaves opposite, sessile; flowers axillary, soli-
tary, sessile or more often very short-pedicellate, minute, ca. 2 mm.
long; calyx of 4 sepals united only at base; corolla dull white, tube
very short, anterior lobes united much farther than the 3 posterior
that are united only half-way, lobes usually flattened; stamens 2,
the anterior-lateral pair, appendaged near the base due to the sharp
infolding of the filament and the free portion knot-like, anthers mi-
nute, divergent, proximate; ovary globose-obovoid about equal to
the style in length, stigma lobes free, somewhat reflexed, pubescent;
capsule globose, a little shorter than the sepals, irregularly dehiscent
and 1-celled by suppression of the septum; seed oblong, apiculate at
one or both ends, light brown, prismatic, finely striate.
A genus of two (or three) species of eastern North and South
America and Cuba; one of these in Central America. Sporadically
collected in South America westward to eastern Peru.
A single species in Peru, M. umbrosum.
Micranthemum umbrosum (Walt.) Blake, Rhodora 17: 131.
1915. Anonymos umbrosa Walt., Fl. Carol. 62. 1788. Globifera orbi-
culatum Michx., Fl. Bor. Amer. 1: 10. t. 2. 1803. Micranthemum
emarginatum Ell., Sketch Bot. S. Car., Ga. 1: 18. 1816.
1 Palate closing the throat developed in M. antirrhiniflora H. & B., a species
unknown in Peru. These two genera are not so easily separable on a world-wide
basis. For a complete analysis of new world material see Munz, 1926 (cited at the
beginning of this genus) and Pennell 1935.
2 Micranthemum Michx., Fl. Bor. Amer. 1: 10. t. 2. 1803. conserved over
Anonymous Walt., Fl. Carol. 62, 1788 and Globifera J. F. Gmel, Syst. Plant. 2:
32. 1791.
FLORA OF PERU 693
Stems and branches striate-striatulate. Leaves orbicular or some-
times very broadly ovate, margins entire or weakly sinuate; sepals
about 1 mm. long and 0.5 mm. wide, ellipsoid or oblong, narrowed
to base, apices usually acute; corolla about 1.5-2.0 mm. long, the
anterior lobe rarely bifid, internally sometimes puberulent at or near
the throat; stamens reaching the lower part of the lobes.
San Martin: Prov. San Martin, near Tarapoto, Spruce 4152.1
M. rotundatum Wright, the outer generally accepted species of
the genus is found mainly in Cuba.
MIMULUS L.
Mimulus L. Sp. PI. 2: 634. 1753. Monavia Adans., Fam. PL 2:
211. 1763.
Annual or occasionally perennial, glabrous, repent herbs of wet
sites, frequently rooting at the lower nodes.2 Leaves opposite, mostly
petiolate, toothed, upper sometimes much reduced ; flowers solitary,
axillary, pedicellate; calyx campanulate, 5-angled, 5-toothed, mostly
plicate, persistent and accrescent (in fruit), teeth very unequal; co-
rolla yellow, 0.7-2.0 cm. long, bilabiate, tubular, tube slender, in-
cluded in the calyx or exserted, gradually expanding into a long, open
throat, upper lip 2-lobed, the lower lip longer, heavily bearded in 2
patches, 3-lobed, median lobe longest; stamens 4, didynamous, in-
cluded or exserted; filaments glabrous, adnate to the lower part of
the corolla; anthers 2-celled, confluent at the apex, bases more or less
divaricate; ovary shorter than the long, slender style, stigma 2-lobed,
lobes equal or unequal, style longer than the stamens; capsule ob-
long, loculicidal to the base along both sutures, placenta united in a
central column; seed numerous, small, longitudinally striate.
A basically temperate zone genus of about 120 species with the
main concentration in California, the western United States, and
Mexico, and a secondary area in Chile and west-coast South America.
Occasional also in most of the temperate areas of the old world,
Europe, and northwestern Africa excepted.
A single species (common) in Peru, M. glabratus.
Mimulus glabratus HBK., Nov. Gen. et Sp. 2: 370. 1817;
Benth., Scroph. Ind. 28. 1835; Gray, in Bot. Mex. Bound. 116. 1859;
1 Specimen in British Museum.
2 The genus Mimulus varies widely and may range from herbaceous annual to
shrubby. For this genus it is particularly important that the reader is aware of
the narrow description.
694 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Conzatti & Smith, Fl. Sin. Mex. 117. 1897. M. andicolus HBK.,
Nov. Gen. et Sp. 2: 370. 1817. M. parviflorus Lindl., Bot. Reg.
t. 874. 1825. M. propinquus Lindl., Bot. Reg. 1: 1330. 1830. M.
luteus var. micranthus Phil., Linnaea 29: 28. 1857. M. kingii Phil.,
An. Univ. Chil. 43: 528. 1873. M. tener Phil., An. Mus. Nat. Chil.
1:70. 1891.1
Stoloniferous or with creeping stems, up to 4 cm. long. Leaves
mostly broadly ovate, mostly 1.5-4.0(5.0) cm. long and 0.8-3.0 cm.
wide, irregularly dentate, sometimes shallow-lobed at the base ; basal
leaves cuneate, subcordate; petioles margined, shorter than the blade;
upper leaves sessile, subcordate; pedicels slender, longer or shorter
than the leaf blade; flowers mostly on upper part of stem; calyx often
red-spotted, campanulate in flower, 5-8 mm. long, broadly oval in
fruit, 1.0-1.2 cm. long, teeth broad, spreading, very short, uppermost
2 or more times longer than the others, broadly rounded, sometimes
mucronate; corolla early deciduous; style glabrous; capsule rounded,
constricted at base, not stipitate, shorter than the calyx tube.2
Apurimac: Prov. Abancay, alt. 3,200 m., C. Vargas C. 759.—
Arequipa: Prov. Condesuyos, Chuquibamba, alt. 7,000 ft., Stafford
1176; Pichu Pichu slopes, alt. 10,000 ft., Sandeman 3759. — Cuzco:
Prov. Canchis, Uyamiri, Sicuani, 12,000 ft., Stafford 959; Cerro de
Colquipata, alt. 3,400-3,700 m., Pennell 14181; San Geronimo, alt.
3,200-3,300 m., Pennell 14204; Oropesa, alt. 3,200-3,300 m., Pen-
nell 14200; La Ruya, wet Distichia moor, alt. 4,200-4,300 m., Pennell
13485; Sin. Loc.: Stafford 1245; Sandeman 3358. — Huancavelica:
Prov. Taycaja, hills N. of Pampas, alt. 3,250 m., Stork & Horton
10246. — Huanuco: Prov. Huanuco, Huacho 8 km. N. of Huanuco,
alt. 1,900 m., Stork & Horton 9408; Muna, shallow brook margin,
in flat mats, alt. about 7,000 ft., Macbride 3983. — Junin: Huancayo,
alt. 3,317 m., Soukup 2945; Mito, sunny, mossy bog, alt. about
9,000 ft., Macbride & Featherstone 1541. — Lima: Canta, corolla
"lemon-chrome" with maroon dots, alt. 3,000-3,200 m., Pennell
14608. — Loreto: Creek edge, alt. about 15,000 ft., Macbride 2999.
— Moquequa: Ilo, sea level, bank of stream, Stafford 929. — Puno:
Chuquibambilla, in rivulet on puna, alt. 3,850-3,900 m., Pennell
13390; Araranca, alt. 4,100-4,200 m., Pennell 13427; San Antonio de
1 Also Mimulus jamesii var. texensis Gray; M. glabratus var. parviflorus (Lindl.)
Grant; M. glabratus fremontii (Benth.) Grant; M. glabratus michigamensis Pennell;
M. inamoenus Greene.
2 Both the generic and specific descriptions largely from Grant, A. L., Mono-
graph of the Genus Mimulus, An. Mo. Bot. Gard. 11: 99-388. 1924.
FLORA OF PERU 695
Esquilache, in streams, alt. 12,000 ft., Stafford 7551; Jesus Maria,
Huancarani Valley, rocky cliffs, alt. 12,000 ft., Stafford 7551; Juncal,
below San Antonio de Esquilache, wet slopes and bogs, alt. 14,000 ft.,
Stafford 111+3? Viso, among rocks in shallow brook, alt. about
9,000 ft., Macbride & Featherstone 604.; near Puno, alt. 4,000 m.,
Soukup lf.59.
