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JUN 1 7 965 



JUL 



FLORA OF COSTA RICA 



BY 

PAUL C. STANDLEY 

CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 




THE LIBRARY OF THE 

DEC 2 4 1938 

UNIVERSITY OF ILLINOIS 
BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 
VOLUME XVIII, PART IV 

NOVKMBKR 30, 1!:!H 

PUBLICATION lj;i 



V 



INDEX 



Achras, 904 
Aciotis, 784 
Acisanthera, 785 
Acnistus, 1036 
Adelobotrys, 785 
Aegiphila, 993 
Allamanda, 930 
Allomarkgrafia, 931 
Alonsoa, 1100 
Amphilophium, 1113 
Anagallis, 902 
Anechites, 932 
Anemopaegma, 1114 
Angelonia, 1100 
Aniseia, 960 
Antirrhinum, 1101 
Apium, 858 
Apocynaceae, 930 
Araliaceae, 851 
Arctostaphylos, 869 
Ardisia, 884 
Arrabidaea, 1115 
Arracacia, 859 
Arthrostemma, 786 
Asclepiadaceae, 949 
Asclepias, 949 
Athenaea, 1036 
Avicennia, 998 
Axinaea, 787 

Bacopa, 1101 
Beaumontia, 932 
Bellucia, 787 
Beureria, 978 
Bignonia, 1116 
Bignoniaceae, 1113 
Blakea, 788 
Blepharodon, 950 
Boraginaceae, 978 
Borago, 979 
Browallia, 1037 
Brunfelsia, 1038 
Buchnera, 1102 
Buddleia, 920 
Bumelia, 905 

Calceolaria, 1102 
Calocarpum, 906 
Calonyction, 960 
Callicarpa, 999 
Callichlamys, 1117 
Calyptrella, 790 
Capraria, 1103 
Capsicum, 1038 
Castilleja, 1104 
Catharanthus, 932 
Cavendishia, 870 
Centaurium, 923 



Centradenia, 791 
Centronia, 791 
Cestrum, 1045 
Chaetolepis, 792 
Chelonanthus, 924 
Chimaphila, 868 
Chrysophyllum, 907 
Citharexylum, 999 
Clavija, 900 
Clerodendron, 1002 
Clethra, 867 
Clethraceae, 867 
Clidemia, 793 
Clytostoma, 1117 
Cobaea, 974 
Coleus, 1015 
Congea, 1003 
Conomorpha, 895 
Conopholis, 1133 
Conostegia, 800 
Convolvulaceae, 960 
Cordia, 979 
Coriandrum, 859 
Cornaceae, 865 
Cornus, 865 
Cornutia, 1004 
Coutoubea, 924 
Crescentia, 1118 
Cufodontia, 933 
Cunila, 1016 
Curtia, 925 
Cuscuta, 961 
Cydista, 1118 
Cynanchum, 951 
Cynoctonum, 921 
Cyphomandra, 1053 

Datura, 1054 
Daucus, 859 
Dermatocalyx, 1105 
Dichondra, 962 
Didymopanax, 851 
Digitalis, 1105 
Diospyros, 912 
Dipholis, 909 
Disterigma, 874 
Duranta, 1005 

Ebenaceae, 912 
Echites, 933 
Ehretia, 984 
Enallagma, 1119 
Enicostema, 925 
Ericaceae, 869 
Eryngium, 860 
Escobedia, 1106 
Evolvulus, 962 
Exolobus, 952 



FLORA OF COSTA RICA 



BY 

PAUL C. STANDLEY 

CURATOR OF THE HERBARIUM, DEPARTMENT OF BOTANY 




THE LI3RARY OF THE 

DEC 24 1938 

UNIVERSITY OF ILLINOIS 

BOTANICAL SERIES 

FIELD MUSEUM OF NATURAL HISTORY 

VOLUME XVIII, PART IV 

NOVEMBER 30, 1938 



PUBLICATION 429 



PRINTED IN THE UNITED STATES OF AMERICA 
BY FIELD MUSEUM PRESS 



D 



FLORA OF COSTA RICA 



PAUL C. STANDLEY 



GESNERIAGEAE 1 

By C. V. Morton 

Herbs or shrubs, often epiphytic; stems often unbranched, fleshy, 
usually pubescent; leaves opposite or rarely alternate, those of a 
pair often unequal, petiolate, the blades usually membranous, entire 
or toothed, pinnately veined; flowers hermaphrodite, axillary or 
rarely terminal, solitary, fasciculate, umbellate, or racemose, often 
bracteate; calyx lobes often colored, free or connate, 5, usually 
valvate, often unequal, sometimes toothed or incised; corolla vari- 
ously colored, gamopetalous, often spurred at base, the tube cylindric, 
ventricose or upwardly ampliate, often hairy, the limb usually bila- 
biate, the lobes rounded, sometimes toothed or fimbriate; stamens 4, 
didynamous, a rudimentary fifth sometimes present, the filaments 
adnate to the corolla at the base, often connate, often contorted, the 
anthers quadrate to oblong, often connate, the cells discrete or con- 
fluent, longitudinally dehiscent; ovary superior or partly or wholly 
inferior, 1-celled, the 2 parietal placentae 2-lobed, the ovules anatro- 
pous, very numerous, borne on the inner surface of the lobes or on 
both the inner and outer surfaces; style simple, elongate; stigma 
' bilobed or stomatomorphic; disk hypogynous or perigynous, annular 
) or of distinct glands; fruit capsular or baccate, the pericarp usually 
coriaceous; seeds minute, very numerous, usually fusiform, spirally 
striate. 

Very little work has been done on this family in the last seventy 

j years. The genera are here adopted in the conventional sense, as 

delimited by Hanstein, but it seems likely that monographic studies 

J will show the necessity for a realignment of the genera, particularly 

in the tribe Columneoideae. The large genera Columnea, Alloplectus, 

< and Drymonia are almost indefinable as at present constituted. A 

& few additional genera are found in Central America. 

