FIELD MUSEUM OF NATURAL HISTORY

FOUNDED BY MARSHALL FIELD, 1893

PUBLICATION 303

BOTANICAL SERIES VOL. XI, No. 3

THE NYCTAGINACEAE AND

CHENOPODIACEAE OF NORTHWESTERN

SOUTH AMERICA

BY

PAUL C. STANDLEY

ASSOCIATE CURATOR OP THE HERBARIUM, DEPARTMENT OP BOTANY

B. E. DAHLGREN

ACTING CURATOR, DEPARTMENT OF BOTANY

EDITOR

-

CHICAGO, U. S. A.

OCTOBER 20, 1931

FIELD MUSEUM OF NATURAL HISTORY

FOUNDED BY MARSHALL FIELD, 1893

PUBLICATION 303

BOTANICAL SERIES VOL. XI, No. 3

THE NYCTAGINACEAE AND

CHENOPODIACEAE OF NORTHWESTERN

SOUTH AMERICA

BY

PAUL C. STANDLEY

ASSOCIATE CURATOR OP THE HERBARIUM, DEPARTMENT OP BOTANY

irpuiy c?

B.E.DAHLGREN 1931

°r ILLINOIS

ACTING CURATOR. DEPARTMENT OF BOTANY

CHICAGO, U. S. A.

OCTOBER 20, 1931

PRINTED IN THE UNITED STATES OF AMERICA

BT FIELD MUSEUM PRESS

CoW

THE NYCTAGINACEAE OF NORTHWESTERN

SOUTH AMERICA

PAUL C. STANDLEY

In connection with the preparation of an account of the Nycta-

ginaceae of Peru for the flora of that country now almost ready for

publication, it was found that the task would be but little greater

if all the plants of the family occurring in northwestern South

America were treated, hence the present paper. The area covered

embraces Bolivia, Ecuador, Colombia, and Venezuela, but most of

the species known from the Guianas also are included.

This account of the South American Nyctaginaceae makes no

pretension to being a monograph, but is rather a preliminary treat-

ment intended to facilitate the determination of current material

received for study. The collections available are altogether inade-

quate to permit the study of specific variations, in which the family

is notoriously prolific.

I know of few groups of plants in which specific differences are

so unstable and so baffling. In most of the genera, but particularly

in Neea, Torrubia, and Mirabilis, no single character seems to be

constant. The natural tendency, with a relatively small series of

specimens, is to accord specific rank to each fairly well-marked form;

yet long experience with the same family, as it is represented in the

United States, indicates that more ample material will prove that

some of these forms now isolated will be connected by other inter-

grading ones. Because of the vagueness of specific limits in such

genera as Torrubia and Bougainvillea, it has been impracticable to

prepare satisfactory keys for separating the species, and it is doubtful

whether it ever will be possible to compile readily usable keys for

them.

The only important genus of the family, as it is represented

in South America, whose species may be recognized by really clear-

cut characters, is Boerhaavia. In Mexico and the southwestern

United States, however, the species of the same genus intergrade

in quite as tantalizing a manner as those of Mirabilis.

There is little reason for expecting that the list of species published

here will be greatly increased by further exploration, because most

of the species already known are rather widely distributed. Very

73

74 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

extensive collections made recently in Peru have contained only

two clearly valid new species of the family. A few additional species

in Torrubia and Neea may be discovered in regions still unexplored.

The study whose results are presented on the following pages

is based primarily on the collections in the Herbarium of Field

Museum (indicated by the letter F in parenthesis). These are rich

in photographs or fragments of types and otherwise authentic

material from the Berlin herbarium, the photographs having been

obtained by Assistant Curator J. Francis Macbride, with the assist-

ance of a fund provided for the purpose by the Rockefeller Founda-

tion. The material includes some representation of almost every

species reported from South America, and it has been possible to

consider most of the species described from Brazil, Paraguay, and

Argentina.

The work has been greatly facilitated by the loan of all the

material of the Nyctaginaceae from the area under consideration in

the United States National Museum (W). To Dr. William R.

Maxon and Mr. Ellsworth P. Killip the writer is deeply indebted

for the opportunity of examining this collection, which is particularly

valuable for its numerous Venezuelan specimens.

KEY TO THE GENERA

Embryo straight; flowers mostly unisexual; trees or shrubs, some-

times armed with spines; leaves all or mostly opposite or

verticillate (Pisonieae).

Stamens included; anthocarp without stipitate glands.

Stamens 5-10; limb of the staminate perianth little or not at

all broader than the tube 1. Neea.

Stamens 25-30; limb of the staminate perianth much broader

than the tube 2. Cephalotomandra.

Stamens exserted.

Anthocarp fleshy, juicy, without glands; flowers cymose-panic-

ulate, unisexual; stems unarmed 3. Torrubia.

Anthocarp dry, coriaceous, bearing numerous stipitate glands.

Flowers unisexual, in cymes; stigma penicillate; stems often

armed with spines 4. Pisonia.

Flowers perfect, in umbels; stigma depressed-capitate; stems

unarmed 5. Pisoniella.

Embryo curved; flowers perfect; chiefly herbaceous plants, sometimes

woody and then often scandent; leaves opposite or alternate.

NYCTAGINACEAE OF SOUTH AMERICA 75

Leaves alternate.

Perianth unchanged in fruit; flowers free from the bracts; plants

herbaceous or nearly so (Salpiantheae) 6. Salpianthus.

Perianth much altered in fruit, the lower part enlarged and

adherent to the fruit; flowers borne in clusters of 3, each

flower adnate to a large colored bract; plants scandent or

erect, woody, usually armed with spines (Bougainmlleae).

7. Bougainvillea.

Leaves opposite.

Perianth lobes valvate. Plants large, sometimes scandent, the

leaves often verticillate; flowers minute, umbellate, not

involucrate (Colignonieae) 8. Colignonia.

Perianth lobes induplicate- valvate (Mirabileae).

Anthocarp lenticular, with recurved, usually dentate margins;

flowers in 3's, subtended by a 3-parted involucre.

9. Allionia.

Anthocarp terete or angled, never margined.

Flowers subtended by distinct bracts 10. Boerhaavia.

Flowers subtended by a gamophyllous involucre.

11. Mirabilis.

1. NEEA R. & P.

Shrubs or trees; leaves opposite or verticillate, rarely alternate,

petiolate, entire, coriaceous to membranaceous; flowers unisexual

and usually dioecious, small, white, reddish, or green, sessile or

pedicellate, commonly tribracteolate at the base, arranged in lateral,

axillary, or terminal cymes, rarely solitary; staminate perianth

urceolate, globose, or tubular, shortly 4-5-dentate; stamens 5-10,

included, the filaments unequal; pistillate perianth urceolate or

tubular, constricted above the ovary, often contracted at the mouth;

stigma penicillate; anthocarp narrowly or broadly ellipsoid, crowned

by the persistent free portion of the perianth.

In this, the largest South American genus of the family, the

species are poorly understood, largely because of the lack of ample

material. Of some species only staminate branches are known, and

of others only pistillate specimens. In most genera of the Nyctagina-

ceae the species are poorly marked, but in Neea it is very difficult

indeed to differentiate them satisfactorily. Only a few species are

listed here, but a large number have been described from the Amazon

Valley of Peru and Brazil. It is probable that many more species

are to be found in the wet forests of Bolivia and Colombia, for the

76 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

parts of those countries in which they may be expected still are

practically unknown botanically. It is noteworthy that no species

of Neea is reported from Ecuador, although several must occur there,

at least on the eastern slopes of the Andes.

Leaves rounded at the apex, firm-coriaceous, closely reddish-tomen-

tulose beneath 1. N. obovata.

Leaves acute or acuminate, comparatively thin.

Leaves permanently tomentose or pilose beneath.

Leaves small, 2.5-4.5 cm. wide. Staminate inflorescence few-

flowered, the peduncle long and slender .... 2. N. Bangii.

Leaves large, 6-15 cm. wide.

Petioles short, 2 cm. long or less 3. N. Woronovii.

Petioles elongate, 4-4.5 cm. long 4. N. mapirensis.

Leaves glabrous beneath at maturity or nearly so.

Inflorescence short-pedunculate, dense, few-flowered, erect.

Leaves green when dried, 6.5-9 cm. wide.

5. N. dimorphophylla.

Leaves fuscous when dried.

Inflorescence small, commonly 2-5.5 cm. broad.

Inflorescence densely ferruginous-puberulent or tomen-

tulose 6. N. divaricata.

Inflorescence sparsely ferruginous-puberulent.

7. N. constrictoides.

Inflorescence large, 10 cm. broad 8. N. Brittonii.

Inflorescence long-pedunculate, lax, usually many-flowered, com-

monly pendent.

Leaves fuscous when dried; inflorescence few-flowered.

9. N. longipedunculata.

Leaves green when dried; inflorescence many-flowered.

Staminate flowers tubular, more than twice as long as broad.

10. N. boliviano,.

Staminate flowers urceolate, less than twice as long as broad.

11. N. anisophylla.

The key here presented for the separation of the species is a most

unsatisfactory one, chiefly on account of the absence of complete

material, and because no specimens at all are available for two of the

species. Those reported from Bolivia seem to be distinct enough

from one another, and it is to be presumed that they are distinct from

N. constrictoides, on geographic grounds if for no other reason.

NYCTAGINACEAE OF SOUTH AMERICA 77

Except in one instance, the Bolivian specimens seem to be different

from the numerous species known from Peru.

1. Neea obovata Spruce ex Heimerl, Beitr. Syst. Nyctag. 38.

1897.

Young branchlets densely reddish-puberulent; leaves opposite,

firmly coriaceous, the stout petioles 5-25 mm. long, very densely

puberulent; leaf blades obovate or elliptic-obovate, 6-11 cm. long,

3-6.5 cm. wide, broadly rounded at the apex, acute or acutish at

the base, when young very densely reddish-puberulent or strigose

on both surfaces but in age glabrate, at least above; inflorescence

erect, pedunculate, open, corymbiform or paniculate, many-flowered,

the branches spreading or ascending, the flowers congested at the

ends of the branches; perianth subcampanulate, 3 mm. long, reddish-

tomentulose or finally glabrate; stamens 6.

Venezuela: San Carlos on the Rio Negro, 1853-54, Spruce 3128

(F, photo, of type ex herb. Berol.). Ad flum. Guainia vel Rio Negro

supra ostiam fluminis Casiquiare, in 1854, Spruce (W).

The species should be enumerated also for the flora of Colombia,

since the type locality is on the boundary between that country and

Colombia. This happens to be one of the few easily recognizable

and well-marked species of the genus, because of its thick leaves,

broadly rounded at the apex, and covered beneath with a minute,

bright reddish tomentum.

2. Neea Bangii Rusby, Bull. N. Y. Bot. Gard. 4: 435. 1907.

A tree 4.5-6 m. high, the young branchlets ferruginous-tomen-

tulose; leaves opposite or ternate, unequal, rather thin, blackish

when dried, the slender petioles 3-8 mm. long, tomentulose or hirtu-

lous; leaf blades elliptic-oblong or mostly oblong-obovate, 6-11 cm.

long, 2.5-4.5 cm. wide, abruptly acute or short-acuminate with

obtuse tip, slightly lustrous above and minutely appressed-pilosulous

or glabrate, beneath sparsely and persistently pilosulous with lax

spreading brownish hairs; pistillate cymes few-flowered, 2-4 cm.

broad, on very slender, sparsely villosulous peduncles 3-5 cm.

long, probably pendent, lax, the bractlets triangular-subulate, 1 mm.

long; perianth yellowish, urceolate, 4 mm. long, sparsely ferruginous-

tomentulose, especially near the base; fruit black, oval, nearly 15

mm. long and half as thick.

Bolivia: At the foot of Mount Uchimachi, Calapampa, July,

1894, Bang 2346 (F, W, type collection). Without locality, Bang

2306 (W). Tumapasa, Williams 585 (W).

3. Neea Woronovii, sp. nov. — Ramuli crassiusculi subdense

adpresse ferrugineo-tomentulosi, internodiis usque ad 10 cm. longis;

folia opposita vel ternata inaequalia tenuiter coriacea, petiolo cras-

siusculo 1-1.6 cm. longo dense ferrugineo-tomentuloso; lamina ob-

78 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

longa vel obovato-oblonga 13-25 cm. longa 5.5-8 cm. lata abrupte

acuminata, basi acuta vel attenuata, supra lucida, in sicco fusca,

glabra vel glabrata, costa prominula, nervis vix elevatis, subtus

paullo pallidior, ubique minute ferrugineo-hirtula, ad nervos breviter

villosula, costa gracili elevata, nervis lateralibus utroque latere c.

11, aliis paullo tenuioribus interjectis, gracilibus, prpminentibus,

arcuatis, remote a margine conjunctis, nervulis prominulis paucis

laxissime reticulatis; inflorescentia feminea axillaris magna laxissime

multiflora c. 13 cm. longa et 15 cm. lata, recurva vel pendula, c. 6.5

cm. longe pedunculata, pedunculo ut rami densissime ferrugineo-

tomentuloso, ramis gracilibus alternis oppositis vel subverticillatis

supra late dichotomis, floribus ad apices ramulorum dense congestis

sessilibus; bracteolae subulatae 1.5-2 mm. longae dense ferrugineo-

tomentulosae; perianthium valde juvenile c. 2 mm. longum ovato-

oblongum acutiusculum densissime tomentulosum. — Colombia: Chin-

gale, Dept. Santander, April 18, 1926, G. Woronow and S. Juzepczuk

4320 (Herb. Field Mus. No. 605,666, type).

The plant is noteworthy for its long, large, and copiously pubescent

leaves, and for the very large and many-flowered inflorescence,

covered with a dense reddish tomentum.

4. Neea mapirensis, sp. nov. — Ramuli puberuli vel glabrati;

folia opposita, petiolo crassiusculo 4-4.5 cm. longo hirtulo vel gla-

brato; lamina crasse membranacea magna anguste elliptico-oblonga

c. 36 cm. longa et 13-15 cm. lata, acuminata, basi acuta vel obtusius-

cula, in sicco subfusca, supra glabra vel glabrata, sublucida, costa

venisque parum prominulis, subtus paullo pallidior, ubique subdense

hirtula vel asperulo-puberula, costa crassiuscula prominente, nervis

lateralibus utroque latere c. 13 gracilibus prominentibus subarcuatis

angulo c. semirecto adscendentibus remote a margine conjunctis,

nervulis paucis laxissime reticulatis parum conspicuis; inflorescentia

feminea cymosa breviter pedunculata pauciflora c. 5 cm. longa et

ut videtur duplo latior, ramis incrassatis glabratis, floribus sessilibus

vel crasse pedicellatis, bracteolis deciduis; anthocarpium oblongum

glabrum 1 cm. longum 5 mm. latum basi obtusum apice calyce

persistente minuto coronatum. — Bolivia: San Antonio near Mapiri,

alt. 850 m., December, 1907, Otto Buchtien 1993 (U. S. Nat. Herb.

No. 1,398,045, type).

Although the plant is known only from too fragmentary material

— two leaves and a detached inflorescence — it seems clearly distinct

from all other species of the genus, especially in its very large leaves

with copious pubescence. The leaves are somewhat broader than

those of N. Woronovii, which differs also in having slender branches

in the inflorescence.

5. Neea dimorphophylla, sp. nov. — Ramuli graciles subteretes

viridescentes, novellis sparse hirsutis cito glabratis, internodiis brevi-

bus vel elongatis; folia ternata vel quaterna, interdum opposita,

NYCTAGINACEAE OF SOUTH AMERICA 79

valde inaequalia, petiolo 5-10 mm. longo gracili hirtello vel glabrato;

lamina crasse membranacea, foliorum majorum oblongo-elliptica

vel ovata, rare obovata, prope medium latissima, 11-21 cm. longa,

5-9 cm. lata, abrupte acuta vel acuminata, basi acuta vel abrupte

contracta et valde inaequalis, in sicco olivaceo-viridis, glabra, supra

vix lucida, costa prominente, venis prominulis et plus minusve

reticulatis, subtus paullo pallidior, interdum ad costam prominentem

sparse pilosa, nervis lateralibus utroque latere c. 10 gracilibus pro-

minentibus angulq lato adscendentibus prope marginem conjunctis,

nervulis prominulis et arete reticulatis; lamina foliorum minorum

rotundata vel ovato-rotundata 3.5-6.5 cm. longa et aequilata, apice

rotundata et abrupte apiculata, basi rotundata; inflorescentiae

femineae axillares et terminates crasse 1.5-3 cm. longe pedunculatae

erectae sublaxe pauciflorae, ramis crassiusculis sparse puberulis et

pilosis suboppositis, floribus sessilibus et splitariis vel pedicellis valde

incrassatis stipatis, bracteolis persistentibus subulatis vix 1 mm.

longis sparse pilosulis; anthocarpium oblongum 8-9 mm. longum

4 mm. latum glabrum basin versus obtusum paullo angustatum,

apice parte superiore calycis persistente coronatum; stylus breviter

exsertus, stigmate penicillato. — Bolivia: Junction of the rivers Beni

and Madre de Dios, August, 1886, H. H. Rusby 2575 (Herb. Field

Mus. No. 164,490, type; duplicate in U. S. Nat. Herb.).

In its foliage the Bolivian plant resembles Neea laxa Poepp. &

Endl., of Peru, but the latter has a conspicuously different inflorescence.

6. Neea divaricata Poepp. & Endl. Nov. Gen. & Sp. 2: 45. 1838.

A medium-sized tree with dark green foliage, the branchlets

glabrous or sparsely puberulent when young; leaves chiefly opposite,

short-petiolate, subcoriaceous, oblong to oblong-elliptic, mostly

8-16 cm. long and 3-7 cm. wide, fuscous when dried, abruptly acute

or short-acuminate with obtuse tip, glabrous or nearly so; inflores-

cences cymose-paniculate, erect, short-pedunculate, usually small

and dense, with opposite or alternate branches, the branches sparsely

or densely puberulent or glabrate; bractlets lance-subulate, 1-1.5

mm. long, puberulent or glabrate; flowers green, congested; pistil-

late perianth oblong-urceolate, 2.5-3 mm. long, sparsely ferruginous-

puberulent, the limb minutely 5-dentate; stamens 5; anthocarp

oblong, glabrous, 8-10 mm. long, obtuse at the base.

Bolivia: Forests of Buenavista, Dept. Santa Cruz, 450 m.,

Steinbach 6443 (F). Also in Peru.

There is in the Herbarium of Field Museum a photograph and

fragment of Buchtien 1765 from Bolivia, received from the Berlin

herbarium, which, so far as I can tell from the available material,

may well be referable to this species, although it is indicated by

Heimerl as the type of a new species.

Steinbach gives the vernacular name as "ajillo," and states that

ashes obtained by burning the wood are employed for making lye

that is used in the preparation of soap.

80 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

7. Neea constrictoides Heimerl, Beitr. Syst. Nyctag. 37. 1897.

Branchlets glabrous; leaves short-petiolate, the petiole 8 mm.

long, the blades oblong-elliptic, 10-14 cm. long, 4-5.5 cm. wide,

acuminate, with obtuse tip, cuneate at the base or attenuate to the

petiole, glabrous, slightly lustrous above; inflorescences pedunculate,

laxly flowered, broadly pyramidal, the branches opposite, sparsely

ferruginous-puberulent, the flowers aggregate; pistillate perianth

subcylindric, 6.5 mm. long, the teeth minute, acutish; immature

fruit oblong-ellipsoid.

Colombia: Villavicencio, Bogota, Karsten.

The plant is known to the writer only from the original description.

8. Neea Brittonii, nom. nov. Neea macrophylla Britton ex

Rusby, Bull. Torrey Club 27: 126. 1900, non Poepp. & Endl. 1838.

