580.5

FB

V.7

1930

cop. 2

Inc.

20 193}

FIELD MUSEUM OF NATURAL HISTORY

FOUNDED BY MARSHALL FIELD, 1893

PUBLICATION 285

BOTANICAL SERIES VOL. VII, No. 2

THE LIBRARY OF 1HE

UNIVERSE Of

THE RUBIACEAE OF ECUADOR

BY

PAUL C. STANDLEY

ASSOCIATE CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY

B. E. DAHLGREN

' ACTING CURATOR, DEPARTMENT OF BOTANY

EDITOR

CHICAGO, U. S. A.

FEBRUARY 5, 1931

Dup.

U. of C.

Oh.

L'ftlVL

URBAN*

FIELD MUSEUM OF NATURAL HISTORY

FOUNDED BY MARSHALL FIELD, 1893

PUBLICATION 285

BOTANICAL SERIES VOL. VII, No. 2

THE LIBRARY OF THE

JUL 241943

UNIVERSITY Of ILLINOIS

THE RUBIACEAE OF ECUADOR

BY

PAUL C. STANDLEY

ASSOCIATE CURATOR OP THE HERBARIUM, DEPARTMENT OF BOTANY

B. E. DAHLGREN

ACTING CURATOR, DEPARTMENT OF BOTANY

EDITOR

CHICAGO, U. S. A.

FEBRUARY 5, 1931

PRINTED IN THE UNITED STATES OF AMERICA

BY FIELD MUSEUM PRESS

Jte. Unir.

V, /'

THE RUBIACEAE OF ECUADOR

PAUL C. STANDLEY

Botanically, Ecuador is least known of any of the countries that

lie along the Andes. Colombia has been explored extensively,

although far from completely, during the past ten years, and huge

collections of plants have been received from Peru during the same

period. The Bolivian flora has been collected somewhat intensively

over an even longer series of years.

In Ecuador, however, few collections, and those small and alto-

gether unsatisfactory ones, have been made recently. Perusal of the

following pages will prove that little has been added to the knowl-

edge of Ecuadorean Rubiaceae by late collections. What we do

know of this family as it is represented in Ecuador is chiefly the

result of the work of the older collectors: Humboldt and Bonpland;

Hartweg; the picturesque Scotchman, Jameson, who resided for

many years in Quito; and the indefatigable Spruce, who spent several

rather painful although botanically profitable years in Ecuador.

Sodiro botanized extensively in the country, but I have seen only a

scattered few specimens of Rubiaceae of his collection. The work of

Baron Eggers along the coast added a substantial number of species

to the list here enumerated, and the handsome specimens obtained

more recently by the Swedish collectors, Holmgren and Heilborn,

have added still others. A good many species are represented only

in the collections of Andre* and Lehmann, both of whom are known

better for their work in Colombia.

The only extensive recent collections from Ecuador are those of

Dr. J. N. Rose and Professor A. S. Hitchcock, made for the United

States National Museum, Gray Herbarium, and New York Botanical

Garden, but neither of these adds much information regarding

the Rubiaceae.

Most of the records of Ecuadorean Rubiaceae are the result of

the labors of Jameson and Spruce. A few years ago I had the

good fortune to travel from New York to Panama in company

with an elderly Ecuadorean gentleman who had studied botany

under Jameson. He related many anecdotes of his teacher to illus-

trate his eccentricities and his all-absorbing interest in botanical

179

180 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

exploration. It is some satisfaction to know that Jameson was

honored by having named in his memory one of the most striking

and curious genera of Andean ferns, a group inhabiting the cold

paramos in which he delighted to botanize.

The Rubiaceae constitute one of the largest and most important

families of the flora of Ecuador. Of particular interest here is the

genus Cinchona. It was from the region of Loja, in southern Ecuador,

that the virtues of these plants were first made known to the world,

and Loja was long the principal center of production of Peruvian bark.

The number of species recorded in this paper is small in com-

parison with the nearly 500 species reported from Colombia, or with

the list already compiled for Peru. There is no doubt that further

exploration will add enormously to the known Rubiaceae of Ecuador,

and probably double them. At the present time nothing is known

of those growing along the Pastasa River and elsewhere on the eastern

slopes of the Andes. Some genera and species are known from

Colombia and Peru, but not from Ecuador, although there is every

reason for assuming that they do grow in the intervening country.

The present enumeration includes the species native in the

Galapagos Islands. Their inclusion is justifiable only on political

grounds, although the few Rubiaceae existing in the Galapagos,

chiefly species of Borreria, perhaps are as closely related to those

of Ecuador as to those of any other region.

This list of the Rubiaceae of Ecuador is the result of the study

of collections in the herbarium of Field Museum of Natural History

(indicated on the following pages by the letter F), and in the herbaria

of the following institutions: Royal Botanic Gardens, Kew (K);

Botanic Garden and Museum, Berlin-Dahlem (B); Riksmuseets

Botaniska Afdelning, Stockholm (S); Jardin Botanique Principal,

Leningrad (L); United States National Museum, Washington (W);

Gray Herbarium of Harvard University (G); New York Botanical

Garden (Y); and the Academy of Natural Sciences of Philadelphia

(P). To those in charge of these collections, who have lent material

for study or have forwarded it for determination, the writer wishes

to express his deep appreciation of the courtesies thus extended.

Of special service in preparing this enumeration have been the

excellent photographs of type specimens of South American plants

obtained at Berlin by J. Francis Macbride through a fund generously

provided for the purpose by the Rockefeller Foundation. Special

thanks must be given to the Director, Dr. L. Diels, and to the staff

of the Berlin Garden for their whole-hearted cooperation in the

THE RUBIACEAE OF ECUADOR 181

successful accomplishment of this project. The writer wishes to

record, also, his special appreciation of the courtesy of Dr. K. Krause,

in charge of the Rubiaceae at Berlin, who aided in selecting the types

to be photographed, and also was responsible for forwarding for

study a large and important shipment of South American Rubiaceae.

KEY TO THE TRIBES AND GENERA1

Ovules 3 or more in each cell of the ovary.

Fruit dry.

Seeds winged, vertically imbricate I. Cinchoneae.

Seeds not winged or, if winged, horizontal.

Corolla lobes imbricate or contorted, never valvate.

III. Rondeletieae.

Corolla lobes valvate.

Seeds horizontal; stipules entire or bifid; plants trees or

large shrubs II. Condamineae.

Seeds peltately attached; stipules often fimbriate; plants

chiefly herbs or very low shrubs IV. Hedyotideae.

Fruit fleshy.

Corolla lobes valvate V. Mussaendeae.

Corolla lobes imbricate or contorted.

Seeds many, minute, pitted or rarely tuberculate.

VI. Hamelieae.

Seeds usually few and large, the testa smooth or fibrous.

VII. Gardenieae.

Ovules 1 or 2 in each cell of the ovary.

Ovules pendulous.

Stamens inserted in the throat of the corolla tube; fruit not

compressed VIII. Guettardeae.

Stamens inserted at the base of the corolla tube; fruit compressed.

IX. Chiococceae.

Ovules erect or ascending.

Corolla lobes contorted X. Ixoreae.

Corolla lobes valvate.

Ovules affixed to the base of the ovary.

JThe key is adapted from that published by Wernham for the tropical

American genera. See Journ. Bot. 54: 326. 1916.

182 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Ovary 1-celled, or 2-celled but with a very thin septum.

XII. Coussareae.

Ovary with 2 or more cells, the septum or septa thick.

Flowers perfect; stamens usually inserted in the throat of

the corolla; plants usually trees or shrubs.

XIII. Psychotrieae.

Flowers usually dioecious; stamens commonly inserted at

the base of the corolla; plants mostly herbaceous.

XIV. Anthospermeae.

Ovules affixed to the septum.

Stipules entire, not leaflike; plants trees or shrubs, the

flowers in heads XL Morindeae.

Stipules fimbriate or leaflike; plants mostly herbaceous.

Stipules fimbriate, not leaflike XV. Spermacoceae.

Stipules resembling the leaves and forming whorls with

them XVI. Galieae.

I. CINCHONEAE

Corolla lobes valvate.

Placenta pendulous from the apex of the cell; inflorescence spikelike.

1. Alseis.

Placenta not pendulous; inflorescence not spikelike.

Placenta ascending or erect from the base of the septum; plants

usually scandent 2. Manettia.

Placenta adnate to the middle of the septum; plants not scandent.

Capsule splitting from below upward 3. Cinchona.

Capsule splitting from above downward.

Capsule loculicidal 4. Macrocnemum.

Capsule septicidal 5. Ladenbergia.

Corolla lobes imbricate or contorted.

Corolla lobes contorted; corolla salverform 6. Hillia.

Corolla lobes imbricate; corolla funnelform 7. Coutarea.

II. CONDAMINEAE

Calyx persistent 8. Chimarrhis.

Calyx deciduous 9. Condaminea.

THE RUBIACEAE OF ECUADOR 183

III. RONDELETIEAE

Corolla lobes imbricate 10. Rondeletia.

Corolla lobes contorted 11. Elaeagia.

IV. HEDYOTIDEAE

A single genus in Ecuador 12. Arcytophyllum.

V. MUSSAENDEAE

Inflorescence terminal.

Inflorescence elongate, spikelike 13. Gonzalagunia.

Inflorescence thyrsiform-paniculate 14. Isertia.

Inflorescence axillary.

Leaves with many close parallel striolae in the meshes between the

ultimate veins; plants not scandent.

Inflorescence 1-3-flowered; leaves small 15. Hippotis.

Inflorescence many-flowered; leaves very large. .16. Pentagonia.

Leaves not striolate between the veins; plants usually scandent.

17. Sabicea.

VI. HAMELIEAE

Ovary 4-5-celled; inflorescences terminal.

Corolla lobes contorted in bud 18. Bertiera.

Corolla lobes imbricate in bud 19. Hamelia.

Ovary 2-celled; inflorescences axillary 20. Hoffmannia.

VII. GARDENIEAE

Corolla tube villous in either throat or base, but not in both; testa

of the seed not fibrous; chiefly shrubs, often armed with spines.

21. Randia.

Corolla tube villous in both throat and base; testa of the seed fibrous;

unarmed trees 22. Genipa.

VIII. GUETTARDEAE

Corolla lobes imbricate in bud.

Fruit dry, separating into 2 small cocci 23. Machaonia.

Fruit drupaceous, indehiscent 24. Guettarda.

Corolla lobes valvate in bud; fruit drupaceous 25. Anisomeris.

184 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

IX. CHIOCOCCEAE

A single genus in Ecuador 26. Chiococca.

X. IXOREAE

Calyx calyculate at the base 27. Coffea.

Calyx simple 28. Ixora.

XL MORINDEAE

A single genus in Ecuador 29. Morinda.

XII. COUSSAREAE

Ovules connate, borne on a common basal column; seed vertical.

30. Coussarea.

Ovules separate, in a 1-celled ovary, collateral, basilar, the single seed

horizontal 31. Faramea.

XIII. PSYCHOTRIEAE

Inflorescence an involucrate head.

Seeds flat on the inner surface; creeping herbs with usually cordate

leaves 32. Geophila.

Seeds deeply furrowed on the inner side; trees or shrubs.

33. Cephaelis.

Inflorescence without an involucre, the flowers usually not in heads.

Stipules usually not pectinate; seeds deeply furrowed or flat on the

inner surface.

Corolla tube not gibbous at the base; inflorescence usually not

thyrsoid; stipules often deciduous 34. Psychotria.

Corolla tube gibbous at the base; inflorescence usually thyrsoid;

stipules persistent 35. Palicourea.

Stipules pectinate or with short subulate dorsal appendages; seed

with inrolled ventral surface 36. Rudgea.

XIV. ANTHOSPERMEAE

Fruit pyriform, leathery, splitting into 2 cocci 37. Corynula.

Fruit a succulent berrylike drupe 38. Nertera.

XV. SPERMACOCEAE

Fruit circumscissile 39. Mitracarpus.

Fruit not circumscissile, separating into cocci.

THE RUBIACEAE OF ECUADOR 185

Cocci more than 2 40. Richardia.

Cocci 2.

Cocci indehiscent 41. Diodia.

Cocci dehiscent at the base or apex.

Cells of the fruit unlike, one opening, the other remaining

closed 42. Spermacoce.

Cells of the fruit alike, both opening 43. Borreria.

XVI. GALIEAE

Flowers surrounded by a calyxlike involucre 44. Relbunium.

Flowers not involucrate 45. Galium.

1. ALSEIS Schott

Alseis Eggersii Standl. Field Mus. Bot. 4: 264. 1929.

El Recreo, Eggers 15738 (F, type; B, K).

A tree; leaves petiolate, elliptic-obovate, acuminate, at the base

obtuse and short-decurrent, soft-pilose beneath; flowers spicate, the

spikes axillary, 13-17 cm. long, the rachises minutely pilose; capsules

cylindric-clavate, 8-11 mm. long, obtuse, puberulent or glabrate.

2. MANETTIAL.

Manettia canescens Schum. in Mart. Fl. Bras. 66: 178. 1889.

Mount Guayrapata, Spruce 5438 (B, G, type collection; photo,

of type, ex Herb. Berol., in F).

Leaves ovate-lanceolate, scaberulous above, densely tomentose

beneath, attenuate-acuminate, acute at the base, the margins

revolute; cymes 2-3-flowered; calyx lobes 4, subulate; corolla 5-6

mm. long; capsule canescent-pubescent.

Manettia echitidea Standl. Field Mus. Bot. 4: 276. 1929.

Malchingui to Pomasqui, Prov. Pichincha, 3,000-3,600 m.,

Hitchcock 20880 (W, Y). Also in Colombia.

Stems sparsely villous; leaves petiolate, lanceolate or ovate-

lanceolate, 4.5-8 cm. long, 1.5-3 cm. wide, long-acuminate, at the

base acute or abruptly contracted and long-decurrent, sparsely

white-pilose beneath; flowers axillary, solitary or fasciculate; calyx

lobes 8, linear or lance-oblong, 2-3 mm. long, recurved; corolla villous,

the tube green, 8 mm. long, the lobes oblong-ovate, white, spreading,

5 mm. long.

Manettia evenia Sprague, Bull. Herb. Boiss. II. 5: 835. 1905.

Type from Pichincha, at 3,650 m., Jameson 74- Sprague reports

also Jameson 352, "in valle License."

186 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Leaves ovate, obtusely acuminate, rounded at the base, 2-4.5

cm. long, 1-1.7 cm. wide, coriaceous, glabrous above, puberulent

beneath; cymes 3-many-flowered; calyx lobes 4, ovate-deltoid, 2 mm.

long; corolla minutely puberulent, the tube 6-7.5 mm. long, the

lobes 3-3.5 mm. long.

Manettia flexilis Brandeg. Univ. Calif. Publ. Bot. 6: 196. 1915.

Chimborazo, at 900 m., Spruce 6185 (ex Wernham, Gen. Manettia

29. 1919). Type from Mexico; also in Guatemala.

Leaves ovate or lance-ovate, 3-7 cm. long, 1-3 cm. wide, acute

or long-attenuate, at the base acute or obtuse, glabrous above or

nearly so, puberulent and short-pilose beneath; calyx lobes 4, linear

to lance-ovate, 1.5-^4 mm. long; corolla rose-colored, glabrous or

sparsely pilose outside, the tube 6-7 mm. long, the lobes 2-3 mm.

long.

To the present writer the occurrence of this Mexican and Central

American species in Ecuador seems very improbable.

Manettia Holwayi, sp. nov. — Scandens, ramis ochraceis nitidis,

novellis pilis gracilibus pallidis villosis; stipulae triangulares obtusae

2-3 mm. longae dentatae saepe recurvae; petiolus 3-6 mm. longus

dense villosulus; lamina subcoriacea, elliptico-oblonga vel oblongo-

ovata, 3-6.5 cm. longa, 1.5-3.3 cm. lata, obtusa vel acuta, basi

obtusa vel subrotundata, supra cinereo-viridis, scaberula, nervis

subimpressis, subtus pallidior, ubique dense velutino-pilosula, costa

gracili elevata, nervis lateralibus utroque latere c. 7, angulo acuto

adscendentibus, leviter arcuatis, margine subreyoluto; flores sessiles

vel pedicellati subcapitati, capitulis densis multifloris 1-1.5 cm. latis

axillaribus et terminalibus, sessilibus vel breviter pedunculatis,

pedicellis dense villosis; hypanthium dense albo-villosum; calycis

lobi 4 ovati vel lanceolati acuminati 3-4 mm. longi villosuli; corolla

extus glabra 7-9 mm. longa, tubo crasso fauce 2.5 mm. lato, lobis

ovatis obtusis 2.5 mm. longis intus glabris. — Ecuador: Huigra, Prov.

Chimborazo, August 4, 1920, E. W. D. and Mary M. Holway 941

(U. S. Nat. Herb. No. 1,050,995, type).

Manettia Lobbii Wernham, Gen. Manettia 26. 1919.

Type, Lobb 97, ascribed to Colombia, but probably from Ecuador.

In valle Lloensi, 2,400 m., Jameson 352 in part (G). Baiios, Prov.

Tungurahua, 1,800 m., Heilborn 385 (F, S).

Leaves thick, elliptic, acuminate, glabrous, 3-5 cm. long, the

lateral nerves obsolete, the margin revolute; calyx lobes 4, ovate-

lanceolate, acute, 2 mm. long; hypanthium glabrous; corolla gla-

brous outside, the tube 7 mm. long, the limb 6-7 mm. broad.

Manettia pichinchensis Wernham, Gen. Manettia 25. 1919.

Mount Pichincha, at 3,000-3,600 m., Couthouy (G, type collec-

tion); Holmgren 279 (F, S); at 3,000-3,200 m., Lehmann 495a (W);

at 3,600 m., Jameson (G); at 3,900 m., Karsten (L). "Quito,"

THE RUBIACEAE OF ECUADOR 187

Jameson (L). Without locality, Jameson (F, W). Wernham reports

also the following collections: Fraser; Jameson 56, 152, 287; Lehmann

495; Hall 80.

Leaves triangular-lanceolate, subcoriaceous, 1.5-2 cm. long, 4-9

mm. wide, acuminate, truncate or cordate at the base, glabrous

above, grayish-tomentellous beneath, the margins revolute; calyx

lobes 4, narrowly lanceolate, 2-3 mm. long; corolla glabrous outside,

the tube 5-6 mm. long, the lobes 2-2.5 mm. long; capsule 5-6 mm.

long, minutely puberulous.

Manettia recurva Sprague, Bull. Herb. Boiss. II. 5: 835. 1905.

Mount Tungurahua, at 3,300 m., Spruce 5835 (G, type collection).

Between Cuenca and Huigra, 2,700-3,000 m., Hitchcock 21666

(W, Y).

Leaves ovate-oblong, obtuse, 1-2.4 cm. long, 5-10 mm. wide,

recurved at the apex, acute at the base, glabrous, the margins

revolute; calyx lobes 4, ovate, 2.5 mm. long; corolla pinkish, glabrous

outside, the tube 5 mm. long, the lobes 2-3 mm. long; capsule 5 mm.

long, sparsely puberulent.

Manettia Teresitae, sp. nov. — Herbacea scandens, ramis graci-

libus subteretibus retrorso-puberulis, internodiis elongatis; stipulae

latissimae truncatae 0.8 mm. longae breviter fimbriato-ciliatae;

petiolus gracilis 5-12 mm. longus puberulus; lamina late elliptica

vel oyato-elliptica, 3-5.5 cm. longa, 1.5-3.5 cm. lata, herbacea, apice

acuminata vel subrotundata et abrupte acuminata, acumine trian-

gulari acuminato, basi acuta vel obtusa, interdum decurrens, supra

viridis, sparse minuteque adpresso-pilosula vel glabrata, nervis non

elevatis, subtus vix pallidior, praesertim ad nervos minute sparseque

adpresso-pilosula, eosta gracili elevata, nervis lateralibus utrpque

latere c. 6, gracilibus, prominulis, angulo acuto adscendentibus,

arcuatis; flores umbellati vulgo 1-2 cm. longe pedicellati, umbellis

axillaribus plerumque longipedunculatis, pedicellis sparse puberulis;

hvpanthium ovale vel obovatum 3 mm. longum subdense cinereo-

pilosulum; calycis tubus 1 mm. longus, lobis 8 lineari-attenuatis

3-5 mm. longis puberulis recurvis; corolla coccinea in alabastro

8 mm. longa pilosula; capsula obovoidea 8-9 mm. longa glabrata. —

Ecuador: Teresita, 3 km. west of Bucay, Prov. Guayas, alt. 270 m.,

July 5-7, 1923, A. S. Hitchcock 20508 (U. S. Nat. Herb. No. 1,195,938,

type; duplicate in Y).

An ally of M. coccinea (Aubl.) Willd., but distinct in the copious

pubescence and small corolla.

Manettia Trianae Wernham, Gen. Manettia 27. 1919.

Woods at the foot of Mount Tungurahua, Spruce 5092 (G).

Also in Colombia.

Leaves firm-chartaceous, subcarnose, elliptic-lanceolate, acumi-

nate at each end, 4-6.5 cm. long, 1.7-2.5 cm. wide, glabrous; calyx

188 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

lobes 4, ovate-lanceolate, 2.5 mm. long; corolla glabrous outside,

the tube 5 mm. long, the lobes 3 mm. long.

3. CINCHONA L.

After the genus Coffea, that produces the coffee of commerce, the

most important group of the family Rubiaceae is the genus Cinchona.

From the trees of this genus is obtained the drug quinine, which is

used in medicine throughout the world. It is singularly appropriate

that tropical America, in which the greatest plague is malaria, should

have produced the best remedy for this malady. One who lives in a

temperate climate can scarcely appreciate the importance of malaria

in relationship to the population of the tropics.

In these regions almost every person sooner or later is stricken

with malaria. Recovery, under proper treatment, may be rapid and

more or less complete, but too often, especially among the poor,

recovery is only partial, and it is followed by further attacks, that

weaken the individual, with many attendant affections, and finally

cause his death. The loss of efficiency due to malaria is incalculable.

Probably no other discovery in medicine could produce so great a

benefit to so large a number of persons as that of a preventive of this

terrible disease.

Since Ecuador is the country from which the efficiency of cinchona

bark as a remedy for malaria was first made known to the world, it is

pertinent to give here a summary of its rather romantic history. The

best account of the history of the drug is that given in a book entitled

Peruvian bark, a popular account of the introduction of Chinchona

cultivation into British India, published in 1880 by Clements R.

Markham, who was instrumental in establishing the Cinchonas in

cultivation in India and Ceylon. The following extract from that

book is slightly condensed:

The aborigines of South America appear, except perhaps in one

locality, to have been ignorant of the virtues of Peruvian bark.

This sovereign remedy is absent in the wallets of itinerant doctors,

whose materia medica has been handed down from father to son,

since the days of the Incas. It is mentioned neither by the Inca

Garcilasso de la Vega, nor by Acosta, in their lists of Indian medicines.

It seems probable, nevertheless, that the Indians were aware of the

virtues of Peruvian bark in the neighborhood of Loja, 230 miles

south of Quito, where its use was first made known to Europeans;

and the local name for the tree, quina-quina, "bark of bark," indicates

that it was believed to possess some special medicinal properties.

The Indians looked upon their conquerors with dislike and suspicion;

it is improbable that they would be inclined to impart knowledge of

THE RUBIACEAE OF ECUADOR 189

this nature to them; and the interval which elapsed between the

discovery and settlement of the country and the first use of Peruvian

bark by Europeans is thus easily accounted for.

La Condamine, Jussieu, and Ruiz, all believed that the Indians

were aware of the medicinal qualities of Peruvian bark, and that they

imparted their knowledge to the Spaniards. Humboldt and Ulloa

were of an opposite opinion. The stories of its virtues having been

discovered by watching the pumas or South American lions chewing

the bark to cure their fevers, mentioned by Condamine; and of an

Indian haying found it out by drinking of the waters of a lake into

which a cinchona tree had fallen, told by Geoffroy, are of modern

and European origin.

The conquest and subsequent civil wars in Peru can not be said

to have been finally concluded until the time of the Marquis de

Canete in 1560; and Jussieu reports that a Jesuit, who had a fever

at Malacotas, was cured by Peruvian bark in 1600. M. de la Con-

damine also found a manuscript in a library of a convent at Loja,

in which it was stated that the Europeans of the province used the

bark at about the same time.

It may be added, however, that though the Indians were aware

of the febrifuge qualifies of this bark, they attached little importance

to them, and this may be another reason for the lapse of time which

occurred before the knowledge was imparted to the Spaniards. This

indifference to, and in many cases even prejudice against, the use of

Peruvian bark amongst the Indians, is very remarkable. Poeppig,

writing in 1830, says that in the Peruvian province of Huanuco the

people, who are much subject to tertian agues, have a strong repug-

nance to its use. The Indian thinks that the cold north alone permits

the use of fever-bark; he considers it as very heating, and therefore

an unfit remedy in complaints which he believes to arise from

inflammation of the blood. Even in Guayaquil the prejudice against

quinine used to be so strong that, when a physician administered it,

he was obliged to call it by another name.

In 1638 the wife of Don Luis Geronimo Fernandez de Cabrera

Bobadilla y Mendoza, fourth Count of Chinchon, and Viceroy of

Peru, lay sick of an intermittent fever in the palace of Lima.

This lady's maiden name was Ana de Osorio, a daughter of the

noble family, whose founder was created Marquis of Astorga by

Henry IV, King of Castille. She was married at Madrid, on Sunday,

the 12th of August, 1621, to the Count of Chinchon. Her husband,

of an old Catalonian family, was the grandchild of the first Count

by Beatriz de Bobadilla, the faithful friend and attendant of Queen

Isabella. He was hereditary Alcaide of Segovia, Lord of the Castle

of Chinchon near Madrid, and of eighteen towns in the kingdom of

Toledo. In 1628 the Count was appointed Viceroy of Peru, and he

and his Countess made their solemn entry into Lima on the 14th of

January, 1629.

190 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

The great event of this viceroyalty was the cure of the Countess

of Chinchon, in the year 1638, of a tertian fever by the use of Peruvian

bark. The news of her illness at Lima reached Don Francisco Lopez

de Canizares, the Corregidor of Loja, who had become acquainted

with the febrifuge virtues of the bark. He sent a parcel of it to the

Vice-Queen, and the new remedy, administered by her physician,

Dr. Don Juan de Vega, effected a rapid and complete cure. It is

known by tradition amongst the bark collectors that the particular

species from which the bark was taken, which cured the Countess

of Chinchon, was that known to them as Cascarilla de Chahuarguera.

These trees are a variety of the C. officinalis of Linnaeus, many

thousands of which are now growing in India.

Madame de Genlis wrote a short novel founded on the cure of

the Countess of Chinchon, which she dedicated to the Comtesse de

Choiseul. It is entitled Zuma, and though erroneous in every

particular so far as all the facts are concerned, it yet proves the deep

and general interest which attaches to the first introduction of quina

bark into Europe by the Vice-Queen.

The Countess of Chinchon returned to Spain in the spring of

1640, bringing with her a supply of that precious quina bark which

had worked so wonderful a cure upon herself, and the healing virtues

of which she intended to distribute amongst the sick on her husband's

estates. It thus gradually became known in Europe, and was most

appropriately called Countess's powder (Pulvis Comitissae). By this

name it was long known to druggists and in commerce. Dr. Juan

de Vega, the learned physician of the Countess of Chinchon, followed

his patient to Spain, bringing with him a quantity of quina bark,

which he sold at Seville for 100 reals the pound. The bark continued

to have the same high value and the same reputation, until the trees

became scarce and the collectors began to adulterate it.

