FB
FLORA OF PERU
BY
J. FRANCIS MACBRIDE
CURATOR, PERUVIAN BOTANY
BOTANICAL SERIES
FIELD MUSEUM OF NATURAL HISTORY
VOLUME XIII, PART V, NUMBER 2
DECEMBER 9, 1960
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FLORA OF PERU
BY
J. FRANCIS MACBRIDE
CURATOR, PERUVIAN BOTANY
BOTANICAL SERIES
FIELD MUSEUM OF NATURAL HISTORY
VOLUME XIII, PART V, NUMBER 2
DECEMBER 9, 1960
PUBLICATION 902
Library of Congress Catalog Card Number: 36-10426
PRINTED IN THE UNITED STATES OF AMERICA
BY CHICAGO NATURAL HISTORY MUSEUM PRESS
FLORA OF PERU
J. FRANCIS MACBRIDB
BORAGINACEAE [Juss.] Lindl.
Various in habit, ligneous or herbaceous, usually pubescent, not
infrequently harshly, the perfect normally 5-merous flowers ordinar-
ily in open and branched or scorpoid inflorescences, rarely solitary.
Calyx never entire, the lobes sometimes uneven. Corolla usually
regular, often plaited, pubescent or, in the throat, somewhat ap-
pendaged. Stamens on the tube and alternate with the lobes. Ovary
superior, bicarpellate, the commonly biovulate carpels free or united,
each remaining entire, parting along a medial false partition or more
or less lobed or separated along the medial line of the partition.
Fruit dry to fleshy in some degree, retaining the composition of the
carpels or, and generally, becoming 2-4 biovulate or uniovulate nut-
lets. Style obsolete to well-developed, simple, bilobed or twice
forked, variously situated, sometimes on an upwardly extended re-
ceptacle (gynobase) ; stigma entire or slightly bilobed.
The numerous papers of I. M. Johnston the outstanding stu-
dent of the family are cited with comment under the genera con-
cerned; the usefulness of this synopsis stems, of course, from his
realistic and thoughtful studies. The cytogenetic studies on the
family by Donald M. Britton, Brittonia 7: 233-266. 1951, will prob-
ably help to an understanding of relationship.
Forget-me-not is probably the best known plant in this family,
certainly one of the most beloved in gardens nearly everywhere.
Otherwise the group is more interesting to taxonomists than to horti-
culturists; however, Echium, Heliotropium and Cordia are often grown
for ornament while a few have popular medicinal repute, notably
borrago (Borago officinalis L.).
Regretfully I here record the passing of my never-to-be-forgotten
friend, Ivan Johnston.
Ovary slightly lobed if at all, sometimes 2-4-grooved laterally.
Style entire, stigma 1; inflorescences or its branches scorpoid or its
flowers solitary.
Clambering shrubs (or, as T. microcalyx, erect) ; fruit pulp aque-
ous, mesocarp fleshy, nutlets 2-4, bony. ... 1. Tournefortia,
539
540 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Herbs or suffrutescent; fruit dry, parting into 2-4 bony nutlets.
2. Heliotropium.
Style 2-parted or early coherent; much-branched prostrate herbs
or ligneous below 3. Coldenia.
Style twice forked, stigmas 4; inflorescence not scorpoid; fruit
drupaceous except C. alliodora.
Flowers sessile, not in foliose panicles; calyx not inflated.
4. Cordia.
Flowers pedicellate in foliose panicles; calyx soon dilated.
5. Saccettium.
Ovary deeply 4-parted, the fruit consisting of 4 (or fewer) nutlets.
Nutlets neither margined nor appendaged.
Corolla lobes contorted in bud; "no-me-olvides" ... 6. Myosotis.
Corolla lobes imbricate in bud.
Flowers ebracteate; leaves crowded toward stout perennial
base 7. Moritzia.
Flowers often bracted or plants annual.
Leaves all alternate; perennials 8. Lithospermum.
Leaves, at least lower, opposite or in a rosette; annuals or
biennials.
Corolla yellow, throat usually smooth .... 9. Amsinckia.
Corolla white at least in large part, throat appendaged.
Nutlets keeled ventrally (or also dorsally).
Leaves mostly in a basal rosette . . 10. Plagiobothrys.
Lower leaves opposite, even the earliest.
11. Allocarya.
Nutlets grooved ventrally, keeled obscurely if at all
dorsally 12. Cryptantha.
Nutlets alate-margined or dorsally appendaged.
Appendages apically barbed; leaves alternate. . . .13. Hackelia.
Appendages or teeth apically uncinate-setose; first leaves oppo-
site 14. Pectocarya.
1. TOURNEFORTIA L.
Reference: Johnston, Contr. Gray Herb. 92: 66-68. 1930.
Lianas or shrubby and then often lax or subscandent, the flowers
in scorpoid racemes or spikes disposed in dichotomous panicles.
FLORA OF PERU 541
Calyx persistent. Corolla limb spreading. Ovary 4-celled; style (or
stigma) terminal, simple, the stigma peltate or conic, usually bifid at
tip, often sessile. Fruit fleshy, obscurely if at all lobed or definitely
consisting of 1-2-seeded irregular nutlets or of 4 consimilar 1-seeded
nutlets; nutlets often with 1-2 empty cavities. Embryo straight
or curved.
The scholarly paper of Johnston, while entitled "Treatment of
Tournefortia," actually concerns only the species of the eastern half
of South America; he divided the genus into section Eutournefortia
Johnst., the obscurely or not lobed fruit bicarpellate, composed of
1-2-seeded uneven (often joined) nutlets, the embryo straight, and
section Cyphocyema Johnst., the evidently lobed fruit obscurely bi-
carpellate, consisting of 4 similar 1-seeded nutlets, the embryo curved
and the corolla lobes very narrow with involute margins. This clari-
fication of the genus is a fine contribution to its taxonomy; most
Peruvian species belong to the latter section (some are incompletely
known) ; at least T. bicolor Sw., T. cuspidata HBK., T. hirsutissima L.,
T. ovalifolia Rusby and T. Ulei Vaupel belong to the former. The
following key attempts to lead to herbarium materials often only
in flower; no doubt several names refer more correctly to variants.
Corolla segments more or less setiform, distinctly longer than wide,
often recurving at tip; calyx segments minute or narrow.
Corolla about 1 cm. long; leaves usually rounded at base; inflores-
cences ordinarily terminal T. maculata.
Corolla to about 5 mm. long; leaves somewhat acute at base; in-
florescences axillary, often also terminal.
Stems as leaves beneath strigose tomentose T. mapirensis.
Stems as leaves green or lightly appressed strigose.
Leaves merely acute at base . . . . T. breviflora, T. psilostachya.
Leaves decurrent at base T. glabra.
Corolla segments rounded to acute, sometimes narrowed above but
not distinctly longer than wide; calyx segments about 1.5 mm.
long or longer, or broad.
Leaves more or less bicolored, tomentose beneath, usually green
and sparsely (rarely densely) strigose above . . T. polystachya.
Leaves concolor, pubescence subequally developed both sides.
Corolla tube 2.5-3 mm. long; calyx minute; leaves oblong-
lanceolate to elliptic, rarely 3.5 cm. long. . . . T. microcalyx.
542 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Corolla tube about 4-12 mm. long; calyx and leaves mostly
larger.
Inflorescence lax, branchlets even in flower divaricate, some
soon several cm. long or if early suberect or short, corollas
about 1 cm. long.
Corollas about 5 mm. long, rather in corymbs; style well-
developed (unless T. glabra).
Flowers sessile; leaves ample, usually drying dark.
T. Ulei, T. glabra.
Flowers pedicellate; leaves drying brownish-green.
Calyx hispidulous; stems scabrous T. ovalifolia.
Calyx glabrate; stems scurfy T. chinchensis.
Corollas 10-12 mm. long, mostly on dichotomous branches;
style short.
Leaves ample, drying black, glabrous T angustiflora.
Leaves rather small, greenish-brown, puberulent.
T. tarmensis, T. Buchtienii.
Inflorescence dense, branchlets even in age rarely 3 cm. long
unless T. hirsutissima, T. undulata, especially if corolla
is long, this usually 5-7 mm. long.
Stems notably spreading hirsute, trichomes long; corolla
about 1 cm. long T. cuspidata.
Stems usually densely, leaves evenly more or less appressed
hispid; corolla densely strigose, shorter.
T. hirsutissima.
Stems glabrous or puberulent as leaves or these rather ob-
scurely pubescent on nerves and veins.
Upper stems and inflorescence closely reddish puberulent.
T. tarmensis, T. Buchtienii.
Upper stems glabrous or glabrate, indument if present
sparse.
Leaf nerves, at least mostly, about 1 cm. distant; peti-
oles about 1 cm. long.
Leaves about twice as long as wide.
Leaves membranous T. setacea, T. bicolor.
Leaves coriaceous T. coriacea.
Leaves about three times longer than wide.
T. longifolia.
FLORA OF PERU 543
Leaf nerves, at least mostly, about 5 mm. distant.
Leaves long-decurrent, about three times longer than
wide T. ternifolia.
Leaves short-decurrent, or subsessile, about two
times longer than wide . . T. virgata, T. undulata.
Tournefortia angustiflora R. & P. Fl. Peruv. 2: 25, pi 151a.
1799.
Spreading, branching, glabrous or nearly, the larger subherba-
ceous glabrous branches about 8 mm. in diameter with ovate-acumi-
nate leaves to 2 dm. long, 8 cm. wide, those of the flowering branchlets
half as large or smaller, narrowly acuminate, minutely hispidulous
both sides; inflorescence branches often arcuate-divaricate; calyx ses-
sile, scarcely 1 mm. long, lobes ovate, subacute; corolla greenish or
yellowish-white, sparsely and obscurely puberulent, the slender tube
1 cm. long, the lobes subrotund, about 5-6 mm. across expanded; fruit
white, succulent, glabrous, 5-6 mm. in diameter. F.M. Neg. 1038.
Hudnuco: Chicoplaya and Pueblo Nuevo, Ruiz & Pavdn, type.
Pozuzo, 4644- Prov. Huamalies, Weberbauer 8637 (det. Vaupel).
Loreto: Yurimaguas, Killip & Smith 28064 (det. Killip). Rio Acre:
Seringal Auristella, Ule 9708. Ecuador.
Tournefortia bicolor Swartz, Prodr. 40. 1788. T. laevigata Lam.
Tab. Encycl. 1: 416. 1791, fide Johnston, Journ. Arnold Arb. 16: 51.
1935.
To several meters tall, finally more or less scandent, its branchlets
glabrate or sparsely appressed pubescent with antrorse trichomes;
petioles 0.5-1.5 cm. long; leaves ovate to elliptic or somewhat lanceo-
late, obtuse-rounded at base, acute, 0.5-1.5 dm. long, 1.3-9 cm. wide,
slightly lustrous, dark green, often obscurely papillate and pubescent
with fragile appressed trichomes, these more abundant beneath; in-
florescence often ample, finally lax but the racemes early crowded,
1-4 cm. long; calyx sparsely strigose, usually sessile, lobes at anthe-
sis usually 1-2.5 mm. long, little accrescent; corolla strigose, tube
4-5 mm. long, limb 6 or 7 mm. across; fruit white, succulent, gla-
brous, elobate; stigmas subsessile. Calyx lobes subfiliform, some-
times 5 mm. long; pubescence nearly lacking or more abundant (var.
calycosa Bonn. Smith, Bot. Gaz. 14: 27. 1889). Common, especially
near Iquitos, only a few collections cited.
San Martin: San Roque, Williams 7783 (det. Killip). Junin:
La Merced, Killip & Smith 23515 (det. Killip). Loreto: Near
544 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Mazan, Mexia 6455 (det. Johnston); Jose Schunke 121; 270 (det.
Killip). Yurimaguas, Williams 4237, 4241 (det. Standley); King
2809 (det. Killip). Near Iquitos, King 735; Killip & Smith 27413;
Williams 1395; 3571. Yarina Cocha, Tessmann 5425. Florida,
Klug 1999. American tropics. "Loblobo" (Mexia).
Tournefortia breviflora DC. Prodr. 9: 520. 1845; 76. T.
xapuryensis Vaupel, Notizbl. Bot. Gart. Berlin: 186. 1914?
Resembles T. mapirensis but green, obscurely if at all strigose,
and the slender divaricately branched inflorescence usually entirely
axillary (or also terminal); stems as petioles somewhat puberulent,
the latter 1-1.5 cm. long; leaves oblong-elliptic, acute at base, rather
abruptly acuminate, to about 1 dm. long, 4 cm. wide, the slender
distant nerves prominent only beneath; inflorescence lax, 5-8 cm. long,
sometimes with a linear bract or two; calyx about 0.5 mm. long, the
puberulent-ciliate lobes lanceolate, acute; corolla 3-4 mm. long,
greenish-yellow, obscurely puberulent, the lobes subulate-subseti-
form (Vaupel). DeCandolle's plant, ex char., may be distinct; his
diagnosis differs in petioles 6 mm. long, leaves about 6 cm. long,
2 cm. wide, corolla 3 mm. long, the short lobes acute. However, the
Vaupel plant probably occurs in Peru like the related T. mapirensis
with in part similar range; both species seem to be scarcely separable
from T. psilostachya HBK. F.M. Neg. 1065.
Rio Acre: Seringa! San Francisco, Vie 9710 (type, T. xapuryensis).
Brazil; Paraguay.
Tournefortia Buchtienii Killip, Journ. Wash. Acad. Sci. 17:
334. 1927.
Branches subterete, ferrugineous-tomentose and scabroid; peti-
oles opposite, to 2 cm. long; leaves oblong or oblong-elliptic, rounded
or subacute at base, acuminate, 7-14 cm. long, 3-5 cm. wide, the 10
or so lateral nerves slightly impressed above, where dark green and
appressed strigillose except the nerves, rusty hirsutulous especially
beneath; peduncles 6-7 cm. long, subternately branched, the branch-
lets 2-3-dichotomous, the subterminal flowers 2.5 mm. distant; sepals
linear, 2-3 mm. long; corolla tube 7-10 mm. long, dilated just be-
low throat, rusty tomentose, the orbicular lobes minute; stamens
near throat, linear; style about 7 mm. long; fruit ovoid-conical,
7-8 mm. long, glabrescent, white. After author, who indicated no
relationship. Leaves of Weberbauer specimen are oblong-elliptic,
FLORA OF PERU 545
to 5 cm. long, 12 mm. wide, while Eyerdam 25123 from La Paz has
leaves to 1.5 dm. long, 5 cm. wide.
Junin: Comas, 3,300 meters, Weberbauer 6607 (det. Killip, 1937,
T. ovalifolia; 1925, T. chinchensis}. Bolivia.
Tournefortia chinchensis Killip, Journ. Wash. Acad. Sci. 17:
331. 1927.
Branches soon terete, scurfy, ferrugineous-hirsutulous; petioles
5-10 mm. long; leaves opposite, ovate-oblong to narrowly oblong,
3-7 cm. long, about 1-3 cm. wide, slightly narrowed at base, acute,
entire, the 6 or 7 nerves impressed above, prominent beneath, sub-
coriaceous, glabrous or slightly hispidulous above, glabrous beneath
except for the rusty hirsutulous nerves and veins; inflorescence 3-4-
dichotomous, the ultimate branches 2-3 cm. long in flower, the
flowers about 2 mm. apart; sepals linear, 2-3 mm. long, glabrous
or a little pilosulous; corolla tube 6-9 mm. long, rusty tomentose,
greenish, the minute orbicular but apiculate lobes glabrous; stamens
1-1.5 mm. long, style 5 mm. long; fruit depressed globose, about
8 mm. in diameter, white, glabrescent. After Killip, who contrasted
two similar species (T. andina Britton, T. ovalifolia Rusby) as follows:
Flowers sessile; corolla tube appressed hirsute; branchlets smooth.
T. andina.
Flowers pedicellate; corolla tube tomentose; branchlets rough.
Calyx lobes 2 mm. long or less; leaves alternate T. ovalifolia.
Calyx lobes longer than 2 mm.; leaves subopposite . T. chinchensis.
However, Johnston included the shrub in T. ovalifolia Rusby.
Ecuadorian specimens by Steyermark, apparently the same, have
leaves 8 cm. long, 3.5 cm. wide.
Huanuco: Hacienda Villcabamba, Rio Chinchao, 5142 (type,
T. chinchensis') . Bolivia, Ecuador.
Tournefortia coriacea Vaup. Bot. Jahrb. 54, Beibl. 119: 3. 1916.
Shrubby, the stout branches smooth; petioles 2 cm. long; leaves
oval or broadly elliptic, short-acuminate, ample, coriaceous, glabrous;
inflorescence small with few branches; flowers about 5 mm. long; calyx
lobes narrowly lanceolate; corolla tube about two times longer than
the calyx, sparsely pilosulous with minute appressed trichomes, the
lobes obtuse; anthers affixed at base of tube, narrowly lanceolate;
pistil shorter than calyx, stigma conical. After the author, who
546 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
merely remarked : at once recognizable by the large leathery leaves.