All of the remaining seven species of Mimulus from western
South America could be expected to occur in Peru. Of the eight
species reported in this area, six have been collected thus far in
Chile. Seven of the eight pertain to the section Simiolus, only
M. bridgesii is placed in section Paradanthus. This latter species is
distributed from Central America to Chile.
ORTHOCARPUS Nutt.
Annual herbs, erect and branching from near the base or un-
branched. Leaves alternate, linear or linear-lanceolate, entire or
pinnatifid, sessile; flowers in dense or interrupted, narrow spikes,
subtended by purplish bracts; calyx tubular-campanulate, 4-cleft,
lobes linear; corolla purplish, narrow, upper lip galeate, erect, en-
tire, little longer than the 3-saccate, lower lip; stamens 4, didyna-
mous included in the galea, anthers glabrous, sacs unequally placed,
upper medianly attached; pistil included in the galea, style filiform,
stigma globose; capsule loculicidal, ovoid, shorter than the calyx;
seeds numerous, testa loose, brown, reticulate.
A genus of 25 species, predominantly western North American,
one of which occurs in the Andes of Chile and Peru, 0. laciniatus.
Orthocarpus laciniatus (H. & A.) Keck, Proc. Calif, Acad.
Sci. ser. 4, 16(17): 548. 1927. Castilleja laciniata H. & A., Bot.
Beech., 40. 1830. Oncorhynchus pinnatifidus Lehm., Ind. Sem. Hort.
Hamb., 1832. 0. tenellus Pisch. & Mey., Ind. Sem. Hort. Petrop. 1 :
35. 1835. Orthocarpus australis Benth. in DC., Prodr. 10: 537. 1846;
S. Wats., Bot. King's Exp., 459. 1871.
Branched or unbranched (rarely) annual with pilose stem and
often pubescent foliage, appearing hazy-purple in mass. Leaves
linear or linear-lanceolate, lower entire, upper pinnatifid with 2-4
linear-filiform divisions, 1.5-4.5 cm. long, usually short-hispid, vary-
1 Duplicated number, both collected on May 19, 1937.
2 Collected November 28, 1937, indicating either a six-month flowering period
or more than one flowering period for this species.
696 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
ing from sparsely hispidulous to densely lax-pubescent; spike narrow;
flowers sessile or almost so, subtended by lanceolate or lance-ovate
bracts with 3-5 divaricate, linear-attenuate divisions, bracts green
and purple or rose-purple tipped, longer or shorter than the flower;
calyx 1.4-2.2 cm. long, soft-pubescent, lobes linear, attenuate, the
tube cleft about halfway medianly and little less laterally; corolla
purplish, narrow, 12-25 mm. long, galea 1.5-2.0 mm. longer than
the lip, slightly falcate, narrowed apically, puberulent to pubescent
externally, the wide, scarious margins excepted, densely pubescent
internally, lip shallowly 3-saccate, teeth 1-2 mm. long, oblong, erect,
basally purple; anthers glabrous, linear, ca. 1.5 mm. long, upper sac
attached dorsally, medianly, lower almost apically, filaments ciliate;
capsule ovoid, 8-14 mm. long, shorter than the calyx, glabrous; seed
oblong-ovoid, narrowed to the funicular end.
High elevations from northern to southern Peru (and into Chile)
in open, dry sites.
Cajamarca: Celendin, ca. 2 km. on track to Balsas. Flowers
purple and yellow, alt. ca. 2,700 m., Hutchison & Wright 5156; gorge
of Rio Maranon, 3-4 km. below summit of Balsas-Celendin road,
overlooking Balsas, alt. 2,950 m., Hutchison & Wright 5267. — Cuzco:
Anta Prov., Huarocondo, alt. 3,200 m., Herrera 3633. — Huanca-
velica: Castrovirreina Prov., above Pisco, above Huaytara, alt.
2,700-2,800 m., Weberbauer 54-16. — Junin: La Aroya, base of lime-
stone cliffs, alt. ca. 12,000 ft., Macbride & Featherstone 965. — Liber-
tad: Huamachuco Prov., summit above Arecapampa, on road to
Huamachuco, alt. 3,970 m., Hutchison & Wright 6266. — Lima: Ma-
tucana, alt. about 8,000 ft., Macbride & Featherstone 152. — Pasco:
Huariaca, alt. about 9,500 ft., Macbride 3114; Yanahuanca, alt.
about 14,000 ft., Macbride & Featherstone 1164- — Sin. Loc.: Weber-
bauer 5J.16.
OURISIA Comm. ex Juss.
Ourisia Comm. ex Juss., Gen. 1: 100. 1789, Dichroma Cav., Icon.
6: 59, t. 582. 1801.
Small erect, repent or caespitose herbs, rarely becoming woody
at base, caulescent or acaulescent. Leaves mostly opposite, narrowly
ovate to almost round, 2-3 mm. to 10-12 mm. long and usually little
narrower, varying from about half as wide as long to sometimes as
wide as long, usually long-petiolate below to sessile above, the lower
petioles sometimes longer than the blades, glabrous to densely long-
FLORA OF PERU 697
pubescent, mostly obtuse at both ends, less often acute at the apex,
infrequently so at base; flowers mostly less than 12 mm. long,
pedicellate, solitary and axillary or terminally forming short racemes,
or subumbelliform and pedunculate; calyx 5-lobed, longer or shorter
than the corolla tube, lobes imbricate to almost valvate, calyx tube
shorter than the oblong or narrowly ovate lobes; corolla 5-lobed usu-
ally red or scarlet, sometimes white, rarely orange or yellow, bilabi-
ate, the lips equal or unequal, tube longer than lobes, throat inter-
nally glabrous and open or densely pubescent and wholly or partly
closed ; stamens 4, didynamous, included or exserted, the pairs almost
equal in length, filaments inserted about halfway from the base1 or
near the base of the corolla anthers divaricate; stigma lobes united,
style longer than the ovary, sometimes equaling or exceeding the
stamens; capsules loculicidal, ovoid or oblong, occasionally orbicular,
placenta thickened, septum semi-persistent, papery; seed numerous,
testa loose, reticulate.
A genus of about 30 species with greatest concentration in north-
ern and western South America, extending to the Antarctic region
and New Zealand (six spp.). This genus exhibits the same trans-
pacific discontinuity as Calceolaria and the Veronica-Hebe complex
(see below) .
1. Corolla internally pilose, the pubescence at least partly closing
the throat; style shorter than the corolla tube.
2. The 3 anterior lobes of the corolla emarginate or obcordate;
calyx subtended by 2 cupular, sheathing bracts. . .0. muscosa.
2. Corolla lobes rounded or obtuse; calyces ebracteate.
3. Pedicels shorter than the subtending leaves; calyx about
equaling the corolla tube in length; lips of the corolla sub-
equal; a staminode sometimes present; plants repent.
0. biflora.
3. Pedicels longer than the subtending leaves and usually longer
than the flowers; calyx much shorter than the corolla tube;
lips of the corolla unequal; staminode always lacking; plants
caespitose, erect 0. pratioides.
1. Corolla internally glabrous, throat open; style longer than the
corolla tube 0. chamaedrifolia.
1 Rare in the family, usually inserted at or near the base of the corolla.
698 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
Ourisia biflora Wedd., Chlor. And. Nov. 2: 117. 1857.
Villose, repent, very short-caulescent, annual herbs. Leaves
ovate, or especially the upper, obovate-cuneate, minute to 8 mm.
long, crenate, apex obtuse, narrowed to base, petiolate; petiole
shorter than the blade; flowers axillary, pedicellate, most often 2 in
each axil, sometimes one, clustered at the apices of the branches;
calyx lobes densely villose, lanceolate or oblong, 4.0-5.5 mm. long,
as long as the corolla tube, united only near the base; corolla white,
reverse pink or violet (fide coll.), 5.0-6.5 mm. long, lobes about 1.0-
1.5 mm. long, lips subequal, anterior lip 3-lobed, lobes broadly ovate,
sinuate, posterior lip (upper) 2-lobed, lobes little narrower than the
anterior, throat closed by dense, soft, white pubescence, tube sub-
funnelform; fertile stamens 4, didynamous, included in the tube,
filament black; style little longer than the ovary, equaling or little
shorter than the filaments; capsule unknown.
Sin. Dept. : Limbani, among rocks, alt. 14,500 ft., Stafford 1112.
Described from Bolivia, Dept. La Paz, no other collections known.
Ourisia chamaedryfolia Benth. in DC., Prodr. 10: 493. 1846.