Glabrous, drying blackish; petioles very stout, 1-2 cm. long;

leaf blades oval, 15-20 cm. long, 7.5-10 cm. wide, short-acuminate,

with acute tip, abruptly narrowed to the petiole, membranaceous,

the veins 12-16 pairs; cymes 10 cm. broad, on a short stout peduncle,

the branches minutely ferruginous-puberulent, stout, the flowers

subsessile, rather densely clustered; bractlets lance-linear, 1-2 mm.

long; staminate perianth almost 1 cm. long, 3-4 mm. broad, the 5

lobes 1 mm. long, ovate, obtuse; stamens 8, the longest less than

half as long as the perianth.

Bolivia: Type collected at the junction of the rivers Beni and

Madre de Dios, August, 1886, H. H. Rusby 2119. A specimen

collected by Pearce at Monterrico also is referred by Britton to the

species, of which I have seen no material.

9. Neea longipedunculata Britton ex Rusby, Bull. Torrey

Club 27 r 126. 1900.

A shrub or small tree 4 m. high with slender branches, the young

branchlets ferruginous-tomentose, soon glabrate; petioles slender,

3-14 mm. long, sparsely pilose or glabrous; leaf blades fuscous when

dried, thick-membranaceous, elliptic-oblong or ovate-oblong, broad-

est at or below the middle, 8-15 cm. long, 3.5-6 cm. wide, acuminate

or long-acuminate, obtuse or narrowly rounded at the base, glabrous,

the veins about 13 pairs; pistillate inflorescences small and few-

flowered, on filiform pendent glabrous peduncles 5-15 cm. long,

2-6 cm. broad, lax, the slender branches densely ferruginous-tomen-

tulose; bractlets subulate, 1 mm. long or less, ferruginous-tomentulose ;

perianth tubular, 1 cm. long, sparsely tomentulose or glabrate, 3 mm.

wide, the lobes 2 mm. long, subrecurved.

Bolivia: Reis, 450 m., June, 1886, Rusby 2709 (F, W, type collec-

tion). Charopampa, near Mapiri, edge of forest, 570 m., Buchtien

1627 (W).

The original description is erroneous in several particulars,

especially in describing the leaves as obovate.

NYCTAGINACEAE OF SOUTH AMERICA 81

10. Neea boliyiana, sp. noy. — Ramuli crassi teretes ochracei,

internodiis elongatis glabris; folia opposita magna subcoriacea in

sicco laete viridia, petiolo 12-18 mm. longp glabrp; lamina oblongo-

elliptica vel obovato-elliptica, prope medium latissima, 15-26 cm.

longa, 7.5-12 cm. lata, subabrupte acuminata vel acuta, acumine

ipso obtuso, basi acutiuscula vel abrupte breviterque contracta,

glabra, supra opaca, costa prominula, venis inconspicuis, subtus

parum pallidior, costa gracili elevata, nervis lateralibus utroque

latere c. 13 angulo fere recto abeuntibus leviter arcuatis gracilibus

prominulis, nervulis vix prominulis paucis laxissime reticulatis;

inflorescentiae masculae terminales et axillares 4 cm. longe peduncu-

latae cymosp-paniculatae, c. 8 cm. longae et aequilatae vel latiores,

ramis alternis vel suboppositis crassiusculis sparse minute tomen-

tulosis vel glabratis, floribus sessilibus vel brevissime crasse pedicel-

latis aggregatis, bracteolis ovato-pblongis dense ferrugineo-tomen-

tulosis vix 1 mm. longis; perianthium tubulosum 7-8 mm. longum

sparse ferrugineo-puberulum vel glabratum versus basin sensim

angustatum sub orem paullo contractum et 2.5 mm. latum, dentibus

minutis late triangularibus vix 1 mm. longis erectis. — Bolivia:

Antahuacana, valley of the Rio Espiritu Santo about 160 km.

northeast of Cochabamba, alt. 750 m., June, 1909, Otto Buchtien

4732 (U. S. Nat. Herb. No. 1,398,053, type).

Well marked by the large and broad, bright green, glabrous

leaves, and by the ample, many-flowered inflorescence.

11. Neea anisophylla Ernst, Flora 57: 215. 1874. N. Wiesneri

Heimerl, Bot. Jahrb. Engl. 11: 89. pi. 2, f. 5. 1889.

A shrub or small tree, usually 3 m. high or less, glabrous through-

out or nearly so; leaves thick-membranaceous, opposite, bright green

when dried, short-petiolate, the slender or stout petiole 2-10 mm.

long; leaf blades elliptic-lanceolate or oblong-lanceolate, sometimes

ovate-lanceolate, 6-23 cm. long, 2.5-9 cm. wide, usually acuminate

or long-acuminate, often merely acute, usually strongly oblique and

acute at the base; inflorescences very lax, many-flowered, on long

slender filiform peduncles, pendent, corymbiform, the flowers pedicel-

late or sessile and glomerate, the bractlets minute; staminate perianth

broadly urceolate, yellow, yellowish white, or green, 5-5.5 mm. long,

glabrous or nearly so; stamens 6-8; pistillate perianth ovoid-tubular,

4.5-5 mm. long; anthocarp ellipsoid, glabrous, 6-7 mm. long.

Venezuela: Type from Catuche, Caracas, Ernst. Chacaito Gorge,

near Caracas, 800-1,000 m., in forest, Pittier 9476 (W). La Guaira,

Moritz 1464 (F, photo, and fragm. ex herb. Berol.; W). Bosque de

Catuche, above Caracas, 1,200-1,800 m., Pittier 6298 (F, W).

Curucuti, between Caracas and La Guaira, 1,100-1,700 m., Pittier

10390 (W), 10391 (W). Hacienda Taborda, near El Palito, Cara-

bobo, Pittier 7667 (W) . Pico Palpma, Eggers 13520 (W) .—Colombia :

Reported by Heimerl from the vicinity of Bogota, Karsten.

82 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

Pittier 11950 from the upper Carrasquel Valley probably repre-

sents an undescribed species. The single specimen that I have seen

is unsatisfactory for purposes of description.

2. CEPHALOTOMANDRA Karst. & Triana

Unarmed trees or shrubs; leaves opposite, petiolate, entire;

flowers dioecious, small, yellowish or reddish, tribracteolate, pedi-

cellate, arranged in terminal long-pedunculate many-flowered cymes;

staminate perianth urceolate-campanulate, abruptly constricted

below the middle, the limb shallowly 5-lobate; stamens numerous,

about 25-30, included, the filaments stout, unequal; pistillate

perianth with a persistent 5-lobed limb; anthocarp woody, oblong,

costate.

One other species of the genus has been described from Panama.

1. Cephalotomandra fragrans Karst. & Triana ex Karst.

Linnaea 28: 430. 1856. Pisonia Cephalotomandra Heimerl, Beitr.

Syst. Nyctag. 34. 1897.

A tree 7.5 m. high; leaves slender-petiolate, the blades elliptic-

oblong to elliptic, mostly 8-16 cm. long and 4.5-9 cm. wide, acute

or obtuse, rounded to acutish at the base, glabrous, subcoriaceous;

inflorescences lax, many-flowered, about 10 cm. broad, the slender

branches minutely fulvous-puberulent; bractlets lance-subulate, less

than 1 mm. long; staminate perianth yellow, 3 mm. long, glabrate

or obscurely puberulent; anthocarp about 13 mm. long (even larger

according to description), 10-12-costate.

Colombia: Western declivity of the mountains of Bogota, at

400-1,200 m., Triana 998 (F; photo, and fragm. of type ex herb.

Berol.).

In the original description it is stated that the fruit is as large

as an English walnut, but this seems altogether improbable, in con-

sideration of the size of the single dried one that I have seen.

3. TORRUBIA Veil.

Unarmed shrubs or trees, glabrous or pubescent; leaves chiefly

opposite and petiolate, entire, membranaceous to coriaceous; flowers

small, dioecious, reddish, greenish, or yellowish, exinvolucrate, 2-3-

bracteolate, sessile or pedicellate, arranged in lateral or terminal,

pedunculate cymes; staminate perianth obconic-campanulate, the

limb 5-dentate; stamens 6-10, exserted, the filaments unequal;

pistillate perianth tubular; anthocarp drupaceous, red to black,

oblong, the exocarp fleshy and juicy.

By most botanists the plants comprising this genus have been

referred to Pisonia, but there are excellent and basically important

characters that distinguish the two groups, consequently it is illogical

and unreasonable to combine them under Pisonia. Although some

NYCTAGINACEAE OF SOUTH AMERICA 83

of the species are not well marked, they are much more clearly differ-

entiated than in the genus Neea. Only one species of Torrubia is

known from Peru.

Inflorescences globose and very dense, headlike. Leaves small,

rounded to subacute at the apex 1. T. ferruginea.

Inflorescences open, sometimes small but distinctly branched.

Leaves densely or sparsely pubescent beneath over the whole sur-

face when young, in age sometimes glabrate except along the

costa.

Leaves rounded or very obtuse at the apex. . . .2. T. floribunda.

Leaves chiefly acute or acuminate.

Branchlets and lower surface of the leaves with a fine close fer-

ruginous tomentum; leaves often glabrate beneath in age.

Inflorescences small and few-flowered, dense; leaves chiefly

acute to attenuate at the base 11. T. salici/olia.

Inflorescences larger, many-flowered and open; leaves mostly

obtuse at the base 3. T. Rusbyana.

Branchlets pilose with spreading hairs.

Leaves chiefly obovate, broadest above the middle.

4. T. suspense/,.

Leaves oblong to elliptic, broadest at or below the middle.

Leaves densely soft-pilose beneath 5. T. pubescens.

Leaves glabrate in age except beneath along the costa.

6. T. boliviano,.

Leaves glabrous beneath or nearly so, even when young.

Leaf blades small, 3 cm. long or less, rounded or very obtuse

at the apex 7. T. microphylla.

Leaf blades much larger, mostly acute or acuminate.

Leaves bright green when dried ; perianth glabrous or nearly so.

8. T. pacurero.

Leaves fuscous when dried; perianth often puberulent or

tomentulose.

Staminate perianth 5-5.5 mm. long, densely rufous-tomen-

tulose; leaves large, mostly 5.5 cm. wide or more.

9. T. uberrima.

Staminate perianth 3.5- 4 mm. long; leaves usually narrower.

Veins of the leaves numerous and close together, con-

spicuous, ascending at a very acute angle; leaves

normally broadest above the middle . .10. T.fragrans.

84 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

Veins few, distant, divergent at a comparatively broad

angle; leaves mostly broadest at or near the middle.

Inflorescence densely rufous-tomentulose, small and

few-flowered 11. T. salicifolia.

Inflorescence glabrate or with sparse short hairs.

12. T.Olfersiana.

1. Torrubia ferruginea (Klotzsch) Standl. Contr. U. S. Nat.

Herb. 18: 100. 1916. Pisonia ferruginea Klotzsch ex Choisy in DC.

Prodr. 132: 445. 1849. T. cephalantha Standl. ex Pittier, PI. Usual.

Venez. 177. 1926.

A deciduous shrub or tree 3-10 m. high, the young branchlets

densely rufous-tomentulose; leaves thick-membranaceous or sub-

coriaceous, small, opposite or verticillate, unequal, fuscous when

dried, the petioles short, rufous-tomentulose; leaf blades oblong-

elliptic to rounded, mostly 2-5.5 cm. long, rounded to acutish at

the apex, acute to rounded at the base, when young densely rufous-

hirtulous beneath but in age usually glabrate; flowers in very dense,

spherical heads about 1.5 cm. in diameter, densely ferruginous-

tomentose, sessile; staminate perianth yellow, 3.5 mm. long; antho-

carp oblong, 7 mm. long, sparsely tomentulose or nearly glabrous.

Venezuela: Without locality, Karsten 148 (F; photo, and fragm.

ex herb. Berol.; presumably the type collection, which was reported

as from Colombia, probably in error). Colonia Tovar, Fendler 1314

(F). Between Caracas and La Guaira, Rose 21829 (W). Zigzag,

November 2, 1916, Rose (W). El Valle, Eggers 13162 (W). Barquisi-

meto, Lara, "not common," Saer 5 (W). Bare hills north of Barqui-

simeto, Saer 168 (W). Above Las Ruinas, near Caracas, on grassy

slopes, 1,000 m., Pittier 9462 (W). Bosque de Catuche, above

Caracas, 1,200-1,800 m., Pittier 6192 (F, W). Lower Catuche wood

above Caracas, 1,000-1,200 m., Pittier 7136 (W). Guaremales,

Carabobo, 10-100 m., Pittier 8837 (W); in forest, Pittier 8857 (W).

Hacienda Panarigua, Valley of Puerto La Cruz, on road to Cagiiita,

dry wooded hills, Pittier 9210 (W).

Vernacular name, "cazabito." Regarding the name, Pittier (loc.

cit.) makes the following statement: Various trees and shrubs of

the family Nyctaginaceae, such as Torrubia Olfersiana and Neea

anisophylla, are given this name, which refers to the characteristic

structure of the wood, reminding one at a distance of dry yuca roots

or cassava bread. This wood, which is light in weight, porous, and

useless, is composed of numerous vascular bundles, derived from the

cambium, which are collected about the original bundles and become

atrophied and dried, leaving in their place numerous pores. Among

the fibers thus formed there are seen many canals and cells that

contain crystals of sodium oxalate.

NYCTAGINACEAE OF SOUTH AMERICA 85

2. Torrubia floribunda (Hook, f.), comb. nov. Pisonia flori-

bunda Hook. f. Trans. Linn. Soc. 20: 193. 1847.

A tree with rough bark and broadly spreading crown, the branch-

lets pale, the young ones densely fulvous-puberulent; leaves short-

petiolate, thick-membranaceous, mostly clustered on short lateral

spurs, the blades oval to elliptic or rounded, about 3 cm. long,

rounded or obtuse at the apex, acutish at the base, hirtulous or

puberulent on both surfaces, more densely so beneath; staminate

inflorescences small and rather dense, many-flowered, short-pedun-

culate, densely puberulent; perianth broadly obcpnic, 3 mm. long,

sparsely pale-puberulent; young anthocarp ellipsoid-oblong, densely

pale-tomentose, costate, 5 mm. long.

Ecuador (Galapagos Islands) : Type from James Island, Darwin.

Duncan Island, at 335 m., Stewart 1462 (W). James Bay, James

Island, 135-500 m., Stewart 1464 (W). Reported also from Abingdon,

Albemarle, Charles, and Indefatigable Islands.

The tree is said to be common on many of the islands, often

forming dense forest or thickets. Sometimes it grows as a low shrub.

3. Torrubia Rusbyana (Heimerl), sp. nov. Pisonia Rusbyana

Heimerl in herb. — Frutex vel arbor, ramulis crassiusculis subteretibus

griseis vel ochraceis, novellis dense minute ferrugineo-puberulis,

internodiis brevibus vel elongatis; folia petiolata crasse membrana-

cea in sicco fuscescentia opposita vel ternata, petiolo gracili 3^40

mm. longo puberulo vel hirtulo; lamina lanceolato-oblonga, elliptico-

oblonga vel elliptica, fr-14 cm. longa, 2-6.5 cm. lata, sensim vel

abrupte acuta vel acuminata, acumine acuto vel obtuso, basi obtusa

vel acutiuscula et saepe obliqua, juvenilis utrinque minute rufo-

totentulosa, adulta glabrata vel subtus sparse hirtula, supra ad

costam prominulam sparse hirtula, opaca, venis obscuris, subtus

concolor, costa gracili elevata, nervis lateralibus utroque latere

c. 9 gracilibus prominulis angulo latiusculo adscendentibus irregulari-

bus remote a margine obscure conjunctis, nervulis obsoletis; inflores-

centia mascula terminalis 1-5.5 cm. longe pedunculata multiflora

sublaxe ramosa 3.5-6 cm. lata erecta, ramulis dense ferrugineo-

puberulis, floribus sessilibus vel brevissime pedicellatis, bracteolis

vix 0.5 mm. longis ovato-lanceolatis; perianthium obconicum 3.5-4

mm. longum basi acutum sparse minute puberulum vel fere glabrum;

stamina c. 8 perianthio duplo longiora; anthocarpium ellipsoideo-

oblongum 7 mm. longum ferrugineo-tomentellum apicem versus

paullo angustatum. — Venezuela: Lower Orinoco, in 1896, H. H.

Rusby and Roy W. Squires 422 (Herb. Field Mus. No. 161,043,

type; duplicate in U. S. Nat. Herb.). Hacienda Puerto La Cruz,

Coastal Range, in 1918, Pittier 8091 (W).

The fruiting specimen collected by Pittier has longer petioles than

the type and a more closely appressed pubescence, but it appears to

represent the same species as the material from the Orinoco.

86 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

4. Torrubia suspensa (Heimerl) Standl. Contr. U. S. Nat.

Herb. 18: 101. 1916. Pisonia suspensa, Heimerl, Med. Rijks Herb.

19: 34. 1913.

A tree with broadly drooping branches, the branchlets densely

rufous-pubescent; petioles 4-8 mm. long; leaf blades broadly elliptic

to obovate-elliptic, 3.5-8.5 cm. long, 2-3 cm. wide, acutish at the

base, short-acuminate, papyraceous, brownish when dried, sparsely

hirtous above, densely subhirsute beneath, the lateral nerves 6-9

pairs; pistillate inflorescence borne on a peduncle 3-5 cm. long, up

to 4.5 cm. wide, lax, many-flowered, umbellately branched at the

base, the flowers subsessile or shortly pedicellate; perianth 3-3.5

mm. long, more or less hirtous.

Bolivia: In forest between Guaridi and Rio Grande, 600 m.,

December, 1910, Herzog 1285 (F, photo, of type ex herb. Berol.).

5. Torrubia pubescens (HBK.), comb. nov. Pisonia pubescens

HBK. Nov. Gen. & Sp. 2: 218. 1818.

A shrub or tree, sometimes 5.5 m. high, the young branchlets

densely tomentose or short-villous; leaves opposite or quaternate,

often very unequal, membranaceous, fuscous when dried, the petioles

seldom more than 5 mm. long; leaf blades oblong to elliptic, mostly

5-9 cm. long and 2-4.5 cm. wide, acute or short-acuminate, at the

base acute or obtuse and more or less oblique, copiously pilose with

weak fulvous hairs on both surfaces, more densely so beneath, some-

what paler beneath; inflorescences chiefly in the forks of the branches,

erect, slender-pedunculate, few-flowered and rather lax, the flowers

pedicellate; staminate perianth yellowish, obconic, acute at the base,

4 mm. long, sparsely and minutely puberulent or glabrate; stamens

8; pistillate inflorescence and fruit unknown.

Venezuela: Type from banks of the Rio Apures, Humboldt &

Bonpland. Without data, Humboldt (F; photo, of specimen in herb.

Willd. ex herb. Berol.; perhaps the type). In hedges, Llanos de La

Rubiera, Guarico, Pittier 12333 (F, B, W). Culebra Lagoon near

San Carlos, Cojedes, in savanna bushes, Pittier 11704 (W).

Heimerl has indicated Pittier 12333 in the Berlin herbarium as a

new species, but this collection agrees perfectly with the original

description of Pisonia pubescens, and I see no reason for distinguish-

ing it. Pittier describes No. 11704 as a vine with green flowers.

Probably there is some mistake in ascribing a scandent habit to the

plant, or else it merely has long branches that are more or less

supported on other shrubs.

6. Torrubia boliviana (Britton) Standl. Contr. U. S. Nat.

Herb. 18: 100. 1916. Pisonia boliviana Britton ex Rusby, Bull.

Torrey Club 27: 125. 1900.