The Williams specimen has petioles 1 cm. long, leaves to 12 cm.
long, 8 cm. wide, nerves 1 cm. apart, lightly impressed above;
calyx lobes subulate, 2 mm. long, corolla puberulent, tube 7 mm.
long, lobes rounded, 1 mm. long. F.M. Neg. 1042.
Loreto: Yurimaguas, Williams 78S9 (det. Killip). Ecuador.
Tournefortia cuspidata HBK. Nov. Gen. & Sp. 3: 83. 1818.
T. obscura A. DC. Prodr. 9: 517. 1845, fide Johnston, Journ. Arnold
Arb. 16: 54. 1935.
Shrub or liana, the branchlets short-pubescent and inordinately
villous-hirsute with slender spreading brown trichomes some 4 mm.
long; petioles to about 1 cm. long or somewhat longer; leaves lance-
olate to subovate, rounded or obtuse at base, acuminate, 7-15 cm.
long, 3-6 cm. wide, closely appressed strigose; spikes approximate
at ends of elongate branches, to 3 cm. long; calyx lobes subulate-
linear, 7-9 mm. long, little longer in fruit, sparsely long-pubescent
and short-strigose; corolla tube 5-8 mm. long, densely strigose,
limb 4-6 mm. across; fruit white, fleshy, somewhat verruculose;
stigma sessile. Common on the lower Rio Huallaga, only a few col-
lections cited.
San Martin : Juanjui, Klug S776. Loreto: Yurimaguas, Williams
3845; 4312; 4776; 4972; 7870. Yarina, Tessmann 3773. Near Iquitos,
Klug 432. Bolivia to Central America and the West Indies.
Tournefortia glabra L. Sp. PI. 141. 1753.
Large scandent or erect shrub or small tree; branchlets puberulent
or glabrate; leaves lanceolate-elliptic to ovate, 0.5-1.5 dm. long,
narrowed to both ends and often long-decurrent on the slender
petioles, membranous, early pilosulous, at maturity nearly glabrous;
spikes frequently many, slender, sometimes greatly elongate, the
inflorescence lax; calyx lobes lanceolate, about 2 mm. long, long-
acuminate as the lobes of the pale green or whitish corolla, its tube
4 mm. long; fruits white, succulent, to about 5 mm. thick, 9 mm.
long, not lobed. Wood bright yellow when cut.
San Martin: Pongo de Cainarachi, Klug 2730 (det. Killip). To
Mexico; West Indies.
Tournefortia hirsutissima L. Sp. PI. 140. 1753. Messer-
schmidtia hirsutissima (L.) Roem. & Schult. Syst. 4: 541. 1819.
FLORA OF PERU 547
Branches subtetragonous, typically densely (not in Peru as deter-
mined) short-hirsute as the petioles and peduncles; petioles stout,
6-9 mm. long or much longer; leaves ovate-oblong or -elliptic, usually
acuminate, rounded to narrowed at base, mostly 6-9 cm. long, about
5 cm. wide to twice as large, membranous, appressed strigose beneath,
scabrous or more sparsely hispid above; inflorescence usually ample,
corymbose or dichotomously divided; spikes dense, narrow, in fruit
3-4 cm. long; calyx hirsute or hispidulous, the calyx teeth linear to
ovate-oblong, acute or acuminate; anthers linear, at middle of tube;
corolla white, at least the tube sericeous, 4-6 mm. long, the lobes
acute or apiculate; style very stout, stigma depressed-conical; fruit
globose, white, hispidulous, about 5 mm. in diameter; nutlets 4,
ovate-oblong. Shrubby or a liana to several meters long; the indu-
ment is sometimes so short that it is inconspicuous; Killip determina-
tions.
San Martin : Juan jui, King 4321 . Tarapoto, Williams 5781 ; 5690;
6629. Huanuco: Puente Durand, 1,900 meters, Weberbauer 9590.
Widely distributed in tropical America.
Tournefortia longifolia R. & P. Fl. Peruv. 2: 25, pi. 150b. 1799.
Glabrous or nearly, erect, the branches sulcate-angled; petioles
about 1 cm. long; leaves oblong-lanceolate or -elliptic, acute both
ends or acuminate, to 2 dm. long, 6-7 cm. wide, submembranous
but somewhat rugose-bullate above in age, much veined; inflores-
cence dichotomous but small, few-flowered; seeds trigonous. Type
in fruit, imperfectly known. F.M. Negs. 12943; 29254.
Huanuco: Pozuzo, Ruiz & Pavdn, type.
Tournefortia macula ta Jacq. Enum. 14. 1760; Sel. Stirp. 47.
1763. T. syringaefolia Vahl, Symb. 3: 23. 1794. T. peruviana Poir.
Encycl. Suppl. 4: 425. 1816. T. volubilis L. as to R. & P. Fl. Peruv.
2: 24, pi. 148b. 1799, fide Urban and Johnston. T. paniculata Cham.
Linnaea 4: 468. 1829?
Shrubby or sometimes more or less clambering or climbing, dark
green and inconspicuously short-strigose or leaves softly pilosulous
beneath, including the lax inflorescence and growing parts; petioles
about 1-1.5 cm. long, leaves ovate or somewhat elliptic-lanceolate,
acute to rounded at base, acuminate, mostly 5-10 cm. long, 3-5 cm.
wide, ordinarily densely and finely tuberculate; calyx 1-1.5 mm. long,
scarcely accrescent but pedicels finally 1-5 mm. long and thickening;
corolla greenish-white, tube 3-4 (-8) mm. long, limb 3-5 mm. across;
548 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
fruit yellow, often blotched with black, the 4 finally separate nutlets
rounded dorsally, angled ventrally. Johnston, Journ. Arnold Arb.
30: 130. 1949, identified the plant of Jacquin. F.M. Neg. 1054
(T. volubilis).
San Martin: Tarapoto, Williams 5414; 6170 (det. Killip). Zepe-
lacio, Klug 3348. Huanuco: Pozuzo, Ruiz & Pav6n (type, T. volu-
bilis). Junin: La Merced, 5473. Loreto: Yurimaguas, Williams
4778; 4971; 4973; 5121. Nanta, Raimondi 9427Cuzco: Valle de
Santa Ana, Herrera 2632. Puno: Gay (as T. voluUlis). To Central
America and the West Indies; Brazil.
Tournefortia mapirensis Lingelsheim, Repert. Nov. Sp. 7: 244.
1909; 76. T. subrotunda Rusby, Bull. N. Y. Bot. Card. 8: 115. 1912,
fide Johnston.
Fulvous strigose-tomentose liana (except leaves above) with axil-
lary (and terminal) leafless lax inflorescences; petioles 1-2 cm. long;
leaves broadly ovate, subcordate or rounded at base, shortly acum-
inate, (4) 8-12 cm. long, (2.5) 4.5-8 cm. wide, dark green but lightly
pubescent above, paler and densely strigose-tomentose beneath, the
nerves 5 or 6 pairs; calyx 2 mm. long, accrescent in fruit; corolla
greenish-yellow, scented, the tube very narrow, pubescent, 4 mm.
long, the subsetiform acute segments 1 mm. long; fruits early yellow,
subglobose, 4 mm. thick, finally purple to black-brown. Mostly
after author; very distinct by its tawny pubescence, definitely axil-
lary inflorescences, the flowering branchlets leafless or the few leaves
much reduced, according to Johnston, who determined the Peruvian
specimen. Inflorescences 4-6 cm. broad, branches 1.5-2.5 cm. long
(Rusby).
Loreto: Yurimaguas, (Spruce 3889). Rio Acre: Seringal San
Francisco, Ule 9712; 9713. Bolivia to Amazonian Brazil.
Tournefortia microcalyx (R. & P.) Johnst. Journ. Arnold Arb.
37: 295. 1956. Heliotropium microcalyx R. & P. Fl. Peruv. 2: 3,
pi. 109b. 1799. Tournefortia hispidula R. & P., in herb.
Open, often stockily branched, a meter or two tall, glabrous or
essentially or the oblong-lanceolate leaves beneath and almost tiny
flowers minutely hispidulous; leaves oblong-lanceolate, attenuate to
the slender petioles, these 5-8 mm. long, moderately acuminate,
5-10 cm. long, 1-3.5 cm. wide, membranous, obscurely venose above,
lightly reticulate-veined beneath; inflorescence axillary, usually
about 3-5 mm. in overall size, the branchlets often arcuate-spreading;
FLORA OF PERU 549
calyx segments sublinear, scarcely 1.5 mm. long, not accrescent in
fruit, usually glabrate as the nearly twice as long corolla, this with
rounded lobes, expanded about 2.5 mm. across, white; fruit composed
of 4 similar nutlets. Shrub but open and with short or longer and
floppy branchlets, white translucent fruit. Determinations by Killip.
F.M. Negs. 1048 (T. hispidula) ; 1051.
Piura: Cerro Viento, Haught 95. Ancash: Tambo de Pariocota,
2550. Lima : Pachacamac, Mexia 8096; 83S3. Lurin, Pennell 1 2205.
Atocongo, Pennell 14758. Prov. Huarochiri, Weberbauer 5250. Near
Lima, Nee; Gaudichaud. Chosica, Grant 7401. Huanuco: Chico-
playa, Isern 2080. Chinchao, Ruiz & Pavdn (type, T. hispidula).
Hills of Huanuco, Ruiz & Pavdn, type.
Tournefortia ovalifolia Rusby, Bull. N. Y. Bot. Gard. 4: 414.
1907.
Branchlets sparsely scabrous; petioles about 1 cm. long, broadly
sulcate; leaves oval, slightly decurrent at base, abruptly and acutely
cuspidate, 6-10 cm. long, 3-6 cm. wide, dark green, the 10-12 pairs
of lateral nerves prominent only beneath where yellowish or purplish;
peduncles 2-3 cm. long, the branches of the dichotomous inflorescence
elongating, flowers 2-3 mm. distant; pedicels 2-3 mm. long; calyx
2 mm. long, truncate at base, the lanceolate lobes erect; corolla tube
5 mm. long, lobes rounded; style 3 mm. long, stigma 1 mm.; fruit
globose-ovoid. Flowers green. Perhaps should be interpreted to in-
clude T. chinchensis Killip.
Peru (cf. note above). Bolivia.
Tournefortia polystachya R. & P. Fl. Peruv. 2: 24, pi. 149,
fig. a. 1799. Heliotr opium oppositifolium R. & P. I.e. 2: pi. 108, fig. b.
T. loxensis HBK. Nov. Gen. & Sp. 3: 81. 1818, at least as to Peru.
An open shrub or tree-like in age, the flowering branchlets puber-
ulent, the ample open terminal inflorescences as the leaves beneath
fulvous-canescent villous; petioles about a fourth as long as leaves,
these, at least the lower, somewhat obliquely rounded to subcordate
at base, shortly acuminate, commonly ovate or the upper sometimes
somewhat elliptic and narrowed to base, those of the flowering
branches 6-15 cm. long, usually about 4-6 cm. wide, but attaining
2.5 by 2 dm., slightly scabrous, glabrate and finally bullate above;
calyx segments sublinear, 2 mm. long, not accrescent; corolla tube
about 4-5 mm. long, lobes rotund; fruit with white translucent pulp
at least 1 cm. in diameter. The flowers, at first greenish-yellowish,
550 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
are white when fully expanded and 5 mm. across, either with a dis-
agreeable odor (2404) or fragrant (4017, 4188; and Stork & Horton);
unnoted determinations by Killip. Type of H. oppositifolium R. & P.
had hispid stems and leaves asperous above but seems to belong here.
F.M. Negs. 1056; 27094 (Mathews); 1052 (T. oppositifolia).
Cajamarca: Below Ocros, Weberbauer 2716; 271 6a. Amazonas:
Chachapoyas, Mathews. La Libertad: Cachicadan, Stork & Horton
9953 (det. Standley, Heliotropium arborescens) . Lima: Ambar, Stork
1 1 433. Huanuco : Chaglla, Weberbauer 6706. Huanuco, 2404; 3098;
Sawada 82. Mufia, 4017; 4188; Ruiz & Pavon (type, H. oppositi-
folium). Mito, 3377. Piedras Grandes, Woytkowski 127. Junfn:
Near Tarma, Huassa-huassi, Ruiz & Pavon, type. Cuzco: Marca-
pata Valley, Weberbauer 7877 (det. Standley). To Colombia. "An-
tagra" (Ruiz & Pavon).
Tournefortia psilostachya HBK. Nov. Gen. & Sp. 3: 78. 1818.
T. floribunda HBK. I.e. 79, at least as to Peru. T. cirrhosa Vaupel,
Bot. Jahrb. 54, Beibl. 119: 3. 1916. T. volubilis L. Sp. PI. 140. 1753,
at least in lit.
Variable in leaf size, form and indument but as interpreted usually
more or less appressed strigillose, especially the younger branchlets
and leaves beneath; petioles a few mm. to 1 cm. long; leaves rather
oblong or narrowly ovate-lanceolate to broadly ovate, obliquely
rounded at acute base, acuminate, sometimes abruptly, usually
5-8 cm. long, about 1.5-2.5 cm. wide or the lower larger; peduncles
in type terminal, the filiform spikes diffuse; calyx strigose, the
lanceolate acuminate segments spreading; corolla long-exserted, seri-
ceous, the throat inflated, the lobes subulate; anthers subsessile near
top of tube, this as long as the filiform style; stigma small, subconic.
T. floribunda HBK. ex char., type from Colombia, has leaves obtuse
or rounded at base, somewhat canescent sericeous beneath, corolla
tube about 3 mm. long, the lobes "short"; Johnston, Contr. Gray
Herb. 92: 88. 1930, suggested that it once may have had a con-
tinuous range to eastern Brazil; however, the single Peruvian collec-
tion referred to it is more probably a form of T. psilostachya, which
name, long in use, is here retained in preference to the perhaps not
identical one of Linnaeus. Finally, Vaupel's plant, type from Ecua-
dor, has ovate leaves 4-5 cm. wide and may be distinguishable; cf.
T. breviflora. Determinations by Killip. F.M. Negs. 38975; 1043
(T. cirrhosa).
FLORA OF PERU 551
Loreto: Mishuyacu, near Iquitos, Klug 1113; 1213. Nancho,
(Raimondi 3414, det. Herb. Dahlem, T. floribunda). Puno: Gay.
Ecuador; northern South America?
Tournefortia setacea Killip, Journ. Wash. Acad. Sci. 17: 332.
1927.
A bushy-branched liana, the younger branches, leaves above and
beneath on the nerves appressed strigillose; petioles 0.5-2 cm. long;
leaves subopposite or in threes, oblanceolate or ovate, 5-14 cm. long,
3-7.5 cm. wide, subdecurrent, acuminate, entire, membranous, dark
green above, paler beneath; inflorescence to about 7 cm. wide, few-
branched, shortly peduncled; flowers sessile; sepals linear-setaceous,
4-5 mm. long, about as long as the strigillose corolla tube, the sub-
orbicular corolla-lobes 1.5 mm. long; stamens near base of tube,
anthers linear, 2.5 mm. long; fruit conical, 3-4 mm. long, strigillose,
white. With the variation that Johnston assigned to T. bicolor Sw.
to which Killip originally referred it; this may not be distinct, but
the author wrote: "the long setaceous sepals distinguish it."
San Martin: Sinchona, Ferreyra 1127. Huanuco: Cushi, 4829.
Rio Pozuzo, Weberbauer 6737. Junin: La Merced, 5579, type.
Colonia Perene", Killip & Smith 24993. Ayacucho: Aina, Killip &
Smith 22733. Estrella near Huanta, Killip & Smith 22668. Cuzco:
Cerro Mascac, Valle de Lares, (Herrera 1779). Bolivia.
Tournefortia tarmensis (Krause) Macbr., comb. nov. Helio-
tropium tarmense Krause, Bot. Jahrb. 37: 632. 1906.
Liana or the branches clambering, these glabrous in age, early
quadrate, sparsely as leaves both sides, reddish-pilosulous; petioles
4-6 mm. long; leaves oblong- or ovate-elliptic, shortly or subacute
both ends, 3-6 cm. long, 1-3 cm. wide, often somewhat undulate,
nearly smooth; spikes at first corymbosely congested, rather lax in
full flower, 2-5 cm. long; flower limb 7 mm. long; calyx lobes linear,
hispidulous, scarcely 2 mm. long; corolla tube greenish, puberulent
only without, the crisped spreading lobes white; anthers linear, acute,
subsessile; style elongate. Distinguished by Krause from T. micro-
calyx (R. & P.) Johnst. especially by the much longer flowers. F.M.