[The Gesneriaceae constitute one of the most characteristic and in 

L0 

,/i many respects most interesting groups of Costa Rican plants, and 
J include also many of the most beautiful and showy ones. While they 
^ are found at all elevations and in almost every locality, they attain 

rt 'Published by permission of the Secretary of the Smithsonian Institution. 

1137 



1138 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

most profuse development at upper elevations, chiefly in the more 
humid forests. While many of the plants are terrestrial, the majority, 
at least as regards number of individuals, are epiphytic in habit. 

Especially beautiful are the Columneas, whose usually large and 
brilliant red blossoms furnish a large part of the bright color seen on 
the upper mountain slopes, where they often are associated with 
Ericaceae and other conspicuous epiphytes. Very handsome, too, 
are some of the epiphytic species of Drymonia and Alloplectus. Most 
curious in their flowers are the Campaneas, whose oddly colored 
blossoms dangle from the branches on cordlike peduncles. Most 
gorgeous of terrestrial Gesneriaceae is Solenophora calycosa, whose 
red-orange corollas are as large as teacups. Paul C. Standley.] 

Ovary superior. 

Leaves alternate; inflorescence a terminal, lax raceme; corolla blue. 

Klugia. 

Leaves opposite or radical; inflorescence axillary, the flowers 
solitary, fasciculate, or umbellate, rarely short-racemose; 
corolla not blue. 

Calyx lobes connate over two thirds their length Tussacia. 

Calyx lobes free or nearly so. 

Plant acaulescent; sepals strongly parallel-nerved . Napeanthus. 
Plants caulescent; sepals not parallel-nerved. 
Anther cells separated by the enlarged connective. Creep- 
ing, herbaceous vine with glabrous, fleshy leaves. 

Codonanthe. 
Anther cells contiguous or confluent. 

Anther cells confluent at the apex; disk annular or rarely 

semiannular Besleria. 

Anther cells discrete; disk of distinct glands or reduced 

to a solitary, posterior gland. 

Plants low, terrestrial, stoloniferous herbs; placentae 
ovuliferous on both surfaces. Corolla lilac, not 
contracted in the throat, the limb widely spreading; 

calyx lobes spatulate Episcia. 

Plants shrubs or rarely herbs, not stoloniferous; 

placentae ovuliferous on the inner surface only. 

Corolla erect in the calyx, not spurred at the base, 

sometimes a little gibbous, not contracted in 

the throat; limb often strongly bilabiate, or if 



FLORA OF COSTA RICA 1139 

not, the leaves of a pair usually unequal, the 

smaller stipule-like Columnea. 

Corolla oblique or horizontal in the calyx (except in 
Alloplectus multiflorus), spurred at the base; 
limb regular or slightly bilabiate; leaves of a 
pair not strongly unequal (except in Alloplectus 
metamorphophyllus) . 

Anthers oblong, sagittate at the base, only par- 
tially dehiscent. 

Corolla ampliate in the throat, usually much 
exceeding the calyx, the limb broad; 
flowers often solitary, ebracteate . Drymonia. 
Corolla usually contracted in the throat, often 
not much longer than the calyx, the limb 
narrow; flowers often fasciculate, bracteate. 

Alloplectus. 

Anthers ovate or quadrate, dehiscent throughout. 

Flowers solitary; corolla tube strongly ventri- 

cose, the limb minute (3 mm. wide), oblique. 

Delicate, herbaceous vine Hypocyrta. 

Flowers usually fasciculate or umbellate; corolla 
tube less strongly ventricose, the limb 
terminal, broader. 

Calyx lobes linear; anthers ovate. 

Centrosolenia. 

Calyx lobes broad; anthers quadrate or 
broader than long Alloplectus. 

Ovary wholly or partly inferior. 

Leaves alternate Gesneria. 

Leaves opposite. 

Corolla rotate. Disk none Pkinaea. 

Corolla tubular or campanulate. 

Disk none; calyx tube cylindric in fruit. Inflorescence 
terminal, open-paniculate Monopyle. 

Disk present; calyx tube turbinate or hemispheric in fruit. 
Disk annular. 

Inflorescence terminal, racemose, the pedicels and bracts 
alternate . . Koellikeria. 



1140 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Inflorescence axillary or terminal, the pedicels and bracts 
(if present) opposite. 

Corolla open-campanulate Gloxinia. 

Corolla tubular. 

Anthers connate, at least when young . . . Achimenes. 

Anthers free Heppiella. 

Disk of 1 to 5 distinct glands or the 2 posterior ones partially 

connate. 

Disk glands 2, posterior, connate, pubescent; calyx tube 
extended long beyond the ovary. Corolla scarlet, 
7-9 cm. long Solenophora. 

Disk glands 5, glabrous; calyx tube not extended beyond 
the ovary. 

Corolla scarlet, tubular Kohleria. 

Corolla yellowish green, open-campanulate. Campanea. 

ACHIMENES P. Br. 

Perennial, terrestrial herbs, increasing by scaly offsets, the stems 
unbranched; leaves opposite, subequal or very unequal, the blades 
toothed, membranous; inflorescence axillary, the flowers solitary or 
several on a common peduncle; calyx tube turbinate, the lobes free, 
elongate, entire; corolla salverform, the tube not ventricose, the 
limb actinomorphic or subbilabiate; filaments inserted near the base 
of the corolla tube; anthers oblong or quadrate, the cells not confluent 
at the apex; ovary entirely inferior; disk annular, entire, glabrous. 
Two or three additional species occur in Central America. 

Stigma stomatomorphic; lamellae of the placentae ovuliferous on the 
inner surface only; disk high, interrupted dorsally; corolla white, 
spotted with red; stems glabrous A. Candida. 

Stigma bilobed; lamellae of the placentae ovuliferous on both sur- 
faces; disk low, entire; corolla red or purple; stems pilosulous or 
pilose. 

Flowers erect in the calyx, not spurred at the base; leaf blades 
cuneate at the base, the veins 3-5 pairs. 

Corolla purple, 5.5-6.5 cm. long; calyx lobes 11-14 mm. long, 
. ciliate, otherwise glabrous A. longiflora. 

Corolla red, 1.5-2 cm. long; calyx lobes 4.5 mm. long, pilosulous, 
not ciliate A. pulchella. 