Creeping, annual, branching herb, stem glabrous or sparsely to
densely beset with multicellular, soft pubescence. Leaves ovate or
oblong-spathulate, glabrous, hirtellus or soft-pubescent on the cre-
nate margins, up to 1.2 cm. long, usually shorter and 2-5 mm. wide,
apices subacute to acute, bases attenuate or obtuse, tissue frequently
punctate, more densely beneath than above, punctations black;
flowers 1.7-2.1 cm. long, solitary, axillary, pedicellate; pedicels up
to 2.8 cm. long, much longer than the subtending leaves, sparsely
pilose with multicellular hair; calyx 6-8 mm. long, sepals glabrous
and ciliolate, oblong, subacute, united up to about a third their
length into a loosely investing tube; corolla glabrous or sparsely
puberulent, scarlet or orange, tube salverform or almost cylindric,
11-13 mm. long, throat open, glabrous, lobes 2-3 mm. long, hemi-
spheric, margins entire, lips equal or unequal; stamens exserted, the
filaments inserted about half way from the base of corolla tube,
didynamous, the pairs subequal in length, anterior barely longer;
style exserted beyond the corolla, 13-16 mm. long, ovary ovoid, ca.
2-5 mm. long, stigma punctiform; capsule ovoid, glabrous, about as
long as the calyx, septum deciduous, placenta little thickened; seeds
irregular or subtrigonous, angles obtuse. Two clearly distinguished
varieties in Peru.
FLORA OF PERU 699
Sepals pilose, eciliolate; corolla lips very unequal; stamens about
equaling the corolla tube in length; leaf blade obtuse at base.
O. chamaedrifolia var. chamaedrifolia.
Sepals glabrous, ciliolate; corolla lips equal or almost so; stamens
usually a little longer than the corolla tube; leaf blade attenuate
at base 0. chamaedrifolia var. elegans.
Ourisia chamaedrifolia Benth. var. chamaedrifolia.
Leaves ovate, base obtuse; petioles terete and differing further
from var. elegans as in the key.
Apurimac: Salcantay, humid rocks, 4,300-4,500 ft., Rauh-Hirsch
PI 409.
Ourisia chamaedrifolia var. elegans Wedd., Chlor. And. Nov.
2: 114-115. 1857.
Leaves oblong-spathulate, base attenuate; petioles flattened.
Cuzco: Marcapata. River bank, in moss under rocks, trailing
growth, 10,000-12,000 ft., Stafford 987.
Ourisia muscosa Benth. in DC., Prodr. 10: 493. 1846.
Repent, trailing herb, rooting at the nodes, glabrous or pilulose,
stems a few cm. long, the flowers on erect pedicels. All parts minute;
leaves 1-3 mm. long on petioles that are thickened and shorter or
more often longer than the ovate or broadly oval, entire blades,
apices rounded, little narrowed to base; flowers axillary, solitary,
pedicellate, 4-6 mm. long; pedicels of mature flowers 1-10 mm. long
with a pair of sheathing, cupular bracts apically, usually glabrous,
thickened and flattened; calyx 2.0-2.5 mm. long, glabrous, tube al-
most lacking, segments ovate, acuminate, 1-nerved; corolla tube
white, glabrous, salverform, ca. 4 mm. long, lips subequal, throat
densely villose internally and at the base of the minute lobes, the
3 anterior lobes apically emarginate or obcordate; stamens included,
barely reaching the throat of the corolla, filaments black; style little
shorter than stamens, longer than the globose, apically grooved
ovary; capsule not seen, ex desc. equaling the calyx lobes.
Arequipa: Pichu Pichu Range in boggy ground, alt. 13,500 ft.,
Stafford 691. — Puno: San Antonio de Esquilache, alt. 16,000 ft., Staf-
ford 800.
This species, beside its very small size, can easily be distinguished
by its emarginate or obcordate anterior corolla lobes, and cupular
700 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. XIII
bracts that sheathe the calyx lobes. Known also from Bolivia and
Ecuador.
Ourisia pratioides Diels in Urban, PI. Nov. And. in Engler,
Bot. Jahrb. 37:428. 1906.
Ex desc. Caespitose, villose herb, stems procumbent, 4-8 cm.
long, apices ascending, 1-flowered. Leaves opposite, lower petiolate,
suborbicular, 3-4 mm. long and wide, almost entire, villose above,
glabrescent beneath; petioles dilated at base, villose, 5-6 mm. long,
upper leaves sessile or subsessile, margins obsoletely crenate; calyx
pilulose, tube about 2 mm. long, the obtusish segments 3-4 mm. long;
corolla pale purple, lips unequal, tube 5-7 mm. long, internally pilose,
the 2 lateral posterior lobes about 4 mm. long, internally barbed,
smaller, and fused farther than the 3 lower lobes that are 6-8 mm.
long and elliptical with the median a little bigger than the lateral;
stamens 6-7 mm. long; ovary glabrous.
Huanuco: Prov. Huamalies, mountains of Monzon, alt. 3,300-
3,400 m., Weberbauer 3724.
Only description and photograph of the type seen. This taxon
appears to be closest to the Bolivian species, 0. pulchellum Wedd.,
where the leaves are larger, not villose above and the stamens and
petioles are longer.
PORODITTIA G. Don
Porodittia G. Don, Gen. Syst. 4: 608. 1838, based on Jovellana
Cav., Icon, et Descr. 5: 32, t. 453. 1799. Calceolaria Vahl, Enum. 1:
81. 1804. Baea Pers., Syn. 1: 15. 1805, non Comm. ex Juss. Trian-
thera Wettst. in Engler & Prantl, Pflanzenf. IV(3b): 155. 1891.
Stemotria Wettst. & Harms in Engler & Prantl, Pflanzenf. Reg. II-
IV: 462. 1898.
Erect, much branched shrub or woody vine, puberulent on wood
and foliage, often glandular-puberulent in the inflorescence, wood
often glabrate with age near the base of the plant, estriate, terete,
usually drying brown. Leaves opposite, variously ovate or elliptic
in outline, margins irregularly deeply toothed or lobed, teeth usually
sharp, sinuses often wider than the teeth, petiolate; petioles often at
least partly, sometimes entirely, surrounding the stem as a thin or
wide collar, rarely only little dilated at base; inflorescences large,
many-flowered, irregularly branching cymes, terminal on branches,
FLORA OF PERU 701
axils subtended by progressively reduced leaves to the apex; pedicels
and peduncles progressively shorter to the terminal pedicel; flowers
zygomorphic, mostly over 1 cm. in diameter; calyx of 4 sepals, united
only near the base; corolla yellow and variously violet- or brown-
striped or -blotched, bilabiate, the lips cut almost to the base, upper
2-lobed, lower 1-lobed, lips little rounded-saccate, tube very short;
stamens 3, inserted at the base of the upper lip of the corolla, 1
median and 2 lateral, filaments short, flattened, exceeded by the sub-
compressed, parallel anthers that dehise by large apical pores ; ovary
ovoid, shorter than the style, stigma lobes united, minutely puncti-
form; capsule ovoid, bilocular and bivalved, valves bifid at apex;
seeds very small.
A monotypic genus (Porodittia triandra (Cav.)), G. Don endemic
in Peru.
Porodittia triandra (Cav.) G. Don, Gen. Syst. 4: 608. 1838.
Jovellana triandra Cav., Icon, & Descr. 5: 453. t. 32. (pro parte) 1799.
Baea triandra Pers., Syn. 1: 15. 1806. Calceolaria triandra Vahl,
Enum. 1: 181. 1804. Trianthera triandra Wettst. in Engler & Prantl,
Pflanzenf. IV(3b): 155. 1891.
Leaves up to 5-6 cm. long and 3 cm. wide, often shorter and nar-
rower, variable in size; petioles 6-15 mm. long; calyx lobes broadly
ovate, acute, ciliolate, subequal, densely puberulent-pubescent,
about 4 mm. long, posterior sepal a little smaller, anterior a little
bigger than the lateral; corolla not or only very little calceolate but
mimicking Calceolaria, lips open, upper, the posterior, 2-lobed about
2 times wider than the 1-lobed anterior lip that is composed of 3
fused petals, posterior lobes free at least half way to the base, tube
very short; filaments about 1 mm. long, anthers about 1.5-2.0 mm.
long and almost as wide, yellow or blue; ovary glandular-pubescent;
capsule ovoid, abruptly narrowed to a long-acute apex, much exceed-
ing the calyx; seeds numerous, septum adherent, placenta much en-
larged at base.