Neg. 1062.
Junin: Comas, Weberbauer 6607, in part? (det. Killip, T. ovali-
folia). Palca, Prov. Tarma, Weberbauer 1750, type. Carpapata,
Prov. Tarma, Soukup 3440. Bolivia.
552 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Tournefortia ternifolia HBK. Nov. Gen. & Sp. 3: 84. 1818.
Branches terete, sparsely appressed pubescent; petioles 1.5-2 cm.
long; leaves ternate, oblong, narrowed to both ends, acute, about
1.5 dm. long, half as wide, membranous, reticulate-veined, glabrous
or appressed pubescent on nerves beneath; spikes terminal, geminate,
peduncled, less than 2.5 cm. long; flowers pedicellate, secund, ebracte-
ate; calyx strigose, teeth linear-lanceolate; corolla strigose, thrice as
long as calyx (after authors). Compare T. longifolia R. & P. to
which the Killip and Smith collection, young, may belong, or pos-
sibly there is only one species.
Cajamarca: Tomependa, Bonpland, type. Amazonas: Chacha-
poyas, Mathews 3009 (det. Killip). Junin: Tarma, Kittip & Smith
21920 (det. Killip).
Tournefortia Ulei Vaupel, Notizbl. Bot. Gart. Berlin 6: 186.
1914.
Scandent, the younger branchlets sparsely puberulent; petioles
1 cm. long or longer; leaves broadly elliptic or ovate, shortly acumi-
nate, to 13 cm. long, 7 cm. wide, glabrous; inflorescence to 2.5 dm.
long, lax, mostly trichotomous with divaricate branches, with sessile
leaves above the first branches; flowers subsessile, crowded, ap-
pressed puberulent; calyx lobes narrowly triangular, 1-2 mm. long,
3-4 times shorter than corolla tube, this 6 mm. long, slightly inflated,
densely yellowish-brown pubescent; flowers yellow-green, about
7 mm. long; anthers below throat; style nearly as long as tube,
stigma conical; fruit glabrous, ovoid, often crowned with pistil (after
author). T. andina Britton, Bull. Torrey Club 26: 148. 1899, of
Bolivia with sessile flowers may be comparable. The Peruvian
specimen has glabrate flowers and no leaves in the inflorescence;
compare T. glabra L.
Loreto: Florida, mouth of Rio Zubineta, King 2083 (det. Stand-
ley). Amazon headwaters (Rio Madeira).
Tournefortia undulata R. & P. Fl. Peruv. 2: 25, pi. 149b. 1799.
Rounded shrub or tree-like, to 3 meters tall, the branchlet tips
and young inflorescences reddish pilose, soon glabrous or nearly;
flowering branches densely leafy, stout; leaves broadest at or slightly
above the middle, attenuate into the short sulcate petioles, rather
abruptly acuminate, usually 8-15 cm. long, 3-6 (8) cm. wide, coria-
ceous, the many arcuate-ascending lateral nerves rather obscure
above, prominent with veins beneath, mostly 5-7 mm. distant,
FLORA OF PERU 553
rugose-bullate in age above; inflorescences and branches densely
flowered, suberect, tardily spreading, to 7 cm. long or usually shorter;
calyx segments subulate, about 3 mm. long, to 4 mm. long in fruit;
corolla tube puberulent, 4 mm. long, lobes subrotund, about 1 mm.
long; fruit not lobed, white, about 5 mm. in diameter, style appar-
ently obsolete. Killip & Smith 21525 from the botanical garden at
Lima was about 3 meters high in 1929. F.M. Neg. 1064.
Huanuco: Mito, 1998. Pillao, Ruiz & Pavdn, type. Arequipa:
Near Chala, Worth & Morrison 15685. Atiquipa, 1,050 meters,
Tqfalla, type; Weberbauer 7192 (det. Killip). Puno: Cerros de Islay,
Isern 2476. Ecuador?
Tournefortia virgata R. & P. Fl. Peruv. 2: 25, pi. 150. 1799.
Similar to T. undulata and apparently not well marked, but juve-
nile pubescence more persisting, canescent; leaves smaller, subcoria-
ceous, the fewer lateral nerves less regularly developed; inflorescence
branches recurving and incurving; calyx lobes about linear, scarcely
2 mm. long, little longer in fruit; corolla tube scarcely 4 mm. long,
rather stout, canescent puberulent, the undulate-crenulate lobes
nearly 1.5 mm. broad; fruit white, not lobed, about 4 mm. in diam-
eter, style obsolete. Description from my collection, the white
flowers (early yellowish) with a heavy, rather unpleasant fragrance;
the many leaves, as in the similar species, are strongly waved or
undulate in life so that they often crease in pressing. The Stork and
Horton specimen in flower is doubtful; corollas about 1 cm. long,
calyx scarcely 2 mm. long.
Huanuco: Mito, 1575 (det. Killip). Junin: Huassa-haussi, Ruiz &
Pavdn, type. Apurimac: Pincos, 2,700 meters, Stork & Horton 10695?
2. HELIOTROPIUM L.
Reference: Johnston, Contr. Gray Herb. 81: 1-73. 1928.
Shrubs, rarely herbs, with alternate leaves and white flowers often
borne in scorpoid racemes or spikes. Ovary undivided, the fruit con-
sisting of 4 nutlets or carpels, these remaining entire, biovulate when
ripe, or breaking into 2 uniovulate nutlets at maturity. Corolla
sinuses more or less plaited in bud. Anthers obtuse or acuminate,
often more or less coherent. Johnston's excellent revision of the
South American species has supplied the following sectional key and
the species key in part, except that vegetative characters have been
emphasized, for convenience. Nidia Gangui has presented an attrac-
554 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
tive account with illustrations of the genus for the species of Argen-
tina, Univ. Nat. Cordova 11: 481-559. 1955.
SECTIONAL KEY TO HELIOTROPIUM (after Johnston)
Carpels not cleft, finally separate biovulate nutlets.
Carpels with several empty cells besides the 2 fertile; corolla gla-
brous within section Tiaridium (Lehm.) Griseb.
Carpels with 2 cells, both usually fertile.
Corolla villous within; fruit adorned vesicularly.
section Schobera (Scop.) Johnst.
Corolla glabrous at least within; fruit (as carpels) smooth.
Plant prostrate-ascending; carpels often 1-seeded.
section Hypsogenia Johnst.
Erect shrubs; carpels normally 2-seeded.
section Cochranea (Miers) Reiche.
Carpels cleft, finally forming 2 uniovulate nutlets.
Tall shrubs; nutlets rough; anthers ciliate toward tip.
section Heliothamnus Johnst.
Herbs or usually less than 5 dm. tall; nutlets smooth, glabrous or
pubescent; anthers, if pubescent, only at tip.
Stigmatic disk thick, narrow on more or less developed style;
usually erect herbs or shrubs. . .section Orthostachys R. Br.
Stigmatic disk very broad, rather thin, sessile; prostrate, succu-
lent herbs section Halmyrophila Johnst.
SPECIES KEY TO HELIOTROPIUM
Prostrate herbs; leaves small, rarely 2 cm. long, or narrow, succulent.
Succulent leaves soon exceeded by spikes (section Halmyrophila).
H. curassavicum.
Herbaceous leaves longer than inflorescences (section Hypsogenia).
H. microstachyum.
Erect or suberect, usually more or less ligneous; leaves never succu-
lent, usually longer than 2 cm., always exceeded by spikes.
Herbs to ligneous, low; leaves mostly or all about 3, usually several
times longer than wide; nutlets smooth; all section Ortho-
stachys except H. Krauseanum (section Cochranea), perhaps
H. lanceolatum (section Heliothamnus?).
FLORA OF PERU 555
Leaves crenulate, pale puberulent beneath, obtuse; carpels not
cleft H. Krauseanum.
Leaves entire, subconcolor, more or less acute; carpels cleft.
Flowers clearly in bractless spikes.
Corolla rather showy, 4-5 mm. across or tube well exserted;
anthers linear, apically pubescent or glandular except
H. procumbens.
Corolla lobes rounded, tube and calyx subequal.
Plant canescent as calyx within H. Lobbii.
Plant green; corolla glabrous within H. toratense.
Corolla lobes more or less acute, the long tube prominent.
Anthers glandular-puberulent; corolla lobes acute.
H. oxylobum.
Anthers crested with antrorse trichomes.
H. lanceolatum.
Corolla small, 1-3 mm. wide, tube little exserted.
Shrub; leaves subcoriaceous; calyx segments suberect;
flowers yellowish, glabrous H. polyanthellum.
Herbs or half-shrubs; leaves herbaceous unless oldest;
calyx segments soon lax; flowers white.
Anthers (as corolla within) pubescent, linear, obtuse;
leaves slightly narrowed apically, acute.
H. pilosum.
Anthers glabrous, subovate, connective extended;
leaves, at least mostly, rounded apically, mucron-
ulate H. procumbens.
Flowers scattered on unevenly foliose branchlets . .H. piurense.
Tall and (or) ligneous; leaves mostly or all about twice longer than
wide except the subovoid H. rufipilum, sometimes H. angio-
spermum and the canescent shrubs H. erianthum, H. lippi-
oides; nutlets rough, lacunose or sculptured except the only
herbaceous species, H. indicum.
Herb, sparsely setulose-hispid; leaves decurrent on long petioles;
fruit lobed, nutlets smooth (section Tiaridium).
H. indicum.
Shrubs or ligneous below; indument, at least in part, pilose or
crinkled; petioles short; nutlets rough (section Heliothamnus
except H. angiospermum) .
556 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Carpels rough with obtuse vesicles, not cleft; leaves membra-
nous, sparsely pilose as stems or glabrate; corolla villous
within, small, tube and calyx subequal (section Schobera) .
H. angiospermum.
Carpels usually lacunose or sculptured, cleft; characters other
than above at least in part (section Helioihamnus).
Leaves membranous, concolored, not rugose, often shorter
than 5 cm.; corolla tube well exserted from the 1-
1.5 mm. long calyx H. adenogynum.
Leaves subcoriaceous to membranous, rugose-bullate espe-
cially when shorter than 5 cm.
Leaves not or obscurely bullate but veins more or less
obvious; corolla tube exserted.
Inflorescence lax, the branches soon several cm. long;
style shorter than stigma H . rufipilum.
Inflorescence congested, branchlets nearly 2-3 cm.
long; style longer than stigma. . . .H. arborescens.
Leaves usually firm and more or less bullate or (and)
gray pubescent beneath; corolla tube and calyx sub-
equal or leaf-character pronounced.
Indument, especially flowering branchlets, somewhat
fulvous; style shorter than stigma.
Calyx to 2 mm. long, longer in fruit.
Leaves 3-6 cm. long, oblong- or rarely subovate-
elliptic, acute or subacute . . . H. Urbanianum.
Leaves ample, mostly ovate-acuminate, often
about a dm. long or longer H. submolle.
Calyx inconspicuous, at most 1.5 mm. long in flower,
little longer in fruit H. Mandonii.
Indument, even of branchlets, canescent; style longer
than stigma.
Leaves lanceolate, 4-5 cm. long, 10-17 mm. wide.
H. erianthum.
Leaves ovate or ovate-elliptic, about twice as long
as wide H. incanum.
Leaves elliptic, 1.5-2 cm. long H. lippioides.
Heliotropium adenogynum Johnst. Contr. Gray Herb. 81 : 42.
1928.
FLORA OF PERU 557
Laxly branched, more or less villous and usually glanduliferous,
even the calyces, these globose, finally 2 mm. long, the elliptic-lanceo-
late erect or connivent lobes somewhat persisting; petioles 3-8 mm.
long; leaves herbaceous, venose, ovate or oblong-lanceolate, 2-5 cm.
long, acute or obtuse at base, acute at apex, 2-5 cm. long, about
0.5-3 cm. wide, paler beneath, entire; spikes 4-10 cm. long; corymb
lax; corolla sparsely villous only without, not rarely glanduliferous,
2-3 times longer than calyx, to 4 mm. across, lobes orbicular; anther
cells pubescent toward tip; ovary glanduliferous; style obsolete,
stigma about 0.8 mm. long; nutlets ovate, rugose, 1-1.5 mm. long,
stipitate-glandular. Remarkably distinct by small flowers, gland-
ular ovary and fruit, scarcely accrescent more or less deciduous
calyces; most closely related to H. Mandonii Johnst. and evidently
also to H. rufipilum (Benth.) Johnst. (author), a species varying in
some if not all of these characters. Section Heliothamnus Johnst.
Lima: Chosica, 495, type; Weberbauer 5312; (Wilkes Exped.}.
Heliotropium angiospermum Murray, Prodr. Stirp. Gott. 217.
1770; 10. H. parviflorum L. Mant. 2: 201. 1771. H. synzystachyum
R. & P. Fl. Peruv. 2: 3, pi. 109a. 1799. Tournefortia synzystachya
(R. & P.) R. & P. Syst. 4: 539. 1819. Synzystachyum peruvianum Raf.
Sylva Tell. 89. 1838. H. simplex Meyen, Reise 1: 436. 1834, fide
Johnston, Journ. Arnold Arb. 18: 18. 1937.
Perennial, sometimes flowering as an annual, rather sparsely
pubescent, erect or becoming lax or prostrate, sometimes a meter
long; petioles often to 1 cm. long or longer; leaves entire, oblong-
lanceolate to elliptic, attenuate to base, acute, small or large (2-3 cm.
long, 1-1.5 cm. wide, or 1.5 dm. long, 5 cm. wide); spikes solitary or
binate, 5-10 (15) cm. long, ebracteate; calyx lobes acute; corolla
white, glabrous but barbate within, about 2 mm. wide, the tube
subincluded in the calyx; fruit depressed, 3-4 mm. across. Section
Schobera (Scop.) Johnst. I.e. 10. Fruit didymous (2-seeded carpels)
and covered with unique vesicular structures.
Piura: Rocky slopes, Cerro Prieto, (Haught & Svenson 11602}.
Talara, Haught 38; Norton 11592. La Libertad: Barranco near
Pacasmayo, (Forbes). San Martin: Tarapoto, Woytkowski 35200.
Lima: Chosica, Mexia 04087. Atocongo, Pennell 14458a. Rio Chil-
lon, Pennell 14458. Lurin, Pennell 12217. Santa Clara, Rose 18626.
Callao to Lima, 52; Watkins 396 (and many others). Chancay,
Vargas 1235; Goodspeed 33033; Ruiz & Pavdn (type, H. synzystachy-
um). Huancavelica: Near Pampano, Weberbauer 5394 (part). lea:
558 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Above Pisco, 1,550 meters, Weberbauer 539 4. Apurimac: Rio Pacha-
chaca, Goodspeed Exped. 10537. Arequipa: Near Arequipa, Meyen.
Murillo Bay, (Macrae). Tacna: Arica, (Meyen, type, H. simplex).
Bolivia to Central America, West Indies, Texas, Florida. "Alacran."
Heliotropium arborescens L. Syst. ed. 10. 913. 1759; 39.
H. peruvianum L. Sp. PI. ed. 2. 187. 1762. H. corymbosum R. & P.
Fl. Peruv. 2: 2, pi. 107a. 1799.
Shrub, sometimes 2 meters tall, usually much lower, with many
short branchlets with lavender-purple flowers; leaves typically ovate-
or oblong-elliptic, more or less decurrent on the short or rarely 1-
2 cm. long petioles, acute, ordinarily 4-8 cm. long, about half as wide
or sometimes considerably larger, membranous or nearly, paler and
venose beneath, more or less pubescent or glabrate above when
smooth or inconspicuously bullate-rugulose in age; inflorescences 3-
10 cm. across, lightly to densely strigose or villous, the short branch-
lets congested even in fruit; calyx segments subulate-linear, about
3-3.5 mm. long, not accrescent, the corolla tube about twice as long,
appressed strigulose, lobes glabrate, rounded, 4-5 mm. across at top;
anthers elongate, crested dorsally by a wavy antrorse trichome
(Johnston); style longer than stigma; carpels pitted (lacunose), ellip-
soid, medially and ventrally affixed by a definite elliptical areola
(Johnston). Var. grisellum Johnst. I.e. 40, differs in the longer,
more canescent indument, the leaves usually rugose and more canes-
cent beneath (all the southern Peru specimens; cf. H. Mandonii).
As Johnston remarks, the Ancash specimens are not typical. An
attractive flowering shrub forming large mounds or clumps. The
native name "Vaynilla" refers to the fragrance (Ruiz & Pavon).
Section Heliothamnus Johnst. Illustrated, Bot. Mag. pi. 1609.
Ancash: Below Ocros, 2,650 meters, Weberbauer 2710?; Soukup (t).
