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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 2782 1 ; 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 10921 1 ;
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. Puno 3 : 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 red 1 ;
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 subshrub 2 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-
late 1 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 Trs 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 directe