After their return from Peru, the Count and Countess usually

resided at the Castle of Chinchon, built by the Count's father in

1590. The Countess administered Peruvian bark to the sufferers

from tertian agues on her lord's estates, in the fertile but unhealthy

vegas of the Tagus, the Jarama, and the Tajuiia. She thus spread

blessings around her, and her good deeds are even now remembered

by the people of Chinchon and Colmenar, in local traditions.

In memory of the great service to humanity performed by the

Countess of Chinchon, Linnaeus named the genus which yields Peru-

vian bark, Chinchona. Unfortunately the great botanist was misin-

formed as to the name of her whom he desired to honor. This is to

be accounted for by his having received his knowledge of the Countess

through a foreign, and not a Spanish source. Thus misled, Linnaeus

spelt the word Cinchona (Gen. PI. ed. 1742) and Cinhona (Gen. PL

ed. 1767), omitting one or two letters. It was still more unlucky

that Linnaeus died before the error was pointed out and corrected.

This was done by the Spanish botanists Ruiz and Pavon, who landed

THE RUBIACEAE OF ECUADOR 191

in Peru in 1778, the year of Linnaeus's death. They advocated the

correct spelling, and their example was followed by Mutis and others.1

After the cure of the Countess of Chinchon, the Jesuits were the

great promoters of the introduction of bark into Europe. In 1670

these fathers sent parcels of the powdered bark to Rome, whence it

was distributed to members of the fraternity throughout Europe, by

Cardinal de Lugo, and used for the cure of agues with great success .

Hence the name of "Jesuits' bark," and "Cardinal's bark"; and it

was a ludicrous result of its patronage by the Jesuits that its use

should have been for a long time opposed by the Protestants, and

favored by Roman Catholics. In 1679 Louis XIV bought the secret

of preparing quinquina from Sir Robert Talbor, an English doctor,

for 2,000 louis d'or, a large pension, and a title. From that time

Peruvian bark seems to have been recognized as the most efficacious

remedy for intermittent fevers.

However, a very strong prejudice was raised against it, which it

took many years to conquer; and the controversies that arose on the

subject, between learned doctors, were long and acrimonious. It

remained a subject of controversy as late as 1714, when two Italian

doctors, Ramazzini and Torti, held opposite views on the subject.

Whilst the inestimable value of Peruvian bark was gradually

forcing conviction on the most bigoted medical conservatives of

Europe, and whilst the number and efficacy of the cures effected by

its means were bringing it into general use, and so consequently

increasing the demand, it was long before any knowledge was obtained

of the tree from which it was taken. In 1682 La Fontaine, at the

solicitation of the Duchess of Bouillon, who had been cured of a

dangerous fever by taking Peruvian bark, composed a poem in two

cantos to celebrate its virtues; but the exquisite beauty of the leaves,

and the delicious fragrance of the flowers of the quinquina-tree,

with allusions to which he might have adorned his poem, were still

unknown in Europe.

The first description of the quinquina-tree is due to that mem-

orable French expedition to South America, to which all branches

of science owe so much. Its members, MM. de la Condamine, Godin,

Bouguer, and the botanist Joseph de Jussieu, sailed from Rochelle

on the 16th of May, 1735, to measure an arc of the meridian near

Quito, and thus determine the shape of the earth. After a residence

at Quito, Jussieu set out for Loja, to examine the quinquina- tree,

in March, 1739, and in 1743 La Condamine visited Loja, and stayed

for some time at Malacotas, with a Spaniard whose chief source of

income was the collection of bark. He obtained some young plants

with the intention of taking them down the river Amazons to

Cayenne, and thence transporting them to the Jardin des Plantes at

Paris; but a wave washed over his little vessel near Para, at the mouth

According to the general rules regarding the spelling of generic names, the

name is now written Cinchona. — P.C.S.

192 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

of the great river, and carried off the box in which he had preserved

those plants for more than eight months.

Condamine described the quinquina-tree of Loja in the Memoires

de I' Academic; he was the first man of science who examined and

described this important plant; and in 1742 Linnaeus established the

genus Cinchona, in honor of the Countess Ana of Chinchon. He,

however, only knew of two species, that of Loja, which was named

C. officinalis, and the C. caribaea, since degraded to the medicinally

worthless genus ofExostema.

For many years the quinquina-tree of Loja, the C. officinalis of

Linnaeus, was the only species with which botanists were acquainted;

and from 1640 to 1776 no other bark was met with in commerce

than that which was exported from the Peruvian port of Payta,

brought down from the forests in the neighborhood of Loja. The

constant practice of improvidently felling the trees over so small an

area for more than a century, without any cessation, inevitably led

to their becoming very scarce, and threatened their eventual ex-

tinction. As early as 1735 Ulloa reported to the Spanish Govern-

ment, that the habit of cutting down the trees in the forests of Loja,

and afterwards barking them, without taking the precaution of

planting others in their places, would undoubtedly cause their

complete extirpation. "Though the trees are numerous," he added,

"yet they have an end"; and he suggested that the Corregidor of

Loja should be directed to appoint an overseer, whose duty it should

be to examine the forests, and satisfy himself that a tree was planted

in place of every one that was felled, on pain of a fine. This wise

rule was never enforced, and sixty years afterwards Humboldt

reported that 25,000 trees were destroyed in one year. Yet the

Jesuits are said to have enforced it as a religious duty that, for every

tree felled, the bark collector should plant a quincunx.

The measures adopted by the Spanish Government towards the

end of the last century, in sending botanical expeditions to explore

the cinchona forests in other parts of their vast South American

possessions led to the discovery of additional valuable species, the

introduction of their barks into commerce, and the reduction of the

pressure on the Loja forests, which were thus relieved from being

the sole source whence Peruvian bark could be supplied to the world.

The region of cinchona trees extends from 19° S. latitude, where

Weddell found C. australis, to 10° N., following the almost semicir-

cular curve of the cordillera of the Andes, over at least 1,500 miles of

latitude. They flourish in a cool and equable temperature, on the

slopes and in the valleys and ravines of the mountains, surrounded

by the most majestic scenery, never descending below an elevation

of 2500, and ascending as high as 9000 feet above the sea. Within

these limits their usual companions are tree-ferns, Melastomaceae,

arborescent passion-flowers, and allied genera of cinchonaceous plants.

Below them are forests abounding in palms and bamboos, above their

highest alpine limits are a few lowly alpine shrubs. But within this

THE RUBIACEAE OF ECUADOR 193

wide zone grow many species of Cinchonas, each within its own

narrower belt as regards elevation above the sea, some yielding the

inestimable bark, and others commercially worthless.

Thus, to give a geographical summary of the cinchona region,

beginning from the south, it commences in the Bolivian province of

Cochabamba in 19° S., passes through the Yungas of La Paz, Larecaja,

Caupolican, Coroico, and Munecas, into the Peruvian province of

Carabaya; thence through the Peruvian forests, on the eastern slopes

of the Andes, of Marcapata, Paucartambo, Santa Ana, Huanta, and

Uchupampa, to Hudnuco and Huamalies, where the gray bark is

found. It then continues through Jaen, to the forests near Loja and

Cuenca, and on the western slopes of Chimborazo. It begins again

in latitude 1° 51' N. at Almaguer, passes through the province of

Popayan, and along the slopes of the Andes of Quindio, until it reaches

its extreme northern limit on the wooded heights of Me>ida and

Santa Marta.

The Cinchonas, when in good soil and under other favorable

circumstances, become large forest trees; on higher elevations, and

when crowded, and growing in rocky ground, they frequently run

up to great heights without a branch; and at the upper limit of their

zone they become mere shrubs. The leaves are of a great variety of

shapes and sizes, but in most of the finest species they are lanceolate,

with a shining surface of bright green, traversed by crimson veins,

and petioles of the same color. The flowers are very small, but hang

in clustering panicles, like lilacs, generally of a deep roseate color,

paler near the stalk, dark crimson within the tube, with white curly-

hairs bordering the laciniae of the corolla. The flowers of C. mi-

crantha are entirely white. They send forth a delicious fragrance

which scents the air in their vicinity.

The region around Loja, on the southern frontier of the modern

republic of Ecuador, is the original home of the cinchona, and nearly

in the center of its latitudinal range of growth. On the lofty grass-

covered slopes of the Andes, around the little town of Loja, and in the

sheltered ravines and dense forests, those precious trees were found

which first made known to the world the healing virtues of Peruvian

bark. They were most plentifully met with in the forests of Uritu-

singa, Rumisitana, Cajanuma, Boqueron, Villonaco, and Monje, all

within short distances of Loja.

Linnaeus named these trees Cinchona officinalis; but, when Hum-

boldt and Bonpland examined them, the discovery of other species

yielding medicinal bark induced them to rechristen this species after

the distinguished Frenchman who had originally described them,

Cinchona Condaminea. Humboldt says that they grow on mica slate

and gneiss, from 6000 to 8000 feet above the sea, with a mean temper-

ature between 60 and 65° Fahr. In his time the tree was cut down in

its first flowering season, or in the fourth or seventh of its age, accord-

ing as it had sprung from a vigorous root-shoot, or from a seed. He

describes the luxuriance of the vegetation to be such that the younger

trees, only six inches in diameter, often attain from fifty-three to

194 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

sixty-four English feet in height. "This beautiful tree," he continues,

"which is adorned with leaves above five inches long and two broad,

growing in dense forests, seems always to aspire to rise above its

neighbors. As its upper branches wave to and fro in the wind, their

red and shining foliage produces a strange and peculiar effect, recog-

nizable from a great distance." It varies much in the shape of its

leaves, according to the altitude at which it grows, and bark collectors

themselves would be deceived if they did not know the tree by the

glands, so long unobserved by botanists. The C. Condaminea

described by Humboldt is the same as the C. uritusinga of Pavon.

It once yielded great quantities of thick trunk bark, but, owing to

reckless felling through a course of years, it is now almost extermi-

nated, and its bark is rarely met with in commerce. The distinguished

botanist Don Francisco Caldas examined the cinchona forests of

Loja after Humboldt, between 1803 and 1809. He says that the

famous quina-tree of Loja grows in the forests of Uritusinga and

Cajanuma, at a height of from 6200 to 8200 feet above the sea, in a

temperature of 41 to 72° Fahr. ; but that it is only found between the

rivers Zamora and Cachiyacu. He describes the tree as from thirty

to forty-eight feet high, with three or more stems growing from the

same root; the leaves as lanceolate, shining on both sides, with veins

a rosy color, a short and tender pubescence on the under side when

young, and when past maturity a bright scarlet color; the bark black

when exposed to the sun and wind, a brownish color when closed in

by other trees, and always covered with lichens; and the rock on

which it grows a micaceous schist.

The Spanish botanists Ruiz and Pavon also examined the cin-

chona-trees of Loja; and the latter described two species, C. uritu-

singa, named from the mountain on which it was once most abundant,

and C. chahuarguera, so called from a fancied resemblance of the bark

to a pair of breeches (huara in Quichua) made from the fiber of the

American aloe (chahuar). To these the botanist Tafalla added C.

crispa. These three species are all included in Humboldt's C.

Condaminea, which is readily known by the little pits, bordered with

hairs, at the axils of the veins on the under side of the leaf. It would

appear that at one period of growth these little pits are wanting

but when the plant is in full vigor they are markedly prominent.

C. chahuarguera is described by Pavon as growing from eighteen to

twenty-four feet in height; although now the trees which yield the

Loja bark of commerce do not attain a height of more than four to

nine feet. It is met with on the grassy open crests of mountain

ridges, in light sandy soil interspersed with rocks, amongst shrubs and

young plants. The barks of Loja were called crown barks, because

they were reserved for the exclusive use of the royal pharmacy at

Madrid; and they originally sold at Panama for five and six dollars,

and at Seville for twelve dollars the pound; but in later times they

were much adulterated, and the price fell to one dollar the pound.

The C. chahuarguera is the rusty crown bark of commerce, and the

C. crispa is the quina fina de Loja or crespilla negra of the natives.

THE RUBIACEAE OF ECUADOR 195

With this rusty crown bark are mixed larger quills particularly rich

in the alkaloid called cinchonidine. It is said that there is a great

difference in the bark, according as it is grown on the sides of moun-

tains most exposed to the morning or evening sun; and its position

is believed to have a great influence on the quality of its alkaloids.

The usual yield of the large quills is 3.5 to 3.6 per cent.

The bark collectors of Loja are said to show some little fore-

thought, a quality which is entirely wanting to most of their fra-

ternity. To save the trees they occasionally cut off the whole of the

bark, with the exception of one long strip, which gradually replaces

its loss; and the second cutting is called cascariUa resecada. This

practice was in use in the days of the botanist Ruiz, who protested

against it, and declared that it was very injurious to the trees, many

having been destroyed by it. Later accounts, however, show that the

bark collectors of Loja are as thoughtlessly destructive as those in

other parts of South America. They often pull up the roots, while

the annual burning of the slopes, and the continual cropping of the

young shoots by cattle, assist the work of destruction.

It is, therefore, well that the C. chahuarguera and C. uritusinga,

the earliest known and among the most valuable of the cinchona-trees,

should have been saved from extinction by timely introduction into

India.

The species yielding "red bark," the richest and most important

of all the Cinchonae, is found in the forests on the western slopes

of Mount Chimborazo, along the banks of the rivers Chanchan,

Chasuan, San Antonio, and their tributaries. So early as 1738

Condamine spoke of "red bark" (cascariUa Colorado) as being of

superior quality; and Pa von sent home specimens of the "red bark

of Huaranda," and named the species C. succirubra. Much light was

thrown upon the history of this valuable species by Dr. Spruce, when

he penetrated into the forests to collect seeds and plants for trans-

mission to India in 1860.

Though little was known of the tree until quite lately, there

was never any doubt concerning the value of the bark. In 1779 a

Spanish ship from Lima, bound to Cadiz, was captured off Lisbon by

the Hussar frigate, and her cargo consisted chiefly of "red bark,"

part of which was imported into England. In 1785 and 1786 Ruiz

states that the collectors began to gather the bark of C. succirubra,

and sell it at Guayaquil, and from that time it continued to be found

in the European markets. It contains a larger proportion of alka-

loids than any other kind, chiefly cinchonidine. Mr. Howard has

procured 8.5 per cent of alkaloids from a specimen of "red bark" from

South America. A large supply of plants of this species is flourishing

in India and Ceylon, and it has become a cultivated plant of great

value and importance, with a much greater yield of total alkaloids

than when in its wild state.

Because the supply of Cinchona bark for the extraction of quinine

was not of dependable regularity, and because there was even a

196 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

possibility of the extinction of the trees as a result of careless cutting,

it was determined to introduce the trees into parts of the East Indies,

where it was believed conditions were suitable for their growth. They

were planted first in Java, but partly because methods for their culti-

vation had not been perfected, and partly on account of the selection

of poor varieties of the plants, the first attempts at cultivation there

did not succeed.

In 1859 definite plans were made under the direction of the

British government for establishing cinchona trees in the mountains

of India. The best available authorities were selected to undertake

the work, which was directed by Clements R. Markham. Plants and

seeds were gathered in all the principal centers of production, from

Bolivia to Colombia. As a result of the precautions taken, and after

great labor in obtaining the seeds and plants, the various species of

Cinchona were successfully established in the Nilgiri Hills and

elsewhere.

Richard Spruce, who from his many years of botanical exploration

along the Amazon and in the Andes was thoroughly familiar with

vegetation of those regions, was entrusted with the collection of

cinchona in the Chimborazo region. An account of his work is given

in Markham's book already quoted, and a much fuller account in

Spruce's Notes of a botanist on the Amazon and Andes. In his work

Spruce was assisted by Robert Cross, who later obtained further

material from the Loja region.

As a result of this undertaking of the British government, cin-

chona cultivation was quickly established in India, and the chief

supply of the drug quinine now comes from the East Indies. Too late

the South American governments realized the importance of for-

bidding the export of living plants and seeds, but it is doubtless

quite as well that they did not attempt too soon to place an embargo

upon their export. If they had done so, seeds or even plants would

have been smuggled abroad sooner or later. Moreover, the drug

quinine is such an essential one in the alleviation of human suffering

that its widest possible distribution is earnestly to be desired.

The literature pertaining to cinchona is an exceedingly extensive

one, for the subject has occupied the attention of many pharmacists

and botanists for two hundred years. In Markham's book thirty

pages are devoted to a list of books and articles treating of the sub-

ject, and many more titles could be added.

Because of the great economic importance of the trees concerned,

and especially because of the varying quinine content in the inhabit-

THE RUBIACEAE OF ECUADOR 197

ants of the various regions, several botanists have devoted a great

deal of time and hundreds of printed pages and plates to a discussion

of the species of the genus. The result of their work has been the

description of a great number of so-called species, based on char-

acters that certainly would not be considered valid or important in

other genera of the Rubiaceae. The writer has examined a large

amount of herbarium material of the genus, as well as photographs

of many of the types involved. He is quite unable to discover any

conservative basis for recognizing the many species to which names

have been given, and he believes that these names relate to variable

races of a few not very clearly marked species which may be recog-

nized for botanical convenience. The mere fact that the variations

of these races are important from the standpoint of the pharmacist

does not justify their recognition, if such were possible, as species in

the ordinary sense of that term.

Cinchona Humboldtiana Lamb. 111. Cinch. 7. 1821. C. villosa

Pavon ex Lamb., loc. cit., as syn. C. conglomerate, Pavon ex Howard,

111. Nueva Quinol. Pav. pi 15. 1862.

Andes of Sigsig, 1,600-1,800 m., Lehmann 6608 (K). Reported

from Loja. Also in Peru and Bolivia.

Branchlets villous; leaves lanceolate to elliptic-oblong, acute at

each end, finally glabrate above, densely villous beneath; panicles

dense, villous; calyx teeth very short, acute; capsule oblong, 2-2.5

cm. long, villous or glabrate.

"Cascarilla colorada."

Cinchona micrantha R. & P. Fl. Peruv. 2: 52. pi. 194. 1799.

Reported from southern Ecuador. Also in Peru and Bolivia.

Branchlets sericeous or glabrate; stipules ovate, obtuse, glabrous;

leaves petiolate, oval, obovate, or rounded, obtuse to rounded at the

apex, acute to rounded at the base, large, coriaceous, nearly glabrous

but usually pilose beneath along the costa; inflorescence paniculate,

terminal, large, open, many-flowered, leafy, minutely tomentose;

calyx minute, 5-dentate, the teeth acute; corolla pink, about 6 mm.

long, densely tomentulose, the lobes much shorter than the tube, vil-

lous within; capsule oblong, acute, about 1.5 cm. long, striate.

It is doubtful whether this is more than an extreme form of C.

pubescens, since the size of the corolla is exceedingly variable in the

forms referred to that species.

Cinchona officinalis L. Sp. PI. 172. 1753 (collected at Loja by

Condamine). C. lancifolia Mutis, "Periodico de Santa Fe," 465.

1793. C. glabra Ruiz, Quinol. 64. 1792. C. nitida R. & P. Fl. Peruv.

2: 50. pi. 191. 1799. C. lanceolata R. & P. Fl. Peruv. 51. 1799. C.

198 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

angustifolia Ruiz, Quinol. Suppl. 14. 1801. C. Condaminea H. & B.

PI. Aequin. 1: 33. pi. 10. 1808 (described from Loja and Ayavaca).

C. colorata Lamb. Bull. Pharm. 294. 1810. C. Condaminea ft lanceo-

lata Lamb. 111. Cinch. 2. 1821. C. stupea Pavon ex Lamb., loc. cit., as

syn. ; Lindl. Fl. Med. 416. 1838. C. academica Guib. Drog. Simpl. 3 : 98.

1822. C. macrocalyx DC. Bibl. Univ. 41: 150. 1829. C. Condaminea

ft chahuarguera DC. Prodr. 4: 352. 1830. C. chahuarguera Pavon ex

DC., loc. cit., as syn. (type from Ecuador). C. macrocalyx ft obtusi-

folia DC. Prodr. 4: 353. 1830. C. obtusifolia Pavon ex DC., loc. cit.,

as syn. 1830. C. macrocalyx 7 lucumaefolia DC., loc. cit. 1830. C.

lucumaefolia Pavon ex DC., loc. cit., as syn. 1830. C. calisaya Wedd.

Ann. Sci. Nat. III. 10: 6. 1848. C. amygdalifolia Wedd., loc. cit. C.

calisaya ft Josephiana Wedd. Ann. Sci. Nat. III. 11: 269. 1849. C.

Condaminea a Condaminea vera Wedd., loc. cit. C. Condaminea ft

Candollei Wedd., loc. cit. C. Condaminea 7 lucumaefolia Wedd., loc.

cit. C. Condaminea 5 lancifolia Wedd., loc. cit. C. crispa Tafalla ex

Howard, Nueya Quinol. Pav. pi. 2. 1862. C. palton Pavon ex

Howard, op. cit. pi. 13. 1862. C. heterophylla Pavon ex Howard, op.

cit. pi. 18. 1862. C. violacea et C. suberosa Pavon ex Howard, op.

cit. 1862. C. uritusinga Pavon ex Howard, op. cit. pi. 19. 1862. C.

calisaya var. rugosa Miq. Ann. Mus. Lugd. Bat. 4. 1868-69. C.

officinalis var. uritusinga Howard, Kept. Intern. Bot. Congr. 201.

1866. C. officinalis var. Condaminea Howard, op. cit. 202. 1866.

C. officinalis var. Bonplandiana Howard, op. cit. 203. 1866. C.

officinalis var. crispa Howard, op. cit. 203. 1866.

Without locality, Humboldt & Bonpland (F, photo, of type? of

Cinchona Condaminea ex Herb. Berol.). Without locality, Pavdn,

(F, photo, and fragm. of type of C. chahuarguera, ex Herb. Berol.).

Loja, Jameson 10 (G). Without locality, Jameson (W). Cerro de

Uritusinga, near Loja, 2,400 m., October, 1835, Jameson (P). Loja,

Rose, Pachano & Rose 23299 (F, W, G, Y), 23300 (F, G, W, Y).

Between Loja and Portovelo, Rose, Pachano & Rose 23319 (G, W,

Y). Pangar, 2,500-2,900 m., Lehmann 5453 (F, K). San Agustin,

Loja, 2,000-2,500 m., Lehmann 7927 (K). Without locality, Lobb

301 (K). Also in Colombia, Peru, and Bolivia.

A large or medium-sized tree, sometimes 24 m. high, with a

trunk 1-1.5 m. in diameter, the bark rugose, the branchlets strigil-

lose-pilosulous; stipules free, lanceolate or oblong, acute or obtuse,

glabrous, deciduous; leaves petioled, lanceolate to ovate, small,

acute to acuminate or obtuse, at the base rounded to attenuate,

subcoriaceous, glabrous above, beneath glabrous, or puberulent or

hispidulous, especially on the nerves, commonly about 10 cm. long

and 4 cm. wide, often scrobiculate beneath; inflorescences terminal,

many-flowered, dense, leafy, cymose-paniculate; hypanthium stri-

gose; calyx reddish, glabrous or nearly so, the teeth triangular, acute;

corolla pink or red, sericeous, the tube about 1 cm. long, the 5 lobes

ovate, acute; capsule oblong, striate-costate, usually 1.5-2 cm. long,

glabrate.

THE RUBIACEAE OF ECUADOR 199

"Cascarilla fina de Uritusinga," "cascarilla colorada del rey,"

"cascarilla amarilla del rey," "cascarilla blanca pata de gallinazo,"

"cascarilla crespilla negra mala," "cascarilla sen-ana acanelada,"

"quina negra," "quina negrilla," "cascarilla hoja de lucma," "quin-

quina payama," "original Loxa crownbark," "rusty crownbark."

Cinchona pubescens Vahl, Skrivt. Naturh. Selsk. 1: 19. 1790.

C. hirsuta R. & P. Fl. Peruv. 2: 51. pi. 192. 1799. C. purpurea R.

& P., op. cit. 52. pi. 193. 1799. C. ovata R. & P., op. cit. 52. pi. 195.

1799. C. cordifolia Mutis ex Humb. Ges. Naturf. Freund. Berlin

Mag. 1: 117. 1807. C. scrobiculata H. & B. PI. Aequin. 1: 165. pi. 47.

1808. C. rotundifolia Pavon ex Lamb. 111. Cinch. 5. 1821 (type

collected at Loja by Pavon). C. Mutisii Lamb., op. cit. 9. 1821

(type collected at Loja by Pavon). C. microphylla Mutis ex Lamb.,

loc. cit., as syn. 1821. C. quercifolia Pavon ex Lamb., loc. cit., as syn.

1821. C. pelalba Pavon ex DC. Bibl. Univ. 41: 152. 1829. C. discolor

Hayne, Arzneigew. 14: sub pi. 14- 1846. C. Delondriana Wedd. Ann.

Sci. Nat. III. 10: 7. 1848. C. rufinervis Wedd., op. cit. 8. 1848.

C. scrobiculata /8 Delondriana Wedd. Ann. Sci. Nat. III. 11: 270.

1849. C. pubescens /3 purpurea Wedd., loc. cit. 1849. C. cordifolia

j8 rotundifolia Wedd., loc. cit. 1849. C. Mutisii a microphylla Wedd.,

loc. cit. 1849. C. Mutisii /3 crispa Wedd., loc. cit. 1849, nomen

nudum. C. succirubra Pavon ex Klotzsch, Abh. Akad. Berl. 1857:

60. 1858. C. erythrantha Pavon ex Howard, Nueya Quinol. Pav.

pi. 12. 1862. C. lutea Pavon ex Howard, Nueva Quinol. Pav. pi. 14.

1862. C. obovata Pavon ex Howard, op. cit. pi. 18. 1862. C. decur-

rentifolia Pavon ex Howard, op. cit. pi. 23. 1862. C. peruviana

Howard, op. cit. pi. 27. 1862. C. viridiflora Pavon ex Howard, op.

cit. 1862. C. Howardiana Kuntze, Monogr. Cinch. 30. 1878.

Western slopes of the Andes of Zaruma, 1,800-2,000 m., Lehmann

7925 (F, K, W). Cuenca, 2,500-3,000 m., Lehmann 4593 (F, K, W).

Chiguinda, 1,600-2,000 m., Lehmann 6495 (F, K). San Cristobal,

western Andes of Guaranda, 1,400-2,000 m., Lehmann 8467 (F).

Western Andes of Cuenca, 2,300-2,800 m., Lehmann 4920 (F).

Huigra, Rose & Rose 22586 (F, G, W, Y), 22481 (F, G, W, Y).

Region of Cuenca, Jameson (P). Chimborazo, 1,800 m., Andre

K528 (F, G, K, Y). Portovelo, Rose & Rose 23428 (W). Without

locality, Jameson 16 (Y); Jameson (W). Cuenca, 2,800-3,000 m.,

Lehmann 6605 (K). Rio Pastaza, 1,500 m., Lehmann 445 (L). Also

in Colombia, and extending southward to Bolivia.

A medium-sized tree, the trunk usually about 30 cm. in diameter,

the branchlets variously pubescent; stipules large, free, ovate, obtuse

or acute, caducous, sericeous or almost glabrous; leaves petioled,

usually orbicular to broadly ovate, large, rounded to acute at the

apex, at the base cordate to acute, often decurrent, usually glabrous

above but sometimes pubescent, beneath densely pilose to nearly

glabrous, often scrobiculate; inflorescence terminal, paniculate, large,

leafy, many-flowered, open, the flowers subsessile; hypanthium seri-

200 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

ceous; calyx 5-dentate, minutely sericeous or tomentulose; corolla

red or pink, sericeous, the tube usually about 1 cm. long, the lobes

lanceolate, 5-7 mm. long, villous within; capsule lanceolate or oblong,

striate-costate, glabrate, usually 1.5-2.5 cm. long.

"Cascarilla amarga," "cascarilla gallinazo," "cascarilla varona"

(Lehmann).