Ancash: Prov. Bolognesi, Matarragra N. of Chiquian, alt. 3,250
m., Ferreyra 5583. — Ayacucho: Coracora, alt. 3,200 m., Weberbauer
5810. — Lima: Prov. Huarochiri, Circa Llacllishi, alt. 2,700-2,800 m.,
Pennell 11+527; Along Rio Chillon, above Obrajillo, alt. 2,700-3,000
m., Pennell 11+362; One league from Obrajillo, Nee s.n.; Near Antai-
cocha, Cerro Colorado, E. of Canta, alt. 3,200-3,400 m., Pennell
11+622— Sin. Depto.: Viso, alt. about 9,000 ft., Macbride & Feather-
stone 578.
702 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
RUSSELIA Jacq.
Plants shrubby or scandent, with terete or angled stems that are
often ridged and striate between the ridges. Leaves opposite or ver-
ticillate; inflorescence axillary, cymose and most often branched;
calyces deeply 5-parted, persistent; corolla tubular or funnelform,
slightly irregular, upper lip emarginate, lower 3-lobed; stamens 4,
didynamous, inserted 1-2 mm. above base of the corolla; staminode
usually short; ovary 2-loculed; fruit a globose capsule, loculicidal,
placentae enlarged, bearing numerous hairs among the many, small
wingless seeds.1
A new world genus of 52 species, native from Mexico to northern
South America including the West Indies. However, several species
are now in cultivation in many parts of the world.
Divided into two subgenera by Dr. Carlson (see footnote for
generic description) . Only the first of these, Juncirusselia, now per-
taining. Its one species, R. equisetiformis S. and C., the only repre-
sentative so far from Peru. This species has an unique habit for
the genus.
Juncirusselia Carlson. Plants junc-like, stems thin, numerous
verticillate. Inflorescence 2-flowered.2 Corolla glabrous internally.
Russelia equisetiformis S. and C., Linnaea 6: 377. 1831. R.
juncea Zucc., Flora 15, Beibl. 2: 99. 1832.
Subshrub up to about 1 m. tall, glabrous or almost so, profusely
branched, stems and leaves verticillate below and opposite above,
stems (in life) green, 4-12-angled with prominent ridges on the angles
and striae between. Leaves caducous, ex desc. lower leaves 3-6 in
a whorl, ovate or elliptical-acute, larger with one or a few teeth on
each side, 8.5-15.0 mm. long and 6-9 mm. wide, upper smaller,
linear, entire, with a few circular resin dots; petioles 3-4 mm. long,
often remaining appressed after leaves fall; inflorescences mostly
apical; bracteoles at bases of peduncle and pedicels, reduced, linear
or narrowly lanceolate, entire, resin-dotted; peduncles short, 3-4 mm.
long or elongate up to 2 cm. long; pedicels shorter; calyx lobes ovate,
acute or mucronate, 2-3 mm. long, much exceeding the short tube;
1 Generic description for this genus is very broad (an exception to the rule that
descriptions are limited to Peruvian material) and follows Carlson, M. C., The
Genus Russelia in Fieldiana: Botany 29(4): 231-292, 6 pi. 1957. This course is
taken because the only species of Russelia thus far reported is exceptional in
the genus (standing as a monotypic subgenus) and it seems likely that at least one
other species will soon be collected in Peru.
2 Sometimes more than two-flowered in cultivation.
FLORA OF PERU 703
corollas red, 1.5-2.5 cm. long, anthers orbicular; staminode subobso-
lete, 0.5-0.7 mm. long; pistil about equal to the shorter stamens,
style much longer than the ovary, stigma with 2, small lobes often
coherent, appearing punctiform or lamellate; capsule ex desc.,1 glo-
bose 3-6 mm. in diameter, lower third in the persistent calyx, style
persisting at apex; seed small oval, testa warty, light brown, among
white hairs that fill the capsule.
Lima: Botanical Gardens in Lima at 150 m. alt., Soukup 2919.—
Loreto: Iquitos and vicinity, alt. 120 m., LI. Williams 3582.
Generally distributed from Mexico through Central America and
the West Indies to the Guianas, Colombia, and Peru. Cultivated in
numerous other parts of the world. Frequently collections from the
natural range are prepared from cultivated material although col-
lectors often neglect to note this.2
Other species which may likely be found in Peru are R. sarmentosa
Jacq. and R. coccinea Wettst., basically Central American but occur-
ring in diverse ecologic sites. These pertain to the subgenus Eurus-
selia Carlson.
SCOPARIA L.
Ref. Chodat, R. Bull. Herb. Boiss. 2(8): 1-16; 85-89. 1907-1908.
Shrub or subshrub woody only at base, subglabrous. Stems te-
rete, branches terete or 4-angled, longitudinally striate, when winged
the wings sometimes strongly ridged; leaves decreasing in size base
to apex, opposite or ternately verticellate, especially the upper;
flowers pedicellate, usually 2 or 3 in each leaf axil, often appearing
clustered; pedicels frequently shorter than the bracts and longer than
the calyx, only rarely exceeding 3 times longer than the calyx, usually
very sparsely puberulent with stiff hairs, occasionally vesture dense;
sepals 4,3 due to loss of the posterior, imbricate, usually united only
at the base, thinly scarious margined and ciliolate; corolla subrotate,
4-lobed due to loss of posterior petal, white, internally beset with
longish hairs at the base of the lobes, petals frequently reflexed, at
least when dry; stamens 4, inserted on the base of the short corolla
tube but alternating with the lobes, filaments filiform, anthers sub-
sagittate, parallel, dorsifixed, proximate; style erect, usually little
1 Both mature lower leaves and capsules wanting in Peruvian collections.
2 Fide Carlson, I.e.
3 Sepals sometimes 5, in extra Peruvian material.
704 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
longer than the ovoid to globose ovary, stigma lobes short, truncate,
free at the tips but appearing united, punctiform or clavate; capsule
septicidal and secondarily barely loculicidal at the apex, ovoid or
globose, seeds numerous, furrowed, ovoid, placenta well developed.
A single species S. dulcis L., well represented and of broad distri-
bution in Peru. The genus is widely distributed in the warm tem-
perate and tropical regions of the world. Its greatest concentration,
about 10 species, is in the Western Hemisphere in Argentina and
Paraguay. Only S. dulcis L. reported from North Africa, India,
China, Philipine Islands, Malaya, and Australia.
Scoparia dulcis L., Sp. PI. 116. 1753; Gen. PL ed. 5. 83. 1754.
S. ternata Forsk., Fl. Aeg.-Ar. 30. 1775. Gratiola micrantha Nutt.,
in Amer. Journ. Sci. 5: 287. 1822. Scoparia grandiflora Nash, in
Bull. Torrey Club 23: 105. 1896.
Largest leaves up to 5 cm. long, blades mostly 1.0-3.5 cm. long,
narrowly lance-ovate to ovate, uppermost sometimes lanceolate,
apex usually acute, cuneately narrowed to the short petiole, margins
deeply, irregularly serrate (the larger leaves) to serrate-serrulate (the
smaller, upper), glandular-punctate; pedicels up to 9 mm. long; calyx
lobes about 1.5-2.0 mm. long, elliptic to elliptic-ovate, 3-nerved;
corolla white, 4-10 mm. wide, lobes almost equal, ovate; stamens
with the filaments about 2 times longer than the anthers; capsule
1.5-2.5 mm. long; seeds yellowish brown.
Cuzco: Valle del Urubamba, 2,200 m., Herrera 3200; Prov. Quispi-
carchis, Quinco Mic, 800-900 m., C. Vargas C. 9759.— Junin: Chan-
chamayo Valley, Carlos Schunke 374; 1,200 m., Carlos Schunke 398;
La Merced, alt. 4,000 ft., Macbride 5669. — Loreto: Along Rio Itaya,
LI. Williams 131 ; Lower Rio Nanay, LI. Williams 450, 726, and 1300;
Santa Ana on Upper Rio Nanay, LI. Williams 1250; On the Amazon
River, San Salvador, LI. Williams 1862; Pro 1982; La Victoria 2653;
Lower Rio Huallaga, 155-210 m., LI. Williams 4362. — Piura: Nigri-
tos,1 Haught F-402; Parinas Valley, Haught F-402; Talara, alt. 500 m.,
Rauh-Hirsch C 26. — San Martin: Prov. Tingo Maria, Monson R.
near Tingo Maria, alt. 625-1,100 m., Allard 21157; Lima, 840 m.,
LI. Williams 6480; San Rogue, 1,350-1,500 m., LI. Williams 7707,
Alto Rio Huallaga, alt. 360-900 m., LI. Williams 5623.— Sin. Depto.:
1 Correctly Negritos
2 Numbers may be erroneous.
FLORA OF PERU 705
Pampayacu, Hacienda at the mouth of Chinachao Rio,1 about 3,500
ft. alt., Macbride 5127.