Young branchlets pilose with short, spreading, fulvous or ochra-

ceous hairs; petioles slender, 1.5 cm. long or less, pilose like the branch-

NYCTAGINACEAE OF SOUTH AMERICA 87

lets; leaf blades membranaceous, fuscous when dried, elliptic to ellip-

tic-oblong or ovate-elliptic, broadest at or near the middle, 10-18

cm. long, 4-8 cm. wide, acute or abruptly acute at the apex, with

very obtuse tip, obtuse to acute at the base, oblique, and often

abruptly contracted and short-decurrent, glabrous above or nearly

so, beneath sparsely pilose over the whole surface with long spreading

hairs, densely pilose along the costa, in age glabrate except along the

costa; staminate inflorescences terminal and axillary, small and few-

flowered, short-pedunculate, only 1-2 cm. long, the branches sparsely

villosulous, the flowers sessile or nearly so; perianth obconic, 3.5

mm. long, glabrous or nearly so; stamens 10; pistillate inflorescence

and fruit unknown.

Bolivia: Junction of the rivers Beni and Madre de Dios, Rusby

2502 (F, W; type collection).

Rushy 2501, mentioned in the original description, is a species

of Neea.

7. Torrubia microphylla (Heimerl) Standl. Contr. U. S. Nat.

Herb. 18: 100. 1916. Pisonia microphylla Heimerl in Urban, Symb.

Antill. 7: 215. 1912.

A shrub or small tree, the young branchlets ferruginous-puberu-

lent; leaves very small, fuscous when dried, subcpriaceous, opposite,

the petioles 1.5-2.5 mm. long; leaf blades varying from oblong or

oblanceolate-oblong to oval or suborbicular, 2.8 cm. long and 2 cm.

wide or smaller, rounded at the apex, rounded to attenuate at the

base, at first ferruginous-puberulent beneath along the costa but in

age glabrous; staminate inflorescences terminal, short-pedunculate,

laxly few-flowered, 3 cm. wide or less, sparsely appressed-hirtulous,

the flowers sessile or nearly so; perianth 4-4.5 mm. long, sparsely

and minutely puberulent or almost glabrous; stamens 7-8.

Venezuela: El Valle, Margarita Island, Miller & Johnston 231

(F, type collection, also photo, of type ex herb. Berol.; W). Around

El Palito, near Puerto Cabello, in the cactus formation, at sea level,

Pittier 6421 (F, photo, ex herb. Berol.; W).

Pittier 6421 has leaves slightly broader than those of the type,

and it has been determined by Heimerl as a new form of Pisonia

microphylla, but it scarcely merits the distinction of a name.

8. Torrubia pacurero (HBK.) Standl. Contr. U. S. Nat. Herb.

18: 101. 1916. Pisonia pacurero HBK. Nov. Gen. & Sp. 2: 218.

1818.

A shrub or small tree, sometimes 7 m. high, the branchlets

glabrous or only obscurely puberulent; leaves opposite or ternate,

membranaceous, bright green or yellowish green when dried, on

slender and often elongate petioles; leaf blades chiefly elliptic-oblong

to elliptic or ovate-elliptic, sometimes oblong-lanceolate, broadest

at or below the middle, 5.5-16 cm. long, 2-6.5 cm. wide, acute to

long-acuminate, at the base acute or obtuse, often oblique, some-

88 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

times abruptly decurrent, glabrous; staminate inflorescences terminal,

on slender, short or elongate peduncles, many-flowered, dense or

open, the branches glabrous or nearly so; perianth green, whitish,

or reddish, obconic, 4 mm. long, glabrous; stamens 8; pistillate

perianth tubular, 2 mm. long; anthocarp narrowly oblong, 1 cm.

long, conspicuously costate, glabrous.

Trinidad: Island of Patos, a small tree with brittle wood, Broad-

way 2714 (F). — Venezuela: Cumana, Humboldt (F; photo, of type

ex herb. Berol.; type locality given in the original description as

"locis umbrosis siccis Novae Andalusiae"). Aricana, on hills, Broad-

way 845 (W) ; near the shore, Broadway 609 (W) ; seashore, Broadway

807 (W). Cristobal Colon, Broadway 691 (W); on hills, Broadway

198 (W); near the sea, Broadway 232 (W). Hacienda Taborda, near

El Palito, Carabobo, Pittier 7667 (F), 7077 (F, W; these two numbers

are mixed with material of Neea anisophylla). Hacienda Puerto

La Cruz, Coastal Range, Distrito Federal, Pittier 8093 (W). Rio

Turbio, near Barquisimeto, Lara, Saer 237 (W). Rio Chico, Miranda,

John 1275 (W). Around El Palito, near Puerto Cabello, Carabobo,

in the cactus formation, at sea level, Pittier 6422 (F, W); on hills,

Pittier 9074 (W). Puerto Cabello, common, Curran & Haman 1180

(F, W); Curran & Haman 1169 (W), 1147 (W). Without locality,

Curran & Haman 1221 (W). Between Adecora and Pueblo Nuevo,

Paraguana, Curran & Haman 563 (W). Isla de San Carlos, Curran

& Haman 784 (W). Santa Lucia, Curran & Haman 1097 (F, W).—

Colombia: "Colombia," Karsten 18 (F; fragm. ex herb. Berol.).

Region of Santa Marta, near sea level, H. H. Smith 2741 (F, W),

2096 (F, W), 397 (F, W).

This is a well-marked species, easy to recognize because of its thin,

bright green leaves, and almost complete lack of pubescence. The

name has been applied sometimes to the forms referable to Torrubia

fragrans, and even to other species not very closely related.

The vernacular names "pacurero," "casabito," and "fruta de

culebra" are reported from Venezuela.

9. Torrubia uberrima, sp. nov. — Ramuli novelli dense minute-

que adpresse ferrugineo-tomentulosi subteretes graciles, internodiis

brevibus; folia opposita firme membranacea in sicco fusco-brunnea

longiuscule petiolata, petiolo gracili 1.5-3 cm. longo dense ferrugineo-

tomentuloso; lamina elliptico-oblonga vel ovato-oblonga, prope vel

infra medium latissima, 13-16.5 cm. longa, 5.5-7 cm. lata, acuminata,

basi valde obliqua et uno latere abrupte decurrens, glabra vel in

statu juvenili ad costam minute ferrugineo-tomentulosa, supra opaca,

costa venisque vix prominulis, subtus fere concolor, cpsta gracili

prominente, nervis lateralibus utroque latere c. 9 gracillimis pro-

minulis angulo semirecto adscendentibus remote a margine laxe

conjunctis, nervulis obsoletis; inflorescentia mascula terminals c.

9 cm. longe pedunculata, pedunculo ut ramuli dense ferrugineo-

tomentello erecto, panicula laxe multiflora 6-8 cm. longa 10-13 cm.

NYCTAGINACEAE OF SOUTH AMERICA 89

lata rotundata, ramis basalibus oppositis vel verticillatis patentibus,

floribus sessilibus vel vix 1 mm. longe pedicellatis in cymulas densas

paucifloras dispositis; perianthium masculum obconicum S-5.5 mm.

longum basi acutum dense ferrugineo-tomentulosum, lobis brevis-

simis; stamina c. 6 perianthio fere duplo longiora. — Colombia: Re-

gion of Santa Marta, near sea level, September, 1898-1901, Herbert

H. Smith 1873 (Herb. Field Mus. No. 138,714, type; duplicate in

U. S. Nat. Herb.).

The type collection was distributed as Pisoniaferruginea Klotzsch,

to which the plant has no close relationship.

Here may belong the following fruiting specimens from Colombia:

San Martin de Loba, Dept. Bolivar, Curran 371 (W), 184 (W).

Curran describes the plant as a tree 6.5 m. high, with a trunk 10

cm. in diameter, vernacular name "clavo cuchillo."

10. Torrubia fragrans (Dum.-Cours.) Standl. Contr. U. S.

Nat. Herb. 18: 100. 1916. Pisonia fragrans Dum.-Cours. Bot. Cult,

ed. 2. 7: 114. 1814. P. eucalyptifolia HBK. Nov. Gen. & Sp. 7: 197.

1825.

A tree 15 m. high or less, the trunk up to 30 cm. in diameter,

the branchlets grayish- or rufous-puberulent; leaves opposite, often

very irregular and unequal, the petioles 3-10 mm. long; leaf blades

rhombic-elliptic to obovate-oblong, obovate, or oblanceolate, 3-9 cm.

long, 2.5-5 cm. wide, broadest above the middle, at the apex abruptly

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

fuscous when dried, firm-membranaceous, glabrous or practically

so, the lateral nerves numerous and conspicuous; inflorescences

mostly terminal, pedunculate, commonly small and rather few-

flowered, 2-6 cm. broad, the branches puberulent or glabrate, the

flowers sessile or on pedicels 1-2 mm. long; staminate perianth puber-

ulent; stamens 6-8; anthocarp ellipsoid-oblong, 7-11 mm. long,

glabrous.

Curacao: Hofje Abau, Curran & Raman 104 (W). Klein Sint

Martha, Curran & Raman 138 (W). — Tobago: Providence Road,

a small tree with greenish flowers, Broadway 3544 (F). Calderhal,

Broadway 3541 (F). — Venezuela: San Juan Mountain, Margarita

Island, 600 m., Johnston 121 (F, W). Between Moron and El

Sanchon, Carabobo, Pittier 12205 (F, W). Cristobal Colon, a small

tree with green flowers, Broadway 740 (W). — Colombia: Region of

Santa Marta, 75 m., H. H. Smith 396 (F, W). Widely distributed

in the West Indies.

The vernacular name "laabra" is reported from Curacao.

Pittier 12205 is indicated by Heimerl, in the Berlin herbarium,

as a new species, but it does not seem to the writer more than a form

of T. fragrans, differing chiefly in the shape of the leaves, which are

unusually narrow, and rounded or very obtuse at the apex.

90 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

The locality of Pisonia eucalyptifolia is given merely as "inter

tropicos Americae." It is possible that it is not really a synonym of

Torrubia fragrans, although the description agrees well with the plant

described here. It seems probable that the proper name for T.

fragrans is Torrubia inermis (Jacq.) Britton (Bull. Torrey Club 31:

614. 1904. Pisonia inermis Jacq. Sel. Stirp. 275. 1763). The type

locality of Jacquin's species is the region of Cartagena, Colombia.

No specimens of Torrubia from that locality have been seen by the

writer, but if any of the species here listed occurs about Cartagena,

it is almost certain to be T. fragrans. Jacquin's description is unsatis-

factory and incomplete but, so far as it goes, it applies satisfactorily

to T. fragrans.

11. Torrubia salicifolia (Heimerl) Standl. Contr. U. S. Nat.

Herb. 18: 101. 1916. Pisonia salicifolia Heimerl in Urban, Symb.

Antill. 7: 216. 1912.

A small tree, the young branchlets densely ferruginous-puberu-

lent; leaves opposite, fuscous when dried, the petioles 1-11 mm.

long; leaf blades chiefly oblong-lanceolate or narrowly elliptic-oblong,

mostly 5-9 cm. long and 1.5-3 cm. wide, subabruptly acuminate

or long-acuminate, cuneately acute at the base, hirtulous at first

but soon glabrate except sometimes along the costa; staminate in-

florescence pedunculate, 3 cm. wide or less, the slender branches

spreading; pistillate flowers yellowish; staminate perianth ferrugi-

nous-hirtulous; stamens 6-7; anthocarp ellipsoid-oblong, 10-11 mm.

long.

Trinidad: Type from Moruga, Broadway 2421. Buenos Ayres,

Erin, May 9, 1919, Broadway (F). — British Guiana: Without

locality, Jenman (F). Pomeroon District, Moruka River, De La

Cruz 4557 (F).

This is probably no more than a form, and not a clearly defined

one, of T. Olfersiana.

12. Torrubia Olfersiana (Link, Klotzsch & Otto) Standl. Contr.

U. S. Nat. Herb. 18: 101. 1916. Pisonia Olfersiana Link, Klotzsch

& Otto, Icon. PL Rar. 36. pi. 15. 1841. P. Cafferiana Casar. Nov.

Stirp. Bras. 68. 1842. P. guianensis Klotzsch in Schpmb. Fauna &

Fl. Brit. Guian. 1131. 1848, nomen nudum. P. minor Choisy in

DC. Prodr. 132: 443. 1849. P. nitida Mart, ex Schmidt in Mart.

Fl. Bras. 142: 356. 1872. P. acuminata Mart, ex Schmidt, op. cit.

357. 1872. P. Eggersiana Heimerl, Bot. Jahrb. Engl. 21: 627. 1896.

P. Schomburgkiana Heimerl, Beitr. Syst. Nyctag. 34. 1897, nomen

nudum. P. Olfersiana var. nitida Heimerl, Beitr. Syst. Nyctag. 35.

1897. Torrubia Cafferiana Standl. Contr. U. S. Nat. Herb. 18: 100.

1916. T. Eggersiana Standl. loc. cit. 1916. T. nitida Standl. op.

cit. 101. 1916.

NYCTAGINACEAE OF SOUTH AMERICA 91

A shrub or tree, sometimes as much as 15 m. high, the branchlets

glabrous or sometimes yillosulous or puberulent; leaves chiefly oppo-

site, fuscous when dried, thick-membranaceous or subcoriaceous,

the petioles commonly about 1 cm. long; leaf blades oblong to elliptic,

usually 9-16 cm. long and 3-6 cm. wide, abruptly acuminate or

acute, rarely obtuse, acute at the base, usually glabrous; inflores-

cences on long or short peduncles, lax and many-flowered, sparsely

puberulent or hirtous or almost glabrous; flowers greenish, the

perianth sparsely and minutely puberulent; stamens 5-7; anthocarp

dark purple, ellipsoid-oblong, 8-12 mm. long.

Tobago: The Widow, Broadway 4155 (F). Roseborough Estate,

Broadway 4532 (F). — Trinidad: Mora Forest via Sangre Grande,

Broadway 5696 (F) . St. Anns, in light forest, June 11, 1922, Broadway

(F). — French Guiana: "Cayenne," Jelski (F; photo, and fragm. ex

herb. Berol.). — Surinam: Kegel 658 and Wullschlaegel 1005 (reported

by Heimerl as Pisonia Eggersiana). — British Guiana: Without de-

finite locality, Schomburgk 1031, 600, 595 (F; photo, and fragms. ex

herb. Berol.). Without locality, Schomburgk 588 (F; photo, and fragm.

ex herb. Berol.). Kabakaburi, Pomeroon District, De La Cruz 3315

(F, W), 3317 (F, W). Kamakusa, Upper Mazaruni River, De La

Cruz 4120 (F). Pomeroon River, De La Cruz 3017 (F). Issorora,

Aruka River, in hill forest, Hitchcock 17550 (W). — Venezuela: Los

Teques, Miranda, 1,400-1,500 m., Pittier 7499 (W). Quebradita de

las Ruinas, Federal District, 800-1,000 m., in light forest, Pittier

9463 (W). Cariaquita, Paria Peninsula, Bond, Gillin & Brown 246

(W). Mesa de El Sombrero, Guarico, in bushes around savanna,

Pittier 12371 (W). Quebrada de Chacaito, Federal District, 800-

1,000 m., near cascade in forest, Pittier 10323 (W).— Brazil: Without

locality, Sello (F; photo, and fragm. ex herb. Berol.). Rio de Janeiro,

Sello (F; fragm. ex herb. Berol.). Without locality, Regnell III.

1021 (F).— Ecuador: Balao, in forest, Eggers 14249 (W).— Bolivia:

Bosques del Rio Surutu, Dept. Santa Cruz, 400 m., Steinbach 7284

(F; photo, and fragm. ex herb. Berol.).

The vernacular name is reported from Venezuela as "salado."

Steinbach 7284 has been determined by Heimerl as a new species, and

it may be distinct from T. Olfersiana, but the incomplete material

available for examination does not seem to be essentially different.

The material here cited is conspicuously variable in leaf form and

in pubescence, but it does not seem practical to separate it into

several species. Other related species probably must be reduced to

synonymy under T. Olfersiana, when more abundant material has

accumulated, because the slight differences upon which they have

been based probably are not constant.

4. PISONIA L.

Trees or shrubs, erect or more often with long, pendent or clamber-

ing branches, often armed with spines; leaves chiefly opposite and

92 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

petiolate, entire; flowers dioecious, small, reddish or yellowish green,

arranged in sessile or pedunculate cymes, exinvolucrate, 2-3-bracteo-

late; staminate perianth obconic-campanulate, the limb 5-dentate;

stamens 6-8, exserted, unequal; pistillate perianth tubular, the limb

5-dentate; anthocarp coriaceous, clavate to oblong or obovoid,

terete and costate or 5-angulate, the angles or costae furnished with

one or more rows of viscid stipitate glands.

Anthocarp with glands only along the upper part 1. P. zapallo.

Anthocarp with glands along the whole length of the angles.

Mature fruit 6-10 mm. in diameter, the glands along the angles

in two or more rows; spines few or wanting on the flowering

branches 2. P. macranthocarpa.

Mature fruit 3-4 mm. in diameter, the glands along the angles in

a single row or nearly so; flowering branches usually armed

with recurved spines 3. P. aculeata.

1. Pisonia zapallo Griseb. Goett. Abh. 24: 39. 1879. P.

aculeata f. inermis Kuntze, Rev. Gen. 32: 265. 1898.

A small tree, unarmed; leaves slender-petiolate, the blades broadly

elliptic to ovate, mostly 5-10 cm. long, acute at the base, acute to

subobtuse at the apex, when young tomentose beneath, in age

sparsely pilose or glabrate; cymes pedunculate, tomentose, the

flowers glomerate; staminate perianth 2 mm. long; anthocarp nar-

rowly clavate, 15-20 mm. long, the glands in 2 or more series along

each angle above the middle of the fruit.

Bolivia: Rio Achira, 1,000 m., Steinbach 8223 (F). Sierra de

Santa Cruz, 1,600 m., May, 1892, Kuntze (F, type collection of

P. aculeata f. inermis). — Also in Argentina.

From Argentina there are reported for this species the vernacular

names "palo de zapallo," "zapallo caspi," and "ombu-ra."

2. Pisonia macranthocarpa Bonn. Smith, Bot. Gaz. 20: 293.

1895. P. aculeata var. macranthocarpa Donn. Smith, Bot. Gaz. 16:

198. 1891. P. aculeata var. pedicellaris Griseb. ex Heimerl, Bot.

Jahrb. Engl. 21: 633. 1896.

A shrub or small tree, 3-4 m. high or sometimes larger, the branch-

lets puberulent when young but soon glabrate; spines few and often

or usually wanting on herbarium specimens, mostly straight; leaf

blades elliptic to oval or obovate, acute at the base, acute to

acuminate at the apex, puberulent or short-villous beneath along

the costa or often glabrate in age; inflorescence pedunculate, the

staminate cymes dense, many-flowered, 2-3.5 cm. broad, the flowers

short-pedicellate; staminate perianth yellowish green, 3-4 mm. broad,

puberulent; stamens usually 8, twice as long as the perianth; antho-

carp woody, 1-2 cm. long, densely tomentulose.

NYCTAGINACEAE OF SOUTH AMERICA 93

Venezuela: La Trinidad de Maracay, Aragua, 440 m., Pittier

5851 (W). Valencia, Rose 21837 (W). Lara, Saer 156 (W).—

Colombia: Region of Santa Marta, H. H. Smith 395 (F, W).—

Ecuador: Guayaquil, Stevens 39 (W). Duran, Rose & Rose 23611

(W). Near Guayaquil, in woods, Mille 206 (W). Daule, Andre

K829 (F). El Recreo, Eggers 15332 (F). Also in Cuba, Central

America, and southern Mexico.

Known in Salvador by the names "espuela del diablo" and

"guaco."