4. MACROCNEMUM L.

Macrocnemum cuencanum, nom. nov. Cinchona ovalifolia

HBK. Nov. Gen. & Sp. 3: 403. 1820, non Mutis, 1807. C. Humboldti-

ana R. & S. Syst. Veg. 5: 13. 1819, non Lamb. 1797. Lasionema

Humboldtianum Wedd. Hist. Quinquin. 99. 1849. Macrocnemum

Humboldtianum Wedd. Ann. Sci. Nat. IV. 1: 76. 1854.

Type from Cuenca.

A small tree; leaves ovate-lanceolate or obovate, acutish, some-

what attenuate at the base, glabrous above, tomentose-hirsute

beneath; calyx lobes triangular, acute; corolla lobes valvate; capsule

ovate-oblong.

5. LADENBERGIA Klotzsch

Ladenbergia crassifolia (Pavon), comb. nov. Cinchona crassi-

folia Pavon ex DC. Bibl. Univ. 41: 150. 1829; Prodr. 4: 354. 1830.

Cascarilla calyptrata Wedd. Ann. Sci. Nat. III. 10: 13. 1848. Buena

crassifolia Wedd. Journ. Linn. Soc. 11: 187. 1869.

Described from Quito and Loja.

Leaves oblong, subobtuse, attenuate at the base, coriaceous,

glabrous, when young villous beneath in the axils of the nerves;

stipules united into a cap; flowers in terminal trichotomous few-

flowered corymbs; calyx lobes triangular-lanceolate; capsules oval-

oblong, 3.5 cm. long, 12 mm. broad.

Ladenbergia macrocarpa (Vahl) Klotzsch in Hayne, Arzneigew.

14: sub pi. 15. 1846. Cinchona macrocarpa Vahl, Skrivt. Naturh.

Selsk. 1: 20. 1790. Cascarilla macrocarpa Wedd. Ann. Sci. Nat. III.

10: 13. 1848. Cinchona magniflora Pavon ex DC. Prodr. 4: 354.

1830, as syn.

Reported from Loja and Cuenca, Pavdn. Also in Colombia and

Peru.

A tree, the branchlets ferruginous-tpmentose; stipules united into

a caducous cap, tomentose; leaves petiolate, oval-elliptic or broadly

oval, large, coriaceous, obtuse, at the base usually rounded or sub-

cordate, sometimes acute, glabrous above, ferruginous-tomentose

beneath or glabrate; panicles terminal, few-flowered, the flowers

sessile or nearly so; calyx tomentose, the teeth short, obtuse; corolla

densely sericeous, 5.5 cm. long, the 5-6 lobes about half as long as

the tube; capsule oblong-clavate, 5-8 cm. long, tomentose or glabrate.

Ladenbergia magnifolia (R. & P.) Klotzsch in Hayne, Arznei-

gew. 14: sub pi. 15. 1846. Cinchona magnifolia R. & P. Fl. Peruv. 2:

THE RUBIACEAE OF ECUADOR 201

53. pi. 196. 1799. Cascarilla magnifolia Wedd. Ann. Sci. Nat. III.

10: 10. 1848. Buena magnifolia Wedd. Journ. Linn. Soc. 11: 186.

1869.

This tree doubtless grows in Ecuador, but I have seen no speci-

mens. The species is known from Colombia, Peru, and Bolivia.

Ladenbergia Pavonii (Lamb.), comb. nov. Cinchona Pavonii

Lamb. 111. Cinch. 8. 1821. C. cava Pavon ex Lamb., loc. cit., as syn.

Ladenbergia cava Klotzsch in Hayne, Arzneigew. 14: sub pi. 15. 1846.

Cascarilla Pavonii Wedd. Ann. Sci. Nat. III. 10: 11. 1848. Buena

Pavonii Wedd. Journ. Linn. Soc. 11: 186. 1869.

Type from the forests of Loja, Pavdn.

Branchlets ferruginous- tomentose; stipules free or short-connate;

leaves ovate or orbicular, obtuse, at the base obtuse or cordate,

subcoriaceous, glabrate above, ferruginous-tomentose beneath; pan-

icles many-flowered, dense, tomentose; calyx large, obsoletely dentic-

ulate; corolla large, sericeous, the lobes ovate-oblong, obtuse, fleshy;

capsule terete, "as large as a finger."

Ladenbergia Riveroana (Wedd.), comb. nov. Cinchona oblongi-

folia Lamb. 111. Cinchon. 12. 1821, non Mutis, 1807. Cascarilla

Riveroana Wedd. Ann. Sci. Nat. III. 10: 11. 1848. Buena Riveroana

Wedd. Journ. Linn. Soc. 11: 186. 1869. Remijia Riveroana Flueck.

Chinarinde 17. 1883.

Type from forests of the mountains of Loja, Pavdn.

Leaves oblong-lanceolate or ovate, acute at each end or rounded

at the base, subcoriaceous, puberulent or glabrate above, pubescent-

tomentose beneath; stipules free or nearly so; inflorescence corym-

biform, scabrous-pilose; calyx teeth triangular-ovate; corolla pilose,

the lobes linear; capsule linear-lanceolate, pubescent.

"Azahar macho," "azahar hembra." Weddell gives the type

locality as "Peruvia," without indication of a specific locality or of

the collector, but the plant is reported by other writers to have been

collected in the Loja region by Pavon.

6. HILLIA Jacq.

Hillia macrophylla, sp. nov. — Frutex subramosus 3-metralis

epiphyticus terrestrisque, ramulis crassis glabris, internodiis c. 5.5

cm. longis; stipulae deciduae ovali-ovatae vel ovato-oblongae, 3-3.5

cm. longae, apice rotundatae vel obtusae, ferrugineae; folia petiolata

opposita, petiolo 1-1.5 cm. longo crasso glabro; lamina crasse mem-

branacea, elliptico-ovata vel late elliptica, 10-19 cm. longa, 6-10 cm.

lata, abrupte breviterque acuminata, acumine anguste triangular!

c. 1 cm. longo obtuso, basi obtusa vel acuta, glabra, supra fusca,

nervis subimpressis, subtus paullo pallidior, costa crassiuscula elevata,

nervis lateralibus utroque latere c. 7, gracilibus, prominentibus,

angulo acuto adscendentibus, subarcuatis, prope marginem con-

202 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

junctis, nervulis inconspicuis laxissime reticulatis, margine piano;

flores terminales pauci odorati sessiles; hypanthium crasse cylindri-

cum 6-7 mm. longum glabrum; calyx non visus; corolla alba extus

glabra, tubo gracili 9.5 cm. longo gracili basi 2.5 mm. lato, superne

sensim dilatato, ore 1 cm. lato, lobis 5 patentibus vel reflexis lineari-

lanceolatis longe attenuatis 3.5-4 cm. longis; capsula 9.5 cm. longa,

valvis 2 cm. latis. — Ecuador: Base of Mt. Chimborazo, alt. 900 m.,

June, 1860, R. Spruce 6186 (Herb. Kew., type; photo, in herb. Field

Mus.). Fruit was collected from the same plant in August, 1860.

From H. parasitica Jacq. this species is easily differentiated by

the very large leaves with conspicuous nerves, and by the larger

flowers.

Hillia parasitica Jacq. Enum. PI. Carib. 18. 1760.

Andes, Spruce 5079 (G). Also in Brazil, the West Indies, Mexico,

and elsewhere in tropical America.

A glabrous epiphytic shrub; leaves petiolate, elliptic, 6-14 cm.

long, cuspidate-acuminate; flowers terminal, solitary, sessile; calyx

6-parted or absent; corolla white, the slender tube 6-10 cm. long.

7. COUTAREA Aubl.

Coutarea Andrei, sp. npv. — Ramuli rigidi subteretes ochracei

rimosi glabri, internodiis brevibus; stipulae ovato-triangulares acutae

2 mm. longae; petiolus glaber 1-1.5 mm. longus; lamina ovato-

elliptica vel late ovata, 1.5-3 cm. longa, 1-1.6 cm. lata, obtusa, basi

obtusa vel subacuta, membranacea, glabra, fere concolor, costa sub-

tus prominente gracili pallida, nervis lateralibus utrpque latere c. 5,

adscendentibus, arcuatis, remote a margine conjunctis; inflprescentia

terminalis 1-4-flora umbelliformis, pedicellis 2-3 mm. longis glabris;

hypanthium obovatum 3 mm. longum glabrum; calyx 5-partitus,

laciniis lanceolato-subulatis inaequalibus 3-6 mm. longis ciliolatis;

corolla tubulosa 17-20 mm. longa glabra, tubo angulatp superne

paullo dilatato, fauce 3-4 mm. lato, lobis erectis late oyatis obtusis

fere 4 mm. longis; filamenta gracilia glabra, antheris linearibus

rectis 7 mm. longis. — Ecuador: Chuquiribamba, November 16, 1876,

E. Andre 4443 (Herb. Field Mus. No. 533,792, type; duplicates

in K and Y).

In its small flowers, narrow corolla, and small leaves this is quite

distinct from all other members of the genus.

Coutarea Lindeniana Baill. Adansonia 12: 300. 1879.

Near Ona, Jameson (K). Also in Colombia and Venezuela.

A small tree; leaves elliptic or oblong-elliptic, petioled, acuminate,

glabrous or sometimes velvety-pubescent beneath; cymes dense,

many-flowered; calyx lobes elongate, linear-subulate; corolla light

purple (according to Jameson), 2.5-3.5 cm. long.

Jameson describes the plant as "a very elegant shrub."

THE RUBIACEAE OF ECUADOR 203

8. CHIMARRHIS Jacq.

Chimarrhis dioica Schum. & Krause, Bot. Jahrb. Engler 40:

312. 1908.

Type from Rio Vargro, at 3,400-3,600 m., Lehmann 7718 (F,

photo, of type ex Herb. Berol.). Also in Peru.

A shrub 3 m. high; leaves short-petiolate, oblong or oblong-

elliptic, 8-20 cm. long, 4-12 cm. wide, acuminate, at the base acute,

subtomentose beneath, especially on the nerves, subcoriaceous;

flowers dioecious, in terminal panicles 3-6 cm. long; calyx lobes ovate,

acute; corolla yellowish green, rotate, 4-5 mm. long, glabrous;

capsule subglobose, 3 mm. long.

9. GONDAMINEA DC.

Condaminea breviflora Standl. Field Mus. Bot. 4: 264. 1929.

Zamora, eastern slopes of the eastern Andes of Loja, 800-1,200

m., Lehmann 5651 (F, type).

Glabrous; stipules 2-3 cm. long, bifid; leaves short-petiolate,

obovate-oblong, 19-27 cm. long, abruptly short-acuminate, truncate

or rounded at the base; corymbs terminal, trichotomous, long-

pedunculate; hypanthium 6-7 mm. long, the calyx truncate, 4 mm.

long; corolla tube 1 cm. long, the lobes oblong, 7 mm. long, obtuse;

anthers exserted.

Condaminea corymbosa (R. & P.) DC. Prodr. 4: 402. 1830.

Macrocnemum corymbosum R. & P. Fl. Peruv. 2: 48. pi. 189. 1799.

Portovelo, Rose & Rose 23^26 (W). Widely distributed in South

America, and in Panama.

A shrub or small tree, usually glabrous, with very large, oblong

or obovate, usually sessile and clasping leaves; calyx 8 mm. long,

truncate or irregularly dentate; corolla 2.5 cm. long; capsule bisulcate,

about 17 mm. long.

10. RONDELETIA L.

Rondeletia chimboracensis, sp. nov. — Frutex tenuis multi-

caulis interdum 6-metralis, ramulis gracilibus subteretibus dense

hirsutis, internodiis brevibus vel elongatis; stipulae persistentes 10-14

mm. longae, parte basali rotundata subscariosa hirsuta, abrupte in

caudam lanceolato-linearem angustam longe attenuatam ciliatam

desinente; folia breviter petiolata opposita, petiolo 5-7 mm. longo

dense hirsute; lamina crasse membranacea, lanceolato-oblonga vel

lanceolata, rarius oblongo-elliptica, 6-11 cm. longa, 2-5 cm. lata,

longissime acuminata, acumine angusto longe attenuate, basi acuta,

supra viridis, dense pilis brevibus fulvis hirsuta, arete bullulata,

nervis nervulisque omnibus profunde impressis, subtus pallida, dense

pilis patentibus et intertextis albidis hirsuta, costa gracili elevata,

nervis lateralibus utroque latere c. 13, gracillimis, prominentibus,

angulo angusto adscendentibus, arcuatis, prope marginem conjunctis,

nervulis arete reticulatis prominentibusque; inflorescentia terminalis

204 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

1.2-2 cm. longe pedunculata cymosa vel cymoso-paniculata laxe

pauciflora, 3-5 cm. longa et aequilata, floribus plerumque secundis

sessilibus vel brevissime pedicellatis, bracteis filiformibus vel lineari-

bus 3-6 mm. longis hirtellis; hypanthium oboyoideum 2 mm. longum

densissime ochraceo-hispidulum; calyx 4-partitus, laciniis patentibus

2.5-3 mm. longis linearibus versus apicem longe attenuatis extus

hispidulis; corolla extus dense albido-hirsuta, tubo albo 16 mm. longo

gracili tereti, lobis 4 rubris rotundatis 3 mm. longis intus minute

papillosis. — Ecuador: At the base of Mt. Chimborazo, alt. 900 m.,

July, 1860, R. Spruce 6187 (Herb. Kew., type; photo, in herb. Field

Mus.).

Rondeletia cupreiflora Schum. & Krause, Bot. Jahrb. Engler

40: 314. 1908.

Rio Badcun, 2,000-2,500 m.,Lehmann 5448 (F). Also in Colombia.

A shrub or small tree 5 m. high; stipules triangular, acute, 5-6

mm. long; leaves oblong, 7-14 cm. long, acuminate, sparsely pilose

beneath; flowers 4-parted, in corymbiform panicles; calyx lobes

lance-subulate, nearly equaling the hypanthium; corolla copper-red,

20-23 mm. long, puberulent.

Rondeletia trichotoma, sp. nov. — Arbor tenuis subramosa 7.5

m. alta, ramulis gracilibus subteretibus vel in sicco compressis,

brunneis, laxe cinereo-tomentosis vel glabratis, internodiis brevibus

vel elongatis; stipulae e basi triangulari lineari-cuspidatae, 5 mm.

longae, erectae, persistentes, tomentulosae vel glabratae; folia oppo-

sita, petiolo crassiusculo 3-5 mm. longo tomentuloso vel glabrato;

lamina ellipticq-oblonga vel lanceolato-oblonga, 6-8.5 cm. longa, 2-3

cm. lata, longiacuminata, basi acuta vel attenuata, crasse mem-

branacea, supra viridis, ad nervos arachnoideo-tomentosa vel fere

glabra, nervis inconspicuis, subtus pallidior, ad nervos cinereo-

tomentosa, costa nervisque gracilibus prominentibus, nervis lateral-

ibus utroque latere c. 9, angulo acuto adscendentibus, leviter arcu-

atis, prope marginem conjunctis, nervulis vix prominentibus arete

reticulatis; flores tetrameri in paniculas terminales trichotomas c.

4 cm. longas et 8 cm. latas 5 cm. longe pedunculatas dispositi, cymulis

paucis dense multifloris, floribus sessilibus vel usque ad 3 mm. longe

pedicellatis, bracteis inferioribus linearibus 1 cm. longis, superioribus

c. 5 mm. longis; hypanthium oblongum fere 3 mm. longum dense

albido-tomentosum ; calyx 4-partitus, lobis 2.5-5 mm. longis lineari-

bus vel anguste oblongis obtusis, extus sparse tomentulosis, intus

glabris; corolla rubra extus sparse minuteque tomentulosa, tubo

gracili 12 mm. longo superne paullo ampliato, lobis 4 rotundatis 2

mm. longis crispulis intus glabris. — Ecuador: In the Andes at the

foot of Mount Tungurahua, alt. 1,950 m., in 1857, R. Spruce 5116

(Gray Herb., type; duplicate in K; photo, in herb. Field Mus.).

Although not marked by any outstanding characters, this plant

differs in some important detail from each of the species known from

Colombia.

THE RUBIACEAE OF ECUADOR 205

PSEUDOHAMELIA Wernham

Pseudohamelia hirsuta Wernham, Journ. Bot. 50: 242. pi.

521. 1912.

Type from Ecuador, Fraser.

Stipules ovate, caudate-acuminate; leaves oval, about 10 cm.

long, acute, ferruginous-pilose; flowers in terminal, dichotomous or

trichotomous, subscorpioid cymes, usually sessile; calyx lobes linear,

2-4 mm. long; corolla pilose, the tube 23 mm. long, the 4 lobes 4.5

mm. long; fruit baccate, 2-celled, ovoid, densely villous, 4 mm. long.

This genus is not included in the key to genera here presented,

because there is much uncertainty regarding its proper position.

Wernham places the plant in the Hamelieae, where it would certainly

constitute a distinct genus, but I doubt that it is referred correctly to

that group. Judging from the illustration, there is little to indicate

that the plant has a fleshy fruit. In dried specimens of the Rubiaceae

often it is exceedingly difficult to determine whether a fruit is dry or

fleshy. In general appearance, as indicated in the illustration, the

plant has a strong resemblance to species of Rondeletia, and although

it does not agree, according to description, with any Rondeletia

described or reported from Ecuador, I suspect that it should be

referred there. ^

11. ELAEAGIA Wedd.

Elaeagia utilis (Goudot) Wedd. Monogr. Cinchon. 94. 1849.

Condaminea utilis Goudot, Compt. Rend. 18: 260. 1844.

In the Kew herbarium there is a specimen from the eastern

Andes of Loja, at 1,400^1,800 m., Lehmann 5446, which may belong

to this species. It consists of leaves only, and the determination is

therefore very uncertain. The leaves are 30-40 cm. long and 18-21

cm. wide. The species occurs in the mountains of Colombia.

12. ARCYTOPHYLLUM Willd.

Arcytophyllum Andreanum, sp. nov. — Frutex erectus dense

ramosus, ramis gracilibus, ochraceis vel substramineis, glabris, lucidis,

dense foliatis; stipulae ovatae vel triangulari-ovatae, 1.5 mm. longae,

acuminatae, coriaceae, adpressae, integrae, dense puberulae, per-

sistentes; folia patentia c. 1 mm. longe petiolata, oblonga, 5-9 mm.

longa, 2-3 mm. lata, obtusa, mucronulata, basi obtusa, coriacea,

glabra, costa supra sulcata, subtus prominente, marginibus revolutis;

flores terminates solitarii sessiles; hypanthium glabrum; calycis lobi

4 oblongi acuti 1.5-2 mm. longi erecti vel apice recurvi, sinu angusto

acutq separati; corolla 5 mm. longa extus glabra, tubo superne

ampliato, lobis 4 tubo aequilongis oblongo-triangularibus acutius-

culis, intus breviter albido-villosis; antherae exsertae oblongo-lineares

1.2 mm. longae; capsula ovalis 3 mm. longa. — Ecuador: Tambo de

206 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Sabanilla, December 18, 1876, E. Andre K. 1101 (Herb. Field Mus.

No. 533,558, type; duplicates in K and Y). Tungurahua, 3,000 m.,

Spruce 511 4 (K).

Related to A. vernicosum and A. Jamesonii, but distinguished

from both by the densely puberulent stipules.

Arcytophyllum aristatum Standl. Field Mus. Bot. 7: 37. 1930.

Paramos 13 miles west of Tulcan, Prov. Carchi, 3,300 m., Hitch-

cock 20923 (W, Y). Hills near Cuenca, August, 1864, Jameson (F,

S, W). Villanaco, near Loja, Jameson (K). Hills near Cuenca,

Jameson 52 (K). Also in Colombia.

A prostrate shrub, the stems glabrous; stipule sheath subtruncate,

obscurely bidentate; leaves sessile, narrowly oblong, about 5 mm. long

and 1.5 mm. wide, obtuse or acute and piliferous, glabrous, shining;

flowers terminal, solitary, subsessile; calyx lobes linear-subulate,

1.8 mm. long; corolla white, glabrous, 4.5 mm. long, the tube nearly

twice as long as the lobes.

Arcytophyllum capitatum (Benth.) Schum. in E. & P. Nat.

Pfl. 44: 28. /. 8. 1891. Rachicallis capitata Benth. PI. Hartw. 195.

1845. Ereicoctis crassifolia (Spruce) Kuntze, Rev. Gen. PI. 1: 281.

1891, nomen nudum.

Andes of Ecuador, Spruce 5829 (G, K). Without locality,

Seemann (S). Andes of Ecuador, 3,600-3,900 m., Pearce (K). Also

in Colombia.

An erect glabrous shrub; leaves oblong, acute, coriaceous; stipules

broadly ovate, ciliate with several long bristles; flowers sessile in

terminal heads; corolla white, the lobes pilose within; calyx lobes

linear.

Kuntze's name evidently is based on an unpublished one of

Spruce, appearing upon the labels of his No. 5829.

Arcytophyllum confertum (R. & P.) Standl., var. bryoides

(R. & S.), comb. nov. Houstonia bryoides R. & S. Syst. Veg. 3: 527.

1818. Hedyotis hypnoides HBK. Nov. Gen. & Sp. 3: 389. 1820.

Anotis hypnoides DC. Prodr. 4: 432. 1830. Hedyotis conferta

R. & P. j8 pusilla Wedd. Chlor. And. 2: 42. 1857. Ereicoctis hypnodes

Kuntze, Rev. Gen. PI. 1: 281, 1891.

Type from the Andes of Assuaya, Humboldt & Bonpland (photo,

of type from Herb. Berol. in F). Loja, Zamora, Andre K. 1100 (F,

G, K, Y). Canar, Rose & Rose 22679 (G, W, Y). Near Cuenca,

Jameson (G).

A very slender, creeping, matted shrub; leaves subulate-linear,

glabrous; flowers terminal, solitary, white.

This form is easily distinguished from the Peruvian type by the

very short, slender leaves, and may deserve specific rank.

THE RUBIACEAE OF ECUADOR 207

Arcytophyllum Jamesonii, sp. nov. — Frutex erectus dense

ramosus, ramis crassis ochraceis vel fuscis glabris, stipulis persistenti-

bus imbricatis fere occultis; stipulae triangulares yel late ovatae,

1.5 mm. Ipngae, glabrae, acutae vel breyiter acuminatae, integrae

vel 3-denticulatae, adpressae, lucidae; folia ovata c. 4 mm. longa et

2.3 mm. lata, involuta, obtusa, mutica, glabra, supra concava, subtus

obtuse carinata, adscendentia vel patentia, coriacea, marginibus

tenuibus; flores terminales, sessiles, subcapitati; hypanthium late

obovoideum 1.5 mm. longum glabrum; calycis lobi 4 anguste tri-

angulares obtusiusculi vel acuti crassi glabri erecti; corolla extus

glabra, tubo crasso 3 mm. longo, lobis 4 patentibus oblongis obtusis

3 mm. longis, intus papilloso-puberulis; capsula subglobosa lucida

2.5 mm. longa. — Ecuador: Vicinity of Cafiar, September 16, 1918,

J. N. & George Rose 22771 (U. S. Nat. Herb. No. 1,022,389, type;

duplicate in Y). Mount Pillzhicui, 3,600 m., Jameson (G).

The plant is much like A. vernicosum in general appearance, but

differs in its broad involute obtuse leaves, numerous flowers, and

broader calyx lobes.

Arcytophyllum juniperifolium (R. & P.), comb. nov. Hedyotis

juniperifolia R. & P. Fl. Peruv. 1: 57. pi. 87, f. c. 1798.

Between Ona and Cuenca, Prov. Aznay, 2,700-3,300 m., Hitch-

cock 21594 (W). Quebrada Chalan, Punin, 2,760 m., Anthony &

Tate 424 (W). Antisanilla, 3,450 m., Anthony & Tate 336 (W).

Cuenca, Rose, Pachano & Rose 22883 (W). Also in Peru.

A densely branched shrub, the branches puberulent; stipules

ovate, 1-3-dentate; leaves linear, subobtuse, muticous, 2.5-3.5 mm.

long, revolute, glabrous; flowers terminal, usually in 3's and short-

pedicellate; calyx lobes triangular-lanceolate; corolla glabrous, white.

Arcytophyllum microphyllum (Willd.), comb. nov. Houstonia

microphylla Willd. ex R. & S. Syst. Veg. 3: 527. 1818. H. serpens

HBK. Nov. Gen. & Sp. 3: 390. pi. 289. 1820. Anotis serpens DC.

Prodr. 4: 433. 1830. Ereicoctis microphylla Kuntze, Rev. Gen. PI.

1:281.1891.

Roemer and Schultes give the type locality as "Quindiu" (Colom-

bia), but Kunth states that the type is from the Volcan de Antisana

(Ecuador) at 3,900 m. Cuenca, Rose, Pachano & Rose 23797 (G, Y).

Guanojo, Andre 3969 (F, G, Y). San Jose", Andre K534 (F, G, Y).

Tungurahua, December, 1865, Spruce (K). Pichincha, Holmgren

214 (F, S). Valle de Turubamba, Prov. Pichincha, 2,800 m., Firmin

646 (W). Weddell reports Hartweg 1065 and 1067 from Pichincha.

Also in Venezuela.

Plants prostrate, the branches filiform; leaves rounded-ovate or

elliptic-ovate, subobtuse, glabrous, the margins scaberulous; flowers

axillary, solitary, short-pedicellate; capsule broadly rounded, com-

pressed; calyx lobes spatulate-lanceolate, acute.

208 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Arcytophyllum Riveti Dang. & Cherm. Bull. Mus. Hist. Nat.

Paris 28: 434. 1922.

Type from Chillacocha, at 3,500 m., Rivet 774. Surucucho, near

Cuenca, July, 1864, Jameson (S, W). Villanaco, near Loja, Jameson

(P). Calaisaca, 2,500 m., Townsend 999 (W). Ecuador (?), Lobb

(K). Without locality, Jameson (F). Cisne, Andre K706 (G).

Hills, Loja, Jameson 50 (K). Loja, Seemann 929 (K). Villanaco,

Prov. Loja, January, 1883, Hubsch (K). Zamora, mountains east

of Loja, October 1, 1861, Cross (K). Also in Peru.

An erect, densely branched shrub, the branches puberulent;

stipules triangular, dentate; leaves linear, revolute, 2 mm. long,

glabrous; flowers solitary, terminal; calyx lobes lanceolate, acute;

corolla 5-6 mm. long, purplish, glabrous.

Arcytophyllum setosum (R. & P.), comb. nov. Hedyotis setosa

R. & P. Fl. Peruv. 1: 57. pi. 88, f. a. 1798.

Andes of Ecuador, Spruce 5887 (G, K, B, S). Without locality,

Lehmann 4683 (W). Quito, Jameson 43 (G), 148 (G). Without

locality, Lobb 166 (K). Loja, Seemann 933 (K). Surucucho, near

Cuenca, Jameson 16 (K). Without definite locality, at 4,200 m.,

Jameson 138 (K). Also in Peru.

An erect, densely branched, nearly glabrous shrub; leaves oblong

to ovate, mostly 5-7 mm. long, acute, mucronate, coriaceous; stipules

ovate, ciliate with several long bristles, glabrous; flowers few, at the

ends of the branches, subcapitate; corolla glabrous within.

Arcytophyllum thymifolium (R. & P.), comb. nov. Hedyotis

thymifolia R. & P. Fl. Peruv. 1: 56. pi. 88, f. a. 1798. H. coarctata

Willd. ex R. & S. Syst. Veg. 3: 527. 1818. H. thujoides Willd. ex

R. & S. loc. cit. 1818. H. thesioides Willd. ex HBK. Nov. Gen.

& Sp. 3: 391. 1820, as syn. Anotis thymifolia DC. Prodr. 4: 432. 1830.

Anotis thymifolia DC. 0 thesioides DC. Prodr. 4: 432. 1830.