SIBTHORPIA L.
Sibthorpia L., Sp. PL 631. 1753; Gen. PI. 320. 1754. Disandra
L. f., Suppl. Gen. PI. 32. 1781.
Prostrate herb, rooting at the puberulent, distant nodes, stems
glabrous or occasionally sparsely puberulent. Leaves alternate, pu-
bescent on both surfaces, almost round, up to about 3 cm. in diam-
eter, margins lobed, lobes rounded; petioles pubescent, frequently
longer than the diameter of the blades; flowers solitary, axillary, pe-
dicellate; pedicels pubescent, longer than the flowers; sepals 4 (3-5,
rarely 8), united into a campanulate calyx; corolla little longer than
the calyx, red or greenish-white, tube lacking or minute, lobes obtuse
subequal ; stamens 4 (3-5) equal, inserted near the base of the corolla,
exserted beyond the tube, anther locules parallel, contiguous at apex;
style about as long as the ovary, the pistil shorter than the stamens;
capsule loculicidal, compressed-globose or broadly compressed-obo-
void, little exceeding the calyx, apex obcordate; seed reticulate, dark
red; placenta thin, septum deciduous.
A genus of about six species in the subtropics of the old and new
worlds. One species with two forms collected in Peru.
Corolla red (rose-purple or pink); capsule glabrous or occasionally
soft-pubescent S. rotundifolia f. rotundifolia.
Corolla greenish-white; capsule stiff-pubescent. .S. rotundifolia f. alba.
Sibthorpia rotundifolia (R. & P.) Edwin, Fieldiana: Botany
31(8): 228. 1967. Veronica rotundifolia R. & P., Fl. Peruv. 1: 6.
1798. Sibthorpia pichinchensis HBK., Nov. Gen. et Sp. 2: 390,
t. 176. 1817.
Flowers up to 4 mm. long; calyx about 3 mm. long; corolla rose-
purple to pink, rotate or subrotate, lobes ovate, entire; capsules up
to about 4.5 mm. long; seed several, compressed.
Junin: Mito, moist, deep humus in a "clearing" in eastern woods,
alt. ca. 9,000 ft., Macbride & Featherstone 1717.
Sibthorpia rotundifolia f. alba Edwin, Fieldiana: Botany
31(8) : 228. 1967. Differing from the typical form as in the key.
1 May be another name for the Rio San Juan, if actually Chincha Rio.
708 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Cuzco: Near Rio Yanamayo, below "Pillahuata," wet mossy
rocks by cascade, alt. 2,100-2,300 m., Pennell 13935.
The typical form having been collected in north-central Peru
and the white-flowered form in southern Peru indicate a genus much
more prevalent than the scanty material indicates.
The genus is well represented by collections from Mexico to
Colombia and Argentina. The species limits, however, are poorly
defined.1
STEMODIA
Stemodia L , Syst. Nat. ed. 10. L 118. 1759. Lendernia Minod., in
Bull. Soc. Geneve II. 10: 240. 1918. Unanuea Pennell, Proc. Acad.
Nat. Sci. 72:160. 1920.
Ref. Minod in Bull. Soc. Bot. Geneve II. 10: 155-252. 1918.
Herb or less often shrub or subshrub, frequently glandular-pubes-
cent or villose, often aromatic. Leaves opposite or verticillate, 3-4
in each verticil ; flowers axillary, pedicellate, with often 1 sometimes
2, rarely more in each axil or forming spikes or spike-like, rarely
racemose inflorescences; pedicels usually short, sometimes longer
than the flower, ebracteolate or 2-bractiolate; calyx lobes 5, basally
connate, narrow, all alike or the posterior lobe a little longer, imbri-
cate; corolla white, blue or purple, with 4 lobes, the posterior pair
fused, the 3 latero-anterior apparent, scarcely to strongly zygo-
morphic, bilabiate, the upper, posterior lip erect or arched, the lower,
anterior, flattened, corolla tube cylindric, usually longer than the
lobes; stamens 4, didynamous, usually included, filaments filiform,
anther locules disjunct, stipitate; ovary ellipsoid to narrowly ovoid,
shorter than the style that is apically dilated, stigma lobes 2, inter-
nally stigmatic; capsule globose, ovoid and usually acuminate, septi-
cidal and often secondarily loculicidal; seeds numerous, minute, testa
longitudinally furrowed.
While the collections bear about ten names it is felt that five spe-
cies are adequate to designate the Peruvian Material. A genus of
about 30 species, distributed throughout much of the temperate and
tropical zones of the world, but basically warm-temperate to tropical
American.
1. Flowers in spiciform inflorescences or spikes; pedicels bibracteo-
late; leaves opposite.
1 Sibthorpia repens (L.) Kuntze based on Willichia repens L. may be conspecific
with rotundifolia. They are maintained as distinct taxa for the present.
FLORA OF PERU 707
2. Corolla violet-blue, strongly zygomorphic, pubescent over the
base of the lower lip; bracteoles and sepals attenuate, linear-
lanceolate, equal in length or bracteoles shorter; annual or less
often perennial S. durantifolia.
2. Corolla white, scarcely zygomorphic, glabrous; bracteoles lin-
ear-acute, longer than the narrowly lanceolate, acute sepals;
perennial S. piurensis.
1. Flowers mostly axillary, occasionally in falsely spiciform inflores-
cences (S. suffruticosa) ; pedicels ebracteolate; leaves verticillate
(opposite and verticillate in S. parviflora}.
3. Leaf blades all or mostly less than 2.0 cm. long; anterior sta-
mens with anthers smaller than posterior, anterior filaments
inserted lower than posterior S. parviflora.
3. Leaf blades more than 2.0 cm. long, up to 6.5 cm. long; anthers
of all stamens equal in size, anterior filaments inserted higher
than posterior.
4. Pedicels 5-6 mm. long; corolla often less than 10 mm. long,
up to 12 mm. ; flowers often more than one per leaf axil.
S. suffruticosa.
4. Pedicels ca. 15 mm. long; corolla ca. 17 mm. long; flowers
usually one per leaf axil S. chodati.
Stemodia chodati Minod., Bull. Soc. Bot. Geneve II. 10: 199.
1918.
Erect shrub, stem striate, pubescent. Leaves up to 6.5 cm. long
and 2 cm. wide, subsessile, ternate, ovate-lanceolate,1 base attenuate,
tissue on upper surface bullulate, very short-pubescent, margins den-
tate, teeth round or obtuse, subglabrous beneath; flowers solitary,2
axillary2; pedicels 15 mm. long; sepals 11 mm. long and 0.5 mm.
wide, subulate; corolla 17 mm. long, globose at base, constricted
medianly, anterior lip 3-lobed, laterals ovate-lanceolate, acuminate,
median broadly ovate, hardly acuminate, posterior lip rounded, little
emarginate; anterior stamens longer and inserted higher on the co-
rolla tube than the posterior the filaments of which are thicker; ovary
ovoid; style thick, the apical stigma clavate; fruit unknown.
1 Photograph of the type (the only) collection displays elliptically ovate leaves,
and attenuate, acute or subobtuse bases.
2 From photograph, Field Museum 8550.
708 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Except as footnoted, this description from the original.
Amazonas: Prov. Chachapoyas, southern part. With neither
collector or number. Presumably the type at the Geneva herbarium.
This species is closely related to S. suffruticosa, differing from this
latter as in the key and having acuminate anterior lobes of the corolla
as well as little different leaf shape and non-rostellate ovary.
Stemodia durantifolia (L.) Sw., Obs. Bot. 240. 1791. Phae-
lypea erecta P. Br., Nat. Hist. Jam. 269. 1756. Capraria durantifolia
L., Syst. Nat. ed. 10. 1116. 1759. Conobea verticillaria Spreng., Nov.
Prov. 13. 1819. Stemodiacre durantifolia (L.) Kuntze, Rev. Gen.:
466. 1891. Stemodia erecta (P. Br.) Mined, in Bull. Soc. Geneve II.
10: 212. 1918.