FLORA OF COSTA RICA 1141 

Flowers oblique or horizontal in the calyx, short-spurred at base; 

leaf blades rounded or subcordate at base, the veins 8-12 pairs. 

Corolla scarlet, spotted, stipitate-glandular in the throat; 

peduncles often 2-3-flowered A. pedunculata. 

Corolla purple, not spotted, glabrous within the throat; pe- 
duncles 1-flowered A. grandiflora. 

Achimenes Candida Lindl. El General. Guatemala. An herb 
45-50 cm. high, the stems glabrate; leaves opposite, usually very 
unequal, the short petioles pilosulous on the upper side only, the 
blades elliptic, up to 8 cm. long and 3.3 cm. wide, acuminate, cuneate 
at the base, serrate (the teeth 10-15 on each side), sparsely scabrous- 
puberulous beneath; common peduncle 4-7 mm. long, 2-3-flowered, 
the pedicels slender (less than 0.5 mm. in diameter) ; calyx about 5.5 
mm. long, the tube 2.5 mm. long, articulate-pilose, the lobes puberu- 
lous on both sides; corolla white, spotted in the throat, 15-17 mm. 
long, erect in the calyx, not spurred, nearly glabrous, the limb 12-13 
mm. wide; filaments glabrous. 

Achimenes grandiflora (Schiede) DC. Central mountains, 
1,200-1,600 meters; region of San Ramon. Northward to Veracruz. 
An herb 30-60 cm. high; leaves opposite, long-petiolate, the blades 
ovate, up to 15 cm. long and 8 cm. wide, acuminate, oblique and sub- 
cordate at the base, coarsely serrate (the teeth 15-25 on each side), 
scabrous-puberulous beneath ; common peduncle absent, the pedicels 
solitary, 5-7.5 cm. long, bulbilliferous; calyx 8-10 mm. long, the tube 
4 mm. long, hirtous-puberulous, the lobes scabrous-puberulous on 
both sides, not ciliate; corolla purple, not spotted, horizontal in the 
calyx, 3.5-4.5 cm. long, spurred at the base, the tube pilosulous, the 
limb slightly or strongly bilabiate, 2.5-3 cm. wide; filaments glabrous. 

Achimenes longiflora DC. Centralamer. Gesner. 17. pi. 9, f. 
1, 2. 1858 (Cerro de Aguacate, Oersted). A. pauciflora Oerst. Atlan- 
tic coast; central mountains; region of San Ramon. Mexico to Pana- 
ma. Herb, 10-30 cm. high; leaves opposite or ternate, short-petiolate 
(6 mm.), the blades ovate to lanceolate, 2-8.5 cm. long, 1.3-3 cm. 
wide, acute, cuneate at the base, serrate (the teeth few), appressed- 
pubescent; common peduncle absent, the pedicel solitary, 4-7 mm. 
long; calyx 13-17 mm. long, the tube 2-3 mm. long, densely pilose, 
the lobes linear-lanceolate, nearly glabrous, long-ciliate; corolla 
purple, 5.5-6.5 cm. long, erect in the calyx, not spurred, the tube 
very slender, glabrous, the limb broad (4-5 cm.), subactinomorphic; 
filaments glabrous. Several minor named forms occur in Costa Rica. 



1142 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Achimenes pedunculata Benth. Central mountains, at 1,000- 
1,200 meters; region of San Ramon. Guatemala. An herb 40-85 
cm. high, the stems short-puberulous upwardly; leaves opposite, 
long-petiolate, the blades broadly elliptic, up to 15 cm. long and 
8 cm. wide, cuspidate-acuminate, oblique and rounded at the base, 
coarsely toothed (the teeth 30 or more on each side), minutely 
puberulous beneath; common peduncle obsolete or up to 7 cm. long, 
bibracteate at the apex, 1-3-flowered, the pedicels elongate, bulb- 
illiferous at middle; calyx 6.5 mm. long, the tube 3 mm. long, scab- 
rous-puberulous, the lobes not ciliate, scabrous-puberulous; corolla 
scarlet, red-spotted in the throat, 30 mm. long, short-spurred, not 
ventricose, short-puberulous, the limb regular, stipitate-glandular 
in the throat; filaments pilosulous. 

Achimenes pulchella (L'He>.) Hitchc. A. coccinea Pers. 
Reported from Costa Rica by Hemsley . Widely distributed in Mexico, 
Central America, and the West Indies. An herb 10-30 cm. high; 
leaves opposite or ternate, short-petiolate, the blades ovate to elliptic, 
up to 5 cm. long and 1.6 cm. wide, acute or obtusish, acute at the 
base, coarsely dentate (the teeth 3 to 5 on each side), pilosulous; 
common peduncle none, the pedicel solitary, up to 2.5 cm. long, 
slender, ebracteate, not bulbilliferous; calyx about 6.5 mm. long, the 
tube 2 mm. long, densely pilosulous, the lobes pilosulous on both sides; 
corolla red or pink, not spotted, 15-20 mm. long, erect, not spurred, 
the tube puberulous, the limb 12-15 mm. wide; filaments glabrous. 

ALLOPLECTUS Mart. 

Shrubs or vines, usually epiphytic; leaves opposite, equal or 
rarely unequal, often rather long-petiolate, the blades entire or 
toothed; flowers fasciculate or rarely solitary or in short, axillary 
racemes, bracteate, the bracts often large and colored; calyx usually 
colored, the lobes free, broad, often toothed or incised; corolla yellow 
or red, erect or horizontal in the calyx, usually not much exceeding 
the calyx, the tube usually ventricose, pilose, contracted in the 
throat, the limb narrow, nearly regular, the lobes rounded, usually 
entire; filaments connate at the base, usually adnate to the corolla, 
the anthers commonly free, oblong to quadrate or broader than long, 
the cells distinct, sometimes sagittate at the base, sometimes only 
partially dehiscent; disk gland solitary, large; fruit baccate. Several 
other species are found in Central America. 

Leaves dimorphic, the smaller ones being stipule-like phyllodes. 

A, metamorphophyllus. 



FLORA OF COSTA RICA 1143 

Leaves not dimorphic, sometimes unequal. 