Guaranda, Andre K535 (F, G). Quitensian Andes, in 1855,

Couthouy (G, P). Andes, Spruce 4984 (G). Quito, Rose & Rose

23543 (G, W, Y). Rio Pita, Chillo Valley, 2,940 m., Anthony &

Tate 216 (W). Bafios, Prov. Tungurahua, 1,950 m., Tate 641 (W).

Ambato, Prov. Tungurahua, Pachano 18 (W, Y), 100 (W, Y); alt.

2,600 m., Hitchcock 21717 (W, Y). Valle de La Magdalena, Prov.

Pichincha, 2,800 m., Firmin (W). Huigra, Rose & Rose 22253 (G,

W, Y) ; at 1,200 m., Hitchcock 20619 (W, Y). Cuenca, Rose, Pachano

& Rose 22883 (Y). Andes, alt. 2,500-3,000 m., Lehmann 150 (W).

Loja, 1,800-2,500 m., Lehmann 7715 (F, W). Valle de Turubamba,

Prov. Pichincha, 2,800 m., Firmin 512 (F). Volcan de Tungurahua,

1,500-2,500 m., Lehmann 5137 (F). Chreabamba, Lehmann B.T.688

(F, G). Pichincha, Holmgren 224 (F, S). Loja, Seemann 930 (K),

757 (K). Andes, in 1857-59, Spruce (K). Rio Chambo, Spruce

4984 (K). Near Quito, Hartweg 1073 (Y). Andes of Quito, 2,400-

3,000 m., very abundant, Jameson 748 (L). Pichincha, Karsten (L).

Prov. Pichincha, 2,700 m., Firmin 648 (W). "Quito," Humboldt

THE RUBIACEAE OF ECUADOR 209

(F, photo, of type of H. coarctata, from Herb. Berol.). Also in

Colombia and Peru.

An erect shrub, the branches puberulent; stipules triangular-

ovate, 1-3-dentate; leaves linear, 5-15 mm. long, revolute; flowers

in small terminal cymes; corolla white, glabrous, 6-8 mm. long.

Arcytophyllum vernicosum, sp. nov. — Frutex erectus dense

ramosus, ramis crassiusculis teretibus ochraceis stipulis persistentibus

imbricatis saepius occultis; stipulae 1.5 mm. longae triangulares

acutae vel acuminatae coriaceae lucidae arete adpressae, integrae

vel apice obscure bidenticulatae, glabrae; folia sessilia 3-4 mm.

longa, 1.5 mm. lata, acuta, basi late rotundata, coriacea, lucidissima,

glabra, adscendentia, marginibus valde incrassatis; flores terminales

sessiles solitarii vel glomerati; calycis lobi lineari-triangulares 2.5

mm. longi approximati erecti attenuati crassi lucidi; capsula subglo-

bosa 2 mm. longa lucida. — Ecuador: Without locality, W. Jameson

(Herb. Field Mus. No. 578,871, type; duplicate in W). Without

locality, Seemann (G). Hills near Loja, September, 1864, Jameson

(F, S). Hills near Loja, Jameson 51 (K).

The species is well differentiated by the usually entire, tightly

imbricate stipules, by the shining leaves and branches, and by the

small size of the leaves.

TEINOSOLEN Hook. f. in B. & H. Gen. PI. 2: 61. 1873.

Schumann (in E. & P. Nat. Pfl. 44: 27. /. 8, J. 1891) states that

the genus consists of "3-4 species in the Andes of Bolivia and Quito,

for example, T. Jamesonii K. Sch." T. Jamesonii is evidently the

plant of Quito, but it has not been published formally. The genus

should probably be reduced to synonymy under Arcytophyllum.

13. GONZALAGUNIA R. & P.

Gonzalagunia cornifolia (HBK.) Standl. Field Mus. Bot. 4:

279. 1929. Gonzalea cornifolia HBK. Nov. Gen. & Sp. 3: 416. 1819.

Peripa, Andre 4228 (F, K). Also in Colombia and Peru.

A slender shrub, the branches often recurved, hispidulous; stipules

subulate; leaves short-petiolate, lanceolate to ovate, long-acuminate,

at the base obtuse or rounded, appressed-pilose, thin; panicles

elongate, spikelike; calyx lobes short, deltoid or rounded; corolla

white, 8-10 mm. long; fruit white, 2-3 mm. broad.

Gonzalagunia dependens R. & P. Fl. Peruv. 1: 56. pi. 86.

1798. Gonzalea tomentosa H. & B. PI. Aequin. 1: 225. pi. 64. 1808.

Gonzalagunia tomentosa Kuntze, Rev. Gen. PL 1: 284. 1891. Duggena

tomentosa Standl. Contr. U. S. Nat. Herb. 18: 126. 1916.

Type of Gonzalea tomentosa collected between Loja and Gon-

zanama, Humboldt & Bonpland. Pichincha, Jameson (Y). Without

locality, Jameson (W). Also in Colombia and Peru.

210 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

A slender shrub; leaves lanceolate or ovate, rugose, glabrous

above, densely white-tomentose beneath; corolla white, the tube 5

mm. long; fruit white, 4-coccous.

Gonzalagunia flexuosa Standl. Field Mus. Bot. 4: 280. 1929.

Valley of Pastaza River, between Banos and Cashurco, Prov.

Tungurahua, 1,300-1,800 m., Hitchcock 21 803 (Y). Also in Colombia.

A shrub or small tree; leaves short-petiolate, ovate-oblong or

narrowly ovate, 10.5-15.5 cm. long, long-acuminate, above sparsely

strigose or glabrate, beneath pilose with spreading hairs and strigose

along the veins; rachis of the elongate panicle flexuous, densely

strigose; calyx lobes oblong-linear, 1-2 mm. long, unequal; corolla

white, 6-8 mm. long; fruit 4-coccous.

Gonzalagunia mollis Spruce ex Schum. in Mart. Fl. Bras. 66:

290. 1889. Gonzalea mollis Wernham, Journ. Bot. 51: 219. 1913.

Duggena mollis Standl. Contr. U. S. Nat. Herb. 18: 125. 1916.

Andes of Ecuador, Spruce 5052 (G, type collection).

Leaves short-petiolate, lanceolate, acuminate, rounded or obtuse

at the base, canescent-tomentose beneath; calyx lobes ovate- tri-

angular; corolla 12-14 mm. long, appressed-hirsute; fruit 4-coccous.

Gonzalagunia pulverulenta (H. & B.) Kuntze, Rev. Gen. PI.

1: 284. 1891. Gonzalea pulverulenta H. & B. PL Aequin. 1: 228.

1808. Duggena pulverulenta Standl. Contr. U. S. Nat. Herb. 18:

126. 1916.

Type from Gonzanama, Humboldt & Bonpland.

Leaves lanceolate, obtuse at the base, pulverulent-pubescent

beneath.

The plant is so briefly described that it is impossible to determine

whether it is identical with any of the species reported from Colombia,

as may very likely be the case.

Gonzalagunia sessilifolia, sp. nov. — Frutex tenuis sarmentosus

parce ramosus 3-metralis, ramulis gracilibus teretibus dense ochraceo-

strigosis, internodiis elongatis; stipulae 10 mm. longae, e basi paullo

dilatato lineares, sparse strigosae, persistentes; folia sessilia, late

ovata, 6-15 cm. longa, 2.5-8.5 cm. lata, abrupte acuminata, acumine

longo angusto caudato-attenuatp, basi late rotundata vel subcordata,

firme membranacea, supra viridia, glabra, nervis non eleyatis, subtus

pallida, praesertim ad nervos dense ochraceo-strigosa, inter nervos

sparse strigosa vel glabrata, costa gracili elevata, nervis lateralibus

utroque latere c. 10, gracilibus, prominentibus, angulo acuto adscen-

dentibus, juxta marginem conjunctis, nervulis vix prominulis arete

reticulatis; paniculae spiciformes simplices flexuosae breviter pedun-

culatae, 22 cm. longae et ultra, rhachi gracili dense ochraceo-strigosa,

floribus glomeratis, glomerulis paucifloris distantibus, bracteis subu-

latis 2-4 mm. longis; hypanthium globosum dense strigosum 1-1.5

THE RUBIACEAE OF ECUADOR 211

mm. longum; calyx 4-lobus, lobis valde inaequalibus, pvalibus vel

oblongis, 1-2.5 mm. longis, obtusis, extus sparse strigosis, intus gla-

bris; corolla alba setoso-strigosa 2.5-3 mm. longa, tubo superne paullo

ampliato, lobis 4 brevissimis late ovatis obtusis tubo duplo breviori-

bus; fructus albus depresso-globosus 4 mm. latus dense strigosus. —

Ecuador: Ad radices mentis Chimborazo, 900 m., July, 1860, R.

Spruce (Herb. Kew., type; photo, in herb. Field Mus.).

The large broad sessile leaves and the minute corollas are char-

acters which enable one to differentiate this species from all others

known from the region.

Gonzalagunia sororia, sp. nov. — Frutex, ramulis gracilibus

teretibus dense strigosis vel hinc inde hispidulis, internodiis elongatis;

stipulae lanceolato-triangulares longiacuminatae 3-6 mm. longae

glabratae erectae persistentes; petiolus crassus 3-4 mm. longus his-

pidulus; lamina lanceolato-oblonga vel oblongo-ovata, 4.5-7 cm.

longa, 1.5-2.5 cm. lata, longiacuminata, acumine obtuso, basi obtusa

vel acuta, coriacea, valde rugosa, supra viridis, ad costam strigosa,

nervis profunde impressis, subtus pallidior, ad nervos neryulosque

pilis brevibus adpressis vel patentibus hispidula, costa gracili elevata,

nervis lateralibus utroque latere c. 11, elevatis, angulo acuto adscen-

dentibus, gracilibus, leviterarcuatis, marginem attingentibus, nervulis

prominentibus rectis parallelis connexis; paniculae spiciformes 3-7

cm. longae graciles multiflorae, simplices vel prope basin breviter

ramosae, rhachi pilis longiusculis patentibus vel adpressis pilosa,

floribus solitariis vel aggregatis 1-2 mm. longe pedicellatis; bracteae

lineares vel oblongae pedicellis aequilongae vel paullo longiores, fere

glabrae; hypanthium subglobosum fere 2 mm. longum strigosum;

calyx 1-1.5 mm. longus 4-lobus, lobis triangularibus acutis inaequali-

bus glabratis; corolla alba in alabastro 5 mm. longa sparse strigosa,

tubo superne paullo sensimque dilatato; fructus 4-coccus 3-3.5 mm.

latus depresso-globosus, pilis brevibus subadpressis pilosulus. —

Ecuador: Between La Chorrita and Portovelo, Prov. Oro, alt.

1,000-2,000 m., August 28, 1923, A. S. Hitchcock 21176 (U. S. Nat.

Herb. No. 1,196,221, type; duplicate in Y).

In its strongly rugose leaves this plant of Ecuador is similar to

G. dependens R. & P., but in that species the leaves are densely white-

tomentose beneath.

14. ISERTIA Schreb.

Isertia hypoleuca Benth. in Hook. Journ. Bot. 3: 220. 1841.

Ayabamba (Ecuador?), Andr6 430 (K). Ranging to the Guianas,

Brazil, and Peru.

A tree 6^-8 m. high; leaves large, long-petiolate, obovate-oblong,

densely white-tpmentose beneath; panicles large, dense, many-

flowered, thyrsoid; corolla tubular, 5-6 mm. long, red, more or less

tomentose; fruit baccate.

212 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

15. HIPPOTIS R. & P.

Hippotis scarlatina Krause, Bot. Jahrb. Engler 40: 324. 1908.

In wet forest near Zamora, Lehmann 7712 (W, type collection;

photo, of type ex Herb. Berol. in F).

A tree 10 m. high; leaves short-petioled, elliptic-obovate, acumi-

nate, sericeous beneath, finely striolate between the nerves; flowers

axillary; corolla dark red, 2.5-3 cm. long, sericeous.

16. PENTAGONIA Benth.

Pentagonia orthoneura, sp. nov. — Arbor tenuis 4.5 m. alta

ramos paucos patulos proferens, ramulis crassis obtuse tetragonis

glabris; stipulae angustae 5 cm. longae acuminatae ferrugineae extus

minute strigillosae; folia opposita, petiolo gracili 4.5 cm. longo minu-

tissime puberulo vel glabrato; lamina subcoriacea, oblongo-elliptica,

44 cm. longa, 19 cm. lata, acuta, basi subacuta, supra fusca, dense

minuteque adpresso-pilosula, nervis prominulis, subtus brunnescens,

obscure minuteque adpresso-pilosula, inter nervulos pulchre et

creberrime lineolata, costa gracili eleyata, nervis lateralibus utroque

latere c. 21, angulo lato divergentibus, gracilibus, elevatis, fere

rectis, non ramosis, prope marginem arcuato-conjunctis, nervulis

vix prominulis reticulatis; bacca globosa 2.5-3 cm. longa, basi et

apice rotundata, glabra, pericarpio 2 mm. crasso; calyx persistens

6 mm. longus 5-lobus, lobis late oblongis obtusis. — Ecuador: At

foot of Mount Chimborazo, along the Rio Chasuan, rare, alt. 900-

1,200 m., August, 1860, R. Spruce 6231 (Herb. Kew., type; photo,

in herb. Field Mus.).

The collector's notes state that "Fruit was abundant, but I

could find no good flowers." The specific name here used was

suggested by Spruce, but under a new generic name still unpublished.

Pentagonia peruviana Standl. Field Mus. Bot. 4: 326. 1929.

Banks of the Rio Pastaza, Spruce 4968 (K, type; photo, in F).

"A scandent shrub;" leaves short-petiolate, obovate-elliptic, 32-

45 cm. long, 16-19 cm. wide, acute, at the base acute or subacumi-

nate, glabrous above, strigose beneath along the veins and elsewhere

strigillose; flowers clustered in the leaf axils, short-pedicellate; calyx

closed before anthesis, in flower spathaceous and cleft along one side,

2 cm. long; corolla 3 cm. long or more, glabrous outside.

It is unfortunate that the specific name peruviana should have

been given carelessly to this plant, but there is apparently no

authority for changing the specific name at the present time.

Pentagonia Sprucei, sp. nov. — Arbor tenuis simplex vel ramos

perpaucos suberectos emittens 4.5-12-metralis; stipulae crassae 14

cm. longae lanceolato-triangulares Ipngiacuminatae, extus dense pilis

brevibus fulvis adscendentibus hispidulae; folia magna, petiolo 9 cm.

THE RUBIACEAE OF ECUADOR 213

longo subtereti dense velutino-pilosulo; lamina coriacea, 27 cm. lata

et ultra, basi subcordata, supra glabra, subtus ubique dense velutino-

pilosula, costa crassa subtereti, nervis lateralibus angulo recto diver-

gentibus crassis elevatis remote a margine dichotomis, inter nervulos

densissime et parallele striato-nervulosa; flores in axillis foliorum

cymoso-paniculati, paniculis petiolo brevioribus laxe paucifloris,

floribus sessilibus yel subsessilibus; hypanthium turbinatum 7 mm.

longum, sparse minuteque puberulum vel glabratum; calyx cam-

panulatus 12 mm. longus glaber 5-lobus, lobis late ovatis obtusis

vel acutis; corolla alba coriacea extus glabra, tubo 4 cm. longo basi

5 mm. lato, superne sensim dilatato, fauce 13 mm. lato, intus ad

insertionem staminum dense albido-barbato, lobis ovatis obtusis 8

mm. longis; stamina c. 7 mm. supra basin tubi inserta, filamentis 1.5

cm. longis filiformibus glabris; bacca subglobosa 2.5 cm. longa,

pericarpio 3 mm. crasso, basi et apice rotundata. — Ecuador: Ad

radices montis Chimborazo, secus flum. Chasuan, frequens, alt.

300-1,500 m., July-September, 1860, R. Spruce 6230 (Herb. Kew.,

type; photo, in herb. Field Mus.).

When growing this must be a handsome plant, as are most of the

other species of the genus. From other South American Pentagonias

it is easily distinguished by the copious soft pubescence of the leaves.

The stipules, also, are remarkable for their large size.

Spruce indicated the plant as a member of a new genus, doubtless

unaware of the fact that similar plants had been described from

Panama. It is fitting that this striking plant should bear the name

of its discoverer, most celebrated and ablest botanical explorer who

has ever worked in western South America.

17. SABICEAAubl.

Sabicea villosa R. & S. Syst. Veg. 5: 265. 1819. S. hirsuta

HBK. Nov. Gen. & Sp. 3: 417. 1820.

Balao, in forest, Eggers 14401 (L, W). Teresita, Stevens 74 (W).

Wernham (Monogr. Sabicea 55. 1914) reports a specimen from Valle

Mindo, Sodiro. The species is widely distributed in tropical America.

A scandent shrub; stipules broadly ovate, brown, reflexed;

leaves petiolate, elliptic-oblong, acuminate at each end, hirsute,

6-12 cm. long; flowers sessile in axillary clusters; calyx lobes linear-

lanceolate, up to 4 mm. long, accrescent; corolla white, the tube

6 mm. long, strigose above, the lobes triangular, minute; fruit violet,

fleshy, 1 cm. long.

18. BERTIERA Aubl.

Bertiera procumbens Schum. & Krause, Bot. Jahrb. Engler

40: 328. 1908.

Balao, in forest, Eggers 14282 (W, type collection).

A shrub 2.5 m. high; leaves very shortly petiolate, oblong or

oblong-lanceolate, 6-12 cm. long, acuminate, pilose; flowers small,

214 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

white, in terminal panicles 8-12 cm. long; berries globose, blue,

glabrous, 2-3 mm. in diameter.

19. HAMELIAJacq.

Hamelia grandiflora Spruce ex Wernham, Journ. Bot. 49: 209.

1911.

Type from Chimborazo, Spruce 61 93 ; also Triana 1 759. Teresita,

Prov. Guayas, 270 m., Hitchcock 20554 (W, Y).

A tree; leaves mostly verticillate, elliptic or elliptic-oblong, 15-25

cm. long, glabrous; corolla 3 cm. long, yellow, funnelform; fruit

1 cm. long.

Hamelia patens Jacq. Enum. PI. Carib. 16. 1760.

El Recreo,Eggers 15449 (F). Ventura, Rose & Rose 23516 (G, Y).

Between Huigra and Naranjapata, Prov. Chimborazo, 600-1,200 m.,

Hitchcock 20685 (W, Y). Chimborazo, 750-900 m., Spruce 6226 (K).

San Ignacio, Prov. Guayas, Heilborn 81 (F, S). Widely distributed

in tropical America.

A shrub or small tree, sometimes 4.5 m. high; leaves mostly

ternate, pubescent or tomentose beneath; corolla short, tubular,

orange-red; fruit a juicy berry.

Hamelia pedicellata Wernham, Journ. Bot. 49: 212. 1911.

Milagro, Prov. Guayas, 50 m., Hitchcock 20291 (W, Y). Also

in Colombia and Venezuela.

A shrub 2 m. high; leaves mostly in 4's, nearly glabrous; flowers

chiefly pedicellate; corolla tubular, 1.5 cm. long; berries red or purple.

20. HOFFMANNIA Sw.

Hoffmannia ecuatoriana, sp. nov. — Frutex 1.8 m. altus, ramis

crassis teretibus glabris, internodiis elongatis; stipulae non visae; folia

opposita, petiolo vero fere nullo; lamina elliptica vel obovato-elliptica,

membranacea, c. 24 cm. longa et 10 cm. lata, abrupte angusteque

acuminata, acumine attenuate acuto, basin versus angustata,

abrupte contracta et longissime decurrens, parte infima petioliformi

et c. 4 cm. longa, supra yiridis, glabra, nervis non elevatis, cqsta

subimpressa, subtus pallidior, ad nervos sparse et minutissime

puberula vel fere omnino glabra, costa crassa elevata, nervis lateral-

ibus utroque latere c. 15, angulo lato adscendentibus, arcuatis, gracili-

bus, prominentibus, prope marginem conjunctis, nervulis prominulis

laxe reticulatis; flores cymosi, cymis ad nodos inferiores defoliatos

insertis, laxe paucifloris, '2 cm. longe pedunculatis, pedicellis 2-4

mm. longis glabris; hypanthium obovoideum 2 mm. longum glabrum;

calyx brevissime 4-dentatus glaber, dentibus latissime triangularibus;

corolla rubra in alabastro oblongo-linearis obtusa, extus glabra, lobis

4 linearibus tubo duplo longioribus. — Ecuador: Province Tungura-

hua, Valley of Pastaza River between Banos and Cashurco, alt. 1,300-

THE RUBIACEAE OF ECUADOR 215

1,800 m., September 25, 1923, A. S. Hitchcock 21836 (Gray Herb.,

type; duplicates in W and Y).

Hoffmannia Sprucei, sp. nov. — Ramuli graciles teretes dense

ferrugineo-villosuli, vetustioribus fusco-ferrugineis, internodiis elon-

gatis; stipulae triangulares acutae 2 mm. longae ferrugineo-tomentu-

losae; folia opposita, petiolo gracili 1.3-2.2 cm. longo villosulo;

lamina subcoriacea, elliptico-oblonga vel ovato-oblonga, 9-13 cm.

longa, 3.5-4.7 cm. lata, abrupte acuminata, acumine angusto obtuso,

basi acuta vel contracta et decurrens, supra sparse villosula, nervis

non elevatis, subtus ferruginea, ubique ferrugineo-villosula, costa

gracili elevata, nervis lateralibus utroque latere c. 10, gracilibus,

angulo lato adscendentibus, prominentibus, prope marginem con-

junctis, nervulis prominulis laxe reticulatis; inflorescentiae axillares

sessiles, e basi ramosae, 1-2.5 cm. longae, multiflorae, floribus cymosis

1-4 mm. longe pedicellatis vel rarius sessilibus, pedicellis ferrugineo-

villosis; hypanthium turbinatum 1.5 mm. longum villosulum; calyx

1-1.4 mm. longus villosulus, dentibus triangularibus acutis erectis;

.corolla extus glabra 6-7 mm. longa, tubo superne vix dilatato, lobis

oblongis obtusis tubo fere duplo brevioribus. — Ecuador: Andes of

Ecuador, 1857-59, R. Spruce 5098 (Gray Herb., type; photo, in

herb. Field Mus.).

Related to H. latifolia (Bartl.) Kuntze, of Peru and Bolivia, but

distinct in the much narrower leaves on long petioles.

21. RANDIA L.

Randia aculeata L. Sp. PL 1192. 1753. R. glomerata Benth.

Bot. Voy. Sulph. 103. 1844.

Type of R. glomerata from Atacames. The species ranges north-

ward to Mexico, and is widely distributed in the West Indies.

A spiny shrub with small, nearly or quite glabrous leaves; corolla

white, 6 mm. long, the throat barbate; fruit globose, 6-12 mm. in

diameter, the pulp blackish and full of large seeds.

Although I have seen no specimens of this species from Ecuador,

the description of R. glomerata indicates that the plant so named is

merely a form of R. aculeata.

Randia aurantiaca Standl. Field Mus. Bot. 4: 327. 1929.

Playas, Peninsula Morro, Mille 190 (W). Also in Peru.

An unarmed shrub 1-4 m. high; leaves oblong-elliptic or obovate-

elliptic, 4-13 cm. long, 2.5-6.5 cm. wide, obtuse or abruptly short-

acuminate, sparsely or densely pilose above, densely grayish-tomen-

tose beneath; flowers perfect, solitary or clustered at the tips of the

branches; calyx lobes linear-subulate; corolla orange, densely grayish-

sericeous, the tube 2 cm. long, the lobes ovate-oblong, long-acuminate,

2.5 cm. long; immature fruit ellipsoid, 1.5 cm. long, puberulent.

216 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Randia calycina Cham. Linnaea 9: 246. 1834. Basanacantha

calycina Schum. in Mart. Fl. Bras. 66: 375. 1889.

Balao, in forest, Eggers 14392 (W). El Recrep, Prov. Manabi,

Eggers 15521 (K, L, W). Also in Bolivia and Brazil.

A spiny shrub 2 m. high; leaves pblong-obovate to rounded-

obovate, nearly glabrous; flowers dioecious; calyx lobes obovate or

oblanceolate; corolla 2-2.5 cm. long, glabrous.

Similar to R. spinosa, but in that species the calyx lobes are linear

or subulate.

Randia formosa (Jacq.) Schum. in Mart. Fl. Bras. 66: 342.

1889. Mussaenda formosa Jacq. Enum. PI. Carib. 16. 1760.

Balao, in savannas, Eggers 14192 (L, W). Guayaquil, in 1852,

Andersson (S). Widely distributed in South America, Panama, and

the West Indies.

An unarmed shrub; leaves more or less pilose beneath; flowers

perfect; corolla white, the slender tube 5-10 cm. long, the lobes

lanceolate to ovate, acuminate; fruit large, oblong.

Randia spinosa (Jacq.) Karst. Fl. Columb. 2: 128. 1869. Mus-

saenda spinosa Jacq. Sel. Stirp. 70. 1763. Gardenia armata Sw.

Prodr. Veg. Ind. Occ. 51. 1788. R. armata DC. Prodr. 4: 387. 1830.

M. pubescens HBK. Nov. Gen. & Sp. 3: 410. 1818. Gardenia

Humboldtiana R. & S. Syst. Veg. 5: 243. 1819. M. Humboldtiana

Steud. Norn. Bot. 540. 1841. G. pubescens Bartl. ex DC. Prodr. 4:

387. 1830, as syn. Basanacantha spinosa Schum. in Mart. Fl.

Bras. 6c: 376. 1889. B. spinosa var. pubescens Schum. in Mart.

Fl. Bras. 66: 378. 1889.

"Guayaquil," Ruiz (B). Type of M. pubescens from Guayaquil,

Humboldt &Bonpland (photo, of type, ex Herb. Berol., in F). Widely

distributed in tropical America.

A spiny shrub or small tree; leaves usually pubescent, large;

flowers dioecious; corolla white, the tube about 2.5 cm. long; fruit

subglobose, 2.5-3.5 cm. long, filled with blackish pulp and few

large seeds.

22. GENIPA L.

Genipa americana L. Syst. Nat. ed. 10. 931. 1759. G. oblon-

gifolia R. & P. Fl. Peruv. 2: 67. pi. 220, /. a. 1799.

Reported by DeCandoHe (Prodr. 4: 378. 1830) from Guayaquil.

Widely distributed in tropical America.

A medium-sized tree; leaves large, obovate, short-petiolate, gla-

brous; flowers yellowish white, 2-4.5 cm. long, in small terminal

cymes; fruit globose, brownish, 6-7 cm. long, the pulp filled with

numerous large compressed seeds.

THE RUBIACEAE OF ECUADOR 217

23. MACHAONIA H. & B.

Machaonia acuminata H. & B. PL Aequin. 1: 101. pi. 29. 1808.

Type from Guayaquil, Humboldt & Bonpland (fragment, ex Herb.

Berol., in F). Daule, Spruce 6303 (S). Banks of Guayaquil River,

Jameson 526 (L). Also in Brazil and Colombia.

A small tree; leaves ovate-elliptic, acuminate, short-petiolate,

densely pubescent beneath; flowers very small, in broad panicles;

sepals ovate, obtuse; fruit dry, densely pubescent.

The vernacular name is given as "ceiba blanca."

24. GUETTARDA L.

Guettarda hirsuta (R. & P.) Pers. Syn. PI. 1: 200. 1805.

Laugeria hirsuta R. & P. Fl. Peruv. 2: 22. pi. 45, f. a. 1799.

Andes of Ecuador, Spruce 5195 (G, Y). Also in Peru.

A shrub or small tree; leaves elliptic, acuminate, fulvous-tomen-

tose beneath, hirsute along the nerves, at least when young, glabrate

above.

The Ecuador collection may be distinct from the Peruvian plant,

but more material from both regions is necessary in order to determine

the specific variations.