Erect, viscid, glandular-pubescent herb or subshrub, woody at
base, branchlets and branches 4-angled or the older terete. Blades
opposite or 3-4-verticillate, sessile often broadly clasping or amplexi-
caul, obovate, or elliptically obovate, acute at apex, long-attenuate
toward the base and abruptly flaring at the stems or branches, up to
9-10 cm. long and 3.5-4.0 cm. wide, usually smaller, much reduced
base to apex, margins serrate or serrate-dentate, glandular-pubescent
beneath, less so above; inflorescences spiciform or spikes; flowers
bracteate and bibracteolate, one in the axil of each bract; pedicels
almost lacking; bracteoles lance-linear, attenuate, glandular-pubes-
cent, clasping the calyx; calyx-lobes 5, lance-linear, attenuate, 4-6
mm. long, viscid, glandular-pubescent, fused at the base into a small
tube, sepals about the length of the corolla tube; corolla light blue,
the lobes very short, lips subequal, shorter than the tube, lower lip
and tube densely pubescent within, upper sparsely so; stamens in-
serted more than halfway up the tube, the anterior pair of filaments
almost twice the length of the posterior or the pairs almost equal in
length; anther locules stipitate, small, about a fourth to a fifth the
length of the filaments, stipe only little if at all thickened; style fili-
form, stigma abruptly bilobed; capsule shorter than the sepals, ellip-
soid or subovoid-ellipsoid; seed when dry, almost black.
Piura: Nigritos,1 Haught f-322: Parinas Valley, Haught F-32.2—
San Martin: Alto Rio Huallaga, 360-900 m., LI Williams 621*2.
1 Correctly Negritos
2 These may be mixed numbers.
FLORA OF PERU 709
One of the most generally distributed species of the genus, occur-
ring from California to Texas through Mexico, Central America,
West Indies, and South America to Chile and Argentina.
Stemodia parviflora Ait., Hort. Kew. ed. 2, 4: 52. 1812. Erinus
verticillatus Mill., Card. Diet. ed. 8. 1768. Capraria humilis Soland.
in Ait., Hort. Kew ed. 1. 2: 353. 1789 (non Pavon). Stemodia are-
naria HBK., Nov. Gen. et Sp. 2: 287, t. 175. 1817. Stemodiacre
verticillata Ktze., Rev. Gen. 2: 466. 1891. Stemodia verticillata
Hassler, Contr. Fl. Chaco 110. 1909. S. neglecta Minod., Bull. Soc.
Geneve II. 10:195. 1918.1
Repent or prostrate viscid herb, sometimes forming mats, rarely
erect or suberect, then less than 1.5 dm. tall. Stems densely pubescent
with lax, mixed glandular and eglandular, often multicellular hair;
leaves opposite or more often verticillate, verticils 3-foliate, glabrous
to pubescent, mostly sparsely pubescent or puberulent, the vesture
frequently restricted to the veins and margins, usually broadly ovate
triangular-ovate, up to about 1.5 cm. long and 9.0 mm. wide, usually
smaller, less than twice as long as wide, acute, base variable,usu-
ally obtuse and the blade narrowly decurrent part way along the
short, pubescent petiole, margins irregularly serrate, serrations an-
trorse, occasionally dentate or denticulate; pedicel very short, pubes-
cent, ebracteolate; flowers solitary, usually 1 in an axil of a scarcely
modified, leaf-like bract; calyx lobes 5, lanceolate, subulate, more or
less than 1 mm. long, margins scarious; corolla 4-6 mm. long, light
blue, lips subequal, the two posterior imperfectly fused, pubescent
within, anterior and lateral, a little shorter, tube longer than the
lobes, pubescent within; anterior pair of stamens with longer fila-
ments and smaller anthers than the posterior, stipes little thickened,
anthers included or little exserted beyond the throat, filaments un-
equally inserted in pairs, the anterior lower on the tube; ovary little
shorter than the 2-lobed style; capsule about 3 mm. long, septicidal
and secondarily loculicidal (hygroscopic?), obovoid or subglobose-
ovoid; seed light brown, broadly ovoid.
Junin: La Merced, Macbride 5673. — Loreto: Near Iquitos, alt.
100 m., Klug 1229; Caballo-Cocha on the Amazon River, LI. Wil-
liams 24.52. — San Martin: San Roque, 1,350-1,500 m./LZ. Williams
7753. Sin. loc. : Pavon.
1 This listing of synonyms is only partial since verification of all names is be-
yond the scope of this paper. The interested reader may wish to check Conobea pu-
mila Spreng.; Poarium veronicoides (Desv.) Ham., and Stemodia pauciflora Rusby.
710 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Widely distributed, reported from Mexico to Argentina including
the West Indies.
Stemodia piurensis Pennell, Not. Nat. 179. p. 2. 1946.
Plant probably perennial, woody at base, stem glandular-pubes-
cent or finely pubescent. Leaves oblong-lanceolate, acute, closely
serrate-dentate with sharp outstanding teeth, glandular-puberulent,
bluish green (at least when dry),1 rounded-clasping at base, largest
3.0 cm. long and 7.5 mm. wide, lower somewhat narrowed proximally,
middle and upper scarcely so and these latter subtending flowers;
pedicels less than 1 mm. long; bracteoles linear-acute, longer than
the sepals; sepals 5, narrowly lanceolate, acute, 4-5 mm. long; corolla
white, glabrous, 4-5 mm. long, lobes subequal, uppermost and lower-
most slightly larger than the others; anther cells slightly stipitate;
capsule 3 mm. long, apparently conic.
Piura: Decumbent. River Valleys, Que"brada Mogollon, Amo-
tape Hills, in flower. March 28-30, 1941. Haught & Swenson 1151*2
(BKL).
The original description is fully sufficient to demonstrate the dif-
ferences between this taxon and S. durantifolia to which it is closely
allied.
Stemodia suffruticosa HBK., Nov. Gen. et Sp. 2: 287. 1817.
S.frutescens Reicheb., Hort. Bot. 1: 1827. (III). Unanuea febrifuga
Steud., Nomen. 2: 729. 1841. Stemodiacre suffruticosa Ktze., Rev.
Gen. 2: 466. 1891. Stemodia suffruticosa f. dentata Minod., Bull.
Soc. Geneve II. 10: 201. 1918. Unanuea dentata (R. & P.) Pennell,
Proc. Acad. Nat. Sci. Phila. 72: 161. 1920, based on Unanuea
R. & P., Ic. Fl. Peru, ined.
Erect shrub or subshrub, pubescent, the hairs usually multicellu-
lar. Stem with internodes up to 10 cm. long, internodes on branch-
lets much shorter, less than 1 cm. long near the tips; leaf blades
ternately verticillate, ovate, sometimes ovate-sublanceolate, bullu-
late, up to 4.5 cm. long and 1.6 cm. wide, usually smaller, pubescent
on veins beneath and on tissue and veins above, apex acute, base
acute or cuneately acute, margins crenate-dentate; flowers mostly
2-3 in each axil of the leaf -like bracts, pedicellate; pedicels 5-6 (-8)
mm.2 long; sepals 5, united at the base only, linear to subulate or
1 Plant unknown to me. Description after Pennell. S. durantifolia appears
yellowish to dark green or brown-green when dry.
2 According to Minod up to 11-12 mm. long — Miers 3018, Paccha Mountains.
FLORA OF PERU 711
very narrowly lanceolate, long-acute, 6-7 mm. long. Corolla when
dry dark brown-red1; 9-13 mm. long, tube pubescent externally,
pubescent or glabrous anteriorly internally, little longer than the
sepals lobes equal, spreading; the anterior pair of filaments longer
than and inserted higher on the corolla tube than the posterior, in-
cluded in the tube, locules elliptic, connective thickened ; style about
twice as long as the rostellate ovary, thickened apically, persisting
in fruit, stigma lobes distinct, internally stigmatic; capsule ellipsoid,
or narrowly ovoid-ellipsoid, shorter than the sepals, septicidal and
secondarily apically loculicidal; seeds oblong.
Piura: Prov. Ay abaca,2 Weberbauer 6351 and 6362.
Widely collected from Mexico through Central America and
western South America to northern Peru, none seen from the West
Indies.
VERONICA L.
Veronica L., Sp. PL 9: 1753; Gen. PL 10. 1754.