Corolla erect in the calyx, not spurred or saccate at the base; 

calyx lobes equal, erect, red, pectinate- toothed. A. multiflorus. 

Corolla oblique or horizontal in the calyx, spurred or saccate at 

the base; calyx oblique or the lobes unequal. 
Anthers quadrate or broader than long, not sagittate at the base. 
Calyx lobes entire, linear or linear-lanceolate. A. congestiflorus. 
Calyx lobes conspicuously toothed, ovate to lanceolate. 
Corolla contracted in the throat; anthers quadrate; pedicels 
about 1 cm. long A. ichthyoderma. 

Corolla enlarged in the throat; anthers broader than long; 
pedicels 3-5 cm. long A. tetragonus. 

Anthers oblong, sagittate at the base. 

Corolla and calyx glabrous A. stenophyllus. 

Corolla and calyx pubescent. 

Leaves peltate A. peltatus. 

Leaves not peltate. 

Calyx lobes entire; flowers short-racemose. 

A. tucurriquensis. 
Calyx lobes toothed; flowers fasciculate. 

Calyx lobes minutely dentate at the apex. 

A. coriaceus. 
Calyx lobes subulate- toothed. 

Corolla not contracted in the throat; calyx lobes 

ovate A. macrophyllus. 

Corolla contracted in the throat; calyx lobes lanceo- 
late A. parviflorus. 

Alloplectus congestiflorus (Donn. Smith) Morton, comb. nov. 
Besleria congestiflora Donn. Smith, Bot. Gaz. 61: 379. 1916. La 
Palma, Prov. San Jose", 1,460 meters, Tonduz 12658. Region of 
San Ramon. Endemic. An epiphytic shrub, 25-60 cm. high, 
the stems tomentose, sparingly branched; leaves equal, petiolate, 
the petiole up to 3 cm. long, the blade oblanceolate, up to 10.5 
cm. long and 3.5 cm. broad, acuminate, attenuate at the base, ser- 
rulate, densely tomentose on both sides; flowers aggregate in the 
leaf axils, the pedicels 3-5 mm. long, the bracts linear-lanceolate; 
calyx lobes red, linear or linear-lanceolate, 7-10 mm. long, equal, 
acuminate, entire, pilose on both sides; corolla white, marked with 



1144 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

rose on the limb, tubular, about 12 mm. long, pilose, the throat con- 
tracted, the limb spreading, narrow, subregular; anthers quadrate, 
persistently connate, the cells orbicular, not confluent; disk gland 
solitary. 

Alloplectus coriaceus (Oerst.) Hanst. Linnaea 34: 363. 1865- 
66. Erythranthus coriaceus Oerst. Centralamer. Gesner. 47. pi. 3. 
1858. A. ventricosus Donn. Smith, Bot. Gaz. 25: 154. 1898 (Atirro, 
Prov. Cartago, 600 meters, J. D. Smith 6727). Turrialba, Oersted. 
Atirro; Tucurrique; Guanacaste. Endemic. An epiphytic shrub 
up to 2.4 meters high, the stems unbranched, minutely strigillose; 
leaves subequal, short-petiolate (1-3 cm.), the blades oblanceo- 
late, up to 25 cm. long and 8 cm. wide, short-acuminate, attenuate 
at the base, subcoriaceous, entire, glabrous, reddish; inflorescence 
congested, the bracts red, numerous, persistent, orbicular-oval, 
about 1.5 cm. long, sparsely strigillose; calyx oblique, red, the 
lobes unequal, obovate-oval, about 1 cm. long, dentate at the apex, 
red-strigillose; corolla red, gibbous at the base, oblique in the 
calyx, 1.6-2.8 cm. long, reddish-tomentose, the tube strongly ven- 
tricose upwardly, contracted in the throat, the limb oblique, 
about 3 mm. wide, subregular, the lobes minute, erect; anthers 
oblong, 3 mm. long. 

Columnea costaricensis Kuntze, Rev. Gen. 2: 471. 1891 (Port 
Limon, Kuntze). Species dubious, doubtless belonging to Alloplectus. 

Alloplectus ichthyoderma Hanst. Linnaea 34: 372. 1865-66. 
A. ruacophilus Donn. Smith, Bot. Gaz. 54: 238. 1912 (forests of 
Volcan de Barba, Prov. Heredia, 2,500-2,700 meters, Tonduz 1997). 
Costa Rica, without locality, Wendland 833. Central mountains, 
1,800-2,700 meters. Panama. A terrestrial shrub, 0.6-1.5 meters 
high, the stems unbranched, hirsute, the epidermis peeling off in 
scales; leaves subequal, long-petiolate, 2.5-8.5 cm. long, the blades 
elliptic, up to 20 cm. long and 9 cm. wide, cuspidate-acuminate, 
cuneate at the base, denticulate, membranous, green and pilose above, 
pale and minutely strigillose beneath; flowers fasciculate, numerous, 
the bracts ovate, red, entire, about 1 cm. long, the pedicels 1.5-2 cm. 
long, appressed-pilose; calyx oblique, the lobes red, or green with red 
margins, ovate to lanceolate, up to 1.8 cm. long, obtuse to acuminate, 
subequal, serrate, white-pilose, especially near the midrib and base; 
corolla yellow, tubular, 15-23 mm. long, gibbous at the base, hori- 
zontal in the calyx, the tube white-tomentose externally, strongly 
ventricose upwardly, contracted in the throat, the limb oblique, 



FLORA OF COSTA RICA 1145 

about 8 mm. wide, subregular, the lobes subequal, erect, 2-3 mm. 
long; filaments glabrous; anthers quadrate, connate; ovary tomen- 
tose; style glabrous; stigma stomatomorphic. 

Alloplectus ichthyoderma var. hirsutulus Morton, var. nov. 
A var. typica foliis subtus hirsutulis nee strigillosis differt. Atirro, 
Prov. Cartago, alt. 600 meters, Donnell Smith 6715 (type in U. S. 
Nat. Herb., No. 1,336,496). 