3. Pisonia aculeata L. Sp. PL 1026. 1753.

A shrub or a small tree, densely branched, sometimes with a

thick trunk, the branches often very long and scandent, or more

frequently pendent, armed with numerous stout recurved spines;

leaves petiolate, the blades elliptic to ovate or even rounded, mostly

4-8 cm. long, acute to rounded at the base, acute or acuminate at

the apex, glabrate beneath or frequently puberulent or short- villous;

staminate cymes pedunculate, loosely or densely cymose, 2-6 cm.

broad, many-flowered; staminate perianth 2-4 mm. long, yellowish

green, densely puberulent or tomentulose; stamens usually 6, twice

as long as the perianth; anthocarp 9-12 mm. long, puberulent or

glabrate.

Colombia: Region of Santa Marta, 45 m., H. H. Smith 1861

(F, W). Ranging through the West Indies to Florida, and in Central

America and Mexico; occurring also in the Old World tropics.

The species has been reported from Venezuela, but the reports

probably relate to P. macranthocarpa, which is closely related but

quite distinct. It is common in Central America at low elevations,

often growing in thickets near the seashore. In Central America it

is known by the names "espino negro" and "cagalero," and in

Mexico and Cuba as "una de gato."

5. PISONIELLA (Heimerl) Standl.

Erect shrubs, dichotomously much branched; leaves opposite,

petiolate, entire; flowers perfect, in headlike, pedunculate, axillary

and terminal, many-flowered umbels, pedicellate, each pedicel sub-

tended by a minute bract; perianth tubular-campanulate, slightly

constricted at the middle, the limb shallowly 5-lobate, the lobes

rounded or subtruncate; stamens 6-11, the filaments exserted;

anthocarp oblong-clavate, slightly curved, coriaceous, obtuse at

the apex, attenuate below, 5-costate, the angles bearing numerous

verruciform glands.

The genus consists of a single species.

1. Pisoniella arborescens (Lag. & Rodr.) Standl. Contr. U. S.

Nat. Herb. 13: 385. 1911. Boerhaavia arborescens Lag. & Rodr.

Anal. Cienc. Nat. 4: 257. 1801. Pisonia hirtella HBK. Nov. Gen.

94 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

& Sp. 2: 217. 1818. Pisonia arborescens Kuntze, Rev. Gen. 32: 265.

1898.

The species grows in central and southern Mexico. In South

America it is represented by the following variety:

la. Pisoniella arborescens (Lag. & Rodr.) Standl., var. gla-

brata Heimerl, Ann. Cons. Jard. Bot. Geneve 14: 231. 1914. Pisonia

hirtella f. glabrata Heimerl, op. cit. 5: 196. 1901. Pisoniella glabrata

Standl. Contr. U. S. Nat. Herb. 13: 386. 1911.

A slender shrub, densely branched, the young branchlets ferrugi-

nous-tomentulose, soon glabrate; leaves on short slender petioles, the

blades thin, ovate to oblong-elliptic, mostly 3.5-8 cm. long, acute

to attenuate at the base, abruptly acute to very long-attenuate at

the apex, glabrous or nearly so; peduncles 2-10 cm. long, the umbels

dense, many-flowered, 12-15 mm. in diameter, the flowers short-

pedicellate; perianth greenish white, 5-7 mm. long, sparsely and

minutely viscid-puberulent; anthocarp 1 cm. long and 1.5-2 mm.

thick, dark brown, glabrous.

Bolivia: Valle de Cochabamba, 2,600 m., Steinbach 8728 (F).

Sorata, 2,400 m., Rusby 2500 (F, W, type collection; cited erroneously

as "2510" in the original publication). Cochabamba, Bang 1063

(F, W). Without locality, Bang 1809 (F, W). Cochabamba, 3,000

m., March, 1892, Kuntze (F). — Argentina: El Rastrojq, Prov.

Catamarca, 1,600 m., Jorgensen 1240 (W). Siambon, Sierra de

Tucuman, Lorentz & Hieronymus 776 (F).

The variety is isolated by a great distance from the area occupied

by the typical form of the species, in Mexico, but the differences

between the type and the variety are so slight that they would

scarcely deserve attention if both forms occupied the same area.

6. SALPIANTHUS Humb. & Bonpl.

Large herbs or small shrubs, pubescent and usually viscid; leaves

alternate, petiolate, the blades broad, entire; flowers perfect, not

involucrate, ebracteate, very small, glomerate or racemose, forming

terminal and axillary, panicled cymes; perianth campanulate, urceo-

late, tubular, or pyriform, green or reddish, persistent, scarcely

accrescent, the limb with 4-5 teeth or lobes; stamens 3-5; fruit

subglobose, sometimes somewhat compressed, coriaceous, minute.

About three additional species of the genus are known from

Mexico.

Leaves bright green, acute or acuminate, large; flowers in panicled

cymes; perianth campanulate, with very short teeth, bearing

numerous uncinate hairs 1. S. purpurascens.

Leaves grayish, usually rounded at the apex, small; flowers in

racemosely arranged cymes; perianth pyriform, at least in age,

NYCTAGINACEAE OF SOUTH AMERICA 95

with conspicuous, more or less spreading lobes, without uncinate

hairs 2. S. pyriformis.

1. Salpianthus purpurascens (Cav.) H. & A. Bot. Beechey

Voy. 308. 1837. Boldoa purpurascens Cav. ex Lag. Gen. & Sp. Nov.

10. 1816. B. ovatifolia Lag. loc. cit. 1816. Cryptocarpus globosus

HBK. Nov. Gen. & Sp. 2: 187. pi. 123. 1818. C. paniculatus HBK.

ex Schlecht. Linnaea 26: 643. 1853.

Plants herbaceous or suffrutescent, 2 m. high or less, much

branched, the branches slender, sparsely puberulent or glabrate,

those of the inflorescence viscid and bearing numerous short uncinate

hairs; leaves long-petiolate, the blades broadly rhombic-ovate to

ovate-deltoid, 5-22 cm. long, acute or acuminate, at the base abruptly

contracted and long-decurrent, finely scaberulous when young but

soon glabrate; flowers sessile or subsessile; perianth 2.5-3 mm. long,

green, 4-dentate; stamens 4, exserted; fruit subglobose, 1.5 mm. in

diameter.

Venezuela: Near Caracas, 900-1,150 m., L. H. & E. Z. Bailey

675 (W). Maracay, Aragua, Cornelia 87 (W). Type of C. paniculatus

apparently Wagener 289 from Maiquetia. Also in Mexico and

Central America.

Heimerl reports the species from Colombia, but without citation

of material to support its occurrence there.

2. Salpianthus pyriformis (HBK.), comb. nov. Cryptocarpus

pyriformis HBK. Nov. Gen. & Sp. 2: 188. pi. 124. 1818. C. cordi-

folius Moric. PI. AmeY. 75. pi. 50. 1830. C. pyriformis var. cordi-

folius Moq. in DC. Prodr. 132: 88. 1849.

Plants herbaceous or suffrutescent, usually more or less scandent,

and 2 m. long, sometimes scandent to a height of 10 m., the branches

densely viscid-puberulent; leaves on stout petioles, the blades

rhombic-ovate to broadly ovate-deltoid or rounded-ovate, mostly

2.5-5 cm. long, usually cordate or deeply cordate at the base but

varying to rounded, rounded or very obtuse at the apex, densely

viscid-puberulent or tomentulose on both surfaces; inflorescence very

leafy or naked, densely viscid-puberulent, the flowers subsessile;

perianth less than 2 mm. long; stamens 4; anthocarp globose, blackish,

1 mm. in diameter.

Ecuador: Reported from Guayaquil, and also from Abingdon,

Albemarle, Barrington, Bindloe, Charles, Chatham, Indefatigable,

James, Jervis, Narborough, and Seymour Islands of the Galapagos

Islands. Occurring also in Peru.

Called "nacupillo" in Peru.

By Heimerl and others this plant has been treated as the type

of a distinct genus, Cryptocarpus. It does not seem to possess any

important characters that distinguish it from Salpianthus, and it

96 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

is therefore preferable to treat it as a member of that genus. By

Moquin this plant, as well as S. purpurascens, was referred to the

family Chenopodiaceae. In volume 13, part 2, of De Candolle's

Prodromus, S. purpurascens appears twice under different names, on

page 88 in the genus Cryptocarpus among the Chenopodiaceae, and

on pages 438 and 439 under Boldoa, among the Nyctaginaceae.

Cryptocarpus cordifolius was described from Guayaquil.

7. BOUGAINVILLEA Commers.

Shrubs or small trees, often scandent, usually armed with simple

or branched spines; leaves alternate, petiolate, entire; flowers perfect,

either solitary and subtended by 3 bracts, or usually in a 3-flowered

axillary inflorescence consisting of 3 large, persistent, often brightly

colored bracts, a flower borne on the inner surface of each bract,

its pedicel confluent with the costa of the bract; perianth tubular,

the limb usually shallowly 5-lobed, the tube subterete or 5-angled;

stamens 5-10, somewhat unequal, connate at the base into a short

cup; anthocarp fusiform, coriaceous, 5-costate.

Bracts brightly colored, purplish red or bright red, retaining the color

when dried, mostly 2.5-4 cm. long.

Perianth tube glabrous 1. B. peruviana.

Perianth tube variously pubescent.

Perianth tube hirsute or villous; leaves usually copiously villous.

2. B. spectabilis.

Perianth tube puberulent or glabrate; leaves glabrate.

3. B. glabra.

Bracts green, or sometimes brightly colored when fresh but losing

their color when dried, usually smaller, and only 1-2.5 cm. long.

Perianth only 6.5-7 mm. long, gradually dilated from base to apex,

glabrate, or hirtous above; bracts 7-12 mm. long.

4. B. campanulata.

Perianth 9-20 mm. long, constricted above; bracts 13-27 mm. long.

Perianth glabrous, 11-14 mm. long; bracts 25-27 mm. long,

13-15 mm. wide, glabrous except on the costa.

5. B. berberidifolia.

Perianth variously pubescent; bracts 13-22 mm. long.

Perianth 9-11 mm. long, densely tomentulose. . 6. B. praecox.

Perianth 12-20 mm. long.

Perianth puberulent 7. B. stipitata.

Perianth pubescent or hirsute 8. B. infesta.

NYCTAGINACEAE OF SOUTH AMERICA 97

1. Bougainvillea peruviana H. & B. PI. Aequin. 1: 174. pi. 49.

1808. Tricycla peruviana Poir. Encycl. Suppl. 5: 359. 1817.

A shrub 3-7 m. high, erect or scandent, armed with numerous

slender spines 1-2.5 cm. long, the branches sparsely puberulent or

glabrate; leaves thin, slender-petiolate, the blades broadly ovate to

suborbicular, 5-7 cm. long, subtruncate at the base, subobtuse or

abruptly acute at the apex, except when very young glabrous or

nearly so; bracts bright rose, 1.5-3.5 cm. long, obtuse or rounded

at the apex, glabrous except along the puberulent costa; perianth

16-20 mm. long, white or whitish, the limb 5-6 mm. broad; stamens

usually 6; anthocarp about 10 mm. long and 2.5 mm. thick, glabrous.

Colombia: Gigante, Dept. Huila, in 1920, Bro. Ariste- Joseph (W).

—Ecuador: Without locality, Townsend 837 (W). Without locality

(perhaps from Colombia), Lehmann B. T. 446 (F). Naranjo, Andre

K.708 (F). Also in Peru.

Townsend reports the vernacular name as "papelillo." This

species is closely related to B. glabra, but the dried specimens, when

compared with others of that species, have a plainly different aspect.

2. Bougainvillea spectabilis Willd. Sp. PI. 2: 348. 1789.

A large woody vine, armed with numerous stout spines 4 cm.

long or shorter, the branches copiously fulvous-villous, rarely glabrate ;

leaf blades broadly ovate to suborbicular or rounded-oval, 5-10 cm.

long, rounded to acutish at the base and often short-decurrent,

abruptly acute or acuminate at the apex, usually densely villous

beneath; bracts purplish red, ovate-oval or broadly ovate, 2-4.5 cm.

long, subcordate at the base, abruptly acute or acuminate or some-

times obtuse at the apex, sparsely puberulent or short-villous;

perianth 15-30 mm. long, the tube green, the limb 6^7 mm. wide,

yellowish; anthocarp 11-14 mm. long, densely short-villous.

Bolivia: Unduavi (doubtless cultivated), Bro. Julio 358 (W).

Native of Brazil.

This Brazilian plant is cultivated for ornament in many parts of

the tropics, but in most regions it is far less common than B. glabra.

From Argentina there are reported the vernacular names "Santa

Rita" and "Tres Marias."

3. Bougainvillea glabra Choisy in DC. Prodr. 132: 437. 1849.

B. spectabilis var. glabra Hook, in Curtis's Bot. Mag. pi. 4810. 1854.

A large woody vine, armed with numerous stout spines, the

branchlets puberulent when young but soon glabrate; leaf blades

broadly ovate to ovate-lanceolate, 4-10 cm. long, rounded to acute

at the base, abruptly or gradually acute to acuminate at the apex,

puberulent when young but quickly glabrate; bracts commonly

purplish red, broadly ovate to oval, subcordate at the base, commdnly

abruptly acute or acuminate at the apex, sparsely puberulent or

98 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

glabrous; perianth 15-25 mm. long; stamens 8; anthocarp turbinate,

7-13 mm. long, with 5 acute angles.

Venezuela: El Valle, Margarita Island, Miller & Johnston 99

(F, W).— Colombia: Medellin, Dept. Antioquia, Archer 763 (F).

Barranquilla, Bro. Paul 34 (W). Cartagena, Bro. Heriberto 153 (W).

La Esperanza, Dept. Cundinamarca, Bro. Ariste- Joseph (W). Be-

tween El Colegio and Tequendama, Cundinamarca, Bro. Ariste-

Joseph 1069 (W).— Ecuador: Guayaquil, Mitte 16 (W). Native of

Brazil.

This is the common Bougainvillea of cultivation, that is grown

for ornament in most tropical regions, often in such abundance that

one becomes excessively bored with its continued display. In Ecuador

and Colombia the vine is called "flor de verano" — summer flower —

because it blooms during the dry months, the "summer" of tropical

America, although the winter months of the north. The name

"trinitaria" also is used for the plant in Colombia, and from Argen-

tina there are reported the names "Santa Rita" and "Tres Marias."

The species is not too distinct from B. spectabilis, although prob-

ably it has as good characters as most other members of the genus.

It might be preferable to treat it as a variety of that species, as was

done by Hooker.

4. Bougainvillea campanulata Heimerl, Med. Rijks Herb.

19: 33. 1913.

A shrub or small tree, unarmed, the branchlets glabrous or when

young minutely tomentulose; leaves short-petiolate, elliptic-lanceo-

late or narrowly oblong-elliptic, 2 cm. long and 12 mm. wide or

smaller, broadest at the middle, obtuse at the apex, acute at the

base, minutely hirtellous beneath along the costa, otherwise glabrous;

bracts elliptic to obovate-elliptic, 7-12 mm. long, 4-6.5 mm. wide,

subobtuse to rounded at the apex, yellowish green, sparsely puberu-

lent or villosulous; perianth 6.5-7 mm. long, yellow, hirtous above,

the limb 9-10 mm. broad; stamens 7-8.

Bolivia: Yuquirenda, left bank of the Rio Pilcomayo, 400 m.,

Herzog 1124, type. Comarapa, Dept. Santa Cruz, 2,000 m., Stein-

bach 8579 (F) . Without locality, Herzog 1137 (photo, ex herb. Berol.) .

Villamontes, Pflanz 696 (fragm. ex herb. Berol.). — Argentina: Bal-

coyna, Dept. del Alto, 1,250 m., Venturi 7240 (F). Reported from

the departments of Formosa, Tucuman, and Jujuy.

The vernacular name is reported from Argentina as "coronillo."

5. Bougainvillea berberidifolia Heimerl, Denkschr. Akad.

Wiss. Wien 70: 121. pi 1, f. 2-3. 1900.

A spiny shrub, the spines mostly 1-2 cm. long, the branchlets

sparsely tomentulose when young, soon glabrate; leaf blades elliptic

to obovate-elliptic, broadest at or above the middle, 1.5-2.5 cm.

NYCTAGINACEAE OF SOUTH AMERICA 99

long, rounded or obtuse at the apex, attenuate to the base, glabrate;

bracts bright rose, but turning greenish when dried, ovate-oblong,

25-27 mm. long, 13-15 mm. wide, subobtuse, sometimes suborbicular

and broadly rounded at the apex, glabrous except along the costa;

perianth 11-14 mm. long, yellowish or red, glabrous, the limb 4-5

mm. broad; stamens usually 5.

Bolivia: Type collected by Cuming, without locality; Heimerl

reports also a specimen collected by Bridges, without number, and

Orbigny 517, from Valle Grande. In spiny thickets of Pulquina and

Comarapa, 1,900 m., Herzog 1799 (photo, ex herb. Berol.).

6. Bougainvillea praecox Griseb. Symb. Fl. Argent. 40. 1879.

B. praecox var. spinosa Chod. & Hassl. Bull. Herb. Boiss. II. 3:

415. 1903. B. praecox var. rhombifolia Heimerl, Verh. Zool. Bot.

Ges. Wien 62: 4. 1912. B. modesta Heimerl, Denkschr. Akad. Wiss.

Wien 70: 118. pi. l,f. 4. 1900.

A shrub or tree, sparsely spiny or unarmed, the spines mostly

5 mm. long or less, the branchlets tomentulose or soon glabrate;

leaf blades ovate to elliptic or ovate-elliptic, 8 cm. long and 4 cm.

wide or smaller, narrowed to an obtuse apex, acute or attenuate at

the base, appressed-tomentose beneath at first but later glabrate;

bracts white or reddish, becoming greenish when dried, broadly

ovate, about 14 mm. long and 12 mm. wide, rounded or subcordate

at the base, rounded or very obtuse at the apex, densely puberulent,

especially on the veins; perianth 9-11 mm. long, densely tomentulose,

the limb 4 mm. broad; stamens 5-6.

Bolivia: Piragwald, Dept. Santa Cruz, 400 m., Steinbach 8122

(F). Near Coroico, Bang 2398 (F, W; type collection of B. modesta).

Huachi, head of Beni River, 540 m., White 973 (F, W). Heimerl

reports also the following collections: Between Ipawassu and Fortin

d'Orbigny, Herzog 1076, 1073 (var. rhombifolia). Left bank of Rio

Pilcomayo, Herzog 1106 (var. spinosa).

The species occurs also in Argentina and Paraguay. The vernacu-

lar name "duraznillo" is reported from Argentina. The two varieties

described by Heimerl are minor forms scarcely worthy of nomencla-

torial recognition, especially in a group in which the individual

variations are so great. Heimerl suggested the possibility of combin-

ing B. praecox and B. modesta, and to the present writer there do not

appear to be good grounds for separating them. The same might

be said also of most of the other species listed here, for the differences

between them certainly are for the most part rather inconsequential

ones. B. modesta is described as a tree 25 m. high, but the description

is probably erroneous in ascribing such a height to the tree.

7. Bougainvillea stipitata Griseb. Symb. Argent. 39. 1879.

B. longispinosa Rusby, Mem. Torrey Club 6: 109. 1896. B. stipitata

var. longispinosa Heimerl, Denkschr. Akad. Wiss. Wien 70: 116.

100 FIELD MUSEUM OF NATURAL HISTORY — BOTANY, VOL. XI

1900. B. stipitata var. Kuntzeana Heimerl, op. cit. 117. 1900. B.

stipitata var. Fiebrigii Heimerl, Bot. Jahrb. Engl. 42: 76. 1908.

A shrub or small tree, unarmed or armed with stout or slender

spines 2.5 cm. long or less, the young branches puberulent or tomen-

tulose; leaves petiolate, the blades ovate, rhombic, deltoid-ovate,

or rarely lanceolate, usually acuminate but sometimes obtuse, at

the base rounded to acute, often decurrent, puberulent or glabrous,

3.5-7.5 cm. long; bracts green or tinged with rose, greenish when

dried, broadly ovate or ovate-elliptic, 14-20 mm. long, 12-15 mm.

wide, subtruncate to shallowly cordate at the base, obtuse or acutish

at the apex, sparsely or rather densely puberulent; perianth 13-20

mm. long, greenish, puberulent or rarely glabrous; stamens 7-8;

anthocarp fusiform, 12-15 mm. long, minutely puberulent or glabrous.