25. ANISOMERIS Presl

Anisomeris ecuadorensis Schum. Bot. Jahrb. Engler 40: 329.

1908.

Hacienda El Recreo, Prov. Manabi, Eggers 15411 (F, W, type

collection; photo, of type from Herb. Berol., in F). El Recreo, April

30, 1897, Eggers (F).

A tree 5-6 m. high; leaves ovate-elliptic or elliptic, 4-6 cm. long,

barbate beneath in the axils of the nerves; corolla tube 20-23 mm.

long.

26. CHIOCOCCA L.

Chiococca alba (L.) Hitchc. Rept. Mo. Bot. Gard. 4: 94. 1893.

Lonicera alba L. Sp. PL 175. 1753. C. racemosa L. Syst. Nat. ed. 10.

917. 1759. C. brachiata R. & P. FL Peruv. 2: 67. pi. 219. 1799. C.

trisperma Hook. f. Trans. Linn. Soc. 20: 219. 1847.

El Recreo, Eggers 15409 (F); reported from the Galapagos

Islands. Widely distributed in tropical America.

A slender shrub, the branches usually long and recurved or fre-

quently subscandent; flowers pale yellow or whitish, small; fruit

compressed, orbicular, white, fleshy.

27. GOFFEA L.

Coffea arabica L. Sp. PL 172. 1753.

Coffee has long been grown in the mountains of Ecuador, and is

an important article of export from regions in which the climate is

218 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

suitable. The coffee of Ecuador is of superior quality, ranking with

that of Colombia, Venezuela, and Central America.

28. IXORA L.

Ixora sparsifolia Krause, Bot. Jahrb. Engler 40: 328. 1908.

In forests and thickets, Entable, near Naranjal, Lehmann 5589

(W, type collection; photo, of type, ex Herb. Berol., in F).

A shrub 3 m. high; leaves nearly sessile, oblong or elliptic-oblong,

14-21 cm. long, 4-8 cm. wide, short-acuminate, acutish at the base,

puberulent beneath along the costa; panicles lax, trichotomous,

pedunculate, the flowers 4-parted, slender-pedicellate; calyx lobes

broadly ovate, obtuse, puberulent; corolla pinkish white, puberulent,

the tube 7 mm. long, the lobes shorter, spreading.

29. MORINDA L.

Morinda Roioc L. Sp. PI. 176. 1753.

Balao, in forest, Eggers 1421 7 (L). Widely distributed in tropical

America.

A shrub about 2.5 m. high, often subscandent; leaves mostly

linear-oblong to oblanceolate-oblong, acute or acuminate, glabrous

or sparsely pubescent; flowers white, 1 cm. long, in small dense heads

on peduncles 3-10 mm. long; fruits fleshy.

30. COUSSAREA Aubl.

Coussarea pilosiflora Standl., sp. nov. — Frutex, praeter corollas

omnino glaber, ramulis gracilibus, obtuse tetragonis, internodiis

elongatis; stipulae 3 mm. longae, rotundatae, erectae, persistentes;

folia opposita, petiolis gracilibus, 1.3-1.6 cm. longis; limbus oblanceo-

lato-oblongus vel obovato-oblongus, 11.5-15 cm. longus, 3.5-5.5 cm.

latus, abrupte acuminatus, acumine anguste triangulari, obtusiusculo,

basi acutus, membranaceus, concolor, costa et nervis lateralibus

utrinque prominentibus, gracilibus, nervis lateralibus utroque latere

c. 10, angulo lato divergentibus, fere rectis, prope marginem irregular-

iter arcuato-conjunctis, nervulis prominulis,reticulatis;inflorescentiae

cymoso-paniculatae, terminales, pauciflorae, c. 8 mm. longe pedun-

culatae, 2-3 cm. longae, cymulis breviter pedunculatis, 3-6-floris,

floribus sessilibus vel brevissime pedicellatis; hypanthium obovoid-

eum, 1.5 mm. longum; calyx 1.2 mm. longus, hypanthio latior, sub-

truncatus et remote 4-denticulatus; corolla alba, extus pilis sparsis

patentibus pilosa, tubo gracili, 11 mm. longo, superne vix dilatato,

fauce 1.5 mm. lato, lobis 4, anguste oblongis, obtusis, 6 mm. longis.

—Ecuador: Valley of Pastaza River, between Banos and Cashurco,

8 hours east of Banos, Prov. Tungurahua, alt. 1,300-1,800 m.,

September 25, 1923, A. S. Hitchcock 2184-8 (U. S. Nat. Herb. No.

1,196,554, type; duplicate in herb. N. Y. Bot. Gard.).

The species is noteworthy because of the pilose corollas.

THE RUBIACEAE OF ECUADOR 219

31. FARAMEA Aubl.

Faramea coerulescens Schum. & Krause, Bot. Jahrb. Engler

40: 347. 1908.

Between Pindilie and Shoray, Prov. Cuenca, 2,800-3.000 m.,

Lehmann 4.935 (F, photo, of type ex Herb. Berol.). Loja, Ruiz &

Pav6nC!;K). Also in Peru.

A shrub 3-5 m. high; stipules triangular, acuminate; petioles

3-5 mm. long; leaf blades oblong or obovate-oblong, 3.5-8 cm. long,

shortly obtuse-acuminate, cuneate at the base, rigid-coriaceous;

corymbs few-flowered; calyx 4-dentate; corolla white, the tube 18-20

mm. long, the lobes one-third as long.

Faramea cuencana, sp. nov. — Ramuli graciles teretes virides

glabri, internodiis elongatis; stipulae cito deciduae, non visae; folia

brevissime petiolata, opposita, petiolo 2-3 mm. longo glabro supra

sulcato; lamina firme membranacea, oblonga vel anguste oblonga,

6-9.5 cm. longa, 1.5^3 cm. lata, abrupte caudato-acuminata, acumine

angusto interdum lineari 1-1.5 cm. longo attenuate, basi acuta yel

obtusa et abrupte contracta, utrinque glabra, supra atro-viridis,

lucida, costa nervisque prominentibus, subtus paullo pallidior, costa

gracili eleyata, nervis lateralibus utrpque latere c. 14, gracillimis,

prominentibus, angulo fere recto divaricatis, remote a margine

arcuato-conjunctis, nervulis prominulis laxe reticulatis; inflorescentia

terminalis sessilis cymoso-paniculata laxe multiflora, basi trichotoma,

ramis adscendentibus gracilibus glabris, bracteis paucis subulatis

2-3 mm. longis, floribus pedicellatis, pedicellis gracilibus rectis ple-

rumque 3-8 mm. longis; hypanthium obovoideum glabrum 1-15.

mm. longum; calyx 2 mm. longus profunde 4-lobus, laciniis anguste

lanceolatis longe angusteque attenuatis; corolla 8-10 mm. longa extus

glabra, tubo gracili superne paullo dilatato, lobis 4 anguste oblongis

acuminatis tubo paullo brevioribus. — Ecuador: Eastern Andes,

Cuenca, alt. 1,500-1,800 m., Pearce (Herb. Kew., type; photo, in

herb. Field Mus.).

Faramea guayaquilensis DC. Prodr. 4: 497. 1830. Tetramer-

ium multiflorum Bartl. ex DC., loc. cit., as syn.

Type from Guayaquil, Haenke.

Leaves oblong, subsessile, acuminate; stipules oblong, acuminate,

subaristate, persistent; panicles terminal, trichotomous.

Perhaps referable to some other genus, for it is stated that the

upper stipules are bifid.

Faramea maynensis Spruce ex B. & H. Gen. PI. 2: 121. 1873,

nomen nudum; Rusby, Mem. N. Y. Bot. Gard. 4: 300. 1907.

Teresita, Prov. Guayas, 270 m., Hitchcock 20449 (W, Y). Also

in Peru and Bolivia.

A glabrous shrub 1-2.5 m. high; stipules aristate-acuminate;

leaves narrowly oblong, caudate-acuminate, thin; flowers cymose-

220 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

corymbose; calyx 4-denticulate; corolla blue, glabrous, 12-17 mm.

long, the lobes equaling or shorter than the tube; fruit about 14 mm.

broad.

32. GEOPHILA Don

Geophila herbacea (Jacq.) Schum. in E. & P. Nat. Pfl. 44: 119.

1891. Psychotria herbacea Jacq. Enum. PI. Carib. 16. 1760.

El Recreo, March 10, 1897, Eggers (F). Without locality, Eggers

14904 (W). Widely distributed in the lowlands of tropical America,

often growing as a weed.

A creeping perennial herb; leaves long-petiolate, rounded to

ovate, cordate; flowers small, white, in terminal long-pedunculate

few-flowered heads; ovary glabrous; fruits fleshy, red or black,

5 mm. long.

33. CEPHAELIS Sw.

Cephaelis jacobinioides, sp. nov. — Frutex 2.5 m. altus dicho-

tome ramosus, ramulis gracilibus subteretibus viridibus, novellis

sparse puberulis, internodiis 1.5-5 cm. longis; stipulae persistentes

erectae, basi in vaginam breyem connatae, glabrae, profunde bifidae,

laciniis lanceolatis attenuatis; folia opposita, petiolo gracili 1-2.3

cm. longo, sparse et minute strigilloso vel fere glabro; lamina crasse

membranacea, elliptico-oblonga vel elliptica, 10-17 cm. longa, 4-7.5

cm. lata, abrupte acuminata, acumine anguste triangulari superne

attenuate apice obtuso, basi acuta vel subobtusa, interdum abrupte

contracta et cuneatim decurrens, supra laete viridis, glabra, costa,

prominente, nervis conspicuis sed non elevatis, subtus paullo palli-

dior, ubique minute strigillosa, costa gracili elevata, nervis lateralibus

utroque latere c. 27, gracilibus, prominentibus, angulo lato adscen-

dentibus, subarcuatis, prope marginem nervum distinctum regularem

collectivum efformantibus, nervis subregularibus fere rectis costis

secundariis subparallelis inter nervos interpositis, nervulis prominulis

laxe reticulatis; inflorescentia terminalis 1.8 cm. longe pedunculata

vel fere sessilis, subcapitata, densissime multiflora, c. 3 cm. longa et

fere aequilata, basi foliis 2 magnis bracteata, e capitulis paucis

secundariis dense congestis subsessilibus efformata, bracteis numero-

sis arete adpresso-imbricatis, exterioribus majoribus basi breviter

connatis vel fere distinctis, ovatis vel lanceolato-oblpngis, 1.5-1.8

cm. longis, acutis vel acuminatis, margine hispidulis, sanguineo-

purpureis, saepe profunde bifidis, bracteis interioribus angustioribus,

lanceolatis vel lanceolato-linearibus, brevioribus; flores inter bracteas

sessiles; hypanthium glabrum vix 1 mm. longum; calyx 5-partitus,

laciniis linearibus viridibus 4-5 mm. longis ciliatis; corolla alba,

tubo 6-7 mm. longo superne paullo dilatato extus glabro, lobis 5

anguste oblongis patentibus acutiusculis.^ — Ecuador: At the base of

Mt. Chimborazo, alt. 900 m., June, 1860, R. Spruce 6188 (Herb.

Kew., type). In forest, 400-900 m., Sodiro 8436 (B).

THE RUBIACEAE OF ECUADOR 221

A species well marked by the many-veined leaves and by the

distinctive form of the inflorescence with its very numerous narrow

bracts irregularly imbricate in numerous series.

Cephaelis Jamesonii, sp. noy. — Frutex subramosus 3-metralis,

ramulis gracilibus primo dense pilis rigidiusculis saepe subretrorsis

villosis, serius glabratis et tuberculatis, internodiis 1.5-5 cm. longis;

stipulae 3-4 mm. longae, breviter connatae, basi yillpsae, ad medium

bilqbae, lobis triangularibus subacutis glabratis ciliatis; folia opposita,

petiolo crasso 4-7 mm. longo villoso; lamina late ovata vel ovato-

elliptica, 4-5.5 cm. longa, 2.3-3.2 cm. lata, acuta vel subobtusa, basi

rotundata, coriacea, supra flavo-viridis, ad nervos sparse villosula,

saepe scaberula, costa nervisque prominulis, nervulis subimpressis,

subtus fere concolor, ad costam validam elevatam villosa, aliter

glabra, nervis lateralibus utroque latere c. 8, prominentibus, gracili-

bus, saepius angulo acuto adscendentibus, rectis vel leviter curvis,

prope marginem conjunctis, nervulis obscuris; inflorescentia axillaris

pauciflora capitata, 2.5-3 cm. longe pedunculata, pedunculo dense

villoso, floribus sessilibus; bracteae involucrantes, late ovatae, c. 1.5

cm. longae, acutae, erectae, parte inferiore villosae, superne glabrae;

corolla infundibuliformis, tubo 2.5 cm. longo, inferne glabro, superne

sparse villoso, fauce 6 mm. lato, lobis ovatis obtusis 6 mm. longis,

extus pilis longis mollibus villosis, intus fere glabris; fructus oblongus

7 mm. longus glabratus; semina facie ventrali profunde angusteque

sulcata. — Ecuador: In declivitate occidentali mentis Pichincha, alt.

2,400 m., in 1864, W. Jameson (U. S. Nat. Herb. No. 534,965, type).

Valle de Pallatenga, 2,700 m., Spruce 5521 (K). Dense forest, Alto

de Panga, near Pallatenga, 2,000-2,400 m., September, lSQ4,Lehmann

K. 143 (K).

A well-marked species, in general appearance somewhat suggesting

the genus Coccocypselum. It is remarkable for the very large corollas

and copious pubescence.

Cephaelis peruviana Wernham, Journ. Bot. 51: 221. 1913.

Type from Guayaquil, Ruiz & Pavdn.

Branches ferruginous-pilose; stipules broadly ovate, 2-aristate;

leaves short-petiolate, broadly elliptic, 6 cm. long and 3.5 cm. wide,

obtuse, rounded at the base, pilose along the nerves; heads oblong,

2.5 cm. long, pedunculate, the 4 bracts nearly glabrous, ovate;

flowers usually 3; corolla glabrous outside, the tube 1.5 cm. long.

The specific name is a misleading one, for the plant is not known

at present from Peru.

Cephaelis Remyana (BailL), comb. nov. Uragoga Remyana

Baill. Adansonia 12: 253. 1879.

Type collected in forest between Guaranda and Bodegas, Remy.

Glabrous; stipules connate into a deciduous sheath; leaves long-

petiolate, ovate-lanceolate, 15 cm. long and 8 cm. wide, acuminate,

222 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

paler beneath; flower heads terminal, in clusters of 2-3, the bracts

ovate, acute, imbricate; corolla tube slender, the 5 lobes ovate, acute,

recurved.

Cephaelis tomentosa (Aubl.) Vahl, Eclog. Amer. 1: 19. 1796.

Tapogomea tomentosa Aubl. PI. Guian. 1 : 160. pi. 61 . 1775. Uragoga

tomentosa Kuntze, Rev. Gen. PI. 1: 301. 1891.

Between Santa Rosa and La Chorrita, Prov. Oro, 100 m., Hitch-

cock 21119 (W, Y). Ranging northward to Mexico and south to

Bolivia.

A hirsute shrub 1-2.5 m. high; leaves short-petiolate, lanceolate

to ovate-elliptic, long-acuminate; flower heads terminal, pedunculate,

large, dense, the large bracts red, longer than the flowers; corollas

yellow; fruits blue.

34. PSYCHOTRIA L.

Psychotria acutiflora DC. Prodr. 4: 506. 1830. Guettarda

acutiflora Bartl. ex DC., loc. cit., as syn.

Type from Guayaquil, Haenke.

Stipules connate, the free portion lanceolate; leaves oblong,

acuminate, ciliate, pubescent beneath, scaberulous above, hirsute-

tomentose on the costa; inflorescence terminal, paniculate, hirsutu-

lous, longer than the leaves.

Psychotria alba R. & P. Fl. Peruv. 2: 58. pi 205, f. a. 1799.

Balao, Eggers 14266 (B, L, W). Rio Tenguel, Prov. Guayas,

Holmgren 70 (F, S). Widely distributed in South America.

A nearly glabrous shrub 2 m. high; stipules large, brown, obtuse,

caducous; leaves petiolate, elliptic-oblong to obovate, acute or

acuminate, minutely puberulent beneath on the nerves or nearly

glabrous; inflorescence terminal, pedunculate, paniculate, many-

flowered, trichotomous or radiately branched at the base, the bracts

minute, deciduous; flowers short-pedicellate; calyx minutely dentic-

ulate; corolla 4 mm. long, white, minutely puberulent or glabrous;

fruit 4.5 mm. long.

Psychotria angustata Anders. Vet. Akad. Handl. Stockh. 1853:

193. 1855.

Type from Charles Island, Galapagos Islands, Andersson.

A glabrous shrub; stipules oval, obtuse, deciduous; leaves petio-

late, broadly lanceolate, acute, narrowed to the petiole, 7.5-8.5 cm.

long, 3.5 cm. wide; inflorescence axillary, pedunculate, thyrsiform,

shorter than the leaves; calyx obscurely 5-dentate; corolla pink,

barbate in the throat, the 5 lobes ovate-oblong, obtuse; fruit 4-6

mm. long, subglobose, 10-costate.

Psychotria brachiata Sw. Prodr. Veg. Ind. Occ. 45. 1788.

Forests of Zamora, Loja, 1,000 m., Lehmann 5653 (Y). Widely

distributed in tropical America.

THE RUBIACEAE OF ECUADOR 223

A shrub with glabrous branches; stipules short, green, persistent,

bilobate, the lobes rounded; leaves petiolate, oblong-obovate to

elliptic-oblong, acuminate, at the base acute to attenuate, glabrous

or nearly so; inflorescence terminal, pedunculate, paniculate, the

branches divaricate, opposite, subtended by long narrow involute

bracts, the flowers in dense bracted clusters; corolla 1 cm. long, pubes-

cent; fruit blue, glabrous.

Psychotria carthaginensis Jacq. Enum. PI. Carib. 16. 1760.

P.foveolata R. & P. Fl. Peruv. 2: 59. pi. 207, f. b. 1799.

Bafios, Prov. Tungurahua, 1,800 m., Heilborn 384 (F, S). Widely

distributed in South America.

A nearly glabrous shrub; stipules obovate, obtuse, brown, cadu-

cous; leaves petiolate, lance-elliptic to oblong-obovate, acute or

acuminate, acute to attenuate at the base, minutely puberulent

beneath or glabrate; inflorescence terminal, pedunculate, paniculate,

usually radiately branched at the base, open, many-flowered, the

bracts minute, deciduous; flowers sessile; calyx denticulate; corolla

white, 4 mm. long, minutely puberulent or glabrous; fruit red, 4 mm.

long, costate, glabrous.

Psychotria chimboracensis, sp. nov. — Arbor 6-metralis patule

ramosus, ramulis crassiusculis subteretibus glabris, internodiis brevi-

bus vel elongatis; stipulae caducae late ovatae 13-14 mm. longae,

obtusae vel subacutae, dense ferrugineo-tomentulosae; folia petiolata

opppsita, petiolo valido 6-12 mm. longo subtus ferrugineo-villosulo;

lamina chartacea, oblongo-elliptica, oblonga vel obovato-oblonga,

plerumque 15-26 cm. longa et 6-12.5 cm. lata, acuta vel breviter

acuminata, basi acuta vel basin versus longius angustata, supra fusca,

glabra, nervis non elevatis, subtus pallidior, brunnescens, ubique vel

tantum ad nervos ferrugineo-villosula, serius glabrata, costa valida

elevata, nervis lateralibus utroque latere c. 17, gracilibus, angulo lato

adscendentibus, fere rectis, prominentibus, prope marginem con-

junctis, nervulis obscuris; inflorescentia terminalis 4 cm. longe

pedunculata late cymoso-paniculata dense multiflora, c. 9 cm. longa

et 14 cm. lata, pedunculo crasso compresso, ramis oppositis

divaricatis glabris vel sparse ferrugineo-villosulis, floribus ad apices

ramulorum dense capitato-glomeratis sessilibus, bracteis plerumque

deciduis 1-2 mm. longis interdum persistentibus et reflexis ferrugineo-

puberulis; hypanthium late turbinatum 1.5-2 mm. longum glabrum;

calyx 2-3 mm. longus late campanulatus glaber breviter 5-lobus,

lobis late triangularibus obtusis pilis minutis ferrugineis ciliatis;

corolla eburnea, tubo 3-3.5 mm. longo obconico extus glabro, in

fauce pilis lilacinis dense barbato, lobis 5 oblongo-triangularibus

tubum fere aequantibus, apice acutiusculis et cucullatis; antherae

exsertae 1.5 mm. longae oblongae. — Ecuador: At the base of Mt.

Chimborazo, alt. 750 m., July, 1860, R. Spruce 6192 (Herb. Kew.,

type; photo, in herb. Field Mus.).

224 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Psychotria cuneifolia DC. Prodr. 4: 507. 1830. P. glabrata

Bartl. ex DC., loc. cit., as syn.

Type from Guayaquil, Haenke.

Glabrous; leaves obovate, long-acuminate, short-cuspidate; stip-

ules small, deciduous; panicles erect, shorter than the leaves, the

branches opposite; fruit ovoid-globose, 10-striate.

The description is too brief to enable one to identify the species.

Psychotria Haenkeana DC. Prodr. 4: 507. 1830. P. peduncu-

lata Bartl. ex DC., loc. cit., as syn.

Type from Guayaquil, Haenke.

Glabrous; stipules bifid, acuminate; leaves oval-oblong, acuminate

at each end; peduncles slightly shorter than the leaves, the cymes

contracted, obconic; flowers and fruit unknown.

The description is too brief for determination, and it appears to

be somewhat misleading as regards the stipules. There is at hand a

photograph of a specimen in the Berlin herbarium which probably

belongs to the type collection of this species. Dr. Urban has indi-

cated on the sheet that the plant is Psychotria marginata Sw., but

it seems to me to belong more probably with P. alba R. & P.

Psychotria hebeclada DC. Prodr. 4: 513. 1830.

Rio de Ventanas, near Guayaquil, Spruce 6350 (K, L). Ranging

northward to Mexico.

A shrub 2.5 m. high, the young branches densely puberulent;

stipules short, green, persistent, bidentate; leaves thin, petiolate,

ovate to oblong-elliptic, long-acuminate, acute or attenuate at the

base, finely pubescent, at least beneath; inflorescence terminal, pedun-

culate, thyrsoid-paniculate, dense, many-flowered, densely pubescent;

calyx lobes short, ovate, acute or obtuse; corolla whitish, 5 mm. long,

pubescent; fruit glabrate, subglobose, 4 mm. long.

Psychotria horizontalis Sw. Prodr. Veg. Ind. Occ. 44. 1788.

El Recreo, Eggers 15410 (F, K, L, W), 15205 (F, K). Ranging

northward to Mexico and the West Indies.

A nearly glabrous shrub; stipules ovate-triangular, 3-8 mm. long,

entire, caducous; leaves short-petiolate, elliptic to lance-oblong,

acuminate, at the base usually acute, glabrous or nearly so; inflores-

cence terminal, pedunculate, cymose-paniculate, small, the branches

puberulent, the flowers sessile, the bracts minute, caducous; calyx

deeply lobate, the lobes linear; corolla 3-4 mm. long, white, glabrous;

fruit red, glabrous, 4 mm. long.

Psychotria justiciaefolia, sp. nov. — Frutex, ramulis gracilibus

obtuse tetragonis dense pilis pallidis adpressis vel subadpressis pilosis,

internodiis elongatis; stipulae 10-14 mm. longae persistentes breviter

connatae dense pilosae, ad medium vel profundius bilobae, lobis

THE RUBIACEAE OF ECUADOR 225

anguste lanceolatis longiattenuatis; folia opposita, petiolo 1-3 cm.

longo gracili dense pilis longis piloso; lamina oblongo-elliptica vel

rarius obovato-elliptica, 9-20 cm. longa, 3-9 cm. lata, longe abrup-

teque acuminata, acumine angusto obtuso vel acutp, papyracea,

supra viridis, sparse setoso-pilosa vel glabrata, nervis prominulis,

subtus pallidior, subdense pilis longis debilibus subadpressis pilosa,

costa gracili elevata, nervis lateralibus utroque latere c. 16, gracilli-

mis, prominentibus, angulo acuto vel lato adscendentibus, arcuatis,

prope marginem conjunctis, nervulis prominulis laxe reticulatis;

inflorescentia terminalis 1.5-5 cm. longa, 1-2 cm. lata, 1-4 cm. longe

pedunculata, thyrsoideo-paniculata, densissime multiflora, interdum

spiciformis, pedunculo crasso dense piloso, ramis primariis brevissimis

vel obsoletis, floribus sessilibus ad apices ramulorum capitato-con-

gestis; bracteae oblongae, ovales vel obovatae, apice acutae vel

rotundatae, 3-4 mm. longae, sparse adpresso-pilosulae, dense et

conspicue ciliatae; calyx extus densissime breviterque hirsutus;

fructus immaturus hispidulus. — Ecuador: Chacayacu, western Andes

of Cuenca, alt. 1,000-1,800 m.,F. C.Lehmann 5602 (Herb. Field Mus.

No. 578,071, type; duplicate in K). Ad radices Chimborazo, alt.

900 m., August, 1860, Spruce 6189 (K) ; a shrub 3 m. high.— Colombia:

Las Dantas to Puerto Canoa, on Rio Esmeralda, Bolivar, 400-800

m., Pennell 4518 (W, Y). Damp forest ravine, by a stream, Val-

paraiso, 1,350 m., H. H. Smith 2796 (Y).

The plant is a close relative of P. daguensis Standl., of Colombia,

but it appears substantially different in the broad thin leaves and

large stipules.

Psychotria luxurians Rusby, Mem. Torrey Club 6: 50. 1896.

Palicourea membranifolia Schum. & Krause, Bot. Jahrb. Engler 40:

333. 1908.

Western Andes of Naranjal, 500 m., Lehmann 6678 (K). Type

of P. membranifolia collected above Naranjal, on the western slope

of the western Andes, in dense wet forest at 200-600 m., Lehmann

6671 (photo, and fragm. of type, ex Herb. Berol., in F). Ranging

southward to Bolivia.

A shrub or small tree 5 m. high; stipules triangular, 4-5 mm. long,

bidentate at the apex; leaves short-petiolate, thinly membranous,

oblong or ovate-oblong, acuminate, at the base cuneate, 9-16 cm.

long and 3.5-6 cm. wide, glabrous, the lateral nerves about 12 pairs;

panicles long-pedunculate, many-flowered, the bracts filiform, 6-8

mm. long; flowers short-pedicellate; calyx lobes broadly triangular,

acute; corolla yellow, glabrous, 6 mm. long.

Psychotria macrophylla R. & P. Fl. Peruv. 2: 56. pi. 202, /. a.

1799. P. anomothyrsa Schum. & Donn. Smith, Bot. Gaz. 35: 3. 1903.

Near Niebli, Sodiro 48-32 (photo, and fragm. ex Herb. Berol., in F) .

El Recreo, Eggers 15119 (F, W). Teresita, Prov. Guayas, 270 m.,

Hitchcock 20416 (W, Y). Near Tamompeque, Eggers 14861 (F).

226 FIELD MUSEUM OF NATURAL HISTORY— BOTANY, VOL. VII

Tandapi, Prov. Pichincha, 1,500 m., Holmgren 822 (S). Ranging

southward to Bolivia and northward to Mexico.

Stems simple, glabrous, suffrutescent, sometimes 2.5 m. high;

stipules short, broad, apiculate, caducous; leaves large, petiolate,

elliptic to lance-oblong, acuminate, at the base acute to attenuate,

glabrous or nearly so; inflorescence glabrous or pruinose-puberulent,

axillary, pedunculate, paniculate, the branches often reflexed, the

flowers sessile; calyx denticulate; corolla whitish, glabrous, 4-5 mm.

long; fruit white, glabrous, 4-5 mm. long.