Alloplectus ichthyoderma var. pallidus Morton, var. nov. 
A var. typica calycibus flavis vel pallide viridibus nee rubris differt. 
Las Nubes, Prov. San Jose", 1,500-1,900 meters, Standley 38793 (type 
in U. S. Nat. Herb., No. 1,228,403). Same locality, Standley 38363, 
38372. Cerro Gallito, M. Valeria 1004. 

Alloplectus ichthyoderma forma rubescens Morton, f . nov. 
A f. typica foliis subtus rubescentibus nee pallide viridibus differt. 
Santa Clara de Cartago, 1,950 meters, Maxon & Harvey 8195 (type 
in U. S. Nat. Herb., No. 1,181,839). 

Alloplectus macrophyllus (Oerst.) Hemsl. Biol. Centr. Amer. 
Bot. 2: 484. 1882. Caloplectus macrophyllus Oerst. Centralamer. 
Gesner. 45. 1858. Naranjo, Oersted. Llanuras de Santa Clara, Donn. 
Smith 6725, 6733; Pittier 13393. Tsaki, Tonduz 9557. Guatemala. 
A shrub 1.8 meters high, the stems subtomentose at the apex; leaves 
subequal, petiolate, the petioles subtomentose, 0.5-6 cm. long, the 
blades elliptic, up to 33 cm. long and 14 cm. wide, short-acuminate, 
rounded and oblique at the base, obscurely serrulate or entire, 
sparsely strigose above, minutely puberulous beneath; flowers fasci- 
culate, few, the pedicels up to 1 cm. long, the bracts subulate, shorter 
than the pedicels; calyx lobes red or green, ovate, up to 15 mm. long, 
thin-membranous, densely strigillose, conspicuously subulate- toothed; 
corolla pale yellow with maroon stripes along the lower side, about 
3 cm. long, spurred at the base posteriorly, horizontal in the calyx, 
densely but minutely puberulous, the tube regularly ampliate up- 
wardly, not contracted in the throat, the limb slightly bilabiate, about 
2 cm. wide; anthers oblong, sagittate. 

Alloplectus metamorphophyllus Donn. Smith, Bot. Gaz. 
52: 52. 1911. La Palma, Prov. San Jose", 1,500-1,600 meters, 
Tonduz 10884. La Hondura; Tucurrique. Endemic. An epiphytic, 
climbing shrub with unbranched, subtomentose stems; leaves 
strongly dimorphic, the larger long-petiolate (up to 13 cm. 



1146 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

long), the blade broadly elliptic, up to 20 cm. long and 11 cm. wide, 
short-acuminate, cuneate at the base, sharply serrulate, sericeous- 
pilose on both sides; smaller leaves stipuliform, linear-lanceolate, 
about 2-4 cm. long, sessile, entire; inflorescence of congested, many- 
flowered, axillary corymbs, the pedicels very short; calyx lobes erect, 
red, oblanceolate, 11-16 mm. long, pilosulous, deeply pectinate- 
incised; corolla erect in the calyx, shorter than the calyx, not gibbous 
at the base, sparsely strigillose, the tube a little ampliate upwardly, 
contracted in the throat, the limb narrow, regular; anthers connate, 
oblong, the cells distinct; disk gland solitary. 

Alloplectus multiflorus (Oerst.) Hanst. Linnaea 34: 361. 1865- 
66. Calanthus multiflorus Oerst. Centralamer. Gesner. 47. pi. 4- 1858. 
Naranjo, Oersted. Central mountains, 1,300-1,700 meters; Atlantic 
coast. Endemic. A vine, the stems branched, minutely strigillose; 
leaves subequal, short-petiolate (2-3 mm.), the blades elliptic-lanceo- 
late, up to 8 cm. long and 3 cm. wide, acuminate, cuneate at the base, 
entire, membranous, glabrous above, strigillose beneath; flowers 
solitary, axillary, the pedicels 1-1.5 cm. long, glabrate; calyx lobes 
red, equal, not oblique, ovate, up to 15 mm. long, deeply subulate- 
toothed, sparingly strigose; corolla yellow with red stripes, dark 
purple within, tubular, about 2.5 cm. long, erect in the calyx, the 
tube slender, strongly ventricose, glabrous, contracted in the throat, 
the throat sparsely pilosulous, the limb subbilabiate; anthers narrowly 
oblong. 

Alloplectus parviflorus (Hanst.) Hemsl. Biol. Centr. Amer. 
Bot. 2: 484. 1882. Polythysania parviflora Hanst. ex Oerst. Central- 
amer. Gesner. 50. 1858. Naranjo, 1,200-1, 500 meters, Oersted Known 
definitely only from the type. Stems villous at the apex; leaves un- 
equal, very short-petiolate, the blades obovate, the larger 7.5-10 cm. 
long, the smaller one-half shorter, acuminate, attenuate at the base, 
serrate, pubescent; flowers solitary, short-pedicellate, the pedicels 
hirsute; calyx lobes lanceolate, unequal, pubescent, laciniate-toothed; 
corolla saccate at the base, horizontal in the calyx, the tube inflated 
above, contracted in the throat, the lobes short; anthers oblong, 
sagittate. Description adapted from the original. 

Alloplectus peltatus Oliver, Bot. Mag. 103: pi. 6383. 1877. 
Known only from material collected by Endres and cultivated at 
Kew. A shrub, the stems glabrous; leaves strongly unequal, the 
larger petiolate (2.5-5 cm.), the blade peltate, oval-oblong, 15-22.5 
cm. long, 4.2-6.2 cm. wide, subcoriaceous, at first ciliolate, soon 



FLORA OF COSTA RICA 1147 

glabrous; flowers fasciculate, few, the pedicels 1.2-1.7 cm. long, the 
bracts lanceolate or ovate-lanceolate, acute, shorter than the pedicels; 
calyx oblique, red-purple, the lobes lanceolate, 2.5 cm. long, unequal, 
the posterior shorter; corolla pale yellow, the limb slightly bilabiate, 
the lobes fimbriate; stamens included, the filaments glabrous, the 
anthers sagittate, oblong. The above description is adapted from 
the original. 