Annual or perennial, glabrous, pubescent or glandular, erect, re-
pent or prostrate herbs. Leaves all opposite or upper alternate,
linear or lanceolate to broadly ovate, petiolate or sessile, entire or
toothed, obtuse to acute or acuminate at one or both ends, about 1-
7 cm. long and 0.3-3.0 cm. wide; flowers most often less than 1 cm.
long, usually in lateral or terminal spikes or racemes, or axillary and
racemose, pedicellate; calyx 4-lobed, tube very short or lacking, lobes
lanceolate to ovate, acuminate, acute or obtuse; corolla rotate, 4-
lobed (the 2 posterior wholly united), white to dark blue or violet,
lobes ovate or sometimes narrower, equal in length and width or
unequal in one or both dimensions; stamens 2, the anterior-lateral
pair, filaments divergent and usually exceeding the corolla, anthers
small, contiguous, parallel; style minute to 2-3 times longer than the
globose or ovoid, often apically notched ovary, stigma lobes united
or scarcely bifid at apex; capsule usually compressed, longer than
wide or wider than long, often distinctly to deeply notched at the
apex, with most often a persisting style or style base in the notch,
when deeply notched the free parts of the capsule often divergent,
longer or shorter than the sepals, loculicidal (secondarily barely sep-
1 Blue in life according to Bentham and Minod; purple-blue according to
Pennell.
2 Often Ayavaca
712 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
ticidal), placenta thin, septum deciduous; seeds few or numerous,
flattened, plane or convex dorsally and ventrally, yellow to yellow-
brown, testa almost smooth to rugose. Six species thus far collected
in Peru.
A genus of about 250 species, mostly old world but well repre-
sented in the new world with about 25 species. Specimens have been
collected in wet to dryish sites at varying altitudes from Alaska to
Chile. The following five species not yet collected may well be en-
countered in Peru, the first three more likely than the latter two:
V. beccabunga L.; V. didyma Tenore; V. salicifolia Forst.; V. ameri-
cana (Raf.) Schwein; V. plebeja R. Br.
1. Main stem indeterminate, without an inflorescence at its apex,
all leaves opposite V. anagallis-aquatica.
1. Main stem determinate, terminated by an inflorescence; the
bract-like, upper leaves alternate.
2. Plants perennial from rhizomes; inflorescences terminal on lat-
eral branches or axillary in the upper leaves of the plant.
3. Stems pubescent throughout with upcurved hairs; corolla
white or whitish with purple markings, 2 mm. long; capsule
3-4 mm. wide, pedicels minutely puberulent.
V. serpyllifolia.
3. Stems distally pubescent with mostly spreading hairs; corolla
pale- violet with darker markings, 3-4 mm. long; capsules
4-5 mm. wide; pedicels pubescent V. humifusa.
2. Plants annual, fibrous rooted; inflorescences or axillary flowers
developed from at or near the base, floriferous over most of its
length.
4. Pedicels shorter than the lanceolate to linear sepals; seeds
numerous, less than 1 mm. long; plants usually erect.
5. Sepals equal; style minute, hidden in the notch of the cap-
sule between the lobes; corolla usually white or whitish,
occasionally pale-blue . . . . V. peregrina subsp. xalapensis.
5. Anterior sepals longer than the posterior; style usually ex-
ceeding the capsule lobes, or if shorter, clearly apparent;
corolla deep violet-blue V. arvensis.
4. Pedicels longer than the ovate sepals; seeds few, 1.3-3.0 mm.
long; plants repent V. persica.
FLORA OF PERU 713
Veronica anagallis-aquatica L., Sp. PL 12. 1753. V. anagallis
latifolia Britt., Bull. Torrey Club 12: 49. 1895. V. brittonii Porter
ex Pennell, Torreya 19: 168. V. anagallis-aquatica brittonii Pen-
nell, Rhodora 23: 36. 1921.
Soft herbs growing in wet sites, erect or sometimes rooting at the
nodes, essentially glabrous, rarely short-puberulent. Leaves sessile,
cordate-amplexicaul or occasionally very short-petiolate, lanceolate,
lance-ovate or lance-oblong, up to 7 cm. long and 2-3 cm. wide, ser-
rate or entire, apex acute, often broadest at or near the base; inflores-
cences many-flowered, axillary racemes; pedicels up to 9 mm. long,
longer than the flowers, subtended by a lanceolate, entire, glabrous
bracteole; calyx lanceolate to narrowly ovate or oblong, 4-6 mm.
long, lobes mostly long-acuminate; corolla little longer than the calyx,
blue, pale-blue or violet; style longer than the ovary; capsule usually
only little notched or rounded at apex; seeds numerous, small, yel-
low-brown, testa rugulose. Stamens as the genus.
Arequipa: Prov. Condesuyos, Huario Valley below Chuquibamba,
alt. 4,000 ft., Stafford 1150; Banks of stream, alt. 7,600 ft., Stafford
628; Lower slopes of Misti, Sandeman 3748.
Collected on all continents except Australia and Arctic areas.
Veronica arvensis L., Sp. PL 13. 1753.
Pubescent, laxly erect, branching or unbranched, annual herb,
hair soft or stiff. Lower leaves ovate, short-petiolate, upper, bract-
like leaves lance-ovate or -oblong, sessile, lower leaves up to 1.3 cm.
long and 7-8 mm. wide, upper much reduced, margins irregularly
few-toothed with rounded or obtuse teeth below to entire above,
ciliate, acute at apex, usually narrowed to base; internodes very
short above; lower flowers axillary, solitary and remote from the
congested upper flowers that form spike-like inflorescences, flowers
usually subsessile, occasionally sessile; sepals lanceolate or narrowly
oblong-lanceolate, anterior pair 3-4 mm. long, posterior 2-3 mm.
long; corolla often shorter than the sepals, dark violet-blue, lobes
broadly ovate, rounded; style about 1 mm. long, little longer than
the ovary; capsule compressed-subglobose, brownish, often glandu-
lar pubescent at least on the darker hardened margins, little longer
than wide to wider than long, usually about equaling or little shorter
than the calyx, the notch at apex up to a third the length of the fruit,
the style little exceeding the lobes; seeds numerous, ovoid, testa
rugulose, light brown; placenta thin septum early deciduous.
714 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Junin: Prov. Yauli, La Oroya, in stones by trail, alt. 12,000 ft.,
Macbride & Featherstone 9^1 ; Mito, alt. about 9,000 ft., Macbride &
Featherstone 1715. — Sin. Dept.: Quinua, potato field, alt. ca. 12,000
ft., Macbride & Featherstone 2028.
Collected on all continents except Australia and the Arctic areas.
Veronica humifusa Dickson, Trans. Linn. Soc. 2: 288. 1794.
V. serpyllifolia humifusa (Dickson) Vahl, Enum. Plant. 1: 65. 1805.
V. serpyllifolia neomexicana Cock., Amer. Nat. 40: 892. 1906. V.
funesta Machr. & Pays., Contr. Gray Herb. II. 49: 68. 1917.
Perennial, rhizomatous, partly downy herb, in dry or wet sites,
erect or repent, stems distally pubescent. Lower leaves broadly
ovate or elliptic, varying to suborbicular, up to 1.8 cm. long and
almost as wide, margins entire to irregularly, subdistantly crenulate,
rounded or obtuse at both ends, usually glabrous, occasionally pu-
berulent, upper bract-leaves lanceolate or narrowly oblong or ovate,
much reduced, margins minutely denticulate, ciliolate; inflorescences
racemes; the lowermost, pubescent pedicels as long as the capsules;
calyx lobes a little unequal, one anterior largest, the other smallest,
posterior equal to each other, elliptically ovate to ovate, 2.5-4.0 mm.
long; petals blue, 2.5-4.5 mm. long, the longest almost as wide, the
posterior pair largest, broadly ovate, one anterior smallest, narrowly
ovate, about half the posterior, remaining anterior petal medium;
filaments thickening toward apex, stamens otherwise as the genus;
style much longer than the often stiffly erect-puberulent, obovate,
apically notched ovary; capsule usually 4-5 mm. wide, obovate,
wider than long, strongly flattened, a little longer than the style,
stiffly, sparsely, often glandular-pubescent at the merely emarginate
apex; seed many, ca. 30-40, broadly ovate to suborbicular, ca. 1.0-
1.3 mm. long, sometimes minutely emarginate at one end, testa light
brown, almost smooth; placenta little thickened medianly, septum
chartaceous, soon deciduous.
Huanuco: Carpish Pass on highway Tingo Maria-Lima, 84 km.
W. of Tingo Maria, alt. ca. 2,750 m., Allard 21047; 15 mi. NE of
Huanuco, very wet, boggy slope, alt. ca. 12,000 ft., Macbride &
Featherstone 2195. — Junin: Prov. Huancayo, moist places near Huan-
cayo, alt. 3,300-3,500 m., Killip & Smith 23353.— Sin. Depto.: Tarn-
bo de Vaca, mossy rocky uplands, alt. ca. 13,000 ft., Macbride &
Featherstone 1769.