Psychotria madida, sp. nov. — Ramuli crassi sparse minuteque

puberuli vel glabrati; stipulae deciduae, non visae; folia opposita,

petiolo gracili 2-3 mm. longo puberulo vel glabrato; lamina obovata

vel oblongo-elliptica, 20-21 cm. longa, 8-10 cm. lata, acuta vel

obtusa, basin versus longe angustata et decurrens, rarius subobtusa,

herbacea, marginata, supra viridis, asperula, nervis manifestis sed

vix elevatis, subtus fusco-brunnescens, asperula, ad nervos minute

puberula, costa crassa, nervis lateralibus utroque latere c. 15, pro-

minentibus, gracilibus, angulo lato adscendentibus, arcuatis, mar-

ginem attingentibus, nervulis fere obsoletis; inflorescentia axillaris

5-6 cm. longe pedunculata late cymoso-paniculata, c. 4 cm. longa et

5 cm. lata, dense multiflora, pedunculo puberulo, ramis oppositis

divaricatis validis dense puberulis, floribus sessilibus dense congestis;

bracteae oblongae vel ovatae 3-5 mm. longae acutae vel obtusae

puberulae; calyx 2.5-3 mm. longus profunde 5-lobus, lobis oblongis

vel obovatis obtusis vel apice rotundatis puberulis vel glabratis;

corolla tubulosa 8 mm. longa sparse puberula vel fere glabra, tubo

2.5 mm. lato, lobis brevissimis; fructus subglobosus compressus

glaber 5-6 mm. longus profunde sulcatus, costis acutis. — Ecuador:

Vicinity of Huigra, September 3, 1918, J. N. andG. Rose 22507 (U. S.

Nat. Herb. No. 1,022,160, type; photo, in F; duplicates in G and Y).

Tandacato, Prov. Pichincha, 2,500 m., Heilborn 496 (S).

Near P. Hartwegiana Standl., but differing in the large calyx,

longer corolla, and well-developed bracts.

Psychotria magnoliaefolia HBK. Nov. Gen. & Sp. 3: 360.

1820, non H. & B. ex R. & S., 1819.

Based on a specimen ascribed doubtfully to Ecuador.

A nearly glabrous shrub; stipules oblong, rounded at the apex,

as long as the petioles, caducous; leaves petiolate, elliptic, acute or

short-acuminate, at the base acute, 15-20 cm. long, 10-12.5 cm. wide,

glabrous; inflorescence terminal, pedunculate, cymose-paniculate,

radiately branched at the base, the bracts minute; flowers sessile;

calyx obsoletely 5-dentate; corolla glabrous, the tube short, the throat

villous.

The authors of this species state that it is close to P. ardisiaefolia

HBK., a synonym of P. alba R. & P. There is at hand a photograph

of an authentic specimen from the Willdenow Herbarium at Berlin.

THE RUBIACEAE OF ECUADOR 227

Although the photograph is a good one, the status of the plant is

doubtful, and it is probably not P. alba. I have not renamed the

species because it is altogether uncertain whether it is a valid one or

not. The doubt regarding the locality from which the material came,

as in the case of other species published in the same work, does not

simplify the matter of identification.

Psychotria microdon (DC.) Urban, Repert. Sp. Nov. 9: 539.

1928. Rondeletia microdon DC. Prodr. 4: 408. 1830. P. pinularis

Sess<§ & Moc. Fl. Mex. ed. 2. 57. 1894.

El Recreo, Eggers 15723 (F, K, L, W). Balao, Eggers 14617

(L, W). Chonana, near Guayaquil, Spruce 6270 (K). Rio Daule,

near Guayaquil, Spruce 6270 (B, L) . Ranging to the Guianas, Colom-

bia, Mexico, and Cuba.

Psychotria racemosa (Aubl.) Willd. Sp. PL 1: 966. 1797.

Nonatelia racemosa Aubl. PL Guian. 1 : 187. pi. 72. 1775.

Teresita, Prov. Guayas, 270 m., Hitchcock 20514 (W, Y). Widely

distributed in South America, and extending to Panama.

A shrub 1.5 m. high, the branches puberulent; stipules biparted,

the segments linear-subulate, green, persistent; leaves short-petiolate,

oblong-elliptic, acuminate, at the base usually abruptly contracted,

nearly glabrous; inflorescence terminal, thyrsiform, short-peduncu-

late, the short branches minutely hirtellous; flowers sessile; calyx

lobes linear-lanceolate; corolla hirtellous; fruit 4-5-celled, sulcate,

4 mm. long.

This species is easily recognized by the 4-5-celled fruit, that of

most species of Psychotria being 2-celled.

Psychotria rufescens H. & B. ex R. & S. Syst. Veg. 5: 192. 1819.

P. micrantha HBK. Nov. Gen. & Sp. 3: 363. pi. 284. 1820.

El Recreo, Eggers 15657 (F, K, L, W). Without locality, Eggers

15176 (F). Ranging northward to Central America; Peru.

A shrub or small tree, the branches rufous-hirsute; stipules large,

brown, bidentate, caducous; leaves petiolate, oblong to oblong-

elliptic, acuminate, acute or attenuate at the base, softly pubes-

cent; inflorescence terminal, pedunculate, open-paniculate, radiately

branched; calyx lobes deltoid, obtuse; corolla white or whitish, 3 mm.

long, pubescent; fruit 4 mm. long, pubescent.

Psychotria rufipes Hook. f. Trans. Linn. Soc. 20: 220. 1847.

Type from James Island, Galapagos Islands; reported also from

Albemarle, Charles, Chatham, and Indefatigable Islands.

Stipules large, broadly ovate, entire; leaves petiolate, obovate-

lanceolate, obtuse, attenuate at the base, glabrous above, rufous-

tomentose beneath; inflorescence terminal and axillary, short, panic-

ulate, rufous- tomentose, the flowers short-pedicellate; corolla villous,

the throat barbate.

228 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Psychotria rugulosa HBK. Nov. Gen. & Sp. 3: 356. 1820.

Loja, Rose, Pachano & Rose 23245 (W, Y). Also in Colombia

and in Peru (?).

A nearly glabrous shrub; stipules oblong, thin, brown, acuminate*

caducous; leaves petiolate, lanceolate or oblong-lanceolate, long-

acuminate, attenuate to the base, finely puberulent beneath along

the nerves, the nerves impressed above; inflorescence terminal, pedun-

culate, cymose-paniculate, radiately branched at the base, the

branches glabrous or minutely puberulent; bracts deciduous; flowers

sessile; calyx denticulate; corolla glabrous outside, 5 mm. long, the

lobes acutish, equaling the tube; fruit globose, glabrous, red, 4 mm.

long.

Psychotria viridis R. & P. Fl. Peruv. 2: 61. pi 210, /. 6. 1799.

P. glomerata HBK. Nov. Gen. & Sp. 3: 362. 1820.

Balao, in forest, Eggers 14476 (L, W). Ranging south to Bolivia

and north to Central America and Cuba.

A glabrous tree 3.5 m. high; stipules large, acuminate, thin,

brown, caducous; leaves short-petiolate, obovate or obovate-oblong,

acute or short-acuminate, cuneate-attenuate to the base; inflores-

cence terminal, pedunculate, spicate-paniculate, the lower branches

verticillate, the minute flowers sessile in distant glomerules; corolla

greenish white; fruit red.

35. PALICOUREA Aubl.

Palicourea angustifolia HBK. Nov. Gen. & Sp. 3: 367. 1820.

Tungurahua, 1,950 m., Spruce 5118 (K). Eastern Andes, 1,800-

2,400 m., collector unknown, No. 347 (K). Also in Colombia, Vene-

zuela, and Peru.

A shrub or small tree, the branchlets densely puberulent or

rarely glabrous; stipule sheath 4-6 mm. long, the lobes subulate,

erect, equaling or longer than the sheath; leaves short-petiolate,

oblong or oblong-lanceolate, 6-15 cm. long, long-acuminate, at the

base obtuse or acute, usually puberulent or pilose beneath, at least

on the veins; panicles pedunculate, thyrsiform, many-flowered, open

or dense, the branches densely puberulent or pilose, the bracts

minute; flowers pedicellate; calyx minute, acutely dentate; corolla

tubular, 10-15 mm. long, violet or purple, commonly puberulent or

short-pilose; fruit 4-5 mm. long, purple-black.

Palicourea aragmatophylla Schum. & Krause, Bot. Jahrb.

Engler 40: 332. 1908.

Type from the western slope of the western Andes, near Cuenca,

in dense forest, at 2,000-2,500 m., Lehmann 6676 (photo, and fragm.

of type, ex Herb. Berol., in F).

A shrub or small tree 5 m. high, the branches puberulent below

the nodes; stipules large, the sheath 8-^10 mm. long, the lobes subu-

late, equaling the sheath; leaves opposite, petiolate, rigid-coriaceous,

THE RUBIACEAE OF ECUADOR 229

oblong or oblong-lanceolate, acuminate, at the base cuneate, 10-17

cm. long, 3-6 cm. wide, glabrous; panicles pedunculate, subpyramidal,

lax, 10-12 cm. long, the branches puberulent, the bracts subulate;

flowers short-pedicellate; calyx lobes oblong, obtuse; corolla blue,

12-14 mm. long, minutely papillose.

Palicourea balnearia, sp. nov. — Frutex, ramulis crassis olivaceis

subteretibus glabris, internodiis 1.5-3 cm. longis; stipulae persistentes

adpressae in vaginam truncatam glabram 2-4 mm. longam connatae,

vagina in lobos 4 remotps triangulares acutos 1-2 mm. longos glabros

desinente; folia opposita, petiolo valido 6-9 mm. longo glabro;

lamina subcoriacea, oblongo-elliptica, 6.5-10 cm. longa, 2.5-3.5 cm.

lata, abrupte breviterque acuminata, acumine triangulari obtuso, basi

acuta vel subobtusa, saepe bullata, utrinque glabra, angustissime

marginata, supra viridis, nervis manifestis sed non eleyatis, subtus

luteo-viridis, costa crassiuscula elevata, nervis lateralibus utrpque

latere c. 11, angulo lato adscendentibus, gracilibus, prominentibus,

arcuatis, prope marginem conjunctis; inflorescentia terminalis 3.5

cm. longe pedunculata, cymoso-paniculata, c. 5 cm. longa et lata,

dense multiflora, multiramosa, ramis basalibus pseudoverticillatis

crassis glabris, patentibus vel subadscendentibus, pedicellis crassis

glabris 1-3 mm. longis, bracteis triangularibus acutis 2-3 mm. longis

glabris; hypanthium cylindraceo-turbinatum 1.5 mm. longum gla-

brum; calyx glaber 1 mm. longus breviter 5-lobus, lobis rotundato-

ovatis inaequalibus obtusis minute ciliolatis; corolla 15-16 mm. longa

extus glabra, tubo crasso basi ampliato, superne dilatato, ore 5 mm.

lato, lobis 5 oblongo-triangularibus obtusis 3 mm. longis intus

minute papillosis; antherae oblongae semiexsertae; fructus subglo-

bosus 6 mm. diam. glaber, pyrenis dorso obtuse costatis. — Ecuador:

Baiios, Prov. Tungurahua, alt. 1,800 m., February 28, 1920, 0.

Heilborn386 (Stockholm Herb., type).

A species with no outstanding characters, but not agreeing with

any other known from Ecuador.

Palicourea calantha, sp. nov. — Frutex, ramulis gracillimis tere-

tibus viridibus glabris, internodiis 5-7 cm. longis; stipulae persistentes

in vaginam glabram truncatam 1.5-2 mm. longam connatae, vagina

in lobos 2 erectos 1-1.5 mm. longos distantes lineari-triangulares

acutiusculos desinente; folia opposita, petiolo gracili 6-23 mm. longo

glabro; lamina membranacea, anguste elliptico-oblonga vel lanceo-

lato-oblonga, 8.5-16 cm. longa, 2.5-4.5 cm. lata, longissime acuminata,

acumine angusto longe attenuato, basin versus angustata et acumi-

nata, supra glabra, laete viridis, costa vix elevata, nervis subimpressis,

subtus fere concplor, ad nervos sparse pilis brevibus albidis patenti-

bus hispidula, aliter glabra, costa gracili elevata, nervis lateralibus

utroque latere c. 16, angulo lato adscendentibus, gracillimis, valde

arcuatis, prominentibus, in marginem desinentibus, nervulis vix pro-

minulis laxe reticulatis; inflorescentia terminalis 3.5-5 cm. longe

pedunculata, late pyramidali-paniculata, 7-11 cm. longa, 12-17 cm.

230 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

lata, laxe multiflora, basi trichotoma, ramis infimis late diyaricatis

gracilibus glabris, superioribus fasciculato-ramosis et suberectis, pani-

cula interdum basi foliaceo-bracteata, pedicellis gracilibus plerumque

5-12 mm. longis glabris, bracteis subulatis glabris 2-4 mm. longis;

hypanthium turbinatum 1 mm. longum glabrum; calyx glaber 0.5-

0.8 mm. longus viridis, lobis late triangularibus obtusis; corolla

pallide lutea extus glabra in alabastro obtusa, tubo crasso 14 mm.

longo superne paullo dilatato, ore 4-4.5 mm. lato, saepe valde curyo

vel e basi recurvo, lobis ovato-oblongis 3-3.5 mm. longis acutiusculis;

stylus apice exsertus. — Ecuador: Tandapi, Prov. Pichincha, in forest

and along roadsides, alt. 1,500 m., July, 1920, 0. Heilborn 777 (Herb.

Field Mus. No. 605,122, type; duplicate in Stockholm herb.). Pulul-

ahua, Sodiro 84-20 (photo, and fragm. ex Herb. Berol. in herb. Field

Mus.).

The plant is one of the most distinct members of the genus

Palicourea, and must be a very handsome shrub when growing. It is

noteworthy for the large and unusually broad panicles with their

peculiar branching, and particularly for the form of the corolla.

The corolla tube is usually strongly curved and often it is abruptly

recurved from the base, the flowers thus appearing nutant.

Palicourea calothyrsus Schum. & Krause, Bot. Jahrb. Engler

40: 334. 1908. P. fragilior Wernham, Journ. Bot. 55: 282. 1917.

P. hedyotoides Wernham, op. cit. 339. 1917.

Eastern slope of the eastern Andes, near Sigsig, 1,800-2,000 m.,

Lehmann 6497 (F, W, type collection; photo, of type, ex Herb. Berol.,

in F). In woods, Talancay, along Rio Chanchan, Spruce 6010 (F,

G, K, L, S, type collection of P. fragilior). In valle Lloensi, inter

arbores, 2,700 m., Jameson 336 (G, type collection of P. hedyotoides).

Carmen, 2,250 m., Tate 479 (W). Niebli, Andre K503 (K, Y).

Tambo de Sabanilla, Andre 4588 (K, Y). Quitensian Andes, in 1855,

Couthouy (G). Huigra, Rose & Rose 22506 (G, W, Y). Guayan, Valley

of Lloa, near Quito, Jameson (K). Tandacato, Prov. Pichincha,

2,500 m., Heilborn 493 (S).

A slender, nearly glabrous shrub 2 m. high; stipule sheath about

2 mm. long, bidentate; leaves opposite, petiolate, firm, small, lanceo-

late or elliptic-oblong, 6-13 cm. long, 1.5-4 cm. wide, long-acuminate,

at the base acute, many-nerved, glabrous or sparsely hirtellous

beneath along the costa; panicles sessile or pedunculate, pyramidal,

lax, the branches ascending, glabrous, the bracts sometimes leaflike

but mostly setaceous and inconspicuous; flowers slender-pedicellate,

often appearing umbellate; calyx lobes oblong, obtuse; corolla yellow,

glabrous, 8-10 mm. long.

Palicourea calycina Benth. PI. Hartw. 133. 1844.

Type from mountains of Loja, Hartweg. Loja, Seemann 889 (K).

A shrub 1.5-2.5 m. high; stipule sheath 4-6 mm. long, truncate,

the lobes subulate, longer than the sheath; leaves elliptic-oblong or

THE RUBIACEAE OF ECUADOR 231

obovate-oblong, 10-15 cm. long, acuminate, at the base acute, pubes-

cent beneath on the nerves; panicles pedunculate, small, dense, many-

flowered, the bracts linear or lanceolate, often exceeding the calyx;

flowers sessile; calyx lobes linear-oblong, foliaceous, 5 mm. long,

hirtellous; corolla tubular, 12-14 mm. long, villous or hirsute.

Palicourea chimboracensis, sp. nov. — Arbuscula 3.5-metralis

vage ramosa, ramulis gracilibus teretibus viridibus glabris, inter-

nodiis c. 3.5 cm. longis; stipulae in vaginam adpressam 4-6 mm.

longam truncatam glabram connatae, vagina in lacinias 4 remotas

anguste triangulares glabras obtusas 1.5-2 mm. longas desinente;

folia opposita, petiolo gracili 12-17 mm. longo glabro; lamina mem-

branacea, elliptico-oblonga vel obovato-oblonga, 14.5-16 cm. longa,

6-7 cm. lata, apice obtusa vel rotundata et breviter acuminata,

acumine triangulari obtuso 5-7 mm. longo, basi acuta vel acuminata,

supra viridis, glabra, costa prominente, nervis non eleyatis, subtus

pallidior, ad nervos et praesertim ad costam pilis debilibus albidis

brevibus sparse pilosula, aliter glabra, costa gracili elevata, nervis

lateralibus utroque latere c. 13, gracillimis, angulp lato adscendenti-

bus, prominentibus, arcuatis, in marginem desinentibus, nervulis

vix prominulis laxe reticulatis; inflorescentia terminalis 3-4 cm. longe

pedunculata thyrsoideo-paniculata, laxe multiflora, 8-12 cm. longa,

c. 7.5 cm. lata, ramis basalibus oppositis gracilibus, angulo lato

adscendentibus, angulatis, glabris, purpureis, floribus laxe cymosis

pedicellatis, pedicellis gracilibus rectis 2-8 mm. longis glabris, bracteis

subulatis vel anguste triangularibus 2-4 mm. longis acutis glabris;

hypanthium subglobosum 2 mm. longum glabrum; calyx glaber 1 mm.

longus brevissime lobatus, lobis latissimis late rotundatis; corolla

lutea extus glabra 22-26 mm. longa, basi valde ampliato 6-7 mm.

lato, superne non dilatato, ore 4 mm. lato, lobis 5 triangulari-oblongis

3 mm. longis obtusis intus minute papillosis; stylus breviter exsertus;

fructus globosus niger 6-7 mm. longus glaber. — Ecuador: At the foot

of Mt. Chimborazo, alt. 900 m., July, 1860, R. Spruce 6182 (Herb.

Kew., type; photo, in herb. Field Mus.).

Palicourea cyclotis, sp. nov. — Frutex 3-metralis, ramulis graci-

libus obtuse tetragonis glabris; stipulae 7 mm. longae fere liberae

profunde bilobae, lobis late rotundatis extus scaberulis ciliolatis; folia

opposita, petiolo gracili 1.5-4 cm. longo glabrato; lamina papyracea,

ovali-elliptica, 15^-18 cm. longa, 9-11 cm. lata, abrupte breviterque

acuminata, acumine lato obtuso, basi obtusa, supra viridis, glabra,

nervis non elevatis, subtus pallida, secus costam hinc inde hirtella vel

glabra, costa gracili elevata, nervis lateralibus utroque latere c. 15,

gracillimis, prominentibus, angulo lato adscendentibus, arcuatis,

prope marginem conjunctis, nervulis obscuris; inflorescentia 5 cm.

longe pedunculata anguste pyramidali-paniculata, 27 cm. longa, 20

cm. lata, laxe multiflora, ramulis gracilibus divaricatis vel adscen-

dentibus angulatis sparsissime minuteque puberulis ; bracteae oblongae

vel anguste triangulares, obtusae vel acutae, 2-4 mm. longae, glabrae,

ciliolatae; flores sparsi graciliter pedicellati, pedicellis 5-11 mm. longis;

232 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

hypanthium turbinatum 1.5 mm. longum glabrum; calyx glaber

2 mm. longus, 3-4.5 mm. latus, profunde 5-lobus, lobis ovalibus vel

rotundatis; corolla intense purpurea extus glabra tubuloso-infundi-

buliformis, tubo basi paullo ampliato superne dilatato, fauce 5 mm.

lato, lobis ovato-oblongis acutis 4-4.5 mm. longis intus glabris;

antherae lineares semiexsertae. — Ecuador: Valley of the Pastaza

River, between Bafios and Cashurco, alt. 1,300-1,800 m., September

25, 1923, A. S. Hitchcock 21805 (U. S. Nat. Herb. No. 1,196,528, type;

photo, in herb. Field Mus.).

In stipule form this is similar to P. vaginata Benth. It differs

from that species in the longer corolla, elongate panicle, and peculiar

calyx. The flowers of P. vaginata are yellow.

Palicourea egena, sp. nov. — Frutex, ramulis validis densissime

pilis brevibus ochraceis hispidulo-tpmentosis, internodiis 1.5-2.5 cm.

longis; stipulae persistentes in vaginam 5^-7 mm. longam truncatam

densissime puberulam connatae, vagina in lobos 4 erectps lineares

puberulos 6-7 mm. longos desinente; folia opppsita, petiolo crasso

7-10 mm. longo hispidulo-tomentoso; lamina elliptico-oblonga, 8-12

cm. longa, 3.3-4.5 cm. lata, acuta, basi cuneatim angustata, rigide

coriacea, supra viridis, dense et minute hispidula, costa prominente,

neryis non eleyatis, subtus luteo-viridis, undique dense pilis brevibus

albidis patentibus hispidula, costa crassa elevata, nervis lateralibus

utroque latere c. 18, gracilibus, elevatis, angulo latp adscendentibus,

arcuatis, marginem attingentibus, nervulis prominulis arete reti-

culatis; inflorescentia terminalis 2.5 cm. longe pedunculata cymoso-

paniculata dense multiflora, 8 cm. longa et lata, e basi trichotoma,

ramis adscendentibus vel suberectis validis dense ochraceo-hispidulis,

floribus congestis, pedicellis 1-3 mm. longis hispidulis, bracteis

oblongis vel linearibus 2-6 mm. longis, obtusis vel acutis, hispidulis;

hypanthium obpvoideum 1.2 mm. longum hispidulum; calyx 1.2

mm. longus, lobis 5 oblongo-oyatis obtusis minute hispidulis; corolla

8 mm. longa extus dense hispidula, tubo validp superne sensim

dilatato ore 3 mm. lato, lobis triangulari-oblongis 2.5 mm. longis

obtusis. — Ecuador: Province of Pasto, September, 1845, Jameson

427 (Herb. Kew., type).

The plant is somewhat noteworthy because of the combination

of thick, multicostate, densely pubescent leaves, short, dense, many-

flowered inflorescence, and densely pubescent corolla.

Palicourea flavescens HBK. Nov. Gen. & Sp. 3: 366. 1820.

Psychotria flavescens Spreng. Syst. Veg. 1: 744. 1825. Palicourea

ochreata Wernham, Journ. Bot. 55: 281. 1917.

Without definite locality, Humboldt & Bonpland (photo, of type,

ex Herb. Berol., in F). Type of P. ochreata from Mt. Tulcan, Prov.

Carchi, 3,000 m., Lehmann 669. Molleturo, Cuenca, 2,500-2,800 m.,

Lehmann 5654 (F, B, K, W, Y). Andes, 1,500-1,800 m., Pearce (K).

THE RUBIACEAE OF ECUADOR 233

Branches densely villous-tomentose; stipule sheath 1-1.5 cm.

long, truncate, the distant subulate lobes 3-4 mm. long; leaves

opposite, short-petiolate, firm, elliptic-lanceolate or oblong-lanceo-

late, long-acuminate, narrowed to the acute base, densely hispidulous-

pubescent on both surfaces; panicles thyrsiform, pedunculate, up to

10 cm. long, the branches spreading, densely pubescent, the bracts

small and inconspicuous; corolla sparsely pubescent, the tube 6-7

mm. long, the lobes ovate, 2 mm. long.

In the original publication of Palicourea flavescens, the locality is

given doubtfully as Peru, and it may be that the specimen came from

that country. Schumann, who had the type before him, referred

Lehmann 5654 to P. flavescens, and this number seems to match

exactly the photograph of the type seen by the writer.

Palicourea graciliflqra, sp. nov. — Frutex 2.5-3-metralis, ramis

gracilibus teretibus glabris vel in partibus novellis sparse puberulis,

internodiis plerumque 2-5 cm. longis; stipulae persistentes glabrae in

vaginam latam 4-6 mm. longam truncatam connatae, vagina in lobos

4 lineares vel anguste triangulares 2-6 mm. longos remotos desinente;

folia opposita, petiolo valido 7-17 mm. longo glabro vel sparse

puberulo; lamina crasse membranacea, oblonga, elliptico-oblonga vel

obovato-oblonga, 11-22 cm. longa, 4.5-9 cm. lata, abrupte acuminate,

acumine 1.5-2 cm. longo angusto attenuate, basi acuta, supra yiridis,

glabra, costa prominente, nervis non elevatis, subtus pallidior, ad

nervos pilis minutis adscendentibus vel adpressis pilosula, costa

gracili elevata, nervis lateralibus utroque latere c. 24, gracillimis,

angulo lato adscendentibus, arcuatis, marginem attingentibus, ner-

vulis prominulis laxe reticulatis; inflorescentia terminalis thyrsoidea

laxe multiflora, 16-20 cm. longa et ultra, 6-16 cm. lata, 5-7.5 cm.

longe pedunculate, ramis breyibus sparse breviterque pilosis patenti-

bus vel saepe recuryis, floribus pedicellatis, pedicellis plerumque

3r4 mm. longis gracilibus, bracteis subulatis vel linearibus 1-3 mm.

longis; hypanthium obovoideum vix 1 mm. longum puberulum vel

glabratum; calyx 0.5 mm. longus minute 5-dentatus, dentibus ovatis

subacutis; corolla purpureo-caerulea gracilis 1.5-2.2 cm. longa, in

alabastro apice obtusa, extus sparsissime puberula vel hirtella, tubo

lineari basi subampliato, superne paullo dilateto, ore 2.5 mm. lato,

lobis oblongis obtusis 2.5 mm. longis; stylus capillaris glaber. —

Ecuador: Andes of Cuenca, alt. 1,200 m., Pearce 335 (Herb. Kew.,

type; photo, in F). — Colombia: Maldonado, Ocana to Pamplona,

Dec. 25, 1878, Kalbreyer 864 (K).

Remarkable for the long and very slender corolla and for the

narrow thyrsoid inflorescence.

Palicourea guianensis Aubl. PL Guian. 1: 173. pi. 66. 1775.

Portovelo, Holway 1001 (W). Without locality, Pearce (K).

Base of Mount Chimborazo, 900 m., Spruce 6181 (K). Widely dis-

tributed in tropical America.

234 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

A shrub or small tree 2.5-6 m. high, nearly glabrous; stipules

8-10 mm. long, biparted, the lobes lanceolate or ovate, obtuse; leaves

short-petiolate, large, thin, elliptic-oblong to ovate or elliptic, acumi-

nate, at the base acute to nearly rounded; panicles pedunculate,

dense, many-flowered, large, the branches ascending or spreading,

glabrous or puberulent, the bracts subulate, inconspicuous; flowers

pedicellate; calyx lobes minute, deltoid, acute or obtuse; corolla

orange-red or yellow, 10-18 mm. long, furfuraceous-tomentose or

glabrate; fruit ovoid, 4-5 mm. long.