Alloplectus stenophyllus Donn. Smith, Bot. Gaz. 25: 153. 
1898. Forests of Suerre, Llanuras de Santa Clara, 300 meters, 
J. D. Smith 6724- Atlantic coast at low elevations; region of San 
Ramon. Endemic. A small shrub, the stems glabrous; leaves 
unequal, often strongly so, petiolate (1-1.5 cm.), the larger blades 
narrowly elliptic to linear, up to 25 cm. long and 4.5 cm. wide, 
acuminate, obtuse at the base, subcoriaceous, often shining, entire, 
glabrous; flowers solitary or paired, the pedicels about 5 mm. long, 
glabrous, the bracts minute, subulate; calyx oblique, the lobes 
lanceolate, unequal, the longer about 16 mm. long, the posterior 
reflexed, about 10 mm. long, all entire, glabrous; corolla white or 
pale yellow, rosy in the throat, about 2.5 cm. long, spurred at the 
base, horizontal in the calyx, glabrous, the tube ventricose, contracted 
in the throat, the limb subbilabiate, about 1 cm. wide; anthers oblong, 
sagittate at the base; ovary glabrous. 

Alloplectus tetragonus (Oerst.) Hanst. Linnaea 34: 368. 1865- 
66. Glossoloma tetragonum Oerst. Centralamer. Gesner. 41. pi. 7. 
1858. Turrialba, 900 meters, Oersted. Central mountains, 1,400- 
2,000 meters. Endemic. A terrestrial or epiphytic shrub, 1-1.5 
meters high, the stems unbranched, yellowish-subtomentose, the 
epidermis not peeling off in scales; leaves subequal, long-petiolate 
(3-9 cm.), the blades elliptic, up to 25 cm. long and 11 cm. wide, 
acuminate, cuneate at the base, serrate or serrulate, membranous, 
green and pilosulous above, pale and minutely strigillose beneath; 
flowers fasciculate, 2-several, the pedicels 3-5 cm. long; calyx oblique, 
the lobes red, or green with red margins, ovate, about 2 cm. long, 
subequal, subulate- toothed, hirsute; corolla red, tubular, 4-5 cm. 
long, gibbous at the base, oblique in the calyx, yellowish-hirsute, 
the tube ampliate upwardly, not contracted in the throat, the 
limb oblique, about 1 cm. wide, the lobes subequal ; anthers broader 
than long. 

Alloplectus tucurriquensis Donn. Smith, Bot. Gaz. 54: 238. 
1912. Forests near Las Vueltas, Tucurrique, Prov. Cartago, 700 



1148 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

meters, Tonduz ISO 42. Known only from the type. An epiphytic 
shrub, the stems unbranched, puberulous at the apex; leaf blades 
broadly elliptic, up to 28 cm. long and 13 cm. wide, acuminate, 
attenuate at the base and long-decurrent into the short, stout 
petiole, serrulate, membranous, green, substrigose above, puberulous 
beneath; flowers in short, dense, axillary, bracteate racemes, the 
common peduncle 5-7 mm. long, the pedicels 13-18 mm. long, the 
bracts scarlet, 26-33 mm. long, entire; calyx green or reddish, slightly 
oblique, the lobes 20-28 mm. long, oblong, entire, minutely puberu- 
lous; corolla spurred at the base, about 3.5 cm. long, horizontal in 
the calyx, densely puberulous, ventricose upwardly, narrowed in 
the throat, the limb about 1 cm. wide, nearly regular; anthers 
narrowly oblong, 5 mm. long; ovary cinereous-strigillose. 

BESLERIA [Plum.] L. 

Shrubs with terete or quadrangular, usually unbranched stems; 
leaves opposite, equal, petiolate, usually elliptic, membranous to 
coriaceous; flowers ebracteate, commonly borne in sessile or pedun- 
culate, axillary umbels, rarely solitary; calyx 5-parted nearly to 
the base, colored, the lobes imbricate; corolla erect or horizontal 
in the calyx, cylindric or spurred at the base, often pilose, the tube 
usually ventricose, usually contracted in the throat, the limb narrow 
or broad, regular or irregular, the lobes entire; filaments not connate, 
usually glabrous, the anthers quadrate, connate when young, the 
cells confluent at the apex; placentae ovuliferous on the inner surface 
only; disk annular or semiannular, glabrous or pubescent, sometimes 
much thickened posteriorly; fruit a fleshy berry. Several other 
species occur in Central America. 

Corolla not spurred at the base, erect or oblique in the calyx; disk 
glabrous, not thickened posteriorly. 

Common peduncle obsolete, the flowers aggregate in the leaf axils 

or solitary. 
Calyx lobes sharply toothed; corolla pubescent. Ovary pilose. 

Stems and leaves strigose; primary leaf veins 12 or 13 pairs; 
calyx lobes ovate B. barbensis. 

Stems and leaves hirsute; primary leaf veins 7-10 pairs; calyx 

lobes lanceolate, acuminate. 

Leaves tuberculate above, the tubercles surmounted by a 
long hair; corolla strongly bilabiate, much exceeding 
the calyx, pilose B. princeps. 



FLORA OF COSTA RICA 1149 

Leaves not tuberculate above; corolla nearly regular, 
scarcely longer than the calyx, copiously red-villous. 

B. columneoides. 
Calyx lobes entire or sometimes a little erose at the apex ; corolla 

glabrous or inconspicuously puberulous. 
Stems and leaf veins beneath hirsute. 

Calyx lobes linear-lanceolate, acuminate; corolla 11-13 mm. 

long, pilose; ovary pilose B. trichostegia. 

Calyx lobes oblong to suborbicular, rounded; corolla 18 mm. 

long, glabrous; ovary glabrous B. hirsuta. 

Stemsand leaves tomentose, puberulous, strigillose, or glabrate. 
Calyx lobes 10-12 mm. long; stems thick, densely tomentose. 

Corolla and ovary glabrous B. robusta. 

Calyx lobes 3-7 mm. long; stems appressed-pubescent or 

strigose. 
Anther cells not confluent; corolla densely long-pilose. 

B. pycnosuzygia. 
Anther cells confluent; corolla glabrous or sparsely pilosu- 

lous or puberulous. 