Lima: Matucana, 2938; Rose 18639. Ambar, Stork 11457. Canta,
Pennell 14352; 14759; Mathews, Savatier. Amancaes, Weberbauer
1637; Soukup 3125. Near Lima, Ruiz & Pavon, (type, H. corym-
bosum); Dombey. Junin: Huancayo, Oxapampa, Soukup 2438.
Ayacucho: Coracora, 2,900 meters, Weberbauer 5799. Arequipa:
Posco, (Cook & Gilbert 40). Mollendo, Weberbauer 1466; Hitchcock
22394; West 8206. South of Chala, Worth & Morrison 15686. Que-
quena, Eyerdam & Beetle 22180. Moquegua: Carumas, Weberbauer
7272, (type, var.). Tacna: Without locality, (Shephard). "Docto"
(Beetle), "vaynilla."
FLORA OF PERU 559
Heliotropium curassavicum L. Sp. PI. 1: 130. 1753; 14. H.
curassavicum L. var. parviflorum Ball, Journ. Linn. Soc. 21 : 227. 1884.
A glabrous glaucous succulent, the branching prostrate stems
sometimes elongating to a meter or so; leaves oblanceolate or obo-
vate, often 2-4 cm. long, 0.5-1 cm. wide; spikes (bractless) often
geminate; corolla typically 1-2.5 mm. long; anthers free, sagittate,
the cells acuminate-appendaged; stigma column in Peruvian plants
less than 0.2 mm. long, fruit 4-lobate, disk prominent, stigma broad.
An unmistakable plant of wet or low places especially when there
is salinity; the white corollas are usually yellow at throat and some-
what purplish in age as sometimes the entire plant. Section Halmy-
rophila Johnst.
Piura: Alto de la Cruz, Stork & Norton 11355. La Libertad:
Toward Salaverry, Worth 8891. Lambayeque: Chiclayo, Stork 11 430.
Lima: Callao, 5881. Rio Rimac, (Ball) and many other collections,
lea: Paracas Bay near Pisco, (Forbes). Arequipa: Lomas, Weber-
bauer 5732. Mollendo, Weberbauer 388; Eyerdam 25165. Tacna:
Near Tacna, Woitschach; Werdermann 711. Chile and Patagonia to
United States; West Indies.
Heliotropium erianthum Johnst. Contr. Gray Herb. 81: 41.
1928.
A canescent shrub, the branches and dense corymbs tomentose-
villous, the leaves strigose-tomentulose above, tomentose beneath;
petioles 5-8 mm. long; leaves lanceolate, acute both ends, 4-5 cm.
long, 1- about 2 cm. wide, tessellate-rugose with impressed nerves
above, obscurely crenulate, scarcely revolute; spikes 2-5 cm. long;
calyx as corolla without appressed villous, the former orange, finally
4-5 mm. long, the lobes lanceolate-subulate, the latter 2.5-3 mm.
long, limb about 3 mm. across, lobes rounded; anthers linear-oblong,
the cells antrorsely pubescent dorsally toward tips; ovary glabrous,
style 1-1.3 mm. long, stigma about 0.8 mm. long. Resembles H. in-
canum R. & P. with harsh ascending indument, larger flowers; also
H. argenteum Lehm., 42, of southern Ecuador, with slightly shorter
style and ovate, less rugose leaves (author). With more collections
it seems possible that Lehmann's species may be found to vary in
these characters and, admitting any variation, the shrub will prob-
ably be shown to be a part of the Ruiz and Pavon species. Section
Heliothamnus Johnst.
Piura: Huancabamba, Weberbauer 6332, type.
560 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Heliotropium incanum R. & P. Fl. Peniv. 2: 2, pi. 108a.
1799; 41.
Shrub, often sprawling or openly branched, a meter or so high,
with rather showy fragrant white or sometimes purplish flowers and
brick red ovate or ovate-elliptic leaves, typically soon very rugose-
bullate and green above, canescently pubescent and reticulate-venose
beneath; leaves narrowed to the scarcely obvious petioles, acute,
ordinarily 4-6 cm. long, about half as wide, thick, more or less sub-
appressed hispid above, glabrate in age, densely and persistently vil-
lous tomentose beneath; corolla about 4 mm. across expanded top
or somewhat broader; calyx lobes subulate-lanceolate, strigose; nut-
lets nearly 2 mm. long, faintly sculptured; otherwise like the similar
H. arborescens which some forms approach closely (Johnston).
Section Heliothamnus Johnst. F.M. Neg. 17326.
Cajamarca: Chota, Stark & Horton 10041 (det. Standley). Huan-
uco: Near Huanuco, 3237; Ruiz & Pavdn, type. Huacho, Stork &
Horton 9405. Ambo, 2423. Junin: Cabello above Huertas, 1327.
La Oroya, Kalenborn 75. Huancavelica: Mejorada, Stork & Horton
10916; Weberbauer 7604. Apurimac: Oropeza Valley, Vargas 9768
(det. Standley). Pincos, Stork & Horton 10674. Rio Pampas, 2,950
meters, Weberbauer 5855. Cuzco: Ollantaitambo, Hen era 231; Pen-
nell 13640. Vilcanota Valley, H err era 1080. Urubamba Valley,
Herrera 1367; 1533.
Heliotropium indicum L. Sp. PI. 130. 1753; 19.
Sparsely hispid-setulose annual but often vigorous, several dm. to
a meter tall, branching above; petioles alate toward leaf -blade, usu-
ally more than half as long as the ovate-elliptic to broadly lanceolate
herbaceous venose leaves, these several cm. to a dm. long or longer,
acute both ends or more or less cordate at base; spikes bractless,
solitary, scorpoid, often becoming 2-3 dm. long; calyx lobes narrow,
2-2.5 mm. long; corolla blue or violet, sometimes white, the tube
2.5-4.5 mm. long, spreading limb 2-4 mm. across; anthers free; style
short but slender; fruit glabrous, prominent lobes divergent, the
eventually free nutlets angled, 2-3 mm. long. Section Tiaridium
(Lehm.) Griseb.; defined by its conspicuously ribbed multicellular
glabrous fruit. Usually a weed in clearings; only a few collections
cited. Illustrated, Univ. Nat. Cordova 11: pi. 8, opp. page 524.
Piura: Near Piura, (Townsend 1393}. San Martin: San Roque,
Williams 7801. Juanjui, Klug 3436. Tarapoto, Woytkowski 33542.
FLORA OF PERU 561
Loreto: Soledad, Tessmann 5260. Near Iquitos, Williams 1312;
139 b. Tropics and subtropics.
Heliotropium Krauseanum Fedde, Bot. Jahresb. 34, pt. 3: 72.
1908 ; 28. H. saxatile Krause, Bot. Jahrb. 37 : 633. 1906, not Brandg.,
1905.
Erect with arcuate-spreading flowering branches, to a meter tall,
the densely leafy branchlets early somewhat pilosulous; leaves sessile
but long and often narrowly attenuate to base, rounded at tip, oblong
or oblanceolate, 2-5 cm. long, 4-8 mm. wide, crenulate, subcoria-
ceous, finely reticulate-veined and glabrate above, canescent puberu-
lent-hispidulous beneath; spikes congested and densely flowered,
3-4 cm. long in fruit; calyx lobes linear, acute, minutely hispidulous,
the nearly twice as long corolla tube sparsely so; corolla lobes white,
glabrous, nearly triangular; filaments ^ery short; style subequaling
calyx. Meritoriously named for a genial botanist of the Dahlem
herbarium. Related to H. taltalense (Phil.) Johnst. of Chile, coarser
with somewhat larger flowers, more pubescent calyces, larger fruits;
may be a variety; resinous and pleasantly scented according to
Johnston, who placed it in section Cochranea (Miers) Reiche. F.M.
Neg. 17327.
Arequipa: Mollendo, Johnston 3533; Weberbauer 1552, type; 147.
Atiquipa to the port of Chala, Weberbauer 7188; Worth & Morrison
15647. Chile.
Heliotropium lanceolatum R. & P. Fl. Peruv. 2: 4, pi. Ilia.
1799; 40.
Shrub, the type with slender strigulose and sparsely hispidulous
flowering branchlets; leaves crowded, oblong-lanceolate, narrowed to
short petioles, acute or acuminate, mostly 2-3.5 cm. long, 4-6 mm.
wide, rather densely appressed asperous hispidulous and rugulose
above, crinkly puberulent and on the prominent nerves hispidulous
beneath; flowers crowded, the inflorescence branchlets 1-2 cm. long;
calyx segments linear-subulate, about 3 mm. long, sparsely strigulose
as the half again as long corolla tube; corolla lobes rounded or sub-
acute, the expanded top about 4 mm. across. Characterized by
small scabrous lanceolate leaves and lax corymb of long-tubed flow-
ers; type locality listed as Huanuco in Ruiz and Pavon's journal but
not found since, so perhaps that name referred to another species, as
H. incanum; the anthers, according to the sectional character (fide
Johnston) are crested dorsally by a row of crowded antrorse trichomes;
562 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
1 have not dissected the one flower on type material seen. Section
Heliothamnus Johnst. F.M. Neg. 17328.
Lima: Lurin, (Ortega); (Wilkes Exped.}; (Nation). Toward Ma-
tucana, Weberbauer 96. Huanuco: Pillao and near Huanuco(?),
Ruiz & Pavdn, type.
Heliotropium lippioides Krause, Bot. Jahrb. 37: 632. 1904; 42.
Suffruticose, the few slender strict erect branches brownish and gla-
brate below, toward the tip as the sparse leaves beneath more or less
ashy tomentose; leaves sessile or essentially, elliptic, shortly narrowed
at base, apically rounded but often mucronulate, 1.5-2 cm. long,
about 8 mm. wide, coriaceous, revolute, grayish pilose above, strongly
rugose by the impressed nerves; spikes 2-3 cm. long, densely flow-
ered, alternately geminate; calyx hispid, the linear lobes somewhat
spreading in fruit; corolla tube hispidulous, white, 6 mm. long or
about 3 times longer than calyx, lobes undulate, subglabrous; fila-
ments almost 1 mm. long; style short, conical, the stigma obtuse.
Seems to be nearest H. incanum R. & P. but leaves much smaller
(author). Type was a meter tall. Section Heliothamnus Johnst.
F.M. Neg. 17330.
Cajamarca: Santa Cruz, 1,650 meters, Weberbauer 4122, type; 189.
Heliotropium Lobbii Johnst. Contr. Gray Herb. 81: 54. 1928.
Canescently villous-hispid shrub; branches ascending, to 1.5 dm.
long; leaves elliptic-oblanceolate, narrowed to petioles (these 1-
2 mm. long), acute, 6-17 mm. long, 2.5-6.5 mm. wide, firm, con-
spicuously pale tomentose beneath, only midnerve prominent; spikes
geminate, ebracteate, 1-2 cm. long; pedicels stout, to 1 mm. long;
calyx (as corolla without) strigose, 2.5-3 mm. long, the subequal
lobes linear-lanceolate; corolla funnelform, 4-5 mm. long, sparsely
long-strigose in throat, cylindric tube 3 mm. long, limb 2.5-3 mm.
across, lobes ovate; anthers sessile, obtuse, glandular; stigma cylin-
dric, to 1 mm. long, 2-3 times longer than style. Habit of H. pilo-
sum R. & P. with corolla smaller and notably different in form
(author). Section Orthostachys R. Br.
Amazonas: Chachapoyas, (Lobb, type).
Heliotropium Mandonii Johnst. Contr. Gray Herb. 81: 43.
1928.
Becoming a meter or two high, the laxly branched branches early
fulvous-villous (trichomes retrorse), finally glabrescent; leaves Ian-
FLORA OF PERU 563
ceolate or oblong-elliptic, reduced into villous petioles about 1 cm.
long, acute, 7-14 cm. long, 2.5-6 cm. wide, scabrous, sparsely villous-
hispid and often by the impressed veins strongly reticulate-rugose
above, paler or canescently velutinous beneath; spikes villosulous, to
a dm. long, in a terminal dichotomous corymb; calyx sessile, slightly
villous, often glandular, globose, finally 2-2.5 mm. long with the
ovate-lanceolate acuminate lobes little plicate in the ovate sinuses;
corolla white, 3-4.5 mm. long, pubescent (often glandular) only with-
out, the cylindric tube to two times longer than calyx, limb 3-5 mm.
across, lobes obtuse; anthers densely pubescent toward apex; stigma
sessile, 0.5 mm. long; ovary eglandular; nutlets 1.5 mm. long, lacu-
nose, compressed, broadly affixed. A coarse species very distinct
from its relatives and suggesting a Tournefortia in gross aspect
(author) ; simulates H. arborescens var. grisellum Johnst. but stigma
sessile and inflorescence in age distinctive; flowers fragrant in my
collections. Type is Mandon 387 from La Paz, Bolivia. Section
Heliothamnus.
Huanuco: Yanano, 1,800 meters, 3660. Rio Santa Domingo,
1,200 meters, 4206 Cuzco: Valle de Santa Ana, (Herrera 899).
Bolivia.
Heliotropium microstachyum R. & P. Fl. Peruv. 2: 3, pi.
llOb. 1799; 12. H. brachystachyum (DC.) Griseb. Gott. Abh. 24:
271. 1879. Heliophytum brachystachyum DC. Prodr. 9: 554. 1845.
A lightly strigulose perennial developing a ligneous caudex, the
slender prostrate stems with glomerate inflorescences in the axils of
the upper branchlets or leaves, these elliptic or suboblong, attenuate
to the slender, usually short petioles, rounded or obtuse at apex,
ordinarily 1.5-2.5 cm. long, 5-10 mm. wide, sometimes smaller or
considerably longer, often glabrous or glabrate in age; nerves slender,
impressed above, rather prominent beneath; inflorescences in fruit
more or less spicate, a cm. or two long; calyx segments linear, about
2 mm. long; corolla 2-3 mm. across at top, glabrous, tube included,
lobes rounded; carpels 2-celled, frequently maturing only a single
seed (Johnston), lacunose, nearly 2 mm. long. Johnston (who dis-
covered the identity of the above names) noted that the upper plant
figured on Ruiz and Pavon's plate 110 is by error labeled H. pilosum,
a confusion in legend occurring elsewhere in their Flora of Peru.
Section Hypsogenia Johnst. Illustrated, Univ. Nat. Cordova 11:
pi 3, opp. page 506. F.M. Neg. 17333.
564 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Lima: Matucana, 202. Huanuco: Near Huanuco, Dombey; Kane-
hira 258. Ambo, 8168. Junin: Tarma, Weberbauer 2370 (distr. as
H. paronichioides) ; 176; Ruiz & Pavdn, type. Huancavelica: Huay-
tara, Weberbauer 54-27, part. lea: Above Pisco, 2,650 meters, Weber-
bauer 5427. Cuzco: Ollantaitambo, Hen era 2254; Pennell 13660.
Near Cuzco, Vargas 654. Arequipa: Cotahuasi, Weberbauer 6871.
Puno: Near Puno, (Shepard 120). Chuquibambilla, Pennell 13358.
Altos de Toledo, 4,650 meters, Meyen. Moquegua: Carumas, Weber-
bauer 7484- Ecuador to Argentina.
Heliotropium oxylobum Johnst. Contr. Gray Herb. 81: 57.
1928.
A shrub with many slender branches, sometimes 6 dm. tall,
sparsely strigose with appressed curved trichomes; leaves lanceolate-
linear or oblanceolate, narrowed to base (petioles 1 mm. long), acute,
1-2.5 cm. long, 1.5-4.5 mm. wide, firm, enervose, pale beneath;
spikes single or paired, ebracteate, 2-6 cm. long; calyx subsessile,
1-2 mm. long, the sublinear lobes subequal; corolla yellowish, densely
strigose without and slightly short-villous within above the middle,
4-5 mm. long, the cylindric tube 3-4 mm. long, lobes ovate-deltoid,
acute; anthers to 1 mm. long, obtuse, glandular adhering; ovary
glabrous or puberulent; stigma about 1 mm. long with thick disk,
to two times longer than style; nutlets 1 mm. long, convex, inner
face angulate. Differs from H. campestre Griseb., known from La Paz
southward, in its more slender habit, smaller leaves, more appressed
indument and pubescence within the corolla. Simulates H. lance-
olatum but apparently differs in anther character. Corolla yellowish
or tube purplish (Goodspeed Exped.). Section Orthostachys R. Br.
Huancavelica: Rio Mantaro below Surcubamba, Weberbauer 6486.
Apurimac: Bridge of Apurimac, (Pearce, type). Rio Pachachaca,
Goodspeed Exped. 10544 (det. Standley, H. pilosum). Pincos, Stork
& Horton 10654 (det. Standley, H. pilosum). Cuzco: Mayhua, Prov.
Paruro, Vargas 890 (det. Standley) ; 9735 (det. Standley, H. pilosum).