Bolivia: Region of Cochabamba, Bang 1123 (F, W, type collec-

tion of B. longispinosa). Tunari, 1,500 m., April, 1892, Kuntze

(F, type collection of var. Kuntzeana). Bermejo, 1,400 m., Fiebrig

2352 (F). Chiquiaca, 1,000 m., Fiebrig 2689 (F). Tatarenda, Gran

Chaco, Fries 1478 (W). Chinchilla apud Paicho a Tarija occidentem

versus in declivibus rupestribus, 3,000 m., Fiebrig 3049 (F, type

collection of var. Fiebrigii). Reported by Heimerl from the following

stations: Villa Montes on the Rio Pilcomayo, 460 m., Herzog 1137.

Between Cumbarute and Itatique, 800 m., Herzog 1181. Samaipata,

1,650 m., Herzog 1724- Quebrada de las Pavas, 1,900 m., Herzog

1834- — Argentina: Ascochinga, Sierra de Cordoba, Lorentz 374 (F,

photo, and fragm. of type ex herb. Berol.). Prov. Cordoba, Lossen

90 (F). Reported from many localities in Argentina, and also in

Matto Grosso, Brazil.

The varieties listed in the synonymy are of minor importance,

and scarcely worthy of nomenclatorial distinction. From Argentina

the following vernacular names are reported for the species: "Tala

falso," "guancar bianco," "alfiler," "alnlerillo," "coronillo bianco."

8. Bougainvillea infesta Griseb. Symb. Argent. 40. 1879.

A shrub about 2 m. high, unarmed or furnished with stout spread-

ing spines 5-15 mm. long, the young branchlets densely pubescent,

the older ones glabrate; leaves petiolate, elliptic to ovate or elliptic-

lanceolate, mostly 2-4 cm. long, broadest at or below the middle,

obtuse to acute, truncate to acute at the base, densely puberulent

or tomentulose beneath; bracts greenish, about 2 cm. long and 11-

13 mm. wide, rounded or subcordate at the base, very obtuse or

rounded at the apex, densely pubescent; perianth 12-14 mm. long,

densely pubescent or hirsute, the limb 5 mm. wide; stamens 5.

Bolivia: Villamontes, Pflanz 2134 (F, W). Espia, head of Bopi

River, 1,050 m., Rusby 139 (W). The following collection is reported

by Heimerl: Dry forest between Embarcacion and Miraflores,

Herzog 1043. — Argentina: Oran, Lorentz & Hieronymus 415 (F,

photo, of type ex herb. Berol.). Sierra de Calitegua, Dept. Ledesma,

NYCTAGINACEAE OF SOUTH AMERICA 101

750 m., Venturi 5404 (F). Reported also from various other localities

in Argentina.

Herzog gives the vernacular name as "palo mataco."

Bougainvillea Herzogiana Heimerl, Med. Rijks Herb. 27: 12.

1915.

The species was based on a sterile specimen from Monte Grande

near Fortin Guarayus, Bolivia, Herzog 127. It is said to be closely

related to B. infesta, but since it is based upon leaf characters alone,

the status of the plant must remain uncertain until complete material

has been collected.

8. COLIGNONIA Endl.

Plants often scandent, weak, shrubby or wholly herbaceous, some-

times with tuberous roots; leaves opposite or whorled, slender-

petioled, broad, entire, those of the inflorescence often wholly or

partly white; flowers very small, perfect, neither involucrate nor

bracteate, umbellate; perianth minute, campanulate or funnelform,

3-5-parted, the tube 3-5-angled; stamens usually 5, the anthers

didymous; style filiform, exserted; anthocarp ellipsoid to fusiform

or obpyramidal, sometimes almost orbicular, with 3-5 wings or

angles.

The genus consists of 10 species, confined to the Andean region

of South America and Panama. The species not listed here are

natives of Peru.

Perianth normally 5-parted; fruit 5-angled, not winged; leaves marked

beneath with numerous raphids.

Perianth slender-stipitate; fruit fusiform 1. C. parviflora.

Perianth not conspicuously stipitate; fruit broadly pyriform or

almost globose 2. C. glomerata.

Perianth usually 3-parted; fruit with normally 3 wings; leaves without

conspicuous raphids.

Leaves densely rufous- tomentose beneath 3. C. rufopilosa.

Leaves glabrous or nearly so beneath.

Leaf blades oblong-ovate to lance-ovate, narrowed to the apex.

4. C. scandens.

Leaf blades mostly orbicular or nearly so, broadly rounded at

the apex 5. C. ovalifolia.

1. Colignonia parviflora (HBK.) Endl. Gen. PL 311. 1837.

Abronia parviflora HBK. Nov. Gen. & Sp. 2: 216. pi. 128. 1818.

Tricratus parviflorus Spreng. Syst. Veg. 1 : 536. 1825.

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

Plants chiefly herbaceous, glabrous throughout, the stems weak

and often subscandent, 2-3 m. long, glabrous; leaves often verticillate,

the slender petioles equaling or shorter than the blades; leaf blades

rounded-ovate to almost orbicular, more or less unequal, 3-4.5 cm.

long, broadly rounded at each end, the blades of the uppermost

leaves often colored white; flowers white, arranged in very numerous

small umbels, the pedicels 2-3 mm. long; perianth 3 mm. long, the

lobes elliptic to obovate-oblong, rounded at the apex; anthocarp

fusiform or fusiform-obpyramidal, 6-10 mm. long, 2.5-3 mm. wide,

conspicuously 5-angled or almost winged.

Colombia: Andes of Popayan near Querchu, Humboldt &Bonpland

(F, photo, of type ex herb. Berol.). Volcan de Cumbal, Stubel 445

(F, fragm. ex herb. Berol.). Heimerl has reported also Triana 995.

Azufral, Andre 3254 (F). — Ecuador: La Rinconada, Prov. Carchi,

3,000 m., Hitchcock 20943 (W). The species is recorded also from

the mountains of Panama, but I doubt that the specimen is correctly

labeled.

2. Colignonia glomerata Griseb. Goett. Abh. 19: 87. 1874.

C. glomerata var. typica Heimerl, Denkschr. Akad. Wiss. Wien 70:

135. 1900. C. glomerata var. boliviana Heimerl, op. cit. 136. 1900.

Plants large and branched, herbaceous, often 2.5 m. high, nearly

or quite glabrous, the coarse stems hollow; leaves very unequal,

the petioles nearly as long as the blades; leaf blades varying from

broadly ovate to ovate-deltoid, or the upper ones lance-ovate and

often white, acutish to rounded at the apex, mostly 5-10 cm. long,

thin, paler beneath; umbels about 8-12-flowered, long-pedunculate,

the pedicels in anthesis 1-2 mm. long; perianth 2-3 mm. long, gla-

brous; anthocarp globose- turbinate to almost pyriform, 3-4 mm.

long, 2-2.5 mm. wide, attenuate toward the base.

Argentina: Cie"naga, Lorentz & Hieronymus 725 (F, photo,

ex herb. Berol.). Siambon, Lorentz & Hieronymus 894 (F, photo, ex

herb. Berol.). Reported by Heimerl also from "Alisowaldern der

Cuesta de Anfama in der Nahe der Cie"naga," Lorentz 313, 754.—

Bolivia: Tunari, May, 1892, Kuntze (F). Yungas, Bang 699 (F,

W). Without locality, Bang 1772 (F, W). Sorata, 3,000 m., Rusby

2705 (F, W); in wet places, Holway 523 (W). Heimerl reports also

Mandon 1007.

Var. boliviana Heimerl was based upon Bang 699 and several other

collections. To the present writer it does not appear to be sufficiently

distinct from the typical form to deserve the distinction of a varietal

name.

3. Colignonia rufopilosa Kuntze, Rev. Gen. 32: 264. 1898;

Heimerl, Denkschr. Akad. Wiss. Wien 70: 132. 1900.

Plants herbaceous or suffrutescent, 1-3 m. long, usually sprawling

or clambering, the stout branches densely ferruginous-tomentose;

leaves chiefly opposite, those of a pair subequal, those of the inflores-

NYCTAGINACEAE OF SOUTH AMERICA 103

cence often white, the slender petioles shorter than the blades; leaf

blades ovate or ovate-elliptic, 3-8 cm. long, narrowed to an obtuse

apex, or the uppermost leaves often acute, green and sparsely tomen-

tulose above, paler beneath and densely rufous-tomentose; umbels

numerous, 10-25-flowered, the pedicels glabrous or nearly so, capil-

lary, 2-4 mm. long, or in fruit up to 5 mm. long; perianth white,

2.5-3 mm. long, glabrous; anthocarp glabrous, suborbicular, 5 mm.

long and broad, rounded at the base, 3-winged.

Ecuador: Huigra, Rose & Rose 22519 (W). Reported by Heimerl

from the Andes of Quito, Jameson 695. — Bolivia: Rio Juntas, 1,000-

2,000 m., April, 1892, Kuntze (F, photo, of type ex herb. Berol.; W,

type collection). Yungas de San Mateo, 2,800 m., Steinbach 84.90

(F). Yungas, Bang 710 (F, W). Unduavi, 2,400 m., Rusby 2706

(F, W); 3,200 m., Buchtien 4736 (W); 3,100 m., Buchtien 2993 (W).

San Felipe, Sur Yungas, Holway 612 (W). Heimerl reports also

collections by Cuming and Bridges, without special locality. The

species occurs in Peru.

Of all the species of the genus, this is the best-marked and the

most easily recognized.

4. Colignonia scandens Benth. PI. Hartw. 148. 1844.

Plants herbaceous or suffrutescent, erect or subscandent, the

stems 1-2 m. long, glabrous or nearly so; leaves often subverticillate,

the small ones of the inflorescence frequently white, those of a pair

subequal, the petioles usually shorter than the blades but sometimes

equaling them; leaf blades elliptic to rhombic-ovate or oblong-ovate,

2-6 cm. long, rounded or very obtuse at the apex, sometimes acute

at the base, sparsely ferruginous-puberulent beneath along the veins

when young, in age glabrous or nearly so; umbels commonly 8-12-

flowered, the pedicels glabrous, filiform, 3-4 mm. long, or in age as

much as 6 mm.; perianth green or greenish, 2.5-3 mm. long, glabrous,

the lobes obovate-elliptic, obtuse or rounded at the apex; anthocarp

suborbicular, 4-5 mm. long and broad, glabrous, rounded at the

base, 3-winged.

Ecuador: Type from the mountains of Loja, Hartweg 828. Be-

tween San Lucas and Ona, Prov. Loja, 2,200-3,100 m., Hitchcock

21558 (W). Loja, December 1, 1876, Andre 4554 (F). Also in Peru.

5. Colignonia ovalifolia Heimerl, Denkschr. Akad. Wiss. Wien

70: 132. 1900.

Plants herbaceous or suffrutescent, erect or subscandent, some-

times 3 m. long, the stems sparsely and closely ferruginous-tomen-

tulose on the younger parts but soon glabrous or practically so;

leaves opposite or subverticillate, rather small, a few of the upper-

most bractlike leaves white, the very slender petioles usually equaling

or often much longer than the blades; leaf blades mostly rounded-

ovate or suborbicular and 2^4 cm. long, broadly rounded at the

base, rounded at the apex, thin, paler beneath, when young sparsely

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

ferruginous-puberulent along the veins but soon glabrous or nearly

so; umbels 12-16-flowered, the slender pedicels glabrous, 3-6 mm.

long; perianth white, 3 mm. long, glabrous; anthocarp suborbicular,

5-6 mm. long and broad, glabrous, rounded at the base, 3-4-winged.

Colombia: Edge of forest, Rosalito, Dept. Tolima, 2,800-3,100

m., Pennell 2984 (F, W). Without locality, Triana 294. (herb. Paris).

Las Escaleretas, Moras Valley, Cauca, 2,500-3,000 m., Pittier 1383

(F, W). Ab9ve Canaan, Mt. Purac^, El Cauca, 3,300-3,400 m.,

Pennell & Killip 6515 (W). Reported by Heimerl from Paramo de

Henre", GoudoL — Ecuador: Type from the Andes of Quito, Spruce

5130. Camino de Chillogallo a San Juan, Prov. Pichincha, 3,000 m.,

Firmin 490 (F). Locality uncertain, Sodiro 128-6 (F, photo, and

fragm. ex herb. Berol.). Western slope of Mt. Chimborazo, 3,000-

3,400 m., Rimbach 4 (F).

9. ALLIONIA L.

Prostrate annual or perennial herbs; leaves opposite, petiolate,

those of a pair unequal, the blades entire or sinuate; flowers perfect,

in axillary clusters of 3, each subtended by a broad green bract,

the bracts cucullate, enclosing the fruit; perianth corolla-like, con-

stricted above the ovary, the limb oblique, 4-5-lobed; stamens 4-7,

the filaments unequal, exserted; anthocarp coriaceous, obovoid or

oval, strongly compressed, 3-costate or cristate on the inner surface,

the outer surface bearing 2 parallel longitudinal rows of stipitate

glands, the thin margins dentate or entire, inflexed.

The genus is represented in South America by a single variable

species. Two other species described from Mexico and the United

States may be distinct, for they are separated on characters which,

for the Nyctaginaceae, are fairly constant.

1. Allionia incarnata L. Syst. Nat. ed. 10. 890. 1759. A.

mendocina Phil. Sert. Mendoc. Alt. 41. 1870. Wedelia incarnata

Kuntze, Rev. Gen. 533. 1891. A. puberula Phil. Anal. Mus. Nac.

Chile Bqt. 1891: 71. 1891. A. Jarae Phil. op. cit. 72. 1891. A.

Bandurriae Phil. Anal. Univ. Chile 41: 274. 1895. Wedeliella in-

carnata Cockerell, Torreya 9: 167. 1909.

Perennial from a slender or thick, woody root, the stems short

or elongate, glandular-puberulent or viscid-villous; leaves on petioles

5-20 mm. long, the blades oval to deltoid-orbicular or oblong, 1-6

cm. long, subcordate or rounded and unequal at the base, rounded

to acute at the apex, paler beneath, glandular-puberulent or viscid-

villous; involucres numerous, on slender peduncles 5 cm. long or

less; bracts obovate-orbicular, 5-8 mm. long, rounded or obtuse at

the apex; perianth 7-15 mm. long, purple-red or rarely white; antho-

carp 3-4.5 mm. long, pale brown or olive, the inner side 3-costate,

the margins with 3-5 low broad teeth, or the teeth more numerous

and slender, the margins rarely entire.

NYCTAGINACEAE OF SOUTH AMERICA 105

Venezuela : Type from Cumana. Savannas of Lagunillas, Me"rida,

1,000 m., John 664 (W). Barquisimeto, in the cactus formation,

600 m., Pittier 6391 (W).— Bolivia: Bolivian Plateau, Bang 928 (F,

W). Rio Tapacani, 3,000 m., March 19, 1892, Kuntze (F). Tarija,

frequent in dry sandy places, Fries 1130 (W). — Argentina: Pampas,

January, 1892, Kuntze (F). Prov. Cordoba, Lossen 175 (F) .—

Chile: Desert of Atacama, Morong 1102 (F). Parca, Cordillera

Quebrada de Quipisca, Prov. Tarapaca, 2,500 m., Werdermann 1056

(F). Tarapaca, Philippi (F, photo, of type material of A. Jarae, ex

herb. BeroL). Prov. Atacama, Philippi (F, photo, of type material

of A. Bandurriae, ex herb. Berol.). Atacama, Philippi (F, photo, of

type material of A. puberula, ex herb. Berol.). The species ranges in

the drier regions northward to the southwestern United States.

The forms referred here are variable as to amount and quality of

pubescence and toothing of the anthocarp wings, but it seems quite

impractical to separate them into recognizable forms or varieties.

10. BOERHAAVIA L.

Annual or perennial herbs, sometimes somewhat frutescent and

subscandent, the stems often with viscous areas in the internodes;

leaves opposite, often unequal, petioled, the blades entire or sinuate;

flowers perfect, small, variously arranged, bracteate, the bracts

usually very small; perianth corolla-like, campanulate, nearly rotate,

or funnelform, the limb shallowly 5-lobate; stamens 1-5, exserted

or included, unequal ; anthocarp cylindric to obovoid or obpyramidal,

terete or 3-10-angulate, sometimes 3-5-winged, glabrous or pubes-

cent, sometimes furnished with stipitate glands.

Fruit terete or 10-ribbed, with stipitate glands near the apex; plants

large and often subscandent; perianth funnelform.

Perianth purple or purplish, 8-10 mm. long; anthocarp 6.5 mm.

long and 2 mm. thick; stamens 3 1. B. tuberosa.

Perianth greenish yellow, 5-8 mm. long; anthocarp 7-13 mm. long

and 1.5-2 mm. thick; stamens 2 2. B. scandens.

Fruit 3-5-angled or 3-5-winged, without stipitate glands; plants

usually low, erect or procumbent; perianth campanulate to

almost rotate.

Flowers in cymes; anthocarp truncate at the apex, glabrous;

perianth white or pink; plants annual 3. B. erecta.

Flowers in heads or glomerules; anthocarp not truncate at the apex,

often pubescent; perianth usually dark red; plants perennial.

Branches of the inflorescence glabrous; flowers mostly in clusters

of 2-4 4. B. coccinea.

Branches of the inflorescence puberulent or glandular-puberulent;

flowers chiefly in many-flowered heads 5. B. caribaea.

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

1. Boerhaavia tuberosa Lam. 111. Gen. 1: 10. 1791. B. excelsa

Willd. Phytogr. 1. 1794. B. scandens Choisy in DC. Prodr. 132: 454.

1849, in part, not L. B. litoralis HBK. Nov. Gen. & Sp. 2: 216. 1818.

Commicarpus tuberosus Standl. Contr. U. S. Nat. Herb. 18: 101. 1916.

Plants much branched, suberect or scandent, sometimes 2 m.

long, glabrous except at the nodes, pale; leaves thick and fleshy,

long-petiolate, the blades broadly ovate or deltoid-ovate, 3.5-5.5

cm. long, acutish to short-acuminate, often subcordate at the base,

glabrous or nearly so; flowers umbellate, the umbels numerous,

4-7-flowered, the pedicels long and slender but stiff; perianth gla-

brous or sparsely hirtulous above; stamens exserted; anthocarp

linear-clavate, truncate at the apex, bearing about 5 glands below

the apex.

Ecuador: James Bay, Galapagos Islands, Stewart 144-7 (W);

common in open woods at 255 rru Albemarle Island, abundant in

open places on the lower parts of the island, Stewart 14-42 (W).

Reported also from Charles, Indefatigable, and Chatham Islands.

Occurring in Peru.

This species as well as C. scandens belongs to a peculiar group of

the genus for which the writer has proposed generic rank, under the

name Commicarpus. The writer is still of the belief that the genus

is quite as good a one as are most of those segregated in the family,

but is willing to suppress it in order to aid in reducing, to as great

an extent as possible, the increasing number of genera. The genus

certainly is quite as good a one as Pisoniella.

2. Boerhaavia scandens L. Sp. PI. 3. 1753. Commicarpus

scandens Standl. Contr. U. S. Nat. Herb. 12: 373. 1909.

Plants large and usually clambering over other plants, suffrutes-

cent near the base, the branches pale, glabrous, obscurely puberulent

about the nodes; leaves thick and fleshy, on petioles 1-2 cm. long,

the blades broadly cordate-ovate to ovate-deltoid, 1.5-6.5 cm.

long, deeply cordate to truncate at the base, acute or obtuse at the

apex, slightly paler beneath, glabrous or when young obscurely puber-

ulent; flowers umbellate, the slender pedicels 5-10 mm. long, gla-

brous; perianth glabrous or rarely puberulent; stamens exserted;

anthocarp glabrous, bearing few or numerous glands irregularly dis-

posed along the costae.