Closely related to Veronica serpyllifolia L. (see below) and per-
haps only varietally distinct.
FLORA OF PERU 715
Widely distributed in both continents and islands of the new
world as well as in Europe. Rarely encountered in east Asia.
Veronica peregrina subsp. xalapensis (HBK.) Pennell,1 Tor-
reya 19: 167. 1919.
V. xalapensis HBK., Nov. Gen. et Sp. 2: 389. 1818. V. sher-
woodii Peck, Torreya 28: 56. 1928.
Erect or rarely prostrate, annual herb, densely to sparsely glandu-
lar-pubescent2 on wood and fruit and sometimes on the leaves, inter-
nodes usually very short, shorter than the leaves. Leaves narrowly
lanceolate or lance-oblong, reduced to linear above, longest up to
3.5 cm., often only 1.5-2.0 cm. long, margin entire or few and dis-
tantly, shallowly toothed, especially on the upper half, acute at
apex, attenuate and decurrent over the very short petiole of the lower
leaves, upper leaves often sessile; flowers axillary, remote and soli-
tary below, inflorescences racemose or subspicate above; the pedicels
shorter than the flower and fruit, up to 2-3 mm. long, most often
shorter; calyx lobes lanceolate, equal, 3-4 mm. long, calyx tube lack-
ing; corolla white, rarely blue, shorter than to about equaling the
calyx in length, lobes ovate, equal or almost so, or the posterior a
little wider; stamens short, only about half the length of the petals,
otherwise as the genus; ovary obcordate, barely notched apically,
style minute, stigma capitate; capsule flattened, ca. 4-5 mm. wide,
wider than long, style hidden in the apical notch, lobes suberect to
divergent; seed numerous, thin ca. 0.7-0.9 mm. long, flattened, testa
smooth; placenta thin, septum deciduous.
Cuzco: Prov. Urubamba, Tun Capata, Santa Rita at alt. 2,800
m., C. Vargas C. 2692; Paucartambo to Tres Cruces, Cerro de Cusil-
luyoc at alt. 3,200-3,400 m., Pennell 14156; Huaracondo, alt. 3,300-
3,400 m., Pennell 13694. — Junin: Mito, eastern wood, alt. about
9,000 ft., Macbride & Featherstone 1720. Lima: Matucana about
8,000 ft., Macbride & Featherstone 288; San Buenaventura, alt. 2,700-
2,800 m., Pennell 14546. — Moquequa: Prov. Moquegua, swamp, alt.
3,100 m., Weberbauer 7480,—Pasco: Cerro de Pasco at alt. 14,000 ft.,
1 Originally published as V. peregrina xalapensis. Pennell (1935) stated that
the subspecies is the intended rank.
2 The variety may be separated from the typical on the basis of vesture.
V. peregrina subsp. xalapensis is the common subspecies of peregrina in Central
and South America.
Plants glabrous V. peregrina L. subsp. peregrina.
Plants densely or sparsely glandular-pubescent. . . . V. peregrina subsp. xalapensis.
716 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XIII
Macbride & Feather stone 2399. — Puno: Lake Titicaca, Capachira
Peninsula, Camjata Hacienda, alt. 12,600 ft., Tutin 1013. — Sin
Dept.: Valle de Pancar at alt. 3,400 m., Herrera 2327; Rio Blanco.
Moist place by deserted house, alt. about 15,000 ft., Macbride &
Feather stone 813.
Widely dispersed in the western hemisphere and generally culti-
vated over the rest of the world.
Veronica persica Poir., Encyc. Meth. 8: 542. 1808. V. tourne-
fortii Gmel, Fl. Ead. 1: 39. 1805.1 V. buxbaumii Ten., Fl. Nap. 1:
7,t.l. 1811. V. precox Raf., Atant. Journ. 79. 1832, non AIL, 1789.
V. diffusa Raf., N. Fl. N. Amer. 4: 38. 1838. V. byzantina (Sibth. &
Smith) BSP., Prelim. Cat. N. Y. PL 40. 1888.
Repent, pubescent herb, the internodes usually longer than the
firm leaves. Leaves petiolate, blades broadly ovate, obovate or
shield-shaped, in the wild state up to 2-3 cm. long and up to 2 cm.
wide, in gardens much larger, margins irregularly, strongly dentate
or lobed, when lobed the lobes acute, entire, blades usually acute at
both ends, varying to subobtuse at apex and obtuse at base, sparsely
pubescent on both surfaces; petioles pubescent below on the groove,
up to 1.5 cm. long, mostly 3-7 mm. long, often winged by the attenu-
ate leaf -base; flowers axillary, solitary, long-pedicellate; pedicels pu-
bescent, up to 3 cm. long, much exceeding the flowers and fruit,
filiform, little reflexed near the apex; calyx lobes united into a short
tube, ca. 1.0-1.5 mm. long, ovate and about 2 times as long as wide,
ciliolate, equal, 5-7 mm. long; corolla light blue to dark violet-blue,
petals broadly ovate, equal or almost so, mostly little shorter than
to equaling the sepals, entire; stamens little shorter than the petals,
anthers large for the genus, about a third to a fourth the length of
the filaments; ovary very broadly obovoid, apically slightly notched,
pubescent on the flattened edges, much shorter than the 2.5-3.5 mm.
style, stigma lobes united, punctiform; capsule wider than long, flat-
tened, lobes widely divergent, pubescent on the edges, style exceed-
ing the lobes; seeds ovoid, brown, narrowed to micropylar end;
septum soon deciduous, placenta thin.
Cuzco: Sacsahuaman above Cuzco, alt. 3,500 m., Pennell 13576;
Calca, Hacienda Urco, C. Vargas C. 3135; Prov. Cuzco, Sacsahua-
man, alt. 3,600 m., C. Vargas C. 15^3. — Huancavelica: Prov. Huan-
1 Epithet predated by V. tournefortii Vill. (1771 ) = V. officinalis L.
FLORA OF PERU 717
cavelica, W. of Huando, alt. 3,650 m., Hutchison 1675; Prov. Taya-
caja, Hills N. of Pampas, alt. 3,250 m., Stork & Horton 10234.—
Junin: La Merced, Hacienda Schunke, alt. ca. 4,000 ft., Macbride
5671. — Lima: Matucana, alt. ca. 8,000 ft., Macbride & Featherstone
125', Canta, open rocky, small canyon, alt. 3,000-3,200 m., Pennell
14598; Huaros, alt. 3,400-3,600 m., Pennell 14733.
An Eurasian species now well established in the western hemi-
sphere from Alaska to Chile, but not yet collected in Central America.
Veronica serpyllifolia L., Sp. PI. 12. 1753.
Repent herb with stems ascending at apices, stems and pedicels
pubescent with short, white, upcurving hair throughout. Lower
leaves ovate-oblong or oval, minute to 2 cm. long and little narrower,
obscurely crenate, obtuse, short-petiolate, the upper, bract-like leaves
narrowly ovate, sessile; flowers in racemes; pedicels up to 3-5 mm.
long; calyx lobes almost free, equal, ovate, ca. 2-3 mm. long; petals
about as long as the calyx, ovate, obtuse, white or whitish or light-
blue with blue or violet markings; stamens little shorter than the
petals; ovary obovoid, barely notched, shorter than the style, stigma
discoid-punctiform; capsule strongly flattened, glandular-pubescent
on the edges, 3-4 mm. wide, little wider than long and little longer
than the style, notched only about a fifth to a sixth its length; seeds
numerous, ovoid, about 1 mm. long, apiculate at micropylar end,
testa light, yellow-brown; septum chartaceous, soon deciduous, pla-
centa thin.
Cuzco: Chichacori Valley, alpine meadows, alt. 10,500 ft., Fisher
3: Lauramarca, alt. 3,800 m., Rauh-Hirsch Pi 129; Paucartambo,
Acanacu, alt. 3,600 m., C. Vargas C. 149; Quispicanchis, Marcapata,
environs of Huaillai, alt. 3,300-3,600 m., C. Vargas C. 9170.— Puno:
Prov. Sandia, Limbani, alt. 3,000 m., C. Vargas C. 1283 and at alt.
3,000-3,400 m., 9637.
HECKMAN
BINDERY INC.
OCT96
UNIVERSITY OF ILLINOIS-URBANfl