Heliotropium pilosum R. & P. Fl. Peruv. 2: 3, pi. HOa. 1799;
56.
Prostrate or spreading-ascending, sometimes a meter high, at
least the tips canescent with more or less appressed pubescence;
caudex ligneous as the prostrate-ascending older stems and few
branches, the herbaceous younger with soft greenish lanceolate-
elliptic leaves, attenuate to short petioles, acute, often to 3 cm. long,
FLORA OF PERU 565
to about a third as wide, the older flowering branches usually with
narrowly oblong-lanceolate leaves, frequently firmer, acuminate, 2-5
cm. long, 5 mm. wide or narrower; inflorescence often composed of
2-3 spikes finally 5 cm. long or longer; calyx segments subulate-
linear, 1.5 mm. long, little longer in fruit, lax; corolla almost minute,
tube within and without strigulose, not at all or slightly exserted,
lobes rounded. Var. albatum Johnst. is merely a form with an
abundant appressed pubescence (author). Plate 11 Ob, not a as cap-
tioned, is obviously this species; details incorrect; nutlets are 1 mm.
long, sparsely hirsute-strigose, convex, the face distinctly acute;
corolla is subsalverf orm with stout cylindric tube not surpassing calyx
lobes. Section Orthostachys R. Br. F.M. Neg. 17339.
Ancash: Lomas de La Chay, Goodspeed Exped. 9214- Lima:
Hills of Lima, (Nation). Rio Rimac, (Ball, type, var.); Metcalf
30249. Surco, 1,950 meters, Weberbauer 5204 (var.). Atocongo,
Pennell 14786. Chosica, Weberbauer 5319. Clays near Barranca,
Weberbauer 1649; 147. Sandy seaside hills, 5922. Amancaes, Ruiz &
Pavon, type. Lurin, 5922. Sayan, Goodspeed 33043. Arequipa:
East of Chala, Worth & Morrison (var.). Laspinas, 2,200 meters,
Eyerdam & Beetle 22159 (var.).
Heliotropium piurense Johnst. Amer. Journ. Bot. 33: 479.
1946.
Slender erect annual 1-2 dm. high, stem at base 1.5 mm. thick,
dichotomously branched above; sparsely appressed strigose with
curved antrorse trichomes; leaves oblanceolate, long-narrowed to
slender petioled (1-2 mm. long), acute, 1-1.5 cm. long, 2-3 mm. wide
above the middle, minutely strigose, enervate; spikes terminal,
slender, 5-10 cm. long or longer, leafy-bracted; pedicels 0.5-1 mm.
long; calyx sparsely strigose, the segments unequal, erect or sub-
erect, lanceolate, acute, 2-2.5 mm. long, in fruit 3-3.5 mm. long;
corolla white, cylindric, 2-4 mm. long, a little strigillose (only out-
side), lobes oblong, erect or nearly, little exceeding calyx; anthers
extended apically, cohering; stigma broadly conical, to 3 mm. high
and broad, subsessile; nutlets about 1 mm. high, ventrally angulate,
pored, well-convex dorsally and pubescent with uncinate trichomes.
"Keys" in the author's revision, Contr. Gray Herb. 81: 49. 1928,
to the southern H. ocellatum Cham., the calyx not uncinate pubes-
cent; in this character resembles the northern H. fruticosum L. that
is less slender and with a definite style (author).
Piura: Among sparse grasses, Cerro Prieto, H aught & Svenson
11553, type. Cabo Blanco, (Haught 256).
566 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Heliotropium polyanthellum Johnst. Contr. Gray Herb. 81:
56. 1928.
Type a lax light green bush about 5 dm. tall, the many slender
branches densely strigose; leaves narrowly lanceolate or oblanceolate,
to about 1 cm. long, 2-4 mm. wide, gradually attenuate into the
1 mm. long petiole, acute, subcoriaceous, not veiny but midrib strong
beneath; spikes single or geminate, slender, ebracteate, 4-6 cm. long;
calyx subsessile, finally 1.5-2 mm. long, the slender lobes unequal;
corolla yellowish, strigose only without, the cylindric tube 2-2.5 mm.
long, less than 1 mm. thick, the limb 2-2.5 mm. across, lobes ovate;
anthers linear, obtuse, sessile, lightly coherent by the glandulosity;
ovary sparsely villosulous; stigma cylindric, about 0.8 mm., style
obsolete; nutlets about 1 mm. long, strigose. Relationship unknown
(author). Section Orthostachys R. Br.
Cajamarca: Between Jaen and Rio Shumba, 750 meters, Weber-
bauer 6184, type.
Heliotropium procumbens Mill. Diet. ed. 8, no. 10. 1768; 52'
H. inundatum Sw. Prodr. 40. 1788.
Flowering as an annual but sometimes more or less enduring and
ligneous toward base, green or subcanescent strigose; leaves often
1-2 cm. long, about half as wide, elliptic or somewhat obovate, nar-
rowed to the much shorter petiole, rounded and mucronate at apex;
spikes usually geminate, soon several cm. long; calyx segments sub-
ulate linear, about 1.5 mm. long, lax in fruit and 2.5 mm. long;
corolla inconspicuous, the strigose tube slightly exserted; nutlets
strigulose but smooth. See Johnston for complete synonymy, as
elsewhere; to it he added, after seeing types (Contr. Gray Herb. 92:
89. 1930), H. americanum Mill., I.e. no. 11, H. brasilianum Roth,
Nov. PL Sp. 103. 1821, and H. rigidulum DC., Prodr. 9: 540. 1845;
he notes species apparently sometimes perennial (Bolivian plateau)
and flowers larger in southern part of range. Section Orthostachys
R. Br. Illustrated, Univ. Nat. Cordova 11: pi. 12, opp. page 538.
Piura: Near Piura, Spruce. Loreto: Middle Rio Ucayali, Tess-
mann 3111. Argentina to United States.
Heliotropium rufipilum (Benth.) Johnst. Contr. Gray Herb.
81: 44. 1928. Tournefortia rufipila Benth. Bot. Sulph. 140. 1844.
Slender, sometimes a meter or two tall, the long stems or branches
especially above more or less fulvous hispid with often somewhat
viscid and usually spreading trichomes; leaves narrowly ovate-
FLORA OF PERU 567
elliptic, acute both ends or more commonly long-decurrent on short
or elongate petioles and acuminate at apex, frequently about a dm.
long, a third as wide or much larger, membranous, obscurely veined
except in age when somewhat rugulose above, more or less hispid-
ulous; inflorescences lax in age, the branches 4-15 cm. long; calyx
segments subulate, 2 mm. long or about 3 mm. long in fruit, the
lightly strigulose corolla tube slightly exserted; corolla lobes rounded,
the flower 3-5 mm. across at expanded top. There is considerable
difference in leaf-form and flower-size between the Haught speci-
mens determined by Johnston and others determined by him from
Ecuador; typically with herbaceous leaves, veins little impressed,
branches lax, spikes elongate, nutlets densely long-stiped glandular;
var. anadenum Johnst., I.e., is a form or race with nutlets sparsely
if at all glandular. Compare H. submolle Klotzsch and H. Urbani-
anum Krause, which will prove, probably, to be a part of this
species; H. adenogynum Johnst. is scarcely more than a variant;
Croton, Lantana and other Andean shrubs respond to local condi-
tions similarly. Section Heliothamnus Johnst.
Piura: Talara, Haught 70. Parinas Valley, Haught 169. Cerro
Viento, Haught 77 (var.). San Martin: San Roque, Williams 7804
(det. Standley). Zepelacio, Klug 3488 (det. Standley). Junin:
Oxapampa, Soukup 1810 (det. Ewan). Bolivia to Mexico.
Heliotropium tora tense Johnst. Contr. Gray Herb. 81: 55.
1928.
Shrubby at base, decumbent-ascending, 1.5-2 dm. high, the many
stems to 3 dm. long, spreading villosulous below, appressed pubes-
cent to tips; leaves oblanceolate, gradually attenuate to base, acute,
3-6 cm. long, 4-8 mm. wide, herbaceous, veinless, obscurely strigil-
lose, narrowly revolute; spikes terminal, geminate or ternate, 3-7 cm.
long; calyx subsessile, 3-4 (finally 5) mm. long, sparsely strigose, the
linear-acuminate lobes unequal; corolla about 4 mm. long, tube
strigose only without, 2-3 mm. long, the limb 4.5 mm. across;
anthers subulate, acuminate, obscurely granulate; stigma 0.3-0.4
mm. long, 4-lobate, subsessile; nutlets villous, 1.7 mm. long, convex,
the inner face angulate. Well-marked species of uncertain relation-
ship (author). Section Orthostachys R. Br.
Moquegua: Torata, Weberbauer 7407, type.
Heliotropium Urbanianum Krause, Bot. Jahrb. 37: 633. 1906;
45.
568 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Flowering branches, especially toward apex and crowded inflor-
escences, conspicuously fulvous villous-hispid as the younger leaves
beneath, the older canescent; petioles 3-5 mm. long; leaves elliptic
or early ovate-elliptic, acute at base, rounded or sometimes sub-
acute at apex, usually 3.5-5 cm. long, about 2 cm. wide; inflorescences
little longer than wide even in fruit, ordinarily only a few cm. long;
calyx segments narrowly ovate-subulate, 2-2.5 mm. long, hispid-
ulous; corolla blue, including the rounded lobes, more or less pilose,
the stout tube barely exserted, the expanded flower about 4 mm.
across; anthers 2 mm. long; style elongate-conical, shorter than
stigma. Distinguished from H. rufipilum at a glance (Johnston)
but, as monographer remarks, characters mostly relatively devel-
oped; spikes and corolla-tubes usually stouter, leaves smaller, usually
rugose and broadest at or above the middle, more pubescent, the
trichomes often bulbous based, and, especially, calyx does not be-
come globose. Section Heliothamnus Johnst. F.M. Neg. 17349.
Amazonas: Chachapoyas, (Mathews); perhaps same locality,
(Lo66). Ecuador.
3. COLDENIA L.
Tiquilia Pers. Syn. 1: 157. 1805.
References: Johnston, Contr. Gray Herb. 70: 58-61. 1924; Journ.
Arnold Arb. 16: 183-185. 1935.
Annual or perennial, sometimes more or less ligneous, often
spreading or prostrate and the veiny or evenose leaves approximate
or subtending the small flowers. Calyx 4-5-parted. Corolla funnel-
form or salverform, appendaged or smooth. Style 2-cleft or 2-parted.
Ovary entire or 4-lobed, in fruit 4, rarely 2, separate nutlets, these
attached basally, usually ventrally.
C. Nuttallii Hook, of western North America has been found in
the Andes of Argentina according to Johnston, I.e. 75: 43. 1925; it is
an annual with smooth ovoid nutlets, appendaged corollas. Most
specimens being in flower, Johnston's key, which follows, is supple-
mented.
Nutlets plano-convex; stamens exserted C. dichotoma.
Nutlets spheroid to ovoid.
Nutlets narrowly ovoid with ventral scar; petioles usually villous-
ciliate C. paronychioides.
Nutlets spherical; petioles not villous.
FLORA OF PERU 569
Corolla about 5 mm. long; stamens included C. elongata.
Corolla about 1 cm. long; stamens exserted C. conspicua.
SUPPLEMENTARY KEY
Flowers very obvious; stamens well-exserted; nutlets subspherical.
C. conspicua.
Flowers more or less obscure; stamens slightly or not exserted.
Leaves about 5-7 mm. long; nutlets narrowly ovoid.
C. paronychioides.
Leaves mostly or all about 1 cm. long or longer; nutlets sub-
spherical to plano-convex.
Stamens included; nutlets spherical C. elongata.
Stamens visible; nutlets plano-convex C. dichotoma.
Coldenia conspicua Johnst. Journ. Arnold Arb. 16: 183. 1935.
Prostrate, the laxly branched stems sometimes a dm. long or
longer, the younger finely hispidulous and more or less glanduliferous
as the setose petioles (2-9 mm. long) and hispid calyces; leaves
broadly lanceolate or elliptic, entire (or essentially), 5-13 mm. long,
2-5 mm. wide, closely and coarsely pustulate-asperous above, finely
and erectly so beneath; calyx lobes linear, subindurate at base, about
9 mm. long, in fruit to 1.5 cm. long; corolla blue, tube 9 mm. long,
limb 10-12 mm. across; throat smooth; stamens exserted; nutlets
early about the top of a subcylindrical gynobase, this finally ex-
panded, turbinate and the nutlets borne basally in the apical tissue
by means of strophiodes. Nutlets quite similar in size, shape and
markings to those of section Sphaerocarya Johnst. Contr. Gray
Herb. 70: 57. 1924. Johnston once referred the Pennell collections
to the Chilean C. grandiflora Phil.; these appear to be annual or possi-
bly biennial; flowers pale to blue-violet (Pennell), leaves 1.5-2 cm.
long.
Arequipa: Sand flat near Mejia and Cachendo, (Guenther &
Buchtien 155, type; det. Bruns, C. dichotoma'); 156. Mollendo,
(Stafford K60); Mexia 94178. Quishuarani, 1,860 meters, Pennell
13044- Sand dunes, Tiabaya, 2,100 meters, Pennell 13059.
Coldenia dichotoma (R. & P.) Lehm. Asperif. 1: 9. 1818; 58.
Lithospermum dichotomum R. & P. Fl. Peruv. 2: 5, pi. lllc. 1799.
C. Dombeyana Juss. ace. to Buck, Index DC. Prodr. 3. 108. 1858.
570 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
C. pentandra Juss. ace. Steud. Nomencl. 212. 1821. Tiguilia dicho-
toma (R. & P.) Pers. Syn. 1: 157. 1805.
Slightly suffrutescent with rugose veiny leaves, bark papery, soon
exfoliating, early shortly glandular pilosulous; petioles 5-10 mm.
long; leaves hispidulous or glabrate, especially above where bullate
by the strongly impressed nerves, mostly oblong-obovate, revolute,
crenulate, about 12-15 mm. long, 5-6.5 mm. wide; corolla 3-4 mm.
wide, bluish-pink, little longer than the sublinear hispid sepals; sta-
mens well-exserted. Nutlets 2, 1.5-2.5 mm. long, strongly convex
dorsally, finely tessellate-granulate, strongly brown-mottled, the an-
terior face flat, spreading to form circular patches sometimes 15 dm.
across (Johnston).
Piura: Talara, Haught 6; Johnston 3517; Eyerdam 9008. Parinas
valley, Haught 158. La Libertad: Salaverry, Johnston 3522. Lima:
On the lomas, Weberbauer 5713; 144. Lurin, Ruiz & Pawn, type;
5946. "Tiquil-tiquil" (Ruiz & Pavon).
Coldenia elongata Rusby, Descr. New Sp. S. Am. PL 107. 1920;
185. C. parviflora Phil, as to Johnston, I.e., fide Johnston, Journ.
Arnold Arb. 16: 185. 1935.
Irregularly branched, shrubby, lower stems glabrate, exfoliating,
younger more or less densely villous-hispid; petioles slender, shorter
than the oblong obtuse leaves, these with 4-5 pairs of veins impressed
above, in type to 7 mm. long, 4 mm. wide, usually larger, oval-ovate,
obtuse both ends, revolute; calyx lobes linear, about 5 mm. long,
long pilose-hispid; corolla about 5 mm. long, the tube about 1.5 mm.
long, the lobes subrotund; stamens included; nutlets 4, spherical or
nearly, attached by a small basal circular scar, smooth, lustrous,
about 0.5 mm. thick or somewhat larger, brown in type, perhaps
black at full maturity. The elongate, usually crenate leaves, tomen-
tose petioles and calyces distinguish this from C. parviflora Phil, of
Chile (Johnston).
Arequipa: Yura, (Williams 2562, type; 2521? nutlets larger);
Karl Schmidt (det. Standley, C. dichotoma). Below Arequipa, Weber-
bauer 6837. Lomas, Prov. Camana, Weberbauer 5730; 5734. Adja-
cent Chile.
Coldenia paronychioides Phil. Cat. Itin. Tarapaca 55. 1891; 60.
Lithospermum aggregatum R. & P. Fl. Peruv. 2: 4. 1799. C. aggregata
Rusby, Descr. New Sp. S. Am. PI. 106. 1920, not based on R. & P.
type.
FLORA OF PERU 571
Prostrate, greenish or canescently appressed to spreading hispid-
ulous with minute pale blue to white flowers, early herbaceous but
soon ligneous toward base and openly sprawling, in age sometimes
several dm. in diameter; petioles 1-3 mm. long; leaves oblong-
oblanceolate, mostly about 5 mm. long, 1.5-2 mm. wide; corolla
nearly hidden in the pubescence of the linear calyx lobes, these only
about 3 mm. long; nutlets narrowly ovoid, lustrous, minutely reticu-
late; habit of C. dichotoma, with which it sometimes grows (John-
ston).