Palicourea Heilbornii, sp. nov. — Frutex, ramulis crassis tere-

tibus glabris, internodiis c. 4 cm. longis; stipulae in vaginam 6-^7 mm.

longam glabram truncatam connatae, vagina in lacinias 4 lineares

obtusas glabras 5 mm. longas erectas desinente; folia opposita,

petiolo crasso 2 cm. longo glabro; lamina crasse chartacea, ovali-

elliptica, 15-18 cm. longa, 8-9.5 cm. lata, abrupte acuta vel breviter

acuminata, acumine triangulari acuto, basi rotundata vel acuta,

supra olivaceo-viridis, glabra, costa prominente, nervis non elevatis,

subtus fere concolor, ad costam crassam elevatam sparsissime pilis

albidis pilosa, aliter glabra, nervis lateralibus utroque latere c. 15,

prominentibus, angulo lato adscendentibus, gracilibus, arcuatis, in

marginem desinentibus, nervulis vix prominulis arete reticulatis;

inflorescentia terminalis 4 cm. longe pedunculata thyrsoideo-pani-

culata, 10 cm. longa et aequilata, laxe multiflora, ramis basalibus

oppositis rigidis crassis glabris, angulo lato adscendentibus, pedicellis

crassis 6-15 mm. longis glabris, bracteis anguste triangularibus 2-4

mm. longis acutis glabris; hypanthium late campanulatum glabrum

2 mm. longum; calyx 3 mm. longus, glaber, laciniis ovalibus vel late

ovatis apice rotundatis vel obtusissimis; corolla coccinea extus glabra

2.3 mm. longa, tubo crasso basi paullo ampliato superne sensim dila-

tato ore 5 mm. lato, lobis 5 anguste triangularibus 5 mm. longis obtu-

sis intus minutissime papillosis; antherae semiexsertae. — Ecuador:

Tandapi, Prov. Pichincha, in silva primaeva, alt. 1,500 m., July,

1920, 0. Heilborn 773 (Stockholm Herb., type).

The plant may be recognized by the combination of large, nearly

glabrous leaves, ample inflorescence, large calyx lobes, and long

stout corolla.

Palicourea Holmgrenii, sp. nov. — Ramuli crassiusculi virides

subteretes subdense pilis debilibus albidis villosuli, internodiis 2.5-6

cm. longis; stipulae in vaginam persistentem 3-4 mm. longam

ferrugineam fere glabram truncatam connatae, vagina in lobos 4

erectos lineares 3-4 mm. longos acutos ciliatos desinente; folia

opposita, petiolo valido 6-15 mm. longo dense villosulo vel hispidulo;

lamina chartacea, oyato-oblonga vel anguste ovata, 6-10 cm. longa,

2.5-4 cm. lata, longissime acuminata, acumine angusto longe attenu-

ate, basi obtusa vel subrotundata, rarius abrupte contracta, supra

olivaceo-viridis, in statu juvenili rhaphidibus minutis brevibus palli-

dis conspersa, ad costam et interdum ad nervos hispidula, aliter

THE RUBIACEAE OF ECUADOR 235

glabra, nervis non elevatis, subtus pallida, ad nervos dense villosa

et hispidula, inter nervos glabra, costa crassa elevata, nervis lateral-

ibus utroque latere c. 16, angulo lato adscendentibus, gracilibus,

prqminentibus, valde arcuatis, in marginem desinentibus, nervulis

reticulatis subimpressis; inflorescentia terminalis sessilis et basi

foliata vel 2.5 cm. longe pedunculata, late pyramidali-paniculata,

dense multiflora, c. 6 cm. longa et 6-8 cm. lata, basi trichotpma, ramis

validis patentibus vel adscendentibus dense hispidulis, floribus cymo-

sis 1-2 mm. longe pedicellatis, pedicellis hispidulis, bracteis linearibus

vel lanceolatis attenuatis 3-4 mm. longis hispidulis et ciliatis viridi-

bus; hypanthium late turbinatum 1.2 mm. longum sparse hispidulum

vel fere glabrum; calyx 1.2 mm. longus, lobis late ovatis acutis glabris;

corolla caeruleo-violacea 9 mm. longa extus sparse villosula, tubo

basi ampliato et 4 mm. lato, versus apicem sensim dilatato et ore

3 mm. lato, lobis oblongis obtusis 2 mm. longis; stylus breviter

exsertus; fructus obovoideo-globosus 5 mm. longus glabratus, pyrenis

obtuse costatis. — Ecuador: Tandacato, Prov. Pichincha, in silva

primaeva, alt. 2,500 m., April 6, 1920, /. Holmgren 469 (Herb. Field

Mus. No. 605,119, type; duplicate in Stockholm herb.).

Palicourea hospitalis, sp. nov.— Frutex 2-3-metralis, ramulis

gracilibus viridibus obtuse tetragonis dense minuteque puberulis,

internodiis 6-7 cm. longis; stipulae virides persistentes 10-13 mm.

Ipngae, basi breviter connatae, dense ochraceo-strigillosae, profunde

bifidae, laciniis oblongo-ovatis obtusis; folia opposita, petiolo gracili

2-2.5 cm. longo dense minuteque sericeo-strigilloso; lamina mem-

branacea, elliptica vel ovato-elliptica, 12-17 cm. longa, 6-8 cm. lata,

acuta vel abrupte acuta, basi acuta vel cuneatim acuta, supra luteo-

viridis, glabra, costa nervisque prominulis, subtus paullo pallidior,

minute adpresso-pilosula, ad costam gracilem elevatam et ad nervos

pilis longioribus subadpressis pilosa, nervis lateralibus utroque latere

c. 20, gracilibus, prominentibus, angulo lato adscendentibus, arcuatis,

in marginem desinentibus, nervulis prominulis arete reticulatis;

inflorescentia terminalis 5 cm. longe pedunculata, late pyramidali-

paniculata, laxe multiflora, 16 cm. longa et aequilata, basi trichotoma

et foliis reductis bracteata, pedunculo erecto strigillpso, ramis basal-

ibus oppositis angulo recto divergentibus, superioribus abbreviatis,

sparse puberulis, floribus plerumque 1-2 mm. longe pedicellatis,

bracteis linearibus vel lanceolatis 2-6 mm. longis extus strigillosis;

hypanthium late obovoideum 1-1.2 mm. longum glabrum; calyx

glaber 1-1.2 mm. longus 5-fidus, laciniis inaequalibus oblongis vel

lanceolato-oblongis obtusis vel acutiusculis; corolla lutea crassa in

alabastro apice rotundata et 2.5 mm. longa, aperta non visa. —

Ecuador (?): Tambo de Sabanilla, December 18, 1876, Ed. Andre

4566 (Herb. Kew., type; photo, in herb. Field Mus.).

Closely similar in general appearance to P. Kalbreyeri Standl., of

Colombia, but with very different stipules and much broader leaves.

Palicourea huigrensis, sp. nov. — Arbor 5.5 m. alta ramosa,

ramulis gracilibus obtuse tetragonis hispidulis vel glabratis olivaceis,

236 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

internodiis brevibus vel elongatis; stipulae in vaginam 5-6 mm.

longam laxam truncatam dense adpresso-pilosulam connatae, vagina

in lobos 4 approximates oblpngo-lineares 5-7 mm. longos obtusos

desinente; folia opposita, petiolo valido 1-2 cm. longo hispidulo vel

glabrato; lamina subcoriacea, elliptico-oblonga, elliptica vel obovato-

oblonga, 8-13.5 cm. longa, 4-5.5 cm. lata, anguste acuminata,

acumine obtuso, basi angustata, supra viridis, ad costam subimpres-

sam pilosa, nervis subimpressis, subtus paullo pallidior, praesertim

ad nervos dense breviterque pilosa vel villosula, costa valida elevata,

nervis lateralibus utroque latere c. 16, prominentibus, angulo lato

adscendentibus, arcuatis, marginem attingentibus, nervulis promi-

nulis reticulatis; inflorescentia 6-7.5 cm. longe pedunculata pyra-

midali-paniculata dense multiflora, 7-8 cm. longa, 4.5-6 cm. lata,

ramulis divaricatis vel adscendentibus sparse puberulis crassiusculis;

bracteae oblongae vel lanceolatae, obtusae vel acutae, 2-3.5 mm.

longae, ciliolatae; flores sparsi, pedicellis crassis 1.5-6 mm. longis

apice vix incrassatis; hypanthium 1.5 mm. longum late turbinatum

glabrum; calyx glaber 1.5 mm. longus profunde 5-lobus, lobis late

ovatis vel rotundatis; corolla violacea vel caerulea tubulosa extus

glabra 16-17 mm. longa, tubo crasso basi paullo ampliato, superne

vix dilatato, fauce 4 mm. lato, lobis ovatis subacutis 2-2.5 mm. longis;

stylus exsertus minute puberulus. — Ecuador: Near Huigra, September

3, 1918, J. N. andG. Rose 22515 (U. S. Nat. Herb. No. 1,022,167,

type; duplicate in Y; photo, in herb. Field Mus.). In sylvis Llalla

ad pedem mentis Azuay, August, 1859, Spruce 6003 (F, K, L).

Palicourea Jamesonii, sp. nov. — Ramuli graciles teretes, vetus-

tioribus ochraceis laevibus, novellis pilis brevibus incurvis basi

dilatatis furfuraceo-puberulis; stipulae virides persistentes in vaginam

1-1.5 mm. longam sparse puberulam vel glabratam truncatam con-

natae, vagina in lacinias 4 erectas lineari-oblongas 1-2 mm. longas

ciliolatas desinente; folia opposita, petiolo 3-5 mm. longo supra

sulcato puberulo ; lamina subcoriacea, anguste elliptica, ovato-oblonga

vel lanceolato-oblonga, 2.5-5.5 cm. longa, 0.8-2 cm. lata, acuminata,

basi acuta vel subobtusa, supra viridis, ad costam prominulam sparse

breviterque pilosa, aliter glabra, nervis non elevatis, subtus pallidior,

ad costam crassiusculam elevatam aciculari-pilosula, aliter glabra,

nervis lateralibus utroque latere c. 7, gracilibus, prominentibus,

angulo angusto adscendentibus, arcuatis, in marginem desinentibus;

inflorescentia terminalis vel pseudo-axillaris 1-3 cm. longe pedun-

culata, plerumque 4-7-flora, densa, pedunculo gracili incurvo-scabe-

rulo, ramis brevissimis suberectis, pedicellis 2-4 mm. longis incurvo-

scaberulis, bracteis linearibus interdum foliaceis 4-7 mm. longis

persistentibus acutis fere glabris ciliolatis; hypanthium glabrum vel

sparse minuteque puberulum 1 mm. longum; calyx glaber 1.8-2 mm.

longus 5-fidus, laciniis late oblongis vel ovalibus obtusis viridibus;

corolla 13-14 mm. longa extus glabra, tubo basi paullo ampliato

superne sensim dilatato, ore 4-5 mm. lato, lobis oblongis 3 mm. longis

obtusis intus minute papillosis; fructus subglobosus glaber 6-7 mm.

THE RUBIACEAE OF ECUADOR 237

longus vix costatus. — Ecuador: "Quito," Jameson (Herb. Kew.,

type).

Palicourea lasiantha Krause, Bot. Jahrb. Engler 40: 341. 1908.

Andes, 2,700 m., Pearce (K). Also in Peru and Bolivia.

A shrub about 3 m. high, nearly glabrous; stipules 4-7 mm. long,

bifid, the lobes rounded; leaves petiolate, firm, oblong to elliptic,

large, short-acuminate, at the base acute to rounded, glabrous or

nearly so; inflorescence paniculate, large, open or dense, many-

flowered, the branches spreading or ascending; flowers pedicellate;

calyx lobes ovate, acutish; corolla purple, about 2 cm. long, densely

lanate, the lobes very short.

Palicourea levis Standl. Field Mus. Bot. 4: 336. 1929.

Andes, in 1857-59, Spruce (K). Pichincha, Jameson (K). Tungu-

rahua, Spruce (K). Also in Peru.

A shrub or tree 2-6 m. high, the branchlets glabrous or minutely

puberulent; stipule sheath 1.5-2.5 mm. long, the lobes narrowly

triangular or broadly linear, 1.5-2 mm. long; leaves opposite, petio-

late, oblong or elliptic-oblong, 5.5-11 cm. long, acute or short-acumi-

nate, obtuse at the base or abruptly contracted, glabrous, or sparsely

barbate beneath in the axils of the veins; inflorescence small, terminal,

densely many-flowered, pedunculate or rarely sessile, thyrsoid-panic-

ulate, 3-7 cm. long, the branches minutely puberulent or hirtellous,

the flowers crowded, sessile; calyx teeth broadly ovate, obtuse or

acutish, 0.6 mm. long; corolla white or yellowish, minutely puberu-

lent, 5-6 mm. long, the lobes one-third as long as the tube; fruit

glabrous, 5 mm. long, smooth.

Palicourea lineata Benth. PI. Hartw. 192. 1845. Uragoga

lineata Kuntze, Rev. Gen. PI. 1: 300. 1891.

Type from forests of Guagua, western slope of Mount Pichincha,

Hartweg 1057 (fragm., ex Herb. Berol., in F). Between El Pichincha

and El Atacaro, 3,300 m., Firmin 484 (F, W). Without locality,

Jameson (W). Malchingui to Pomasaqui, Prov. Pichincha, 3,000-

3,600 m., Hitchcock 20889 (W, Y). Cuenca and coast cordillera,

2,100-2,700 m., Pearce 379 (K). Western side of Pichincha, 2,700

m., Jameson 83 (K). Tigua, Sodiro 8427 (F, a fragm. ex Herb. Berol.).

A shrub 1-2 m. high, nearly or quite glabrous; stipule sheath 3-4

mm. long, the lobes short, oblong, obtuse; leaves petiolate, elliptic

or oblong-elliptic, 7-13 cm. long, acuminate, at the base acute or

obtuse, thick, glabrous or nearly so, paler beneath; panicles pedun-

culate, small, as broad as long, dense, the branches mostly spreading;

flowers pedicellate; calyx lobes ovate, obtuse; corolla blue, glabrous,

slender, 12-18 mm. long; fruit 5-6 mm. long.

Palicourea Lobbii, sp. nov. — Frutex, ramis vetustioribus cras-

siusculis rimosis olivaceis glabratis, novellis gracilioribus dense

scaberulis, internodiis elongatis; stipulae in vaginam scaberulam

238 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

truncatam 1-1.5 mm. longam connatae, vagina in lobos 4 lineares

scaberulos remotos 2-3 mm. longos desinente; folia opposita, petiolo

crassiusculo 2-4 mm. longo dense scaberulo; lamina lanceolato-oblonga

vel ovato-oblonga 2.5-4.5 cm. longa, 1-1.5 cm. lata, longissime

attenuato-acuminata, acumine angusto acuto saepe falcato, basi

obtusa vel subacuta, subcoriacea, supra viridis, scaberula, costa

paullo prominente, nervis obscuris, subtus pallidior, ad nervos minute

incurvo-hispidula, costa gracili elevata, nervis lateralibus utroque

latere c. 9, angulo lato adscendentibus, arcuatis, marginem revolutum

fere attingentibus, nervulis non elevatis arete reticulatis; inflores-

centiae terminates plerumque sessiles et e basi trichotomae, pauci-

florae, f olioso-bracteatae, ramis suberectis dense minuteque hispidulis,

floribus sessilibus vel 1 mm. longe pedicellatis, bracteis minoribus

linearibus vel subulatis scaberulis 2-5 mm. longis; hypanthium his-

pidulum; calyx minute hispidulus, lobis ovato-oblongis subacutis

1 mm. longis; corolla caerulea extus glabra vel hinc inde hispidula,

in alabastro apice anguste rotundata, 1 cm. longa, tubo gracili ore

2 mm. lato; fructus ovalis glabratus apice rotundatus 5-6 mm. longus

tenuiter et obtuse costatus. — Ecuador: Without definite locality,

Lobb (Herb. Kew., type).

In size and shape of the leaves as well as in general appearance

this Ecuadorean plant is similar to P. Lechleri Standl., of Peru, but

the pubescence of the two is clearly distinct.

Palicourea lugubris Schum. & Krause, Bot. Jahrb. Engler 40:

337. 1908.

Dense forest near Naranjal, Lehmann 5652 (F, type collection;

photo, of type, ex Herb. Berol., in F). Balao, in forest, Eggers 14133

(F, L). San Ignacio, Prov. Guayas, Heilborn 8 (F, S). Also in

Colombia.

A nearly glabrous shrub 2 m. high; stipules lobed almost to the

base, 7-8 mm. long, the lobes lance-oblong, acute; leaves short-

petiolate, thin, blackish when dried, elliptic-oblong or oblong-lanceo-

late, 8-17 cm. long, 4-6 cm. wide, acuminate, at the base cuneately

narrowed, glabrous; inflorescence pedunculate, cymose-corymbose,

as broad as long, dense, the branches ascending, puberulent, the

bracts linear; flowers short-pedicellate; calyx lobes linear-oblong,

acute; corolla tubular, white or purplish, sparsely puberulent or

glabrous, 8-10 mm. long, the lobes short, acute; fruit subglobose,

4 mm. in diameter.

Palicourea macrobotrys (R. & P.) R. & S. Syst. Veg. 5: 194.

1819. Psychotria macrobotrys R. & P. Fl. Peruv. 2: 57. pi. 203, /. o.

1799. Palicourea Williamsii Rusby, Descr. N. Sp. S. Amer. PL 142.

1920.

No Ecuador specimens have been seen by the writer. Rusby,

in describing P. Williamsii, states that that species is represented by

a specimen collected in Ecuador by Pearce. Since P. macrobotrys

THE RUBIACEAE OF ECUADOR 239

occurs in both Peru and Colombia, it is probable that it is native also

in Ecuador. The species extends to Bolivia and Brazil.

A shrub, the branches densely pubescent; stipule sheath 4 mm.

long or less, the lobes subulate, usually longer than the sheath ; leaves

short-petiolate, lance-oblong or narrowly elliptic-oblong, thin, long-

acuminate, acute at the base, finely soft-pubescent beneath over the

whole surface; inflorescence thyrsoid, pedunculate or sessile, elongate

and narrow, with very short branches, the flowers pedicellate; calyx

minute, the lobes broadly ovate, acute; corolla yellow, tubular, 1 cm.

long, pubescent; fruit 3.5-4 mm. long.

Palicourea myrtifolia Schum. & Krause, Bot. Jahrb. Engler

40: 334. 1908.

Eastern Andes near Loja, in dense forest along Rio Sabanilla,

1,000-1,500 m.,Lehmann 4934 (F, W, type collection; photo, of type,

ex Herb. Berol., in F). Near Matala, 2,500-3,000 m., Lehmann 7931

(F). Without locality, Andre K504 (K).

A slender, nearly glabrous shrub 3 m. high; stipules 3-4 mm.

long, the lobes subulate; leaves petiolate, small, firm, lanceolate to

elliptic-oblong, 3.5-6 cm. long, 1-2.5 cm. wide, long-acuminate, at

the base acute, glabrous, paler beneath; panicles pyramidal, lax,

sessile or pedunculate, many-flowered, 7-10 cm. long, puberulent,

the bracts small, subulate; calyx lobes ovate, acute; corolla purplish,

minutely puberulent, 1 cm. long, the lobes short, obtuse.

Palicourea oreadium, sp. nov. — Ramuli crassiusculi subteretes

densissime pilis brevibus patentibus hispidulo-tomentosi, internodiis

elongatis; stipulae persistentes in vaginam 2 mm. longam truncatam

dense hispidulo-tomentosam connatae, vagina in lacinias 4 oblongo-

triangulares obtusas 2 mm. longas puberulas approximatas desinente;

folia opposita, petiolo crassiusculo 1-1.5 cm. longo dense hispidulo;

lamina subcoriacea, elliptico-oblonga vel ovato-oblonga, 7.5-10 cm.

longa, 3.5-4 cm. lata, acuminata, acumine triangulari acuto, basi

obtusa vel subrotundata, supra viridis, sparse scaberula, ad nervos

hispidula, nervis non elevatis, subtus paullo pallidior, ubique dense

molliterque hispidula, costa crassa elevata, nervis lateralibus utroque

latere c. 11, prominentibus, angulo acuto adscendentibus, arcuatis,

in marginem desinentibus, nervulis prominulis arete reticulatis;

inflorescentia terminalis 4.5 cm. longe pedunculata thyrsoideo-pani-

culata, 6 cm. longa et aequilata, dense multiflora, ramis basalibus

subverticillatis angulo lato adscendentibus, dense puberulis vel his-

pidulis, floribus pedicellatis, pedicellis 2-4 mm. longis puberulis,

bracteis oblongis vel lanceolatis acutis vel obtusis puberulis 2-6 mm.

longis; hypanthium late turbinatum 1.5-2 mm. longum puberulum

vel fere glabrum; calyx 1.5-2 mm. longus minute puberulus, laciniis

late oblongis vel ovalibus apice rotundatis; corolla 14 mm. longa

extus versus apicem scaberula vel fere glabra, in alabastro apice

obtusa, tubo crasso basi paullo ampliato, superne sensim dilatato,

fauce 4 mm. lato, lobis 5 ovatis obtusis suberectis 2.5 mm. longis intus

240 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

papillosis; antherae semiexsertae. — Ecuador: Ravine of Tabacay

near Azogues, alt. 3,000 m., Jameson (Herb. Kew., type; photo, in

herb. Field Mus.).

Apparently a close relative of P. lineata Benth., which is very

similar in general appearance, but P. oreadium may be distinguished

at once from that species by the dense soft pubescence of the lower

leaf surface.

Palicourea pasti Wernham, Journ. Bot. 55: 283. 1917.

Without locality, Jameson (F, W). Andes, Spruce 5466 (B, G,

K, L). Also in Colombia.

A glabrous shrub or small tree, sometimes 6 m. high; stipules

6-11 mm. long, the lobes broad, rounded at the apex, ciliate; leaves

petiolate, elliptic-oblong to elliptic, 6.5-14 cm. long, 3-6 cm. wide,

acute or obtuse, at the base acute or obtuse, sometimes sparsely

hirtellous beneath along the costa; inflorescence subsessile or pedun-

culate, lax, many-flowered, broadly paniculate, 8-16 cm. long and

broad, the pedicels 1-6 mm. long; calyx 1-1.5 mm. long, the lobes

ovate or rounded, acutish or rounded at the apex; corolla 8-10 mm.

long, dark purple and glabrous outside, pale lilac within, the lobes

2 mm. long.

Palicourea pluricostata, sp. nov. — Ramuli crassiusculi sub-

teretes, fere laeves glabri, internodiis elongatis; stipulae in vaginam

6-9 mm. longam adpressam truncatam margine ciliolatam connatae,

vagina in lacinias 4 oblongas vel lineares obtusas vel acutiusculas

6-7 mm. longas desinente; folia breviter petiolata opposita, petiolo

valido 5-10 mm. longo dense puberulo; lamina crasse membranacea,

elliptico-oblonga, usque ad 20 cm. longa et 7.5 cm. lata, Iqnge

acuminata, acumine angusto longe attenuate, basi acuta, conspicue

pallido-marginata, supra fusco-viridis, ad costam hirtella vel pube-

rula, aliter glabra, nervis prominulis, subtus pallida, luteo-viridis,

praesertim ad nervos hirtella vel hispidula, costa gracili eleyata,

nervis lateralibus utroque latere c. 27, gracilibus, prominentibus,

angulo lato adscendentibus, arcuatis, prope marginem conjunctis,

nervulis prominentibus; inflorescentia terminalis sessilis cymoso-

paniculata 10 cm. longa et 8.5 cm. lata, dense multiflora, ramis basali-

bus opppsitis angulo lato adscendentibus, validis, dense puberulis,

superioribus saepe divaricatis vel reflexis, bracteis subulatis plerum-

que 2-4 mm. longis, floribus dense congestis sessilibus; hypanthium

dense puberulum 0.6 mm. longum; calyx 0.6 mm. longus dense

puberulus 5-lobus, lobis minutis late ovatis apice rotundatis; corolla

6 mm. longa extus dense fulvo-pilosula, in alabastro apice obtusis-

sima, tubo crassiusculo superne paullo ampliato, lobis oblongis obtu-

sis 2.5 mm. longis; antherae exsertae lineares 2 mm. longae.—

Ecuador: Tambo de Sabanilla (?), E. Andre K1112 (Herb. Kew.,

type).

Palicourea Seemannii, sp. nov. — Ramuli crassiusculi in sicco

fusco-olivacei minute papillosi, vetustioribus ochraceis; stipulae laxae

THE RUBIACEAE OF ECUADOR 241

persistentes 7 mm. longae glabrae breviter bilobae, lobis triangulari-

bus acuminatis sinu lato separatis; folia opposita, petiolo gracili

7-16 mm. longo supra sulcato glabro; lamina elliptico-oblonga, rarius

obpvato-oblonga, 7.5-12 cm. longa, 2.2-5 cm. lata, crasse papyracea,

apice abrupte breviterque acuminata, acumine obtuso, basi acuta,

supra viridis, sublucida, ad costam prominentem sparse pilosula,

nervis manifestis sed vix elevatis, subtus vix pallidior, ad nervos pilis

longis sparse pilosa, costa gracili elevata, nervis lateralibus utrpque

latere c. 11, prominentibus, angulo lato vel acuto adscendentibus,

gracilibus, leviter arcuatis, marginem attingentibus; inflorescentia

sessilis late paniculata, c. 3 cm. longa et 5 cm. lata, laxe multiflora,

basi trichotoma, ramulis divaricatis dense puberulis; bracteae subu-

latae yel anguste triangulares 1.5-3 mm. longae glabrae; flores

conferti sessiles vel subsessiles; hypanthium turbinatum glabrum

1 mm. longum; calyx 0.8 mm. longus glaber 5-lobus, lobis late tri-

angulari-ovatis obtusis vel subacutis; corolla infundibuliformis 6 mm.

longa extus glabra, tubo basi paullo ampliato superne dilatato, lobis

oblongis obtusis fere 2 mm. longis; antherae semiexsertae. — Ecuador:

Without locality, B. Seemann (Gray Herb., type). Loja, August,

1847, Seemann 890 (K).

The plant is not obviously related to any other species of Palicou-

rea occurring in western South America. It is distinguished chiefly

by the small flowers and the small sessile inflorescence, hidden among

the leaves.

Palicourea stenosepala, sp. nov. — Frutex, ramulis gracilibus

olivaceis obtuse quadrangularibus glabris, internodiis 2-4.5 cm.

longis; stipulae persistentes 3-4 mm. longae bipartitae, lobis ovatis

obtusis extus strigillosis; folia opposita, petiolo 1.2-3.3 cm. longo

gracili glabro vel sparse hispidulo; lamina elliptico-oblonga vel

obovato-oblonga, 9-14 cm. longa, 3.5-5.5 cm. lata, abrupte acuminata

vel longiacuminata, acumine angusto attenuate, basi acuta, crasse

membranacea, supra viridis, glabra, nervis non elevatis, subtus

pallidior, praesertim ad nervos pilis rigidiusculis patentibus hirtello-

pilosa, costa valida elevata, nervis lateralibus utroque latere c. 12,

gracilibus, angulo lato adscendentibus, arcuatis, prominentibus, in

marginem desinentibus, nervulis reticulatis vix prominulis, margine

anguste revoluto; inflorescentia terminalis thyrsoideo-paniculata laxe

multiflora 1.5-4 cm. longe pedunculata, 4.5-5.5 cm. longa, c. 5 cm.

lata, ramis divaricatis vel adscendentibus, infimis oppositis vel

verticillatis, glabris, flpribus superioribus racemosis, pedicellis ple-

rumque 6^12 mm. longis glabris, bracteis oblongis vel linearibus 2-4

mm. longis obtusis vel acutis saepe ciliatis; hypanthium late pbp-

voideum glabrum 2 mm. longum; calyx 4-5.5 mm. longus laciniis

oblongis vel anguste oblongis glabris scaberulo-ciliatis; corolla alba

extus glabra, tubo crasso 13 mm. longo superne sensim dilatato ore

5 mm. lato basi paullo ampliato, lobis ovato-triangularibus obtusis

3 mm. longis subpatentibus intus minutissime papillosis; fructus

subglobosus 6-7 mm. longus glaber grosse costatus, calyce persistente

242 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

coronatus. — Ecuador: Tandacato, Prov. Pichincha, in silva pri-

maeva, alt. 2,500 m., April 5, 1920, /. Holmgren 465 (Stockholm Herb.,

type).