Colombia: Region of Santa Marta, near sea level, H. H. Smith

571 (F, W). — Venezuela: Between Estanques and Puente Real,

Me*rida, 500 m., Pittier 12839 (W). Sabanas de Lagunillas, M<§rida,

1,000 m., Jahn 663 (W). Rangjng northward to the southern

borders of the United States, and widely dispersed in the West Indies.

The plant is said to be called "pegapega" in Curasao.

3. Boerhaavia erecta L. Sp. PL 3. 1753. B. virgata HBK. Nov.

Gen. & Sp. 2: 215. 1817. B. discolor HBK. loc. cit.

NYCTAGINACEAE OF SOUTH AMERICA 107

Plants annual, erect or decumbent, 1 m. high or less, usually

much branched at the base, the slender branches finely puberulent

below, the middle internodes often with brown viscous bands, the

upper ones glabrous or minutely puberulent; leaves slender-petiolate,

the blades broadly ovate-rhombic or deltoid-ovate, varying to oval

or oblong, 2-6 cm. long, truncate to rounded at the base, broadly

rounded to obtuse or rarely acute at the apex, pale beneath and

usually brown-punctate, glabrous or sparsely puberulent; flowers

on pedicels 1-5 mm. long; perianth 1-1.5 mm. long, glabrous, some-

times glandular-punctate; stamens 2-3, exserted; anthocarp narrowly

obpyramidal, 3-3.5 mm. long, 1-1.5 mm. wide, glabrous, 5-angled.

Venezuela: El Valle, Margarita Island, Miller & Johnston 204

(F, W). Between Caracas and La Guaira, Rose 21642 (W). Near

La Guaira, Otto 449 (W). — Colombia: Region of Santa Marta, 30

m., H. H. Smith 1321 (F, W). Barranquilla, Bro. PaulB-33 (W),

C-39 ( W) . In fields, Sevilla, Dept. Magdalena, Salt S ( W) . Hacienda

de Coloncito, near Turbaco, Dept. Bolivar, Killip & Smith 14378

(W). Cartagena, Bro. Heriberto 408 (W). Girardot, Dept. Cundina-

marca, 350-400 m., Rusby & Pennell 84 (W). — Ecuador: Charles

Island, Galapagos Islands, common to 180 m., Stewart 1439 (F, W).

Narborough Island, Galapagos, occasional in lava crevices, Stewart

1441 (W). Tagus Cove, Albemarle Island, Galapagos, common in

open sunny places in tufaceous soil on the lower parts, Stewart 1440

(F, W). Distributed as a weed through most of tropical America,

and extending to the southern United States.

The plant is one of the most abundant weeds of many parts of

tropical America. Boerhaavia virgata was described from Quetepe

and Cumana, Venezuela, and B. discolor from Guayaquil, Ecuador.

4. Boerhaavia coccinea Mill. Gard. Diet. ed. 8. Boerhaavia

No. 4. 1768. B. paniculata Rich. Act. Soc. Hist. Nat. Par. 1: 105.

1792. B. adscendens Willd. Sp. PI. 1: 19. 1797. B. decumbens Vahl,

Enum. PI. 1: 284. 1804. B. diffusa var. laxa Kuntze, Rev. Gen.

533. 1891.

Plants perennial from an often fusiform, fleshy root, the stems

few or numerous, ascending or procumbent, 1 m. long or less, minutely

puberulent below or often sparsely villous, especially at the nodes,

glabrous above; leaves petioled, somewhat fleshy, the blades rhombic-

orbicular to rhombic-ovate or oval, 2-5.5 cm. long, subcordate to

broadly rounded at the base, rounded or obtuse at the apex, pale

beneath, not punctate, glabrous or obscurely puberulent, rarely

villous along the veins; flowers subsessile in glomerules of 2-4 at

the ends of filiform glabrous peduncles 3-10 mm. long; perianth

2 mm. broad, minutely glandular-puberulent; stamens 2, short-

exserted; anthocarp narrowly obovoid, 3-4 mm. long, rounded at

the apex, densely glandular-puberulent or glandular-pilose, 5-sulcate.

French Guiana: Without locality, Melinon (F). — Surinam:

Groningen, May 10, 1916, Samuels 122 (F). — British Guiana:

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

Georgetown, sandy places, roadside, Persaud 163 (F). — Venezuela:

Ciudad Bolivar, 35 m., Holt & Gehriger 23 (F). El Valle, Margarita

Island, Miller & Johnston 203 (F). La Rubiera near Calabozo,

Guarico, Grisol 7 (W). La Guaira, Monte 1143 (W). Cristobal

Colon, on hills, Broadway 119 (W). In savannas, Santa Rosa de

la Tierra, near Maracaibo, Pittier 10687 (W). Barquisimeto, 400 m.,

Saer 590 (F). — Colombia: Santa Rosa to Cisneros, Dept. El Valle,

on banks, 250-350 m., Kittip 5377 (W). Region of Santa Marta,

on seashore, H. H. Smith 1323 (F, W). Canabetal, Dept. Bolivar,

100 m., in sand along river, Pennell 3876 (W). Near Bello, Antio-

quia, Archer 229 (F). — Ecuador: Caraques Bay, Anthony 97 (W).

Huigra, Rose & Rose 22626 (W). — Bolivia: Guanai to Tipuani,

Bang 1431 (F, W). Bolivian Plateau, Bang 957 (F, W). Junction

of rivers Beni and Madre de Dios, Rusby 904 (F, W). Tarija, in

campo arenoso, Fries 1150 (W). Milluguaya, Nord-Yungas, 1,300

m., Buchtien 4135 (W). — Paraguay: Central Paraguay, Morong 93

(F). Cordillera de Altos, Fiebrig 247 (F). Also in Peru, and widely

distributed in tropical America.

This is a common weedy plant of tropical America, but in some

regions, such as Central America, it is much less frequent than

B. erecta or B. caribaea.

4a. Boerhaavia coccinea Mill., var. leiocarpa (Heimerl),

comb. nov. B. paniculata f . leiocarpa Heimerl, Oesterr. Bot. Zeitschr.

56: 252. 1906. B. paniculata var. guaranitica Heimerl, loc. cit. 1906.

B. Friesii Heimerl, op. cit. 253. 1906. B. paniculata var. leiocarpa

Heimerl, Ann. Cons. Jard. Geneve 17: 225. 1913.

Like the species, but the anthocarp glabrous.

Colombia: Region of Santa Marta, 75 m., H. H. Smith 1320

(F, W). — Bolivia: Tarija, in rupibus siccis apricis, Fries 1206 (W,

type collection). Bermejo, Fries 2315 (F, photo, and fragm. ex herb.

Berol.).— Paraguay: Cordillera de Altos, Fiebrig 465 (F), 465a (F).—

Uruguay: Cerrito, Dept. Montevideo, Herter 61 A (F). Also in

Peru and Argentina.

I do not find any important or essential difference between Boer-

•haavia Friesii and the forms referred by Heimerl to his varieties

leiocarpa and guaranitica. They all agree in having a glabrous antho-

carp, but otherwise they differ little if at all from B. coccinea. It may

be that the lack of pubescence on the fruit is sufficient to give the form

specific rank but this is purely a matter of personal opinion.

5. Boerhaavia caribaea Jacq. Obs. Bot. 4: 5. 1771. B. poly-

morpha Rich. Act. Soc. Hist. Nat. Par. 1: 185. 1792. B. hirsuta Willd.

Phytog. 1: 1. 1794. B. viscosa Lag. & Rodr. Anal. Cienc. Nat. 4: 256.

1801. B. glandulosa Anderss. Svensk. Vet. Akad. Handl. 1853: 171.

1854. B. diffusa var. hirsuta Kuntze, Rev. Gen. 533. 1891. B. diffusa

var. viscosa Heimerl, Beitr. Syst. Nyct. 27. 1897. B. patula Domb. ex

Vahl, Enum. PL 1: 287. 1805.

NYCTAGINACEAE OF SOUTH AMERICA 109

Plants perennial, from a thick woody root, the stems decumbent

or prostrate, below viscid-puberulent and often hirsute or villous,

above densely glandular-puberulent or often merely puberulent and

slightly viscid; leaves thick, petiolate, the blades suborbicular to oval

or oblong, 1.5-5.5 cm. long, truncate to rounded at the base, broadly

rounded or obtuse at the apex, paler beneath and sometimes brown-

puncticulate, glabrous or often puberulent or densely villous or

hirsute; flower heads axillary or usually in open cymes, the branches

of the inflorescence puberulent or glandular-puberulent, the flowers

sessile or short-pedicellate; perianth purplish red, 2 mm. broad, puber-

ulent or glandular-puberulent; stamens 1-3, short-exserted ; anthocarp

narrowly obovoid, 2.5-3 mm. long, 5-sulcate, densely glandular-

puberulent.

Venezuela: San Pablo de Mendoza, Trujillo, in fields, Pittier

13311 (W). In cactus formation, Cabo Blanco, D. F., Pittier 10239

(W). El Sombrero, Guarico, Pittier 11457 (W). Macuto, L. H. &

E. Z. Bailey 1204 (W). La Trinidad de Maracay, Aragua, 440 m.,

Pittier 5761 (W). La Guaira, Moritz 1143 (F). Hacienda El Volcan,

near Santa Lucia, Miranda, Pittier 8248 (W). Valencia, Carabobo,

in savannas, 400-800 m., Pittier 9029 (W). — Colombia: Dagua,

El Valle, 700-900 m., Pennell 5643 (W). Barranquilla, Bro. Paul

C-26 (W); Bro. Elias 236 (W). Rio Frio, Dept. Magdalena, Salt F

(W). Region of Santa Marta, 45 m., H. H. Smith 1499 (F, W).

Santa Rita, Andre 4183 (W). Without locality (possibly from Ecua-

dor), Lehmann 4712 (W). Cartagena, Bro. Heriberto 379 (W).—

Ecuador: Duran, Rose & Rose 23605 (W). Without locality, Eggers

14928 (F). Milagro, Prov. Guayas, 50 m., Hitchcock 20208 (W).

Oil camp between Guayaquil and Salinas, Prov. Guayas, Hitchcock

20009 (W). Caraques Bay, Anthony & Tate 107 (W). Daphne

Island, Wheeler, Rose & Beebe 9 (W). North Indefatigable Island,

Galapagos, Snodgrass & Heller 670 (W). Charles Island, Galapagos,

April 8, 1888, Lee (W); beach to 180 m., in light tufaceous soil,

Stewart 1452 (W). Hood Island, Galapagos, on hillsides at 75 m.,

Stewart 1456 (W). — Bolivia: Cotana am Ilimani, 2,400 m., Buchtien

3169 (W). Also in Peru; generally distributed in tropical America.

This species is one of the most abundant weeds of waste and

cultivated ground in the lowlands of most parts of tropical America.

Through its wide range it varies greatly, at least as regards the

amount and nature of the pubescence, but it seems impractical to

recognize any well-defined varieties or forms.

The vernacular name "tripa de gallina" is reported from Colombia.

Boerhaavia glandulosa was described from the Galapagos Islands.

11. MIRABILIS L.

Perennial herbs, erect or procumbent, often viscid-pubescent, the

branches commonly swollen at the nodes; leaves opposite, petiolate or

sessile, the blades entire or undulate; flowers perfect, involucrate, the

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

involucre 1-several-flowered, 5-lobate, in fruit often accrescent and

becoming almost rotate; perianth corolla-like, tubular to campanulate,

often oblique, the limb 5-lobed, the perianth soon withering and

deciduous; stamens 3-5, unequal, usually exserted; anthocarp cori-

aceous, smooth or 5-angled or 5-sulcate, often constricted at the base,

mucilaginous when wet.

The genus as here treated includes Oxybaphus and several other

groups that have been treated at one time or another as distinct.

If only the species of North America are considered, such genera as

Oxybaphus, Quamoclidion, and Hesperonia seem to be differentiated

by good and constant characters; but, as so often happens, when

extralimital species are taken into account, the characters supposed

to separate the groups break down. It seems necessary, therefore,

to follow Heimerl in considering all the plants of the group as repre-

senting a single genus, Mirabilis.

Perianth very large, 3-5.5 cm. long 1. M. Jalapa.

Perianth small, less than 2 cm. long.

Anthocarp sometimes pubescent, constricted at the base, 5-angu-

late or 5-sulcate; involucre accrescent after anthesis.

Stems densely viscid-villous throughout; leaf blades thick, nearly

all deeply cordate at the base; involucre strongly accrescent

in fruit and with almost entire margin 2. M. viscosa.

Stems glabrous or glabrate below; leaf blades thin, mostly trun-

cate or only shallowly cordate at the base; involucre slightly

accrescent in fruit and remaining deeply lobed.

3. M. violacea.

Anthocarp glabrous, not constricted at the base, terete; involucre

scarcely at all accrescent after anthesis.

Leaves thin, the lower ones on long slender petioles and even the

uppermost usually with conspicuous petioles, the blades

mostly truncate or shallowly cordate at the base.

4. M. prostrata.

Leaves rather thick, the lower ones on comparatively short and

thick petioles, the upper mostly sessile or nearly so, the

blades mostly rounded to acute at the base . . 5. M. expansa.

1. Mirabilis Jalapa L. Sp. PI. 177. 1753. M. odorata L. Cent.

PI. 1: 7. 1755. M. dichotoma L. Sp. PI. ed. 2. 252. 1762.

A stout bushy-branched perennial 1 m. high or less, the branches

glabrous, puberulent, or rarely short- villous; leaves slender-petiolate,

the blades ovate-deltoid to lance-oblong, 5-14 cm. long, subcordate

to rounded at the base, acute to acuminate at the apex, glabrous or

NYCTAGINACEAE OF SOUTH AMERICA 111

rarely puberulent, usually ciliate; flowers in dense clusters at the ends

of the branches; involucre short-pedunculate or subsessile, 7-15 mm.

long, glabrous, puberulent, or short-villous, the lobes linear-lanceolate

to lance-ovate, longer than the tube, acute or acuminate; perianth

3-5.5 cm. long, purplish red or white, yellow, or variegated; stamens

5; anthocarp 7-9 mm. long, 5-angled, verrucose or rugose, dark

brown or black, glabrous or puberulent.

British Guiana: Kamakusa, upper Mazaruni River, De La Cruz

4182 (F). Hyde Park, Demerara, in 1922, G. B. Warren (F).— Vene-

zuela: Caracas, Rose 21626 (W). — Colombia: Region of Santa Marta,

30 m., H. H. Smith 1324 (F).— Bolivia: Yungas, Bang 480 (F).

Chiquiaca, 1,000 m.,Fiebrig 2702 (F). — Paraguay: Central Paraguay,

Morong 622 (F).— Argentina: Prov. Cordoba, Lossen 277 (F), 278 (F).

This species, the common four-o'clock or marvel of Peru, is well

known to horticulturists, being cultivated commonly in most tropical

and even in temperate regions. It is one of the plants most frequently

seen in the gardens of the United States, and thrives in almost any

kind of soil or under neglect. The very sweet-scented flowers open

in late afternoon and close the following morning. There are endless

color varieties, those in which the perianth is striped in two or more

colors being especially striking.

Although the plant is doubtless a native of America, it is not

known where it grew originally. At the present time it probably is

not found anywhere in a truly wild state, although often it persists

in waste ground about settlements, sometimes as an abundant weed.

In these respects it is like numerous other American plants that are

known only in a cultivated state. No doubt the four-o'clock has been

in cultivation for hundreds of years, and it may well have originated

in Mexico, whose inhabitants had a great love for flowers. In the

older botanical works the plant is assumed to be a native of Peru,

but the fact that its nearest relatives grow wild in Mexico, while none

at all occur in Peru, points to its nativity in Mexico

Among the vernacular names reported are the following: "Jazmin

Colorado" (Venezuela); "maravilla," "buenas tardes," "buenas

noches," "Don Pedro de noche," "cuambu" (Argentina); "dengue"

(Chile); "maravilha," "boa noite," "bonina," "purga de nabica"

(Brazil).

2. Mirabilis viscosa Cav. Icon. 1: 13. pi. 19. 1791. Nyctago

parviflora Salisb. Prodr. 57. 1796. Calyxhymenia viscosa R. & P.

Fl. Peruv. 1: 46. 1798. Calymenia viscosa Pers. Syn. PI. 1: 36. 1805.

Vitmania viscosa Turra ex Steud. Nom. Bot. 140, in syn. 1821.

Oxybaphus viscosus L'He>. ex Choisy in DC. Prodr. 132: 430. 1849.

Allionia viscosa Kuntze, Rev. Gen. 533. 1891.

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

A coarse herb, 0.5-1.5 m. high or even larger, the stems some-

times as much as 5 cm. thick, the branches densely viscid-pilose;

leaves rather thick, long-petiolate, the blades broadly cordate-ovate

or ovate-deltoid, 3-10 cm. long, usually cordate and abruptly short-

decurrent at the base and acute to acuminate at the apex, short-

villous or puberulent beneath and often also on the upper surface,

the leaves of the inflorescence reduced and bractlike; inflorescence

large, openly paniculate, the branches chiefly opposite, densely viscid-

villous; involucres at anthesis 3-5 mm. long, in fruit 15-25 mm. broad,

densely viscid-pilose; flowers solitary or rarely 2-3 in the involucre,

the perianth 8-20 mm. long, purplish red, pink, or white, sparsely

pilose, the limb 15-25 mm. broad; stamens 3; anthocarp obovoid,

5 mm. long, glabrous, densely covered with large coarse tubercles.

Colombia: Open limestone slopes below Dagua, Dept. El Valle,

700-900 m., Pennell 5652 (W).— Ecuador: Huigra, Rose & Rose 22146

(W). Also in Peru and Mexico.

3. Mirabilis violacea (L.) Heimerl, Beitr. Syst. Nyctag. 23.

1897. Allionia violacea L. Syst. Nat. ed. 10. 890. 1759. Oxybaphus

violaceus Choisy in DC. Prodr. 132: 432. 1849. 0. violaceus var.

parviflorus Choisy, loc. cit. Allionia craterimorpha Rusby, Descr.

N. Sp. S. Amer. PI. 15. 1920.

Plants annual or perennial, much branched from the base, the

slender branches usually procumbent, bifariously puberulent or gla-

brate, sometimes sparsely viscid-villosulous; leaves on long slender

petioles, the blades broadly ovate-deltoid, rarely elongate-deltoid,

3-11 cm. long or larger, subcordate or truncate at the base, acuminate

or long-acuminate at the apex, rarely obtuse, thin, bright green,

sparsely puberulent or short-pilose or glabrate; inflorescence cymose,

usually dense and congested, or in age open, often leafy, the very

slender branches viscid-pilose; involucres 3 mm. long in anthesis, in

fruit 5-6 mm. long, green, viscid-pilose, the lobes triangular-ovate,

unequal, mostly acute or acuminate; perianth purplish red, 6-8 mm.

long, viscid-pilose or glabrate; stamens usually 3; anthocarp obovoid,

3.5-4 mm. long, dark brown or blackish, short-pilose, sparsely

tuberculate.