Piura: Talara, Haught S3; Pennell 12202; Goodspeed Exped. 9004;
Horton 11555; West 3572; Beetle 26204. Near Piura, Weberbauer 5932.
Negritos, (Haught & Svenson 17606). Paita, Pennell 14819; (Saf-
ford 9; Ball; Williams 2913, type, C. aggregata Rusby); Weberbauer,
154. La Libertad: Salaverry, Johnston 3523. Ancash: Los Zorros,
Goodspeed Exped. 9197. Lima: Quive, Pennell 14305. Sta. Eulalia,
Goodspeed 33092. Sayan, Goodspeed 17358. Junin: Near Tarma,
Ruiz & Pavdn (type, L. aggregatum) . Arequipa: Tingo, 2,100 me-
ters, Pennell 13102. Puno: Pisacoma, (Meyen, distr. as Heliotropium
humifusum, fide Johnston). Northern Chile; Bolivia. "Flor de
arena" (Stork).
4. CORDIA L.
Trees or shrubs, in Peru broad-leaved and the inflorescence vari-
ous. Calyx mostly persisting. Corolla white or yellowish, red to
orange in C. sebestena (cultivation), the lobes 5-9 (-15). Stamens
5-10. Ovary usually 4-celled, ovules 1-4. Style terminal, well-devel-
oped, 2-lobed or 2-parted, the branches each 2-lobed; stigmas 4.
Fruit usually a drupe with dry or more or less mucilaginous exocarp,
or carpoid wall, chartaceous and fibrous (section Gerascanthus) .
Genus first described in detail anatomically by Mez, Bot. Jahrb. 12:
526-588. 1890. Johnston's "The Boraginaceae of Northeastern
South America," Journ. Arnold Arb. 16: 1-64. 1935, is an invaluable
contribution to the knowledge of this family and I hope I have
correctly compiled his painstakingly observed and recorded observa-
tions pertaining to the Peruvian species.
The red or orange flowered C. sebestena L. is probably grown in
Peru for ornament; it is native to the northern coasts of South
America and the West Indies.
Corolla persisting (marcescently) ; calyx costate.
Calyx teeth obscure; fruit enclosed.
572 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves lanceolate, about three times longer than wide.
C. iguaguana.
Leaves oval to ovate in some degree and usually about twice
as long as wide.
Leaves usually longer than wide.
Flowers about 1.5 cm. long C. alliodora.
Flowers about 4 cm. long C. insignis.
Leaves suborbicular C. lutea.
Calyx with long subulate teeth; fruit exserted . . . . C. varronifolia.
Corolla deciduous after drying; calyx (in Peru) not tubulate nor
strongly costate nor ovate and subulate-lobed.
Nodes inflated; stems notably long-hispid C. nodosa.
Nodes not inflated; stems glabrous to variously pubescent.
Flowers showy, in globose heads; peduncles to 1 dm. long; calyx
teeth filiform C. macrocephala.
Flowers small, sometimes capitate but calyx lobes obscure, rarely
subulate-appendaged; peduncles often short.
Branches remotely spurred or nodular by persisting petiole-
bases; flowers usually congested in spikes or heads, dis-
posed rather laxly only in C. Poeppigii; fruit often shorter
than 5 mm.
Inflorescences usually simple, in any case densely flowered
(cf. C. Poeppigii).
Leaves evenly serrate, ample, with subulate teeth to
2 mm. long, 3-5 mm. apart C. macrodonta.
Leaves entire, minutely or densely or unevenly serrate,
usually only a few cm. wide (key characters below
variable, expedient).
Inflorescences soon clearly longer than broad, spiciform.
Leaves coarsely and unevenly serrate . . C. peruviana.
Leaves entire to more or less serrate but closely or
evenly C. cylindristachya, C. spinescens.
Inflorescences usually forked or capitate, especially in
flower.
Leaves more or less acuminate and acutely, mem-
branous or firm, not or obscurely rugose unless
in age, usually glabrate or sparsely pubescent;
cymes small, early dense or capitate.
FLORA OF PERU 573
Inflorescence axillary C. patens.
Inflorescence extra-axillary C. bifurcata.
Leaves if acuminate shortly and not or scarcely
acutely, usually rounded to subacute, subcori-
aceous, reticulate-rugose, often rather softly
pubescent beneath; flowers often capitate even
in age.
Leaves subentire or remotely serrulate.
Flower heads often several on more or less leafy
branchlets.
Calyx lobes subulate-appendaged; branchlets
spreading hirsute C. buddleoides.
Calyx lobes acute; branchlets lanate.
C. mollissima.
Flower heads or small forked cymes solitary;
branches soon glabrate C. scaberrima.
Leaves obviously even if rather minutely, serrulate
or crenate.
Leaves oblong- or ovate-elliptic unless young
shoots, slightly rounded to subacute.
C. Krausiana.
Leaves rather ovate, more or less obtusely and
broadly acuminate, scabrous above.
C. scaberrima.
Leaves subrotund, hispidulous both sides.
C. lantanoides.
Inflorescences mostly compound or foliose, the branches
rather laxly flowered C. Poeppigii.
Branches smooth or in any case inflorescences soon lax or
sometimes simple but laxly flowered or open and flowers
crowded, usually not simple or not typically small; fruit
often 5 mm. long or longer.
Leaves rather ovate, often acutely acuminate; inflorescence
branches few, elongate, simple or subsimple, suberect.
C. Poeppigii.
Leaves rather elliptic, cuspidate or acute or if acuminate,
abruptly or shortly; inflorescence branches short, often
many.
574 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Leaves glabrous or essentially, even beneath any indu-
ment minute and obscure; ovary glabrate or glabrous.
Calyx long-obconic, 6 mm. long in bud; corolla to
12 mm. long; ovary glabrous C. Mexiana.
Calyx campanulate or pyriform, 2.5-4 mm. long; cor-
olla to about 5 mm. long.
Petioles 2-3 cm. long, geniculate at base; calyx lobes
uneven C. collococca.
Petioles shorter or in any case straight; calyx lobes
obscure, equal C. lomatoloba.
Leaves strigose or scabrous at least in part; ovary pubes-
cent except C. hebeclada, C. Ulei, C. ucayaliensis,
sometimes C. ripicola.
Flowers much congested; leaves (as to types) clearly
pubescent beneath.
Inflorescence lightly pubescent.
C. laevior, C. ripicola.
Inflorescence reddish tomentose C. Sprucei.
Flowers not congested or inflorescence branchlets long;
leaves minutely pubescent or scabrous.
Petioles about 1 cm. long; leaves mostly or all nar-
rowed to acute base.
Branchlets puberulent; calyx about 4 mm. long.
Leaves scabrous, nearly three times longer than
wide; ovary pubescent C. scabrifolia.
Leaves pubescent both sides, about twice as long
as wide; ovary glabrous. . .C. ucayaliensis.
Branchlets hirsutulous; calyx about 5 mm. long.
C. Ulei.
Petioles about 1.5-2 cm. long; leaves broadly round-
ed or shortly acute at base or very dissimilar.
Ovary pubescent; larger leaves often acute or nar-
rowed to base C. toqueve.
Ovary glabrous; all leaves rounded or truncate at
base C. hebeclada.
Cordia alliodora (R. & P.) Oken, All. Naturgeschichte, Bot. 2:
1098. 1841; 13. Cerdana alliodora R. & P. Fl. Peruv. 2: 47, pi 184.
1799.
FLORA OF PERU 575
More or less densely stellate-pubescent with small or minute
forked trichomes, especially the ample inflorescences; petioles 1-3 cm.
long; leaves rather broadly elliptic, obtusely acute both ends or
acuminate apically, usually 1-5.5 dm. long, about half as wide, entire,
coriaceous, glabrous above where slightly marked by the impressed
nerves and veins, these rather prominent beneath; inflorescence ter-
minal, laxly branched, the flowers crowded; calyx nearly globular,
strongly 10-ribbed, canescent-puberulent, truncate, 6-7 (14) mm.
long, the teeth minute, the exserted corolla tube with oblong rounded
lobes, these 5-10 mm. long, usually about 3 mm. wide; fruit fibrous,
chartaceous, 5 mm. long, about 3 mm. thick, falling within the per-
sisting calyx tube and corolla.
A distinctive feature of the plant is its myrmecophily (cf . Wheeler,
Bull. Mus. Comp. Zool., 90: 9-41. 1942); the leafy twigs and fre-
quently the axis of the inflorescence develop as irregular swellings
which serve as ant domatia; see also C. nodosa. Small ants some-
times totally destroy the leaves, cutting them into triangular pieces
and storing them symmetrically; each piece may be six times the
size of the ant that carries it by the shortest side, apparently like a
sail (Ruiz & Pavon). See Johnston, I.e., for extra-Peruvian synon-
ymy; he remarks that there is only moderate variation in the species,
in size of flowers and degree of pubescence. In Peru sometimes
attains 20 meters or more, the trunk (bark grayish, fissured) 3 dm.
or more in diameter; timber used for construction and for vehicles
(Williams); leaves and bark used as a condiment, serving as garlic
(Ruiz & Pavon) ; flowers fragrant.
Cajamarca: Hualgayoc, Soukup 3839. San Martin: Tarapoto,
Williams 6751. Hudnuco: Pozuzo, Ruiz & Pavon, type. Loreto:
Lower Itaya, Williams 318. Lower Huallaga, Williams 4306.
Cuzco: Urubamba Valley, Weberbauer, 280. Rio Acre: Upper Rio
Jurupary, Krukoff 5220. Seringal Auristella, Ule 9719. Bolivia to
Mexico; West Indies.
Cordia bifurcata Roem. & Schult. Syst. 4: 466. 1819. Varronia
dichotoma R. & P. Fl. Peruv. 2: 23, pi. 146. 1799, not C. dichotoma
Forst., 1786.
Slender, shrubby below, often openly branched above, the leafy
flowering branchlets minutely and sparsely appressed strigillose and
(or) scabrous as the green leaves at least above, these sometimes also
hispidulous on the veins beneath; petioles about 3 mm. long; leaves
usually ovate- or oblong-elliptic-lanceolate, obliquely acute at base,
576 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
acutely acuminate, 6-7 cm. long, about 2 cm. wide near the middle
or sometimes (as freely interpreted) broadly ovate, rounded at base,
3-4.5 cm. wide and only about twice as long, subentire to remotely
serrate above the middle, membranous, paler beneath; peduncles 1-2
(4) cm. long, terminal on leafy branchlets or from the leaf-axils of
branchlets, the small inflorescences capituliform in flower, obviously
and subumbellately 2-4-dichotomously branched only in fruit when
sometimes about 3 cm. across, half as long; calyx appressed strigil-
lose, glabrate and reddish in fruit, with broad almost minute lobes,
about 3 mm. long or about half as long as the fleshy usually white
corolla; fruits bright red, ovoid-elliptic, 3.5-4 mm. long. Probably
a species variable in pubescence, leaf -form, flower color and perhaps
in position and form of inflorescence (compare C. patens, C. scaber-
rima and others). Johnston, in 1930 (Contr. Gray Herb. 92: 30-34),
under the name C. corymbosa (L.) G. Don, commented at length on
apparent variability and, as regards pubescence, proposed forma
typica, leaves finely tomentulose beneath, coarse trichomes few or
lacking; f. intonsa, coarse trichomes abundant; f. detonsa, pubescence
fine, appressed; and f. hirsuta, pubescence coarse, spreading. In 1949
(Journ. Arnold Arb. 30: 90) he wrote, using the name C. bifurcata
Roem. & Schult.: flowers minute, orange, in terminal or internodal
cymes, these usually forked even in bud. Material determined by
Johnston after 1949 had white flowers, according to the collectors.
Johnston, I.e. 30: 90 et seq. 1949, has argued, apparently correctly,
for the validity of this species, confused with C. lineata (L.) Roem.
& Schult., doubtfully in South America, and C. polycephala (Lam.)
Johnst., also known only north of Peru; both have the axillary
inflorescence of C. patens HBK.
Confusion in interpretation of the shrub's characters has been
equaled by the nomenclature; professional taxonomists apparently
ought to be reminded of the old truism that whatever name used the
rose is still a rose or words to that effect; Tournefortia syringaefolia
Vahl is a case in point, apparently with another name, T. maculata
Jacq. in spite of its retaining the same "spots;" however, if a botan-
ical compiler overlooks or misapplies a rediscovered or reinterpreted
name may heaven forgive him!
Piura: Canchaque, Prov. Huancabamba, Stork 11417 (det. John-
ston, C. macrodonta) . San Martin : Tarapoto, Ule 6478; Woytkowski
35074; Williams 5408; 5571; 6177; 6426; 6684A Huanuco: Cha-
cahuasi, Ruiz & Pavdn (type, V. dichotoma). Tingo Maria, Allard
20494 (det. Lyman Smith, C. polycephala). Rio Santo Domingo,
FLORA OF PERU 577
4210. Pozuzo, 4560. Junfn: La Merced, Killip & Smith. Prov.
Tarma, Woytkowski 37003. Argentina to Costa Rica.
Cordia buddleoides Rusby, Mem. Torrey Bot. Club 6: 83. 1896;
34.
Branchlets elongate, spreading hirsute; petioles stout, very short
to nearly 1 cm. long; leaves ovate, abruptly contracted at base, acute,
7-10 cm. long, 3-5 cm. wide, obsoletely serrulate, firm-membranous,
scabrous and the 6 or so lateral nerves impressed above, strigose be-
neath, the veins strongly reticulate; inflorescence ample, leafy, in
type the flowers capitate; calyx tube 1.5 mm. long, lobes subulate-
linear, 3 mm. long, reflexed; corolla little exserted, stamens included;
ovary slightly 4-lobed, style nearly 2 mm. long, 4-cleft. After author
except that he describes branchlets as harshly tomentose; the small
heads are on long peduncles and form a somewhat leafy open inflores-
cence. Ule 9714 from Rio Acre in Bolivia has flowers in a cylindric
spike, probably due to greater maturity than type with globose
inflorescences from La Paz, matched by Lechler; the Poeppig collec-
tion lacks the apical calyx lobe appendages (Johnston) ; this northern
shrub may rather be C. scaberrima HBK. or affine. Name originally
spelled as above.
Loreto: Maynas (Yurimaguas), (Poeppig 2471). Puno: San Go-
van, (Lechler 2424)- Bolivia.
Cordia collococca [Sandmark] L. Amoen. Acad. 5: 377. 1759
(as C. callococca). C. glabra of authors, not Linnaeus; cf. Johnston,
Journ. Arnold Arb. 21: 345; 1940, I.e. 30: 123-124. 1949.
Essentially glabrous, the indument of the open, dichotomously
branched inflorescence and leaves beneath obsolete or extremely
minute; petioles 2-3 cm. long; leaves elliptic-obovate, obliquely
rounded or subacute at base, broadly acuminate or rounded and
apiculate at apex, submembranous, finely reticulate-veined both
sides, the 7-9 lateral nerves slender; calyx obscurely puberulent,
cupulate, about 4 mm. long, unevenly lobed; corolla about 5 mm.
long; ovary glabrous; drupes bright red. Deciduous, dioecious, the
flowers precocious, the leaves beneath pubescent with many slender
erect bristles (Johnston); umbrella-shaped, 5 meters tall, with pen-
dent ("weeping") habit (Mexia).
Loreto: Rancho Indiana, Mexia 6403 (det. Johnston). To Cuba
and Mexico. "Camungo-tiarina" (Mexia).
578 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Cordia cylindristachya (R. & P.) Roem. & Schult. Syst. 4: 459.
1819. Varronia cylindristachya R. & P. Fl. Peruv. 2: 23, pi. 147a.
1799. V. macrostachya R. & P. I.e. as to plate. C. tarmensis Krause,
Bot. Jahrb. 37: 630. 1906, fide Killip. C. subserrata Krause, I.e. 627.
Branchlets and leaves beneath more or less fulvous or cinereous
tomentose-pilose; petioles 4-10 mm. long; leaves elliptic or ovate-
elliptic, acute both ends, varying in size from about 3 cm. long, 2 cm.
wide, to a dm. or so long, 4 cm. wide or under, submembranous to
coriaceous, entire or subserrate toward tip, rugose, sparsely hispid
both sides or tomentulose beneath; spikes terminal and axillary,
usually 3-5 cm. long; calyx densely pilose, lobes ovate, acute, the
slightly longer glabrous or subglabrous corolla tube about 4 mm. long,
the lobes short; filaments affixed about medially, dilated and villous
toward base; style subequaling calyx. According to Johnston, Journ.
Arnold Arb. 30: 121. 1949, C. curassavica (Jacq.) Roem. & Schult.,
ranging north of Colombia and similar, differs constantly in the com-
plete lack of axillary spikes.