This species is noteworthy for the unusually long and narrow

calyx lobes.

Palicourea stipulates Benth. PI. Hartw. 133. 1844. Psychotria

stipularis Rusby, Mem. Torrey Club 33: 47. 1893.

Type from mountains of El Sisma, region of Loja, Hartweg 749

(photo, of type collection, ex Herb. Berol., in F). Chagal, Andes of

Cuenca, 2,200-2,700 m., Lehmann 4933 (F, W). Between La Chorita

and Portovelo, Prov. Oro, 1,000-2,000 m., Hitchcock 21183 (W, Y),

21186 (W, Y). Loja, Rose, Pachano & Rose 23295 (G, W, Y). Andes,

1,80072,100 m., Pearce (K). Tungurahua, Spruce (K). Also in

Bolivia.

A shrub or small tree, nearly glabrous; stipule sheath 4-5 mm.

long, the lobes linear, distant, usually shorter than the sheath;

leaves short-petiolate, elliptic-oblong, 7-20 cm. long, acuminate, at

the base acute or attenuate, glabrous or pilose beneath on the nerves;

panicles thyrsiform-pyramidal, pedunculate, many-flowered, dense,

the branches mostly ascending, glabrous, the bracts minute; flowers

pedicellate; calyx lobes minute, acute or obtuse; corolla glabrous,

12-20 mm. long, tubular, stout; fruit 5 mm. long.

The color of the corolla is reported variously as "blue," "purple,"

and "pink." It is not certain that all the specimens cited are referred

correctly to P. stipularis, but they agree fairly well with the incom-

plete description of that species.

Palicourea tectoneura Schum. & Krause, Bot. Jahrb. Engler

40: 336. 1908.

Type collected on the western slope of the western Andes near

Cuenca, in dense forest near Chagal, 2,000-2,700 m., Lehmann 7930

(photo, of type, ex Herb. Berol., in F). Chagal, 2,400-2,600 m.,

Lehmann K131 (K).

A tree 10 m. high, the trunk 30 cm. in diameter, the branches

glabrous; stipule sheath 7-8 mm. long, the lobes very short; leaves

short-petiolate, rigid-coriaceous, elliptic or ovate-elliptic, acuminate,

at the base acute to rounded, 8-13 cm. long, 4.5-7 cm. wide, puberu-

lent beneath especially on the nerves; panicles pyramidal, many-

flowered, 7-12 cm. long; the branches sparsely pilosulous, the bracts

subulate, 3-4 mm. long; flowers short-pedicellate; calyx lobes ovate,

acute; corolla yellowish, puberulent, 5-6 mm. long, the 5 lobes

subacute.

36. RUDGEA Salisb.

Rudgea fimbriata (Benth.) Standl. in Standl. & Cald. Lista

PI. Salvador 274. 1925. Psychotria fimbriata Benth. in Hook. Journ.

Bot. 3: 226. 1841. R. micrantha Muell. Arg. Flora 59: 454. 1876.

THE RUBIACEAE OF ECUADOR 243

El Recreo, Eggers 15471 (F), 15571 (B, K, L, W). Also in Bolivia,

Peru, Brazil, the Guianas, and Central America.

A glabrous shrub; stipules triangular-ovate, incised-laciniate;

leaves subsessile, broadly elliptic to elliptic-oblong, 8-15 cm. long,

acuminate, at the base obtuse; flowers sessile, cymose-paniculate,

the panicles small, broad, many-flowered; calyx minutely 5-dentate;

corolla 5-7 mm. long, the lobes longer than the tube; fruit ellipsoid,

5-8 mm. long.

37. CORYNULA Hook. f.

Corynula pilosa (Benth.) Hook. f. Icon. PI. pi. 1123. 1872.

Mitchella pilosa Benth. PI. Hartw. 194. 1845.

Type from Minasbamba, near Quito, Hartweg 1066.

A creeping herb; leaves rounded, acute, 6-12 mm. long, truncate

at the base, pilose beneath along the nerves; flowers 4-5-parted,

solitary, pedicellate, the pedicels longer than the petioles; calyx lobes

linear; corolla 8 mm. long, the lobes narrow, acute, pilose; fruit slightly

fleshy, containing 2 ribbed nutlets.

38. NERTERA Banks & Soland.

Nertera depressa Banks & Soland. ex Gaertn. Fruct. 1 : 124. pi.

26. 1788.

Without locality, Jameson (F). Ravines of Pichincha, Jameson

747 (L). Corazon, 3,000 m., Andre K515 (F). Mt. Pichincha, in

paramos, 3,850 m., Heilborn 282 (F, S). Huigra, Rose & Rose 22491

(G). Widely distributed in the mountains of South and Central

America.

A slender creeping herb, often forming dense mats over banks

and rotten logs; leaves small, petioled; fruits fleshy, bright red.

MITCHELLA OVATA DC. Prodr. 4: 452. 1830. Nertera tetrasperma

HBK. 3: 379. 1820.

The type was collected by Humboldt and Bonpland on the

Volcano of Tungurahua. The name is probably synonymous with

Nertera depressa.

39. MITRACARPUS Zucc.

Mitracarpus hirtus (L.) DC. Prodr. 4: 572. 1830. Spermacoce

hirta L. Sp. PI. ed. 2. 148. 1762.

Tumbaco, April, 1873, Andre (K). Near Ibarra, 2,400 m.,

Jameson (K). A widely distributed weed of tropical America.

An erect, usually much-branched annual, hispidulous or glabrate;

leaves lanceolate to ovate-oblong, acute, scabrous or villous; flowers

in dense axillary clusters; corolla white, not exceeding the calyx.

244 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

40. RICHARDIA L.

Richardia brasiliensis Gomez, Mem. Ipecac. 31. pi. 2. 1801.

Huigra, 1,200 m., Hitchcock 20308 (W, Y). Widely distributed

as a weed in South America.

A prostrate, pilose or hispid annual, often forming mats; leaves

oblong to elliptic, petiolate; flowers white, in dense terminal clusters;

corolla 4-6 mm. long; carpels of the fruit 3, muricate, carinate on the

inner face.

Known in Brazil as "poaya," "poaya branca," and "poaya del

campo."

Richardia scabra L. Sp. PI. 330. 1753.

Olaya, December, 1864, Jameson (W). Widely distributed as a

weed in tropical America.

A pilose or hispid annual, the flowers white, in dense terminal

clusters; carpels of the fruit 3, sulcate on the inner face.

41. DIODIA L.

Diodia dichotoma (HBK.) Schum. in Mart. Fl. Bras. 66: 11.

1888. Spermacoce dichotoma HBK. Nov. Gen. & Sp. 3: 348. 1819.

D. glabra Willd. ex R. & S. Syst. Veg. 3: 532. 1818, non Pers. 1805.

Borreria dichotoma C. & S. Linnaea 3: 340. 1828. Triodon laxum

Spruce ex Schum. loc. cit., as syn.

Type from Ayavaca and slopes of Mount Tungurahua, Humboldt

&Bonpland (photo, of type, ex Herb. Berol., in F). Andes of Ecuador,

Spruce 5997 (G) . Near Loja, September, 1864, Jameson (S) ; October,

1864, Jameson (S). Roadsides near Quito, Holmgren 522 (F, S).

Schumann reports also Jameson 792. Also in Peru.

Plants small, erect, suffrutescent; leaves lanceolate or oblong-

lanceolate, 1-2 cm. long, acuminate, scabrous above, pilosulous

beneath along the nerves or glabrous; flowers in small terminal cymes;

fruit glabrate, the cocci scarcely carinate.

Diodia prostrata Sw. Prodr. Fl. Ind. Occ. 30. 1788.

Puna, Andersson 73 (S), 74 (S). Chanduy, Spruce 6466 (S).

Also in Brazil, Central America, and Jamaica.

Plants annual, sometimes somewhat woody at the base, the stems

hispid; leaves linear, acuminate, mucronate, 1.5-2.5 cm. long, 2-3

mm. wide, pilose above, aculeolate-serrulate; flowers axillary, solitary

or geminate, short-pedicellate; corolla 3 mm. long; fruit 3 mm. long,

obscurely costate dorsally, hispidulous.

Diodia radula (Willd. & Hoffmannsegg) S. & C. Linnaea 3: 342.

1828. Spermacoce radula Willd. & Hoffmannsegg ex R. & S. Syst.

Veg. 3: 531. 1818.

Reported from Albemarle, Charles, Chatham, and James Islands

of the Galapagos group. Also in Brazil.

THE RUBIACEAE OF ECUADOR 245

A perennial herb, the stems prostrate or ascending, elongate,

puberulent, the younger ones sometimes grayish-tomentose; leaves

broadly elliptic to oblong-lanceolate, 2.5-5 cm. long, acute at the

base and apex, sessile, scabrous above, pilose beneath, plicate-nerved;

flowers in 6-10-flowered clusters; hypanthium puberulent and pilose,

the 4 calyx lobes unequal, subulate; corolla 1 cm. long; fruit 3 mm.

long, 4-5 mm. wide, the cocci subglobose, 3-sulcate.

The plant is reported to grow in the Galapagos at elevations of

270-1,050 m.

Diodia rigida (Willd.) C. & S. Linnaea 3: 341. 1828. Spermacoce

rigida Willd. ex R. & S. Syst. Veg. 3: 531. 1818.

Playas, Prov. Guayas, Mille 193 (W). Widely distributed in

tropical America, growing usually in open grassland.

An erect or ascending annual or perennial with linear-lanceolate

leaves 2-4 cm. long; leaves scabrous or hispidulous; flowers lavender,

sessile in the leaf axils, usually solitary; fruit hispidulous or glabrate,

3-costate dorsally.

42. SPERMACOCE L.

Spermacoce glabra Michx. Fl. Bor. Amer. 1: 82. 1803.

Between Guayaquil and Salinas, Prov. Guayas, 100 m., Hitch-

cock 19982 (W, Y). Widely distributed in tropical America, but

seldom if ever very plentiful.

A glabrous annual weed; leaves lanceolate to elliptic; flowers

minute, white, sessile and clustered in the leaf axils; fruit glabrous.

Spermacoce tenuior L. Sp. PI. 102. 1753.

Between Guayaquil and Salinas, Prov. Guayas, 100 m., Hitchcock

20082 (W). Puna, Andersson 72 (S); reported from the Galapagos

Islands. Widely distributed in tropical America.

An annual weed, more or less pubescent, the leaves mostly lanceo-

late or oblong; flowers white or pinkish, in sessile axillary clusters;

fruit pubescent.

43. BORRERIA Mey.

Borreria Anderssonii, sp. nov. — Herba ut videtur annua, cauli-

bus crassiusculis, decumbentibus vel suberectis, subteretibus, pauci-

ramosis, glabris, internodiis elongatis; stipulae in vaginam 4 mm.

longam connatae, vagina truncata brunnescenti glabra apice aristis

pluribus 2-3.5 mm. longis glabris onusta; folia opposita, interdum in

axillis fasciculata, lanceolato-linearia vel anguste oblongo-lanceplata,

2-4.5 cm. longa, 3-8 mm. lata, acuta vel longe attenuato-acuminata,

basin versus angustata et breviter petiolata, membranacea, supra

viridia, dense vel sparse scaberula, subtus pallidiora, ad nervos sparse

scaberula, aliter glabra, costa gracili elevata; flores ad apices ramu-

lorum dense capitati, capitulis multifloris 12-15 mm. latis, foliis

4-6 foliis caulinis conformibus involucratis; folia involucralia basi

246 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

ad vaginam sparse albo-villosa; calycis laciniae 4 fere liberae lineari-

lanceolatae attenuato-acuminatae ciliatae 1.8 mm. longae; corolla

2.5 mm. longa alba extus sparse puberula; capsula 4.5-5 mm. longa

oblonga 2.2 mm. lata sparse villosula. — Ecuador: Puna Island near

Guayaquil, in 1852, N. J. Andersson 71 (Herb. Stockholm, type).

Between Guayaquil and Salinas, border of swamp, Hitchcock 20082

(Y). Near Guayaquil, Mille 92 (Y).

The present plant, which is represented by five sheets of the same

collection in the Stockholm herbarium, is much like Borreria laevis

(Lam.) Griseb. in general appearance, but it differs from that

species in its larger flower heads and much larger capsules. The

coarse stems seem to be somewhat succulent, rather than firm and

stiff, as in B. laevis.

Borreria basalis Anderss. Vet. Akad. Handl. Stockh. 1853: 191.

1854.

Type collected on Chatham Island, Galapagos Islands, by

Andersson.

Plants glabrate, about 30 cm. high, woody, the branches ascending,

glabrate; leaves sessile, linear-lanceolate, mucronate, revolute, gla-

brous, 4-6 mm. long; calyx teeth ovate, longer than the corolla.

Borreria Baurii Rob. & Greenm. Amer. Journ. Sci. 50: 140. 1895.

Known only from the type, collected on Chatham Island, Gala-

pagos Islands, by G. Baur.

Stems woody at the base; leaves ovate to ovate-lanceolate, his-

pid-ciliate, scabrous or smooth above, glabrous beneath, 5-7 mm.

long; flowers 3-4 in each of the upper axils; calyx lobes ovate, acute;

corolla slightly longer than the calyx lobes; fruit hispidulous.

Borreria dispersa Hook. f. Trans. Linn. Soc. 20: 217. 1847.

Reported from Albemarle, Charles, Chatham, Indefatigable, and

James Islands of the Galapagos.

Root woody; stems diffuse, prostrate, dichotomous, acutely tetrag-

onous, shining, glabrous or very minutely puberulent; leaves sub-

sessile, broadly elliptic, mucronate, glabrate above, glabrous beneath,

the margins subrecurved, ciliate; corolla scarcely exceeding the calyx.

Borreria divaricata Hook. f. Trans. Linn. Soc. 20: 219. 1847.

Known only from Charles Island of the Galapagos; collected by

Darwin and Baur.

Stems prostrate (?), woody, the branches terete, divaricately

branched, virgate, remotely leafy, the epidermis pale ashy, the young

branchlets 4-angled, scaberulous; leaves fasciculate, small, oblong,

obtuse, coriaceous, rigid, ciliate, hispidulous on the upper surface;

flowers inconspicuous; calyx lobes acuminate, subciliate; corolla tube

short, barbate with long hairs in the throat; fruit glabrous.

THE RUBIACEAE OF ECUADOR 247

Borreria ericaefolia Hook. f. Trans. Linn. Soc. 20: 218. 1847.

James Bay, James Island, Galapagos, Stewart 3487 (F). Described

from Chatham Island. Reported also from Abingdon, Albemarle,

Charles, Indefatigable, James, Jervis, and Narborough Islands of the

Galapagos.

Glabrous throughout; stems stout, woody, the numerous branches

nearly simple, strict, erect, slender, terete, the young branches

4-angled; leaves small, fasciculate on the short lateral branchlets,

linear-subulate, slightly curved, pungent, rigid-coriaceous, the mar-

gins revolute; fruit glabrous.

Borreria falcifolia Hook. f. Trans. Linn. Soc. 20: 219. 1847.

Known only from the type, collected on Albemarle Island of the

Galapagos group.

Stems terete, stout, woody, the branches elongate, strict, slender,

sparsely branched, the branchlets divaricate, angulate, glabrate;

leaves small, linear, obtuse, subattenuate at the base, scaberulous

above, sulcate along the costa, the margins scaberulous, revolute;

fruit glabrous.

Borreria galapageia Rob. & Greenm. Amer. Journ. Sci. 50: 140.

1895.

Duncan Island of the Galapagos group, the type collected by G.

Baur.

Stems slender, woody, the branches approximate, ascending,

densely pubescent, leafy; leaves ovate-oblong, 4-6 mm. long, 2-3

mm. wide, with fascicles of smaller leaves in their axils, obtuse,

densely grayish-pubescent on both surfaces, the margins strongly

revolute; flowers about 2 in each of the upper axils; calyx lobes acute,

about equaling the corolla, rough-pubescent; fruit rough-pubescent.

Borreria laevis (Lam.) Griseb. Goett. Abh. 7: 231. 1857. Sper-

macoce laevis Lam. 111. 1: 273. 1791.

Without locality, Eggers 15029 (F). La Chonta, Rose, Pachano

& Rose 23471 (G, W, Y). Huigra, 1,500 m., Rose & Rose 22287

(G, W, Y). Bucay, Rose & Rose 22441 (G, Y). Cerro Maglar Alto,

360 m., Anthony & Tate 2 (W). Caraques, Anthony & Tate 82 (W).

Caraques Bay, Anthony & Tate 126 (W). Near Guayaquil, Mille

174(W). Balao, Eggers 1 4095 (L, W) . Loja, October, 1864, Jameson

(S). San Ignacio, Prov. Guayas, Holmgren 17 (S). Widely dis-

tributed as a weed in tropical America.

An annual; leaves petioled, usually more or less pilose beneath;

flowers white, in dense, terminal and axillary clusters; calyx lobes 4,

minute.

Borreria linearifolia Hook. f. Trans. Linn. Soc. 20: 217. 1847.

Type collected on James Island, Galapagos Islands, by Darwin.

Stems elongate, slender, procumbent, divaricately branched, sub-

terete and somewhat woody, the branchlets acutely 4-angled, gla-

248 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

brous, shining; leaves spreading, linear-lanceolate, acute, mucronate,

flat, sparsely hispid-pilose above, the margins recurved, ciliate,

glabrous beneath.

Borreria ocimoides (Burm.) DC. Prodr. 4: 544. 1830. Sperma-

coce ocimoides Burm. Fl. Ind. 34. pi. 13, f. 1. 1768.

Near Ibarra, 2,400 m., Jameson 673 (K), 674 (K). Widely

dispersed in tropical America, and in Asia and Africa.

A slender branched annual, nearly or quite glabrous; flowers

white, minute, densely clustered in the leaf axils; sepals 4; stamens

included; fruit glabrous or puberulent.

Borreria ovalis Anderss. Vet. Akad. Handl. Stockh. 1853: 192.

1854.

Charles Island, Galapagos group, the type collected by Andersson.

Plants 30 cm. high, woody below, the branches glabrate; leaves

sessile, elliptic, spreading, 6-8 mm. long, 4-5 mm. wide, mucronate,

revolute, ciliate-hispidulous, glabrous beneath, coriaceous; calyx

lobes linear, acute; fruit large, hirtous or glabrate.

Borreria ovalis Anderss. f. abingdonensis Robinson, Proc.

Amer. Acad. 38: 205. 1902.

Type collected on Abingdon Island, Galapagos, Baur 142.

Leaves as much as 1.5 cm. long, more or less acute.

Borreria pacifica Rob. & Greenm. Amer. Journ. Sci. 50: 140.

1895.

Indefatigable Island, Galapagos, the type collected south of

Conway Bay by G. Baur.

Woody, the stems terete, the branches numerous, ascending or

widely spreading, mostly simple, scabrous-puberulent, 4-angled;

leaves oblong-lanceolate, sessile, acute, mucronate, scabrous above,

glabrous beneath or nearly so, 7-12 mm. long, with smaller leaves

fascicled in the axils, the margins revolute; flowers 2-3 in an axil;

calyx lobes lanceolate, acute, mucronate, 2-3 mm. long, about equal-

ing the corolla; fruit with scattered grayish pubescence.

Borreria parvifolia Hook. f. Trans. Linn. Soc. 20: 218. 1847.

Type collected on Albemarle Island, Galapagos, by Macrae.

Branches slender, virgate, ramulose, subterete or angulate below,

the branchlets short, leafy; leaves small, fasciculate, coriaceous,

linear-obovate, obtuse, the margins scarcely recurved, hispid above,

glabrate beneath; fruit glabrous.

Borreria perpusilla Hook. f. Trans. Linn. Soc. 20: 218. 1847.

Type collected on James Island of the Galapagos by Darwin.

Plants hispidulous, the root woody, the branches divaricate,

acutely 4-angled; leaves linear-oblong or elliptic-lanceolate, acute,

THE RUBIACEAE OF ECUADOR 249

hispid-pilose above, glabrous beneath, the margins revolute, ciliate;

flowers rather large; corolla campanulate, twice as long as the calyx;

fruit pilose.

Borreria rotundifolia Anderss. Om Galap. Veg. 77. 1857.

Type collected on Indefatigable Island of the Galapagos by

Andersson.

Borreria suaveolens Mey. Prim. Fl. Esseq. 81. pi. 1. 1818.

Spermacoce tenella HBK. Nov. Gen. & Sp. 3: 345. 1819. B. tenella

C. & S. Linnaea 3: 317. 1828.

Hills near Loja, September, 1864, Jameson (S, W) ; October, 1864,

Jameson (S). Perucho, on dry hills, Jameson 646 (P). Widely

distributed in tropical America.

An erect perennial, nearly glabrous, with linear to lanceolate

leaves; flowers white, in dense, chiefly terminal heads; calyx lobes 4.

Borreria suberecta Hook. f. Trans. Linn. Soc. 20: 217. 1847.

Type from Albemarle Island, and reported from Barrington

Island, Galapagos Islands.

Plants hispid-pilose, the stems woody below, the branches striate,

suberect, virgate, acutely 4-angled, leafy; leaves linear, pungent,

spreading or curved, the margins revolute, hispid-pilose on both

surfaces, coriaceous; calyx lobes ovate-lanceolate, acuminate, about

equaling the corolla; fruit pilose.

44. RELBUNIUM Hook. f.

Relbunium ciliatum (R. & P.) Hemsl. Biol. Centr. Amer.

Bot. 2: 62. 1881. Galium ciliatum R. & P. Fl. Peruv. 1: 59. 1798.

G. involucratum HBK. Nov. Gen. & Sp. 3: 335. 1819. Rubia hirta

HBK. Nov. Gen. & Sp. 3: 338. 1819. G. sessiliflorum Willd. ex

Schult. Mant. 3: 179. 1827. Rubia tuberculosa Benth. PI. Hartw.

195. 1845. G. Benthamianum Walp. Repert. Bot. 6: 17. 1861. G.

Kunthii Wedd. Chlor. And. 2: 38. 1857. Relbunium tuberculosum

Ball, Journ. Linn. Soc. 22: 42. 1885.

Antisana, November 3, 1858, Jameson (P). Cuenca, Rose,

Pachano & Rose 22830 (G, W, Y). Urbina, Chimborazo, 3,420 m.,

Anthony & Tate 382 (W). Quitensian Andes, 1855, Couthouy (G).

Guapulo, Prov. Pichincha, 2,500 m., Holmgren & Heilborn 182 (S).

La Magdalena, Prov. Pichincha, 2,800 m., Firmin 372 (F, W), 675

(F). Type of Galium involucratum from Quito, Humboldt & Bon-

pland. Type of R. tuberculosa from Hacienda de Antisana, Hartweg

1 076. Ranging to Colombia and southward to Peru.

Stems elongate, glabrous; leaves oblong to linear-lanceolate, cili-

ate or sometimes eciliate; fruit glabrous.

Relbunium hirsutum (R. & P.) Schum. in Mart. Fl. Bras.

66: 116. 1888. Galium hirsutum R. & P. Fl. Peruv. 1: 59. 1798.

250 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. VII

Without locality, Jameson (F, W). Guanojo, Andre 3968 (F).

Quito, Rose & Rose 23931 (W). Mount Pichincha, October, 1863,

Jameson (W). Andes of Ecuador, Spruce 5215 (G). Near Tulcan,

2,500 m., Hitchcock 20970 (W, Y). Also in Colombia, and extending

southward to Peru.

Stems hirtellous; leaves lance-oblong, acute, hirtellous; fruit

glabrous.

Relbunium hypocarpium (L.) Hemsl. Biol. Centr. Amer. Bot.

2: 63. 1881. Valantia hypocarpia L. Syst. Nat. ed. 10. 1307. 1759.

Riobamba, Sodiro 84-41 (B). La Rinconada, Prov. Carchi,

3,000 m., Hitchcock 20801 in part (W, Y). Andes of Quito, in 1855,

Couthouy (P). Cumbe, Rose, Pachano & Rose 22981 (G, Y). Huigra,

Rose & Rose 22492 (Y). Without locality, Seemann (G). Quitensian

Andes, 1855, Couthouy (G). Banos, Prov. Tungurahua, 1,800 m.,

Heilborn 364 (F, S). La Planta del Chillo, Prov. Pichincha, 2,700

m.,Firmin 696 (F). Reported from the Galapagos Islands. Widely

distributed in tropical America.

Stems pilose; leaves oval or elliptic, rounded to acute at the apex,

usually pilose or hispidulous on both surfaces; flowers greenish yellow,

the peduncles 1-flowered; fruit orange-red, hispidulous or pilose.

Relbunium nitidum (HBK.) Schum. in Mart. Fl. Bras. 66:

114. 1888. Rubia nitida HBK. Nov. Gen. & Sp. 3 : 339. pi. 280. 1819.

Galium quitense Wedd. Chlor. And. 2: 38. 1857.

Type collected at the base of Mount Pichincha, near Quito,

Humboldt &Bonpland (photo, ex Herb. Berol. in F). Weddell (loc.

cit.) reports Jameson 438, from Pichincha at 3,650 m., and a specimen

collected by Remy. Also in Colombia.

Stems pilose; leaves elliptic, obtuse, glabrous above, pilose beneath,

ciliate; fruit fleshy, glabrous, orange or red.

45. GALIUM L.

Galium canescens HBK. Nov. Gen. & Sp. 3: 336. 1819.

Type from Quito, Humboldt & Bonpland (photo, ex Herb. Berol.

in F). Andes of Quito, in 1859, Jameson (G). Guapulo, Prov.

Pichincha, 2,500 m., Holmgren & Heilborn 183 (F, S). Quito, Karsten

(L). Mountains of Quito, Jameson 196 (L). Also in Colombia and

Peru.

Stems pilose; leaves in 4's, ovate, acute or acuminate, hirsute

above, pubescent beneath; flowers solitary, geminate or ternate;

fruit uncinate-hispid.

Galium flaccidum Wedd. Chlor. And. 2: 39. 1857. Rubia

debilis HBK. Nov. Gen. & Sp. 3: 340. 1819, non G. debile Hoffm. &

Link, 1820.

THE RUBIACEAE OF ECUADOR 251

Type collected near Chillo and Falls of Ichubamba, Andes of

Quito, Humboldt & Bonpland.

Stems uncinulate; leaves in 4's, oblong or oblong-lanceolate,

acuminate, the margins and costa retrorse-aculeolate; peduncles

1-flowered; fruit glabrous.

Galium Fraserii Wernham, Journ. Bot. 50: 244. 1912.

Type from Ecuador, Fraser.

Stems minutely uncinulate on the angles; leaves in 4's, 3-nerved,

oval, short-acuminate, revolute, rigid, scabrous; cymes 2-3-flowered;

fruit blackish, fleshy, glabrous.

Galium piliferum HBK. Nov. Gen. & Sp. 3: 337. 1819.

La Rinconada, between Ibarra and Tulcan, Prov. Carchi, 3,000

m., Hitchcock 20801 in part (W, Y). Without locality, Sodiro 44-45

(B). Prov. Pichincha, 3,000 m., Firmin 644 (F). Also in Colombia.

Stems retrorse-hispidulous; leaves in 6's or 8's, linear-lanceolate,

setose-acuminate, retrorse-hispid on the margin and costa; peduncles

3-flowered; fruit uncinate-hispid.

THE LIBRARY OF THE

JUL241943

UNIVERSITY OF ILLINOIS