Corolla tube 5-6 mm. wide, obviously gibbous at the 
base; calyx lobes suborbicular, broader than long, 
strongly imbricate; leaf blades obtuse or rounded 

at the base B. Standleyi. 

Corolla tube not over 3 mm. wide, not gibbous at the 
base; calyx lobes oblong, scarcely imbricate; leaf 
blades cuneate at the base. 

Stems thick (commonly 5-11 mm. in diameter); leaf 
blades subcoriaceous, 6-10 cm. wide. 

B. notabilis. 

Stems slender (usually 2-3 mm. in diameter); leaf 
blades 5.5 cm. wide or less. 

Corolla and ovary glabrous B. solanoides. 

Corolla and ovary hairy. .B. solanoides var. tenera. 
Common peduncle present, usually elongate, the pedicels umbellate 
or subcorymbose, rarely solitary (in B. formosa). Pubescence 
appressed. 

Corolla with a hairy ring within at the insertion of the filaments; 
calyx lobes 7.5-14 mm. long, ovate to lanceolate, acute or 
acuminate. Corolla glabrous externally; ovary glabrous. 



1150 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Leaf blades, peduncles, and pedicels entirely glabrous; corolla 
yellow; calyx lobes narrowly lanceolate, membranous. 

B. macropoda. 

Leaf blades, peduncles, and pedicels at least slightly strigillose; 
corolla orange or red; calyx lobes ovate to ovate-lanceo- 
late, thicker B. laxiflora. 

Corolla lacking a hairy ring within; calyx lobes 5-6 mm. long, 
suborbicular, obtuse or rounded. 

Corolla and ovary puberulous; calyx lobes white. . .B. triflora. 
Corolla and ovary glabrous; calyx lobes green. . . .B. formosa. 

Corolla spurred at the base, horizontal in the calyx; disk pubescent, 
strongly thickened posteriorly. Flowers in a pedunculate umbel. 

Calyx lobes slightly toothed; ovary sericeous; stems and leaf veins 
beneath lanate-tomentulose B. Wendlandiana. 

Calyx lobes entire; ovary glabrous; stems and leaf veins beneath 

strigillose. 

Corolla yellow, bilabiate, purple-spotted in the throat, 2.3-3 cm. 
long, pilosulous externally; calyx lobes ovate-lanceolate, 
12-14 mm. long, pilosulous; primary leaf veins about 8 pairs. 

B. acropoda. 

Corolla scarlet, the limb regular, not spotted, 1.5 cm. long, 
glabrous; calyx lobes suborbicular, 7 mm. long, glabrous; 
leaf veins 16 or 17 pairs. 

Leaf blades entire; corolla spur not uncinate. . . .B. imbricans. 

Besleria acropoda Donn. Smith, Bot. Gaz. 54: 240. 1912. 
Known only from the type, Tsaki, Prov. Limon, 200 meters, Tonduz 
9554. A shrub; stems terete, sparsely strigillose; petioles strigillose, 
1-2 cm. long; leaf blades elliptic, the larger 11 cm. long and 5.5 cm. 
wide, abruptly acuminate, attenuate and decurrent at the base, 
membranous, deeply serrate, glabrous above, strigillose on the veins 
beneath, the primary veins about 8 pairs; common peduncles axillary 
but sub terminal by reduction of the main axis, 1.8-2.3 cm. long, 
glabrous, the pedicels umbellate, few, about 1 cm. long, glabrous; 
calyx lobes ovate-lanceolate, unequal (the larger 14 mm. long), 
free, entire, membranous, oblique, sparsely pilosulous on both sides; 
corolla yellow, 23-30 mm. long, horizontal in the calyx, spurred at 
the base (3 mm. or more), the tube ampliate upwardly, a little 



FLORA OF COSTA RICA 1151 

ventricose, sparsely pilosulous externally, the limb bilabiate, 15 mm. 
wide, the lobes spotted with purple; ovary glabrous; disk enlarged 
posteriorly, pubescent. 

Besleria barbensis Hanst. Linnaea 34: 319. 1865-66. In forest, 
Volcan de Barba, Hoffmann 45. Central mountains at middle 
elevations. Endemic. An herb. 1-1.2 meters high; stems angled, 
sparingly strigose; petioles 2-2.5 cm. long; leaf blades broadly 
elliptic, the largest 15 cm. long and 8.5 cm. wide, short-acuminate, 
obtuse or cuneate at the base, oblique, membranous, irregularly 
serrate, substrigose on the veins beneath, the primary veins 12-13 
pairs; common peduncle absent, the pedicels solitary, axillary, 1.5-2 
cm. long, sparingly pilosulous; calyx lobes ovate, 12-15 mm. long, 
free, sharply serrate, sparsely pilosulous externally, puberulous 
within; corolla orange, erect, not spurred or saccate at the base, 
the tube about 15 mm. long, 8 mm. wide, sparsely pilose externally, 
glabrous within, the limb bilabiate, the lobes spreading, the largest 
9 mm. long and broad; androecium glabrous; ovary pilose at the 
apex; disk annular, thick, glabrous. 

Besleria columneoides Hanst. Linnaea 34: 322. 1865-66. San 
Miguel, Wendland 762. Endemic. Stems densely hirsute; petioles 
2.5 cm. long, hirsute; leaf blades elliptic, up to 22.5 cm. long and 
8.5 cm. wide, cuneate at the base, hirsute beneath, especially on 
the veins, the primary veins 8-9 pairs; pedicels 0.8-1.4 cm. long; 
calyx yellow, 10 mm. long, the lobes free, lanceolate, acuminate, 
externally long-pilose, glabrous within, bearing 2-4 subulate teeth; 
corolla scarcely longer than the calyx, 4 mm. wide, copiously red- 
villous, the lobes glabrous externally, long-ciliate; ovary villous at 
the apex; disk subentire. 