Cajamarca: Socota to Tambillo, Stork & Horton 10178 (det.
Standley). Huanuco: Acomayo and Pillao, Ruiz & Pavdn, type;
Woytkowski 34002. Chaglla, 3641 . Junin : Near Palca, Weberbauer
1758; Killip & Smith 23372. Cuzco: Marcapata, near Cachu-
pampa, Vargas 9696. Arequipa: Mollendo, Weberbauer 1530.
Cordia hebeclada Johnst. Journ. Arnold Arb. 31: 176. 1950.
Tree to 15 meters tall with regular dichotomous branches, dimor-
phic leaves and strongly heterostyled flowers, unisexual by abortion;
branchlets and leaves beneath densely puberulent; petioles often
1-2 cm. long; leaves laxly or appressed subscabrous above, the
larger elliptic-lanceolate or -ovate, rounded at base, more or less
acuminate, 1.5-2.5 dm. long, 7-15 cm. long, the smaller suborbicular
or orbicular-ovate, frequently 8-10 cm. long, often truncate or sub-
cordate at base, rounded or obtuse at apex; inflorescence corymbose,
1-3 dm. across; male flowers with cupulate calyx 3-4.5 mm. long,
densely puberulent but strigose within, the unequal lobes acute;
corolla 6-8 mm. long, oblong recurved lobes and tube 3-4 mm. long;
filaments about 4 mm. long, villous basally; ovary rudimentary,
glabrous; female flowers similar but calyx subcylindric, 2.5-3 mm.
long, filaments glabrous or very sparsely villous; style exserted,
3.5-4 mm. long, bifid; drupe glabrous, depressed globose, strongly
rugose, to 1 cm. long, 8 mm. broad, the expanded fruiting calyx
4-5 mm. across. Probably most closely related to C. panamensis
FLORA OF PERU 579
Riley, but differing in indument and in its regular dichotomous
branching; in C. hebeclada the indument consists of a great abundance
of minute trichomes that give a thin downy uniform very soft
cover (Johnston), scarcely soft in age; perhaps an Andean variant
of C. tetrandra Aublet in which the author included it, Contr. Gray
Herb. 92: 55. 1930.
San Martin: Morales near Tarapoto, Williams 5676. Chazuta,
Klug 4050. Bolivia to Colombia. "Tahnampa-caspi" (Williams);
"quitasol" (Little), or "jutumbe" (Tafalla) in Ecuador. "Rena-
quillo" (Klug).
Cordia iguaguana Melchior ex Johnst. Journ. Arnold Arb. 33 :
63. 1952.
Tree, 8-20 meters tall, branchlets finely fulvous tomentulose, gla-
brate in age, petioles to about 2 cm. long; leaves broadly lanceolate,
obtuse or rounded at base, acute or somewhat attenuate at apex,
1-1.5 dm. long, 4-5 cm. wide, glabrous or minutely and sparsely
appressed pubescent, the 5-7 lateral nerves and many veins lightly
impressed above, rather prominent beneath; paniculate or corymbose
inflorescences terminal on young leafy branches, about 1.5 cm. across;
calyx clavate-cylindrical, prominently 10-costate, about 11 mm. long,
3 mm. thick toward the apex, obscurely appressed-puberulent, the
3-5 often unequal obtuse lobes about 1 mm. long, the sinuses obtuse;
corolla white, 23 mm. long, limb about 2.5 cm. broad, lobes 6-8 mm.
long, 8-10 mm. wide, often emarginate, tube 8 mm. long, 1-1.5 mm.
thick, tomentulose within below the middle, the throat 10-15 mm.
across, about 5 mm. deep; filaments glabrous, affixed near apex, 1-
3 mm. long, the anthers also unequal, 1.5-3 mm. long; ovary with
style about 13 mm. long (nectary lacking), forked about 1 cm. above
the fusiform base. Section Gerascanthus with three other species
known from western South America; C. alliodora (R. & P.) Oken has
stellate indument and is myrmecophilous, the other species not and
furthermore their indument is simple; C. Goeldiana Huber, Amazonian
Brazil to Colombia, has long oblong corolla lobes while C. macrantha
Chodat, of western Ecuador, with the broad rounded corolla lobes of
C. iguaguana, has corollas 3.5-4 cm. long, larger elliptic leaves
evidently villous aspidulose beneath, and differs otherwise according
to Johnston, who, agreeably, took up the well-chosen herbarium
name of Melchior. F.M. Neg. 975.
Cajamarca: Near Jaen, 700-800 meters, Weberbauer 6218, type.
"Iguaguana."
580 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Cordia insignis Cham. Linnaea 8: 122. 1833. C. Haenkeana
Mez, Bot. Jahrb. 12: 560. 1890, fide Johnston, Journ. Arnold Arb.
16: 179. 1935.
Branchlet tips densely more or less ferrugineous tomentulose with
simple trichomes as the terminal corymbose-paniculate inflorescences,
the ultimate branches of these few-flowered; petioles to 4 mm. long;
leaves ovate or suboval, rather rounded at base, basally narrowed to
finally rounded apex, about 14 cm. long, 9 cm. wide, subglabrous and
lustrous above, strigose beneath where prominently reticulate-
veined; calyx costate, 2.5 cm. long, anthers often with few irregular
teeth; corolla to 4 cm. long, coriaceous, persisting, the 5 lobes
emarginate; stamens nearly 2 cm. long, affixed near base, joined
annulately, included; ovary glabrous. Description after Mez, who
remarked: evidently differs from C. insignis in form of leaves and
inflorescence. I greatly doubt that the Haenke collection came from
Peru; C. insignis is known only east and south of the Amazon
Basin, from eastern Brazil to eastern Bolivia, and is certainly not
to be expected in Peru (Johnston). There are apparent exceptions
known to this range unless determinations have been wrong; C.
insignis may have ovate-oblong leaves, flowers in dense cymes.
Illustrated, Mart. Fl. Bras. 8, pt. 1, pi. 1 (C. insignis}.
Peru(?) : See note above; without locality except "Peru: Haenke"
(type, herb. Monac.).
Cordia Krausiana Killip, Journ. Wash. Acad. Sci. 17: 330. 1927.
C. pauciflora Krause, Bot. Jahrb. 37: 629. 1906, not Rusby, 1896.
Flowering branches somewhat ferrugineous tomentose as the
3-5 mm. long petioles; leaves elliptic or ovate-elliptic, acute both
ends, in type 2.5-3 cm. long, 12-16 mm. wide, apparently often
larger, subcoriaceous, rugose, hispidulous and the veins impressed
above, tomentose and the veins conspicuous beneath, clearly, even
coarsely serrate; flowers capitately congested, few in type or in
vigorous examples many; calyx 5-6-parted, the pilose subacute seg-
ments 3 mm. long; corolla tube glabrous, little exserted, the lobes
rounded; filaments affixed just above middle, short, villous at dilated
base; style slightly exceeding calyx, stigma globose. Related to C.
scaberrima HBK. but with smaller, more strongly serrate leaves
(Krause); if the shrub is specifically distinct it seems vegetatively
at least to differ only in the serrate leaves, these in my collections
to 8 cm. long, 3 cm. wide, the flower-heads in fruit 2 cm. thick, the
calyx and drupe about 6 mm. long. On the other hand this and the
FLORA OF PERU 581
plant of HBK. are doubtfully distinct specifically from the Tarapoto
shrub here referred to C. bifurcata but not typical. F.M. Neg. 991.
Ancash: Tamanco to Huaraz, 3,000 meters, Weberbauer 3161,
type. Lima: Matucana, 180; 288k (det. Killip, C. lantanoides) .
Gordia lantanoides Spreng. Syst. 1: 653. 1825. Varronia cre-
nata R. & P. Fl. Peruv. 2: 23, pi. 146b. 1799, not C. crenata Delile,
1813.
Branches tortuous, the many leafy flowering ones minutely hir-
sutulous; petioles slender, to about 6 mm. long; leaves subrotund to
oval-elliptic, rounded both ends, 2-3 cm. long, 1.5-2 cm. wide,
crenate-serrulate nearly to base, submembranous, spreading hispidu-
lous both sides, slightly asperous and rugulose above, reticulate
venose beneath; peduncles axillary, 2-3 cm. long, to 4-dichotomous
in fruit but the flowers early in small heads; calyx glabrate in fruit,
about 4 mm. long, the ovate lobes acute, the white corolla little
exserted; drupe red, seed ovoid, rarely 2-celled (Ruiz & Pavon).
Apparently not found in typical state since the original collection
but possibly should be drawn to include C. Krausiana Killip. F.M.
Neg. 980. *
Huanuco: Ruiz & Pavdn, type.
Cordia laevior Johnst. Journ. Arnold Arb. 16: 180. 1935.
Branchlets soon glabrescent, only toward tips shortly pubescent
with appressed or ascending trichomes; petioles 5-10 mm. long;
leaves similar, oblong-lanceolate, acute at base, long-acuminate,
1.5-2.5 dm. long, 4-10 cm. wide, glabrous above except the hir-
sutulous midrib and 7-8 lateral nerves (tertiary obscure), paler
beneath where puberulent on the many veins; cymes in forks of
branches, the lower branches spreading; calyx in bud obovate,
4-5 mm. long, 2-3 mm. thick, rounded apically, obscurely costate,
densely puberulent, the triangular lobes somewhat uneven; corolla
tube 4 mm. long, lobes 2 mm. wide and long; style sparsely pilose,
the filaments toward base, the ovary evidently toward apex. Less
pubescent than the related more eastern C. Sprucei Mez and cyme
more loosely branched; leaves not roughened above by the many
veinlets; distributed as C. Ulei Johnst. with glabrate ovary, coarser
pubescence (author). It seems desirable to study these species
with more material before their true status (as that of C. ripicola)
can be discovered. Type a tree 6 meters high.
San Martin: Pongo de Cainarachi, (Klug 2756, type).
582 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Cordia lomatoloba Johnst. Journ. Arnold Arb. 18: 12. 1937.
Arborescent, to 25 meters tall, nearly glabrous; leaves similar,
lanceolate or elliptic-lanceolate, narrowed to petiole this 8-15 mm.
long acute or acuminate, 8-14 cm. long, 2-5.5 cm. wide, glabrous
or obscurely strigose, reticulate both sides, lustrous above, paler and
opaque beneath, entire; primary nerves 6-8 pairs; cymes often
somewhat brown-pubescent in the forks of the branches, these lax,
slender; calyx sessile, pyriform, 2.5 mm. long, abruptly contracted
to base, the deltoid equal teeth only marginally obvious; corolla
white, 4 mm. long, tube 1 mm. long, lobes about 1.8 mm. wide,
oblong, recurved, rounded at tip; filaments villous below; ovary
glabrous. Related to the pubescent C. naidophila Johnst. of the
Amazon, distributed as C. ecalyculata Vahl with much narrower
leaves, while C. diospyrifolia Cham, as to type by Sellow has cori-
aceous broader leaves; the characters may prove to be variable.
San Martin: Chazuta, King 4050? Loreto: Mouth of the Rio
Santiago, Tessmann 4454 (det. Melchior, C. diospyrifolia). Rio Acre:
Mouth of Rio Macauhan, Krukoff 5497, type; also 5345. Brazil.
Cordia lutea Lam. 111. 1: 421. 1791. C. marchionica Drake, 111.
Fl. Ins. Pacific 240 (1892), fide Johnston, Journ. Arnold Arb. 33: 65.
1952. C. rotundifolia R. & P. Fl. Peruv. 2: 24, pi. 148a. 1799.
Flowering branchlets many, canescent-lanate to puberulent in-
cluding the conspicuously ribbed calyces of the showy pale yellow
flowers; petioles 1-2.5 cm. long; leaves subrotund to broadly ovate-
elliptic, mostly 4 or 5 cm. wide and little longer, sometimes much
larger, weakly crenulate, early canescent short-pilose beneath, harshly
scabrous above; calyx soon obconic, about 1 cm. long, teeth subulate;
corolla a clear light yellow, 3 cm. long or longer, the throat dilated,
the limb 6-8 lobed; filaments villous at base; fruit ovoid, white.
Johnston, I.e., has presented arguments that seem irrefutable that
the common occurrence of this species in the Marquesas is due to
its introduction by the French during the nineteenth century, prob-
ably as an ornamental. A decoction is used with good results for
jaundice (Ruiz & Pavon); an attractive yellow-flowered bush along
streams. F.M. Neg. 994.
Tumbez, fide Diels. Piura: Near La Brea, (Haught & Svenson
11596}; Weberbauer 5955. Cajamarca: Huambos, Stork & Norton
10205. Lambayeque: Chiclayo, West 3575. Ancash: Tambo de
Pariacota, 2557. Talara, Horton 11516. Pabur, Stork 11422.
Lima: Lurin, Chan cay, Ruiz & Pavon (type, C. rotundifolia}; Dom-
FLORA OF PERU 583
bey, type. Huanuco: Maranon Valley, Weberbauer, 155. Loreto:
Mishuyacu, near Iquitos, Klug 1482. Western Ecuador; Galapagos
Islands. "Tina" or "membrillejo" (quince); "oberal" (Weberbauer).
Cordia macrocephala (Desv.) HBK. Nov. Gen. & Sp. 3: 77.
1818. Varronia macrocephala Desv. Journ. de Bot. 1: 273. 1808.
C. polyantha Benth. PI. Hartw. 121. 1843? C. guayaquilensis DC.
Prodr. 9: 496. 1845.
Often many-stemmed, matted or erect and sometimes 5 meters
tall, the flowers in well-peduncled heads, the coarsely crenate or ob-
tusely serrate leaves greenish but more or less scabrous both sides,
conspicuously veined, to about 6 cm. long, 2.5 cm. wide, often smaller,
ovate- or oblong-elliptic, obtuse or subacute. Johnston, Journ. Ar-
nold Arb. 30: 89. 1949, remarks that the similar C. polyantha Benth.,
PI. Hartw. 121. 1843, "of Ecuador and northern Peru," and C. macro-
cephala (Desv.) HBK., of Peru, have a relative in C. Pringlei Robins,
of more northern range. A pubescent or glabrate shrub striking by
the globose inflorescences of white flowers. Pubescence of unfolding
leaves during dry period may consist of closely packed stellate scales
while in the wet period stellate and single hairs are greatly elongate
and forked hairs are on some specimens (Svenson, Am. Journ. Bot.
33: 479. 1946). F.M. Neg. 984.
Piura: Amotape Hills, (H aught & Svenson 11523). Near Frias,
Weberbauer 6433. Serran, Stork 11373 (det. Johnston, C. polyantha).
Lambayeque, fide Weberbauer. Lima: Chosica, 542; Mexia 04009.
Sta. Eulalia, Goodspeed 33018; Norton 10994. Sayan, Goodspeed
33042; 33046. Rio Rimac Valley, Goodspeed, Weberbauer & Metcalf
30217. Junin: Huacapistana, Killip & Smith 24518 (det. Killip,
C. floribunda). To Colombia.
Cordia macrodonta Killip, Journ. Wash. Acad. Sci. 17: 328.
1927.
Branchlets quadrate, ferrugineous-puberulent and finely pilosu-
lous (as leaves beneath), scabrous; petioles 5-10 mm. long; leaves
broadly ovate, narrowed to base, abruptly short-acuminate, 6-12 cm.
long, 4-6.5 (8) cm. wide, coarsely and acutely serrate except in lower
third, membranous, shortly strigillose-hispid above, lateral nerves
6-7 pairs; peduncles 4 cm. long; flowers solitary (or few) at ends of
the inflorescence branchlets; calyx globose-turbinate, 2 mm. long,
appressed rusty-strigillose, the acute teeth 0.5 mm. long; corolla tube
cylindric, 3.5 mm. long, stamens in throat; fruit ovoid-conic, 5 mm.
584 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
long, glabrous. After author, who remarked: distinguished from
other members of the Corymbosae DC. by the coarsely serrate
leaves; may prove to be a part of the variable C. bifurcata Roem. &
Schult. as suggested by Stork 11417.
Piura: San Antonio, Prov. Huancabamba, Weberbauer 6015, type.
Cordia Mexiana Johnst. Journ. Arnold Arb. 18: 12. 1937.
Tree, 5-7 meters, with puberulent branches; leaves elliptic or
oblong-elliptic, abruptly contracted to petiole, this 1-2.5 cm. long,
obtusely acuminate, 1.5-4 dm. long, 8-20 cm. wide, coriaceous, retic-
ulate both sides, glabrous, lustrous and minutely puncticulate above,
subpuberulent beneath; nerves 5-7 pairs; cymes laxly much branched,
2.5 dm. across, the flexuose branchlets strigose puberulent; calyx
(bud) about 6 mm. long, 2 mm. thick, stouter above, densely brown
strigose, puberulent within above the middle, the 5 deltoid erect lobes
about 1 mm. long; corolla white, 12 mm. long, tube 1.5 mm. thick,
4 mm. long, throat obvious, 2.5-3 mm. long, gradually expanded to
about 3 mm. across, lobes oblong, 4 mm. long, recurved, rounded;
filaments exserted, villous below; ovary glabrous; style deeply forked.