Venezuela: La Victoria, John 212 (W). Lower Catuche Wood

above Caracas, 1,000-1,200 m., Pittier 7203 (W), 7127 (W). La

Trinidad de Maracay, Aragua, 440 m., Pittier 5762 (W). Between

Antimano and Las Adjuntas, D. F., Pittier 12410 (W). Los Chorros,

900 m., Eggers 13042 (W). Hacienda de las Cuadras near Caracas,

Eggers 13007 (W). Near Caracas, 900-1,150 m., L. H. & E. Z. Bailey

713 (W).— Colombia: Near Bonda, 45 m., H. H. Smith 569 (F, W,

type collection of Allionia craterimorpha). Between Surata and

California, Santander, in thicket, 1,740-2,000 m., Killip & Smith

16795 (W). California, Santander, open hillside, 2,200 m., Killip &

Smith 16855 (W). — Ecuador: Between Huigra and Naranjapata,

Prov. Chimborazo, 600-1,200 m., Hitchcock 20650 (W). Ambato,

Pachano74(W). Huigra, Rose, Pachano& Rose 23878 (W). El Recreo,

NYCTAGINACEAE OF SOUTH AMERICA 113

January 30, 1897, Eggers (F). Ranging northward to southern

Mexico.

The var. parviflorus was based on Moritz 216 from Colombia.

The following additional Venezuelan collections are reported by

Knuth (Fl. Venez. 317. 1928) : Palmar, D. F., 1,000 m., Wagener.

San Mateo, D. F., 800 m., Wagener. Nueva Barcelona, Bermudez,

Humboldt & Bonpland.

4. Mirabilis prostrata (R. & P.) Heimerl in E. & P. Nat. Pfl. 3,

Abth. Ib: 24. 1889. Calyxhymenia prostrata R. & P. Fl. Peruv. 1: 46.

pi. 75, f. c. 1798. Oxybaphus prostratus Vahl, Enum. PI. 2: 40. 1806.

0. micranthus Choisy in DC. Prodr. 132: 432. 1849. Mirabilis

micrantha Heimerl in E. & P. Nat. Pfl. 3, Abth. Ib: 24. 1889. M.

prostrata var. pubigera Heimerl, Bot. Jahrb. Engl. 34: Beibl. 78: 10.

1904. Allionia micrantha Molfino, Physis 7: 51. 1923.

Plants much branched, erect or procumbent, reported to be some-

times as much as 2 m. high, the slender branches puberulent, viscid-

villpsulous, or often glabrate, with elongate internodes; leaves slender-

petioled, thin, the uppermost short-petioled, the blades ovate or

ovate-deltoid, often elongate-deltoid, mostly 3-7 cm. long, commonly

truncate or subcordate at the base, acute to long-acuminate,

rarely obtuse at the apex, sparsely villosulous or puberulent or

more often glabrate; inflorescence cymose-paniculate, usually large

and open, often leafy-bracted; involucres 4-5 mm. long, narrow,

densely viscid- villosulous, the lobes narrowly triangular, mostly acute,

subequal, equaling or shorter than the tube; perianth purplish red or

pink, 7-10 mm. long, sparsely viscid-villosulous; stamens 3, usually

exserted but sometimes included; anthocarp obovoid, 3 mm. long,

glabrous, fuscous, nearly smooth.

Ecuador: Huigra, 1,200 m., Hitchcock 20312 (W); Rose & Rose

22407 (W), 22411 (W), 22296 (W). In sepibus interandinis prope

Riobamba, Mille 19 (W). Type of M. prostrata var. pubigera from

Quito, Sodiro 128-1. — Bolivia: La Paz, Camino a Obrajes, 3,550 m.,

Buchtien 377 (F), 140 (W). Cotaiia am Ilimani, 2,450 m., Buchtien

3168 (F, W), 262 (F). Cochabamba, March, 1892, Kuntze (F); at

3,000 m., March 26, 1892, Kuntze (F). Without locality, 3,000 m.,

April, 1892, Kuntze (F). Tarija, Fiebrig 2436 (F, fragm. ex herb.

Berol.). Cochabamba, Bang 1069 (F, W), 1070 (F, W). Near La

Paz, 3,000 m., Bang 36 (W); Claude-Joseph 1131 (W); Rusby 2868

(W). Near Yungas, 1,200 m., Rusby 2688 (F, W).— Argentina:

Quebrada de Choya, Prov. Catamarca, Schickendantz (F, photo, and

fragm. ex herb. Berol.). — Chile: Valparaiso, Stewart (F). Also in Peru.

The species was described from Peru, and 0. micranthus was based

on Bertero 848 from Quillota, Chile.

5. Mirabilis expansa (R. & P.), comb. nov. Calyxhymenia

expansa R. & P. Fl. Peruv. 1: 45. pi. 75, f. a. 1798. Oxybaphus

expansus Vahl, Enum. PI. 2: 41. 1806. Allionia expansa Kuntze,

Rev. Gen. 533. 1891.

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

A coarse herb, suberect or procumbent or even subscandent, the

stems sometimes 2.5 m. long but usually much shorter, the branches

rather stout, very sparsely puberulent or villosulous or practically

glabrous, the internodes mostly longer than the leaves; leaves rather

thick, the petioles stout and relatively short; leaf blades varying from

ovate-rounded to rhombic-ovate, broadly ovate, elliptic, or oblong-

ovate, mostly 2.5-5 cm. long, truncate to acute at the base, often

abruptly decurrent, very rarely cordate, obtuse to rounded at the

apex, the uppermost leaves sometimes acute, puberulent, pilosulous

or glabrate; inflorescence cymose-paniculate, usually rather sparse,

the involucres mostly in small dense clusters; involucre 1-flowered,

4-5 mm. long, not accrescent, densely viscid-villosulous, sometimes

densely covered with short hairs tipped with red glands, the lobes

ovate- triangular, obtuse or acute, much shorter than the tube;

perianth red-purple, 6 mm. long, sparsely puberulent or glabrous;

stamens 3, usually not exserted; anthocarp ellipsoid, 3 mm. long,

obtuse at each end, dark olivaceous, glabrous, almost smooth.

Venezuela: Mucuruba, 2,500 m., on stream bank, Gehriger 187 (F).

— Ecuador: Alausi, Prov. Chimborazo, 2,500 m., Hitchcock 20703

(W), 20702 (W). Ambato, Pachano 34 (W). In fruticetis interandinis

ad Cotocollao, Mille 413 (W).— Chile: Valparaiso, November, 1924,

Behn (F). Also in Peru, from which country the species was

described.

The species is a somewhat variable one, and not sharply distinct

from M. prostrata. The specimen from the isolated Venezuelan

locality is remarkable for the abundant hairs with bright red glands

at the tip covering the involucres and peduncles, but similar pubes-

cence is found also in material from other regions. Some of the

Ecuador specimens are noteworthy for their almost reniform, small

leaves. Although they are appreciably different in appearance from

Peruvian and other collections, they do not seem to be clearly

separable from them.

THE CHENOPODIACEAE OF NORTHWESTERN

SOUTH AMERICA

PAUL C. STANDLEY

The species of Chenopodiaceae occurring in northwestern South

America are so few, and of so little general interest, that it is ques-

tionable whether it is worth while to publish an enumeration of them.

The only genera, indeed, with any considerable number of species

in the area under consideration are Chenopodium and Atriplex, and

several of the representatives of the former group are widely dispersed

weeds.

The region covered embraces Venezuela, Colombia, Ecuador, and

Bolivia, treatment of the Peruvian Chenopodiaceae being reserved

for publication elsewhere. It has been found necessary to describe

but one new species, an Atriplex. While few further plants of this

somewhat "weedy" family may be expected in northwestern South

America, it would not be surprising if a few additions in Atriplex and

Chenopodium should appear in later collections.

The present paper is based upon the material in the Herbarium of

Field Museum, and upon that of the United States National Museum,

lent for study through the courtesy of Dr. William R. Maxon and

Mr. Ellsworth P. Killip.

KEY TO THE GENERA

Embryo spirally coiled. Flowers all axillary; leaves terete or semi-

terete, very fleshy and succulent 1. Suaeda.

Embryo not spirally coiled.

Flowers in fleshy spikes or sunken in the joints of the stems; leaves

much reduced and scalelike.

Flowers in conelike spikes, the bracts free and deciduous.

2. Heterostachys.

Flowers sunken in the joints of the stems, the bracts united and

persistent 3. Salicornia.

Flowers usually glomerate, or solitary in the axils; leaves usually

well developed.

Flowers usually perfect, ebracteolate 4. Chenopodium.

Flowers unisexual, the pistillate ones subtended by 2 bracteoles

which enlarge in age and enclose the fruit 5. Atriplex.

115

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

Besides the genera listed formally, there are cultivated in the

region under consideration the beet, Beta vulgaris L., and spinach,

Spinacia oleracea L.

1. SUAEDAForsk.

Annuals or perennials, erect or prostrate, glabrous or pubescent,

herbaceous or suffrutescent; leaves alternate, terete or semi terete,

rarely flat, entire, fleshy; flowers minute, chiefly perfect, solitary or

glomerate in the leaf axils; perianth fleshy, 5-lobed; stamens 5; fruit

a compressed or depressed utricle, enclosed in the perianth; seed

horizontal or erect, smooth or roughened.

1. Suaeda foliosa Moq. in DC. Prodr. 132: 156. 1849.

Apparently perennial, glabrous or nearly so, much branched, the

stout branches roughened by the persistent leaf bases of fallen leaves;

leaves mostly 5-8 mm. long, glaucous, very thick and fleshy, obtuse

or acutish, semiterete, glabrous; flowers minute, green, solitary or in

clusters of 3.

The species occurs in Peru and Chile. The only report of the genus

Suaeda from the countries here covered is the publication of Suaeda

fruticosa var. crassifolia Moq. (in DC. Prodr. 132: 157. 1849), which

was based upon a specimen collected in Bolivia by D'Orbigny. No

Bolivian material has been seen by the writer, but it seems probable

that any Suaeda occurring there will prove to be S. foliosa, although

there is a possibility that it might be the Argentine S. divaricata Moq.

2. HETEROSTACHYS Ung.-Sternb.

Tall erect much-branched shrubs with stiff woody branches;

leaves minute, opposite or subopposite, free, suborbicular, obtuse,

closely imbricate; flowers perfect, solitary in the axils of opposite

deciduous bracts, the flower spikes resembling small cones; perianth

orbicular, membranaceous, complanate, broadly winged on each side,

4-lobate; stamens 2; utricle compressed, oblong-obovoid; seed ellip-

soid, compressed, smooth.

The genus consists of a single species.

1. Heterostachys Ritteriana (Moq.) Ung.-Sternb. Atti Congr.

Bot. Firenze 332. 1876. Halocnemum Ritterianum Moq. Chenop.

Enum. 109. 1840. Halostachys Ritteriana Moq. in DC. Prodr. 132:

148. 1849. Spirostachys Ritteriana Ung.-Sternb. Vers. Syst. Salicorn.

100. 1866.

A slender, densely branched, glabrous shrub; leaves about 1.5 mm.

long and broad, closely overlapping; flower spikes sessile or short-

pedunculate, 4-12 mm. long, obtuse, many-flowered; bracts broader

than long, concave, clasping, very obtuse.

CHENOPODIACEAE OF SOUTH AMERICA 117

Colombia: Wind-swept plains of Rio Hacha, one-half mile back

from the coast, February 6, 1914, J. G. Sinclair (W). — Venezuela:

La Vela de Coro, Curran & Haman ^2 (W). — Argentina: Totorale-

jas, November, 1892, Kuntze (F). Also in Hispaniola.

3. SALICORNIA L.

Annuals or perennials, sometimes suffrutescent, fleshy, glabrous,

with jointed branches, the joints dilated at the apex into a short

sheath; flowers perfect or polygamous, immersed in groups of 3-7 on

opposite sides of the joints, the flowering joints forming cylindric

terminal spikes; perianth obpyramidal, fleshy, 3^4-dentate; stamens

1-2; fruit a minute utricle, included in the perianth; seed erect, com-

pressed, minutely hairy.

1. Salicornia fruticosa L. Sp. PL ed. 2. 5. 1762. S. peruviana

HBK. Nov. Gen. & Sp. 2: 193. 1818. S. Gaudichaudiana Moq.

Chenop. Enum. 115. 1840. S. biloba Kunze ex Fenzl in Mart. Fl.

Bras. 51: 158. 1864, as syn.

An erect or prostrate perennial, suffrutescent at the base, much

branched, the secondary branches ascending or erect, usually branched,

the joints mostly 1-2 cm. long; sheaths rounded or with acutish

lobes; flower spikes about 2 cm. long and 3 mm. thick, the flowers in

groups of 3; seed yellowish brown, covered with short conic hairs.

Venezuela: Paraguana Peninsula, Curran & Haman 557 (W).—

Ecuador: Between Guayaquil and Salinas, Prov. Guayas, Hitchcock

19996 in part (W; part of this number isBatis maritima L.). — Bolivia:

Pazna, 4,000 m., salt-steppes, Buchtien 1385 (W; material immature

and imperfect and determination therefore uncertain). — Chile: Desert

of Atacama, Morongll59 (F). Arauco, salt marsh, Pennell 12931 (F).

Ancud, Prov. Chiloe, moist sand in rock crevices on beach, Pennell

12579 (F). Widely dispersed on seashores of both hemispheres.

4. CHENOPODIUM L.

Annual or perennial herbs, rarely suffrutescent, often strong-

scented, usually either glandular or covered with a meal-like pubes-

cence of small whitish inflated hairs; leaves alternate, mostly petiolate,

entire to pinnatifid; flowers chiefly perfect, small and green, usually

glomerate; perianth normally 5-lobate; stamens 5 or fewer; fruit an

utricle, erect or depressed, the pericarp free from or adherent to the

seed; seed horizontal or vertical.

The American species of the genus have been treated in detail

by Aellen, Repert. Sp. Nov. 26: 31-64, 119-160. 1929. That author

has given an excellent account of the synonymy of the American

species, with citation of specimens available to him for study.

Seed vertical 1. C. macrospermum.

Seed horizontal.

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

Embryo not completely encircling the endosperm; leaves and

inflorescence bearing numerous glands.

Pericarp gland-dotted ; flowers in spikes 2. C. ambrosioides.

Pericarp not gland-dotted; inflorescence dichotomous, some of

the flowers pediceled 3. C. incisum.

Embryo completely encircling the endosperm; plants without

glands.

Seed white or nearly so 4. C. quinoa.

Seed black or blackish.

Leaves shining on the upper surface, green but sparsely fari-

nose. Inflorescences chiefly axillary and rather few-

flowered, lax 5. C. murale.

Leaves dull on the upper surface.

Plants low, 50 cm. high or less, with spreading branches;

leaves small, 2 cm. long and broad or less, shallowly

or rather deeply 3-lobed.

Seed irregularly punctate 6. C. pallidicaule.

Seed smooth or nearly so, not punctate . . 7. C. carnosulum.

Plants tall and erect; leaves commonly much more than

2 cm. long.

Leaves chiefly deltoid, entire except for the hastate lobes

at the base, commonly obtuse, finely farinose, usually

pale; seed scarcely 1 mm. broad 8. C. petiolare.

Leaves mostly rhombic and sinuate-dentate; seed 1.3-1.8

mm. broad.

Mature seed deeply punctate; plants ill-scented.

9. C. hircinum.

Mature seed minutely puncticulate or almost smooth;

plants without a distinctive odor . . . 10. C. album.

1. Chenopodium macrospermum Hook, f., var. halophilum

(Phil.), comb. nov. C. halophilum Phil. Anal. Univ. Chile 18: 67. 1861.

C. murale var. farinosum Wats. Proc. Amer. Acad. 9: 97. 1874.

C. farinosum Standl. N. Amer. Fl. 21: 28. 1916. C. macrospermum

subsp. halophilum Aellen, Repert. Sp. Nov. 26: 42. 1929. C. macro-

spermum subsp. halophilum f. subviride Aellen, Repert. Sp. Nov. 26:

43. 1929. C. macrospermum subsp. halophilum f. farinosum Aellen,

loc. cit. C. macrospermum subsp. halophilum f. latifolium Thellung &

Aellen, loc. cit. C. macrospermum subsp. halophilum f. angustius

Thellung & Aellen et f. nanum Aellen, op. cit. 44. 1929.

An annual, branched from the base, suberect or spreading, some-

times 50 cm. high but often much reduced, with branches only 2-4

CHENOPODIACEAE OF SOUTH AMERICA 119

cm. long; leaves petiolate, rhombic or deltoid-rhombic, 1-6 cm. long,

green and glabrate on the upper surface, usually densely whitish-

farinose beneath, sinuate-dentate to subentire; flowers spicate, the

spikes axillary and forming narrow dense leafy terminal panicles; seed

dark reddish brown, 1 mm. long.

Bolivia: Guaqui, Lake Titicaca, 3,820 m., Buchtien 2828 (W).

Near La Paz, 3,000 m., Bang 199 (F, W).— Chile: Arauco, at sea

level, salt marsh, Pennell 12919 (F). — Argentina: General Roca, Rio

Negro, 250-360 m., W.Fischer 276 (F).— Paraguay: Rio Pilcomayo,

Morong 918 (F). Also in Mexico and southern California, and ad-

ventive at various places in the United States and in Europe. The

typical form of the species occurs in the Falkland Islands.

The species is a variable one in stature and leaf form, but the

forms to which names have been given seem to be of little significance.

The plant has been confused by various authors with C. rubrum L.

and C. glaucum L. In general appearance it suggests a large and

overgrown plant of the latter species. Aellen lists the Chilean

Blitum salsum Phil, as a synonym of var. halophilum.

2. Ghenopodium ambrosioides L. Sp. PI. 219. 1753. C.

anthelminticum L. Sp. PI. 220. 1753.

An ill-scented annual or perennial, erect or ascending, usually

much less than 1 m. high, conspicuously glandular, commonly more

or less villous or puberulent; lower leaves petiolate, narrow, sinuate-

dentate or sinuate-pinnatifid, the upper ones mostly entire or nearly

so; flowers densely glomerate, forming slender or stout, naked or leafy,

often much elongate spikes.

Colombia: Mutiscua, Norte de Santander, 2,900 m., Killip &

Smith 19665 (F). Santa Marta, 750 m., H. H. Smith 542 (F). High-

lands of Popayan, 1,600-2,000 m., Lehmann BT543 (F).— Bolivia:

Buenavista, Dept. Santa Cruz, 500 m., Steinbach 5135 (F). Hacienda

Casana, Tipuani Valley, 1,400 m., Buchtien 7281 (W). Yungas, Bang

281 (F). La Paz, 3,400 m., Buchtien 4137 (F).— Argentina: In distr.

urb. Posadas, Misiones, Lillieskold (F). General Roca, Rio Negro,

250-360 m., W. Fischer 240 (F). Siambon, Sierra de Tucuman,

Hieronymus 763 (F). Cuesta del Cerro, Sierra Achala de Cordoba,

Hieronymus 533 (F). La Cie*naga, Sierra de Tucuman, Lorentz

& Hieronymus 718 (F). Prov. Cordoba, Lossen 160 (F), 207 (F).—

Chile: Desert of Atacama, Morong 1156 (F). — Uruguay: Casavalle,

Dept. Montevideo, Herter 55a (F). Widely distributed as a weed in

tropical America; naturalized in the United States and in Europe.

Called "paico" in the Amazonian region of Peru. The name

"baico," evidently a corruption or misspelling of paico, is recorded

from Colombia by Killip, who reports the plant as a remedy for

"tropical anemia." Its seeds are, of course, a well-known remedy

for intestinal parasites.

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

2a. Chenopodium ambrosioides L., subsp. chilense (Schrad.)

Aellen, Repert. Sp. Nov. 26: 36. 1929. C. chilense Schrad. Ind. Sem.

Hort. Goett. 1832: 2. 1832. C. vagans Standl. N. Amer. Fl. 21: 26.

1916. C. querciforme Murr, Mag. Bot. Lap. 3: 1. 1904.

Similar to the species; leaves usually smaller and more deeply

incised, even the uppermost deeply sinuate-lobate; spikes usually

elongate, very dense, and naked or nearly so.

Bolivia: La Paz, 3,700 m., open stony banks, Pennell 14258 (F);

3,000 m., Bang 52 (F). — Chile: Arauco, at sea level, sandy knolls

along salt marsh, Pennell 12934 (F). Alto del Carmen, Vallenar, Prov.