Distributed as C. colombiana Killip of Ecuador and Colombia but
distinct because of the elongate corolla with well-developed throat;
this character, and the long tube, are very unusual in the Pilicordia
section (author).
Loreto: Rio Maranon near Rancho Indiana, Mexia 6459, type.
Adjacent Colombia.
Cordia mollissima Killip, Journ. Wash. Acad. Sci. 17: 330. 1927.
Densely lanate-tomentose even the contracted or early subelliptic
few-flowered inflorescences; branches terete; petioles to 1 cm. long;
leaves ovate-lanceolate, rounded or subacute at base, acute or acumi-
nate, to 7 cm. long, 3.5 cm. wide, serrulate, reticulate-veined, dark
green above, paler beneath; calyx campanulate, about 2 mm. across,
the deltoid lobes acute; corolla 3-3.5 mm. long; stamens and style
about 1.5 mm. long, the ovate-oblong anthers scarcely exserted.
Related to C. corymbosa (L.) Don. (C. bifurcata Roem. & Schult.)
but differing in the white indument and the closely serrulate leaves
(author).
Cajamarca: Jaen, Weberbauer 6202, type. Amazonas: Chacha-
poyas, (Mathews 3134). San Martin: Juanjui, King 4385 (det.
Standley, C. scaberrima} .
FLORA OF PERU 585
Cordia nodosa Lam. Tab. Encycl. 1: 422. 1791; 46 and 13. C.
hispidissima DC. Prodr. 9: 475. 1845.
Distinctive by enlarged nodes below each fork, these hollow, ordi-
narily inhabited by ants; marked also by the divaricate rufous or
brownish bristles that are usually more or less developed about the
apiculate calyx tip; petioles 2-5 mm. long; leaves usually subopposite
or whorled, somewhat dissimilar, often elliptic, 1-3 dm. long, obtuse
at base, acuminate, lustrous, more or less bullate at limb in age, the
6-10 pairs of nerves impressed, reticulate; paniculate cymes sessile or
peduncled, up to a dm. across, usually puberulent with crinkly curved
trichomes or the papyraceous scarcely ribbed calyx strigillose, 5 mm.
long in bud, unevenly lobed, ordinarily persisting and becoming
fibrose; corolla white, 4-6 mm. long; style and ovary pubescent
(filaments basally) ; fruits yellow, usually more or less bristly, about
1.5 cm. long. Shrub or tree sometimes 10 meters high, few branches,
bark fissured with many small lenticels (Williams, Field Mus. Bot.
15: 433. 1936), nearly smooth except at nodes to densely bristly
(forma hispidissima). I. W. Bailey, Bot. Gaz. 77: 32-49, pis. 6-7.
1924, gave an account of myrmecophilism here, a curiosity occurring
among Peruvian plants, notably in Melastomaceae and Moraceae;
this intriguing development was investigated first by A. F. W.
Schimper, Bot. Mitteil. Trop. Heft 1: 53-. 1888, and by Schumann,
Pringsheim's Jahrb. 382-. 1888.
San Martin: Tarapoto, Williams 6516. Juanjui, King 3775.
Huanuco: Tingo Maria, 700 meters, Stork & Horton 9519. Valley
of the Monzon, Prov. Huamalies, Weberbauer 3679; 285. Junin:
Rio Pichis, Killip & Smith 26731. Hacienda Schunke, 1,400 meters,
Killip & Smith 24582. Perene" Bridge, Killip & Smith 25359
Loreto: Florida, King 2158; 2305. Rio Nanay, Williams 330 A;
1229. Caballo-Cocha, Williams 2334- Rio Mazan, Jose Schunke 10.
Yurimaguas, Killip & Smith 27688; 27728; Williams 4305; 4677.
Iquitos, Williams 3748; 7974; Killip & Smith 27022; King 480; 510;
803. La Victoria, Williams 2679. Rio Itaya, Killip & Smith 29602.
Bolivia to the Guianas. "Almenchillo," "analliocaspi" or "ayahuas-
co" (Williams); "ibrachoreri" (Huitoto); "pucacurocaspi" (Schunke).
Cordia patens HBK. Nov. Gen. & Sp. 3: 75. 1818.
Young branchlets and leaves beneath densely flavescent tomentu-
lose; petioles 4 mm. long; leaves ovate-oblong, rounded at base, acute
or acuminate, to 7.5 cm. long or longer, rather remotely serrulate
toward tip, firm, reticulate and strigose-scabrous above, the veins and
586 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
nerves prominent beneath; spikes subglobose, axillary and terminal
on slender peduncles 5 cm. long; calyx strigose-hispid with minute
subulate teeth, villous in the throat as the white twice as long corolla;
stamens exserted, exceeded by the style. After HBK. Inflores-
cences axillary, not extra-axillary as in C. bifurcata (Johnston) ; Pe-
ruvian collection was referred to C. bifurcata (C. corymbosa (L.) Don)
by Standley and Killip and the leaves are not yellowish tomentulose
beneath as are those of the type from Venezuela, which in this char-
acter resembles Tarapoto material. F.M. Neg. 38979.
Loreto: Yurimaguas, Killip & Smith 28262; Williams 3898; 3947;
4469; 4787. Contamana, Rio Ucayali, Killip & Smith 26876. To
the Guianas.
Cordia peruviana Roem. & Schult. Syst. 4: 459. 1819. Varronia
obliqua R. & P. Fl. Peruv. 2: 24, pi. 147b. 1799, not C. obliqua Willd.,
1797.
Similar to C. spinescens but leaves coarsely and irregularly serrate,
obliquely decurrent at base, mostly 5-6 cm. long, 2.5-3 cm. wide
somewhat below the middle, submembranous, lightly scabrous and
in age rugulose above, hispidulous on the reticulate veins beneath;
spikes puberulent, long-pedunculate in the upper axils, several cm.
long in fruit; calyx lobes acute. Determinations by Killip; seems to
have little to recommend it as a distinct species but the leaf-indenta-
tion is rather more than an extreme variation of that of C. spines-
cens L.
Ancash: Tambo de Pariacota, 2546. Lima: Chosica, 2862. Ma-
tucana, 302. Junin: Tarma, Ruiz & Pavdn, type.
Cordia Poeppigii DC. Prodr. 9: 492. 1845.
Open or rather compact shrub, the long flowering branches often
supported in thickets, early more or less hispidulous with spreading
subappressed or incurved trichomes; petioles 5-10 mm. long, decid-
uous above the base; leaves ovate-elliptic, more or less rounded at
base and acutely acuminate, a dm. or so long, 4-6 cm. wide, those in
the inflorescence reduced, all except type ex char, sparsely scabrous
with incurved trichomes above, hispidulous beneath especially on
the slender nerves and rather obscure veins, submembranous, not or
obscurely rugulose; inflorescence when well-developed composed of a
number of slender branchlets from upper leaf-axils, and finally 3 dm.
long or longer; calyx subpapyraceous, glabrous or nearly, at least
typically, about 2.5 mm. long, the ovate lobes acute; corolla greenish-
FLORA OF PERU 587
white, campanulate, 4.5 mm. long, the subglobose anthers barely
exserted from the subtruncate limb, and well-exceeded by the slender
forked style; fruit ovoid, 4 mm. long, finally red, the calyx not or
tardily splitting. Type, "Central Peru, Poeppig." Determinations
by Killip. F.M. Neg. 7735.
San Martin: Near Tarapoto, Williams 6969. Huanuco: Chico-
playa, Ruiz & Pav6n. Pozuzo, 456 1. Pampayacu, 5042. Junin:
Colonia Perene", Killip & Smith 24990. Loreto: Iquitos, Tessmann
3560 (det. Melchior, C. floribunda); Williams 3543; 8031; Killip &
Smith 271 27; King 1 344; Mexia 6507. Caballo-Cocha, Williams 2060
(toward C. scaberrima?). "Barillas" (Mexia).
Cordia ripicola Johnst. Journ. Arnold Arb. 16: 180. 1935.
Resembles C. laevior; branchlets scabrous with short trichomes;
petioles 2-6 mm. long, leaves uniform, more oblong or somewhat
obovate, 8-14 cm. long, 3-7 cm. wide, sparsely appressed strigillose
above, closely and appressed hispidulous-puberulent beneath; calyx
(type) strigose, subpapyraceous, unevenly rupturing into 2-5 lobes;
corolla lobes about 2.5 mm. long; style and ovary sparsely pubescent
or the latter glabrous. Calyx character, appressed leaf pubescence
and pubescent pistil separates this from C. Ulei Johnst. and the
scattered appressed trichomes and much appressed ones beneath
from C. Sprucei Mez. (author) of southwestern Brazil. The Schunke
specimen is nearly glabrous, or only scabrous; cf. C. laevior Johnst.
San Martin: Juanjui, Klug 4231 (det. Standley, C. Sprucei).
Loreto: Mouth of Rio Zubineta, (Klug 2262, type). Rio Mazan,
Jose Schunke 292 (det. Killip). "Chore-ey."
Cordia scaberrima HBK. Nov. Gen. & Sp. 3: 71. (55). 1818.
Branches glabrous, younger tetragonous, scabrous as the leaves
both sides, these rugose, serrate, oblong, narrowed to petioles (4 mm.
long), rigid, reticulate- veined, acuminate or acute, the nerves and
midrib strigose scabrous above, veins scabrous or pilose-hispidulous
beneath; corymbs terminal, dichotomous, the heads sometimes 2 cm.
long, 1.5 cm. thick, often tardily forked; calyx striate, hispid or more
or less hispidulous, 5 mm. long in fruit with broad ovate lobes.
The Junin specimen has leaves in shape toward var. ecuadorica Killip
but subentire, and an ellipsoid tardily forked inflorescence. The
Williams specimen with peduncles 2 cm. long, heads globose, 7 mm.
thick, calyx 3 mm. long may be the same as one by Poeppig referred
by Johnston with doubt to C. buddleoides Rusby, the calyx without
588 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
the subulate tips (appendages) that are very evident on that species.
The Huanuco plant, var. ecuadorica Killip, in herb, foliis plus mi-
nusve serratis dense piloso-hirsutis, may be distinct and indeed has
been so designated under a similar name by Johnston. The pedun-
cles are leafy, the globose heads to 2.5 cm. in diameter, calyx in fruit
to 7 mm. long. F.M. Neg. 997.
Piura: Ayavaca, Bonpland, type. Huanuco: Mito, 3328. Ya-
nano, 3789. Junin: Huacapistana, Killip & Smith 24518 (det.
Killip, C. floribunda). Loreto: Santa Rosa, Yurimaguas, Williams
4779 (det. Killip). Ecuador.
Cordia scabrifolia A. DC. Prodr. 9: 485. 1845; 53.
Branchlets and inflorescences minutely puberulent; leaves elliptic-
lanceolate, acute at base, acutely short-acuminate, not markedly
dissimilar, shortly appressed scabrous-strigose above or pustulate,
sometimes smooth and glabrous(?), beneath green, on the nerves
pubescent with some very short, usually inwardly directed stout tri-
chomes; calyx cupulate, about 3 mm. long, nearly as broad, the lobes
about even; corolla about 5 mm. long; anthers nearly 2 mm. long,
exserted; ovary pubescent. No doubt occurs in Peru.
Rio Acre: Seringal Auristella, Ule 9717. Mouth of Rio Macauhan,
Krukoff 5336; 5672. Amazonian Brazil; British Guiana.
Gordia spinescens L. Mant. 2: 206. 1771. Varronia ferruginea
Lam. Tab. Encycl. 1: 418. 1791; Poir, Encycl. 4: 263. 1797. C. fer-
ruginea (Lam.) Roem. & Schult. Syst. 4: 458. 1819. C. pyrifolia
Willd. ex Roem. & Schult. I.e. 802. C. Thibaudiana DC. Prodr. 9:
489. 1845. C. pauciflora Rusby, Mem. Torr. Bot. Club 6: 83. 1896.
Petiole bases persisting spur-like on the upper branches, the
branchlets, petioles and peduncles more or less tomentulose or red-
dish-hirsute; petioles 5-10 mm. long, leaves often broadly ovate-
elliptic, 7-15 cm. long, about half as wide, acute both ends or more
or less abruptly acuminate at apex, scabrous above, more or less
villous-hispidulous beneath, entire to obsoletely to coarsely serrulate,
firm to subcoriaceous, the nerves and reticulate veins correspondingly
conspicuous; peduncles on the upper petioles or several in a terminal
inflorescence, 1-several cm. long; spikes often few-flowered, some-
times ellipsoid, calyx somewhat campanulate, to about 4 mm. long,
the ovate lobes acute, corolla 5-8 mm. long, tube glandular (Worth &
Morrison). The persisting indurated spur-like petiole bases sug-
gested the name according to Johnston, who worked out the synon-
FLORA OF PERU 589
ymy, Journ. Arnold Arb. 30: 104. 1949; includes also the Colombian
C. laxiflora HBK. and C. riparia HBK. Nov. Gen. & Sp. 3: 72. 1818,
and 71, pi. 207 (C. riparia). But it is not clear that the breaking
down of the petiole cells does not occur irregularly and its taxonomic
significance is therefore debatable. F.M. Neg. 1020 (C. pyrifolia).
Arequipa: South of Chala, Worth & Morrison 15697. Bolivia
to Mexico.
Cordia Sprucei Mez, Bot. Jahrb. 12: 549. 1890; 53. Lithocar-
dium Sprucei (Mez) Ktze. Rev. Gen. 2: 977. 1891.
Branchlets apically strigose, subferrugineous-tomentulose, gla-
brate, terete or nearly; petioles to 7 mm. long, geniculately incurved
at base; leaves suboval or obscurely obovate, to 1.5 dm. long, 7 cm.
wide, attenuate to rounded base, acutely acuminate, subglabrous
and very lustrous above, pubescent beneath, prominently reticulate
both sides; inflorescence terminal, laxly paniculate, ferrugineous to-
mentulose, much shorter than leaves; calyx before anthesis entire,
urceolate, ecostate, irregularly much lobed, very slightly strigose
within; corolla with stamens to 8 mm. long, more than three-fourths
divided, the 5 rounded lobes finally reflexed; filaments affixed nearly
at throat, long pilose at base, anthers glandular at tip; ovary yellow-
ish pilose as style at base; stigma discoid; drupe tomentulose, to
12 mm. long, 7 mm. thick, the calyx scarcely enlarged. After author,
who placed it in section Pilicordia; species confused by Fresenius,
in Mart. Fl. Bras. 8, pt. 1: 16, with C. umbraculifera A. DC.
Peru (perhaps). Rio Negro, Brazil.
Cordia toqueve Aublet, PI. Guian. 1: 228, pi. 90. 1775; 52.
C. heterophytta Poir. Diet. Sci. Nat. 10: 409. 1818, fide Johnston.
Branchlets, slenderly branched inflorescence and leaves beneath
softly puberulent-tomentulose; leaves strongly dissimilar, those on a
single branch suborbicular to broadly ovate, alternate or opposite;
larger, long-elliptic, narrowly ovate or lanceolate-oblong ones, often
acute or narrowed to base, rounded or acute at apex, finely pubes-
cent, especially on midrib and lateral nerves or the younger velvety
(Schunke); calyx short conic or proculiform, about 3.5 mm. long,
opening unevenly; corolla about 4 mm. long; anthers oblong, scarcely
1 mm. long. At once distinct from all relatives with strongly hetero-
morphic foliage by the obviously pubescent ovary, fruit and leaves
(Johnston).
590 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XIII
Loreto: Lower Rio Nanay, Williams 578. Mouth of Rio Santi-
ago, Tessmann 4125. Rio Mazan, Jose Schunke 360 (det. Killip).
To eastern Brazil and northern South America. "Bacuri" (Wil-
liams).
Cordia ucayaliensis Johnst. Journ. Arnold Arb. 16: 181. 1935.
C. Ulei Johnst. var. ucayaliensis Johnst. Contr. Gray Herb. 92: 57.
1930.
Resembles C. Ulei but leaves more obovate-elliptic, about 1.5 dm.
long, 8-9 cm. wide, and, especially, finely, sometimes sparsely ap-
pressed strigose and scabrous above, rather softly hispidulous to
puberulent beneath; fruiting calyx scarcely lobulate, 8 mm. across,
the ellipsoid fruits 12 mm. long, about 7 mm. thick. The validity
of the above name may be questioned as the place of publication of
the synonym is not given; readily distinguished from C. ripicola by
the scabrous-strigose upper leaf surface (author). Type a 6-meter
tre 