Atacama, 800 m., Werdermann 155 (F). Reported from many local-

ities in Chile; Santa Catharina, Brazil; naturalized in California.

3. Chenopodium incisum Poir. in Lam. Encycl. Suppl.

1: 392. 1811. Teloxys Mandoni Wats. Proc. Amer. Acad. 9: 91.

1874. C. incisum var. Bangii Murr, Bull. Herb. Boiss. II. 4: 991.

1904. C. rigidum Lingelsh. Repert. Sp. Nov. 7: 241. 1909.

An erect annual, strong-scented, 60 cm. high or less, simple or

branched from the base, the branches sparsely puberulent or glabrate;

leaves slender-petiolate, the blades deltoid-ovate to oblong, sinuate-

pinnatifid or laciniate-pinnatifid, bearing numerous yellow glands

beneath; inflorescence of numerous loosely few-flowered, axillary

cymes; flowers sessile in the forks of the branches and solitary at

the ends of the slender lateral branches, the pedicellate flowers

usually abortive, their pedicels spinose; seed 0.5-0.8 mm. broad,

dark brown.

Bolivia: La Paz, 3,500 m., Buchtien 33 (F). Talca Chugiaguillo,

Bang 799 (F, W). La Paz, 3,000 m., Bang 3 (W). Pazfia, 4,200 m.,

Buchtien 1383 (W). Cotana am Ilimani, 3,450 m., Buchtien 6305

(W). Cochabamba, Bang 1004 (W; type collection of var. Bangii).

La Paz, am Wege nach Obrajes, 3,500 m., Buchtien 4562 (W). Valle,

Prov. Cercado, Dept. Cochabamba, 2,750 m., Steinbach 9710 (F).

Also in Argentina and Peru, and ranging from Mexico to the

southwestern United States.

Chenopodium rigidum was based upon Bolivian material. The

type of Teloxys Mandoni is Mandon 1026 from Bolivia.

4. Chenopodium quinoa Willd. Sp. PI. 1: 1301. 1797. C.

purpurascens Jacq., var. punctulatum Moq. in DC. Prodr. 132: 67.

1849. C. Nuttalliae Safford, Journ. Wash. Acad. Sci. 8: 523. 1918.

C. quinoa f. purpureum Aellen, Repert. Sp. Nov. 26: 124. 1929.

A coarse erect annual, often 1 m. high or more, sparsely and finely

farinose, green or somewhat purplish (f . purpureum) ; leaves slender-

petiolate, the blades large, broadly rhombic, sinuate-dentate, some-

times obscurely lobate at the base; inflorescence erect, stout and

dense, compact, very leafy; seeds whitish, about 1.5 mm. broad.

Ecuador: Ambato, Pachano 40 (W); Rose & Rose 22355 (W).—

Bolivia: La Paz, 3,750 m., Buchtien 552 (F, W), 516 (F). Polo-Polo,

CHENOPODIACEAE OF SOUTH AMERICA 121

North Yungas, 1,100 m., Bucktien 6304 (W). Yungas, Bang 281 (W).

La Paz, Claude- Joseph 1172 (W). Also in Peru and Argentina.

"Quinoa" (Bolivia, Peru). The plant is cultivated commonly in

the Andes for its edible seeds.

5. Chenopodium murale L. Sp. PI. 219. 1753.

An erect or ascending annual, 60 cm. high or less, deep green

but sparsely farinose, often much branched from the base; leaves

slender-petiolate, the blades ovate or ovate-rhombic, 3-8 cm. long,

sinuate-dentate; flowers in small glomerules, these arranged in lax

or dense, chiefly axillary, mostly leafless cymes or panicles; seed

1.2-1.5 mm. broad, dull, finely puncticulate.

Curacao: In 1917, Curran & Haman 12 (W). — Colombia: Pam-

plona, Norte de Santander, 2,300 m., in field, Killip & Smith 20744

(W). Mutiscua, Norte de Santander, 2,900 m., Killip & Smith

19664 (W).— Venezuela: Caracas, 900 m., Bailey 19 (W); "a com-

mon weed." Between Antimano and Las Adjuntas, Distrito Federal,

Pittier 12263 (W). Timotes, MeYida, 2,000 m., waste places, Pittier

12691 (W). El Colorado, Tacagua Valley, Distrito Federal, Pittier

11102 (W). Petare, Miranda, 800 m., Pittier 9691 (W). Without

locality, Curran & Haman 735 (W). — Ecuador: Ambato, Pachano

23 (W). — Bolivia: Cotana, near Ilimani, 2,450 m., Buchtien 3161

(W). — Chile: Coquimbo, 20 m., Werdermann 129 (F). Desert of

Atacama, Morong 1155 (F). — Argentina: Cordoba, October 10,

1877, Hieronymus (F). — Brazil: Tijuca, Prov. Rio de Janeiro,

Glaziou 9566 (F). A native of Europe and other parts of the Old

World, but widely naturalized as a weed in temperate and tropical

regions of the Americas.

The vernacular name is reported from Curacao as "picante

sjimaron."

6. Chenopodium pallidicaule Aellen, Repert. Sp. Nov. 26:

126. 1929.

An erect or spreading annual, 60 cm. high or less, branched from

the base, with pale stems, sparsely farinose; leaves slender-petiolate,

the blades small, 1-2 cm. long and broad, trilobate, the lobes entire

or nearly so, the blades of the uppermost leaves sagittate or entire;

flower spikes short and dense, usually shorter than the leaves; seed

dark brown, minutely and irregularly punctate.

Bolivia: Comanche, 4,000 m., Buchtien 6296 (W). Also in the

mountains of Peru.

7. Chenopodium carnosulum Moq. in DC. Prodr. 132: 64.

1849.

An ascending or spreading, much-branched annual, 30 cm. high

or less, sparsely farinose when young, becoming glabrate; leaves

petiolate, the blades mostly 1 cm. long or less, entire or shallowly

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

trilobate, thick and fleshy; flowers in small dense axillary glomerules

shorter than the leaves; seed 1 mm. broad, nearly black.

Ecuador: Ambato, 2,600 m., Hitchcock 21730 (W).— Bolivia:

Uyuni, 3,700 m., Asplund 6295 (W). La Paz, Claude-Joseph 1115

(W). Also in Chile, Argentina, and southern Mexico.

8. Chenopodium petiolare HBK. Nov. Gen. & Sp. 2: 191.

1818. C. paniculatum Hook. Bot. Misc. 2: 237. 1831. C. sparsi-

florum Phil. Anal. Univ. Chile 91: 419. 1895. C. bolivianum Murr,

Mag. Bot. Lap. 1: 359. 1902. C. Collae Phil, ex Aellen, Repert. Sp.

Nov. 26: 148. 1929, as syn. C. parvulum Phil, ex Aellen, loc. cit.,

as syn. C. petiolare f. incanum Aellen, Repert. Sp. Nov. 26: 150.

1929. C. petiolare f . hastatum Aellen, loc. cit. C. petiolare L scutatum

Aellen, op. cit. 151. 1929. C. petiolare f. trilobum Aellen, loc. cit.

Plants erect or spreading, sometimes 1 m. high, slender and much

branched, pale, the branches rather densely and finely farinose; leaves

slender-petiolate, the blades thin, densely farinose or rarely green

and glabrate, very variable in outline, small, commonly more or

less deltoid and distinctly hastate-lobate at the base, otherwise entire

or remotely sinuate-dentate; inflorescence large and open, much

branched, the spikes slender and much interrupted, naked or with

few reduced leaves; seed black or blackish.

Ecuador: Type collected at Riobamba by Humboldt & Bonpland.

Ambato, Rose & Rose 22387 (W); Tate 537 (W).— Bolivia: La Paz,

3,000 m., Bang 56 (W; type collection of C. bolivianum); Rose 18905

(W); at 3,500 m., roadside, Pennell 14214 (F, W); at 3,500 m.,

Buchtien 2827 (W); at 4,000 m., Claude- Joseph 1173 (W). Without

locality, Bang 2897 (W). La Paz, 3,800 m., sunny slopes, Buchtien

144 (W). Pazna, 4,100 m., among rocks, Buchtien 1384 (W). Cotana

am Ilimani, 2,450 m., Buchtien 227 (F), 2827 (W).— Chile: Vina del

Mar, August 3, 1930, Behn (F). Tacna, 650 m., Werdermann 733

(F). Desert of Atacama, Morong 1230 (F). La Serena, Prov.

Coquimbo, 20 m., Werdermann 374 (F). Also in Peru.

The species has been reported from Bolivia as C. Fremonti Wats,

and as C. Fremonti var. incanum. The forms named by Aellen are

based upon leaf variations of slight systematic importance. Brother

Claude-Joseph reports from La Paz as the Aymara name "canagira."

9. Chenopodium hircinum Schrad. Ind. Sem. Hort. Goett.

1833: 2. 1833. C. quinoa var. orbicans Murr, Mag. Bot. Lap. 3: 37.

1904. C. hircinum subsp. eu-hircinum Aellen, Repert. Sp. Nov. 26:

120. 1929. C. hircinum subsp. eu-hircinum var. typicum Ludwig &

Aellen, op. cit. 121. 1929. C. hircinum subsp. eu-hircinum var.

rhombicum Aellen, op. cit. 122. 1929. C. hircinum subsp. Milleanum

Aellen, loc. cit.

Plants annual, tall and coarse, often 1 m. high or more, erect,

the pale stems more or less striate; leaves long-petiolate, large, broadly

rhombic, coarsely sinuate-dentate and often shallowly trilobate,

CHENOPODIACEAE OF SOUTH AMERICA 123

green but sparsely and minutely farinose; inflorescence very dense,

narrow, leafy, the branches short and erect or ascending.

Colombia: Eastern paramos of Guasca, toward Gacheta, Ariste-

Joseph in 1921 (W). Reported by Aellen from the Rio Magdalena,

Stubel 476.— Ecuador: Turubamba, Prov. Pichincha, 2,800 m., Fir-

min 228 (W). Type of subsp. Milleanum from Riobamba, Mille

6a. — Bolivia: La Paz, Bang 61 (F, W). Also in Peru, Chile, Argen-

tina, Paraguay, and Uruguay.

The local name is reported from Ecuador as "sacha-quinua."

10. Chenopodium album L. Sp. PI. 219. 1753.

An erect annual 1-2 m. high, usually much branched, pale green

and finely farinose; leaves petiolate, the blades broadly rhombic to

lanceolate, often shallowly trilobate, sinuate-dentate; inflorescence

large and open or small and condensed; seed 1.3-1.5 mm. broad,

nearly smooth, black and shining.

Ecuador: Reported by Aellen (Repert. Sp. Nov. 26: 129. 1929)

from Ambato, Pachano 38. A native of the Old World, but exten-

sively naturalized in North America. Reported in South America

also from Chile and Brazil.

5. ATRIPLEX L.

Shrubs or herbs, more or less furfuraceous, farinose, or canescent

with inflated hairs; leaves alternate or opposite; flowers monoecious

or dioecious, usually glomerate, the glomerules axillary or disposed

in solitary or panicled spikes; staminate perianth 3-5-parted; stamens

3-5; pistillate flowers bibracteolate, the bractlets accrescent, free or

united, enclosing the fruit, the perianth usually none; seed erect or

inverted, rarely horizontal.

In the countries of South America covered in the present paper

there are few species of Atriplex, but in Argentina and Chile the genus

is amply represented, although not so extensively as in western

North America.

Leaves sessile, cordate at the base 1. A. imbricata.

Leaves petiolate or, if subsessile, acute at the base.

Leaf blades rhombic or broadly ovate, broadest near the base,

undulate-lobate, long-petiolate; a tall shrub, 1-2 m. high.

2. A. Rusbyi.

Leaf blades oblong to elliptic or obovate, broadest at or above the

middle, entire or dentate, the petiole usually short and not

sharply differentiated from the blade; herbs or low shrubs.

Fruiting bracts not crested or tuberculate dorsally.

United portion of the bracts short-tubular, abruptly dilated

into the free portion 3. A. oestophora.

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

United portion of the bracts broadly obovoid, gradually

expanded into the free portion 4. A. Asplundii.

Fruiting bracts cristate or tuberculate dorsally.

Plants coarse and large, procumbent or ascending; leaves

mostly 10-15 mm. long, often dentate. . . 5. A. pentandra.

Plants small, prostrate, forming dense mats; leaves chiefly

4-7 mm. long, entire 6. A. Herzogii.

1. Atriplex imbricata (Moq.) D. Dietr. Syn. PL 5: 536. 1852.

Obione imbricata Moq. in DC. Prodr. 132: 108. 1849.

A low shrub with stout pale branches, densely furfuraceous-

canescent throughout; leaves thick, sessile, broadly deltoid-cordate,

as much as 1 cm. long but usually much smaller, obtuse or acutish,

entire; fruiting bracts axillary, about 2 mm. long, densely tuberculate

dorsally.

Bolivia: Chiguana, 3,700 m., Asplund 6297 (W). Uyuni, 3,700

m., Asplund 6298 (W). Type collected in Bolivia by D'Orbigny.

The two collections cited do not agree perfectly with the original

description, although they are the only specimens examined that can

possibly represent the species. Moquin describes the leaves as only

2-3 mm. long. The two Asplund collections show rather young

branches with decidedly larger leaves, but it may well be that mature

branches of the same plants would have minute leaves.

The specimens cited above evidently are close to Atriplex axillaris

Phil., a Chilean species, and also to A. microphylla Phil., of the same

country. The specimens of A. microphyUa that I have seen agree

perfectly with the original description of Obione imbricata.

2. Atriplex Rusbyi Britton, Mem. Torrey Club 4: 250. 1895.

A stout shrub 1-2 m. high, pale and densely furfuraceous through-

out; leaves thick, petiolate, the blades mostly rhombic or deltoid

and 2-3.5 cm. long, short-decurrent to truncate at the base, obtuse,

coarsely undulate-lobate or undulate, often crispate; flowers monoe-

cious, the staminate spikes axillary or forming terminal panicles;

fruiting bracts broadly flabelliform, 4-5 mm. long and somewhat

broader, obscurely dentate on the free margin, smooth or short-

tuberculate dorsally near the base.

Bolivia: Near La Paz, 3,000 m.,Bang 181 (F, W, type collection),

134 (W); 3,750 m., Buchtien 575 (F, W). Huaricana, 2,800 m.,

alkaline soil, Buchtien 145 (W). La Granja, 2,600 m., Julio 139 (W).

3. Atriplex oestophora Blake, Contr. Gray Herb. 53: 32. 1918.

Described as a shrub 60 cm. high, but probably merely suffru-

tescent or herbaceous, pale and minutely furfuraceous throughout,

copiously branched; leaves subsessile, obovate or oblong-obovate,

CHENOPODIACEAE OF SOUTH AMERICA 125

1.5-3 cm. long, entire, retuse at the apex, attenuate to the base;

flowers monoecious: bracts united to the middle, 4-5 mm. long, the

free portion broadly deltoid, obtuse, entire, 4-5 mm. wide.

Venezuela: Vela de Coro, Curran & Haman 451 (W, type

collection).

4. Atriplex Asplundii, sp. nov. — Herba perennis e radice lignoso

crasso, ramis gracilibus 10-25 cm. longis subteretibus dense lepidoto-

furfuraceis simplicibus vel parce ramosis, internodiis foliis bre-

vioribus; folia alterna breviter petiolata parya, petiolo crassiusculo

2-4 mm. longo; lamina ovalis vel rotundata, interdum ovato-ovalis,

10-17 mm. longa, 5-14 mm. lata, apice rotundata, basi abrupte

contracta et breviter decurrens, integra, crassiuscula, utrinque canes-

cens, dense minute lepidoto-furfuracea; floras monqeci, masculis cum

femineis intermixtis in glomerulis paryis axillaribus; calyx floris

masculi 5-fidus, laciniis oblongis obtusis, antheris exsertis luteis;

bracteae fructiferae axillares sessiles viridescentes minute furfuraceae

late rhombeae c. 2 mm. longae et paullo latiores, integrae vel supra

remote obscure tridenticulatae, dorso inappendiculatae, basi late

rotundato-cuneatae. — Bolivia: Ulloma, alt. 4,000 m., February 26,

1921, E. Asplund 6295 (U. S. Nat. Herb. 1,157,661, type).

5. Atriplex pentandra (Jacq.) Standl. N. Amer. Fl. 21: 54.

1916. Axyris pentandra Jacq. Sel. Stirp. Amer. 244. 1763. Atriplex

cristata Humb. & Bonpl. ex Willd. Sp. PI. 4: 959. 1806. Obione

cristata Moq. Chenop. Enum. 73. 1840.

An annual or perennial, sometimes suffrutescent at the base,

much branched, pale and minutely furfuraceous throughout; leaves

alternate, subsessile, the blades oblong to obovate, 1-3 cm. long,

rounded to acute at the apex, attenuate to the base, the lower leaves

usually dentate but the upper often entire; flowers monoecious, the

staminate in short dense inconspicuous terminal spikes, the pistillate

fascicled in the upper leaf axils; fruiting bracts sessile, broadly

cuneate-orbicular, 3 mm. long and usually somewhat broader,

dentate, much thickened at maturity, dorsally densely cristate or

muricate.

Venezuela: Type from Cumana, Humboldt & Bonpland. Cabo

Blanco, Distrito Federal, trailing on beach, Pittier 12424 (F, W);

10322 (W). Without locality, Curran & Haman 1248 (W).—

Curacao: Beaches and lagoons, Curran & Haman 74 (W). Also in

the West Indies and Florida.

The species has been reported vaguely from Colombia, but I

have seen no specimens from that country, where it may well be

expected to occur. Reports of the occurrence of this species farther

southward in South America are based upon incorrect determinations.

6. Atriplex Herzogii, nom. nov. A. serpyllifolia Herzog, Med.

Rijks Herb. 27: 11. 1915, non Bunge, 1877. A. cristata Humb. &

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

Bonpl., var. pulvinata Kuntze, Rev. Gen. 32: 266. 1898. A. cristata

var. depauperata Kuntze, loc. cit.

A perennial from a thick woody root, much branched from the

base, the stems prostrate and forming dense mats, 10-40 cm. long,

the whole plant pale and minutely furfuraceous or sometimes

greenish; leaves very numerous, minute or small, alternate, subsessile,

oblong to obovate, usually 4-7 mm. long but often somewhat larger,

obtuse or rounded at the apex; flowers monoecious, the staminate

glomerules small, terminal, solitary or short-spicate, inconspicuous,

the pistillate flowers solitary or fasciculate in the leaf axils; fruiting

bracts cuneate-orbicular, about 2 mm. long and often slightly broader,

obscurely dentate, usually densely tuberculate dorsally.

Bolivia: Type from dry cliffs of Cerro de Oruro, at 3,800 m.,

Herzog 2520. Oruro, 4,000 m., March 14, 1894, Kuntze (F, type

collection of var. pulvinata). La Paz, 3,750 m., Buchtien 514 (F, W).

Obrajes, La Paz, 3,300 m., Buchtien 5303 (W). La Paz, 3,000 m.,

Bang 84 (W). Viacha, 3,900 m., Asplund 6302 (W). Chiguana,

3,700 m., Asplund 6301 (W). Uyuni, 3,700 m., Asplund 6300 (W),

6299 (W). Also in Peru.

It is possible that more than one species is represented by the

specimens listed, which exhibit some variation. I have seen no

specimen of var. depauperata, based upon material collected by

Kuntze at Cochabamba, but presumably it is a synonym or form

of A. Herzogii. Kuntze reports also a var. parvifolia of A. cristata,

which he collected at Carcaje, at an elevation of 3,000 m. Presum-

ably it also is referable here.

THE UBRWY C? 7SE

NOV 2 ' 1931

UNIVERSITY OF ILLINOIS