Besleria formosa Morton, sp. nov. Frutex 1.5-2.5 m. altus, 
caulibus angulatis strigosis, mox glabris; petioli 1-2 cm. longi; lamina 
foliorum lanceolata, usque ad 8 cm. longa et 2.7 cm. lata, acuminata, 
basi attenuata, supra viridis, glabra vel sparse strigosa, subtus 
pallida, glabra, venis strigosis exceptis, venis primariis ca. 5-jugis; 
pedunculus communis brevis, usque ad 7 mm. longus, pedicello 
solitario, 2.5-4 cm. longo, strigoso; calycis lobi coriacei obtusi, pallide 
virides, glabri vel sparse strigillosi; corolla aurantiaco-rubra, crassa, 
18-19 mm. longa, ecalcarata, basi paullo gibbosa, tubo utrinque 
glabro, limbo patente angusto, lobis obtusis; androeceum glabrum, 
antheris connatis; ovarium glabrum; stylus puberulus; discus uno 
latere interruptus. Yerba Buena, Prov. Heredia, alt. 2,000 meters, 



1152 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Standley & Valeria 49926 (type in U. S. Nat. Herb., No. 1,306,250). 
Same locality, Standley & Valeria 49839. Cerros de Zurqui, Prov. 
Heredia, Standley & Valeria 50527, 50695. Cerro de Las Caricias, 
Prov. Heredia, Standley & Valeria 52112. Cerro Gallito, M. Valeria 
1216. 

Besleria hirsuta (Oerst.) Hanst. Gasteranthopsis hirsuta Oerst. 
Siquirres, Prov. Limon, Donn. Smith 6714- Mexico. A shrub; stems 
hirsute; petioles up to 6.5 cm. long, hirsute; leaf blades oblong, up 
to 23 cm. long and 9.5 cm. wide, attenuate at the base, obsoletely 
serrulate, hirsute on the veins beneath, the primary veins 11-13 
pairs; pedicels hirsute, about 1 cm. long; calyx about 5 mm. long, 
the lobes free, oblong or suborbicular, rounded, membranous, short- 
pilosulous; corolla about 18 mm. long, glabrous on both sides: 
ovary glabrous; disk annular. 

Besleria imbricans Donn. Smith, Bot. Gaz. 25: 155. 1898. 
Known only from the type, Shirores, Prov. Limon, sea level, Tonduz 
9192. Stems minutely puberulous, soon glabrous; petioles up to 
4 cm. long; leaf blades elliptic, up to 23 cm. long and 11 cm. wide, 
cuneate and oblique at the base, membranous, entire, strigillose on 
the veins beneath, the primary veins 16-17 pairs; common peduncle 
glabrous, about 8.5 cm. long, the pedicels short, few; calyx lobes 
orange, suborbicular, about 7 mm. long, imbricate, membranous, 
glabrous, entire, not ciliate, unequal; corolla scarlet, horizontal in the 
calyx, 15 mm. long, spurred at the base (1.5 mm.), the tube glabrous, 
"ventricose upwardly, the throat contracted, the limb small, terminal; 
androecium glabrous; ovary glabrous; disk pubescent, enlarged 
posteriorly. 

Besleria imbricans var. uncinata Morton, var. nov. A var. 
typica foliis minoribus serratis, corollae calcare longiore paullo un- 
cinato differt. Las Vueltas, Tucurrique, Prov. Cartago, alt. 900- 
1,000 meters, Tonduz 13038 (type in U. S. Nat. Herb., No. 1,336,615). 

Besleria laxiflora Benth. Besleria chiapensis Brandeg. Low 
elevations, Atlantic, and Pacific coasts. Mexico to Brazil and Colom- 
bia. A shrub or suffrutescent herb; stems densely strigose when 
young; petioles up to 5 cm. long; leaf blades elliptic or narrowly 
oblong, up to 21 cm. long and 8.5 cm. wide, usually smaller, serrulate, 
cuneate at the base, sparsely strigillose on the midrib and veins 
beneath, the primary veins 5-10 pairs; peduncles up to 4.5 cm. long, 
commonly equaling or exceeding the umbellate or subcymose pedi- 
cels; calyx exceedingly variable, 7.5-14 mm. long, orange or red, 



FLORA OF COSTA RICA 1153 

one-third to one-half the length of the corolla, the lobes ovate or 
ovate-lanceolate, subcoriaceous, nearly free, attenuate into a mucro; 
corolla red or orange, erect, not saccate or spurred at the base, up to 
22 mm. long, slightly ventricose, glabrous without, bearing a hairy 
ring within at the insertion of the stamens, glandular within the 
throat; ovary glabrous; disk annular. 

Besleria macropoda Donn. Smith, Bot. Gaz. 25: 155. 1898. 
Damp woods, El Recreo, Pittier 572. Low elevations, Atlantic 
coast. Endemic. An herb; young stems closely strigillose, the older 
ones glabrous; petioles 5-15 mm. long; leaf blades oblong to elliptic, 
10-14 cm. long, 3.5-5 cm. wide, cuneate at the base, denticulate, 
glabrous, the primary veins about 11 pairs; peduncle 6-13 cm. long, 
the pedicels 5-9, about 2 cm. long; calyx lobes subulate-lanceolate, 
10-12 mm. long, glabrous, free; corolla yellow, slightly ventricose, 
15-18 mm. long, glabrous without, with a hairy ring within at the 
insertion of the filaments, glandular-pilose within the throat; ovary 
glabrous; disk annular. 

Besleria notabilis Morton, sp. nov. Frutex 1 m. altus, caulibus 
1 cm. diam., angulatis, appresso-pubescentibus; petioli 3 cm. longi, 
crassi, pubescentes; lamina foliorum elliptica, usque ad 22 cm. longa 
et 10 cm. lata, breviter acuminata, basi cuneata, subcoriacea, integra, 
subtus puberula, venis marginibusque subtomentosis, venis primariis 
6-8-jugis; calycis lobi oblongi, 3 mm. longi, 1.5 mm. lati, obtusi 
liberi ciliolati, extus parce puberuli intus glabri; corolla flavescenti- 
aurea, 9-10 mm. longa, erecta ecalcarata, tubo 2 mm. lato, utrinque 
glabro, limbo terminal! angusto, lobis parvis obtusis aequalibus; 
androeceum glabrum; ovarium apice minute puberulum; stylus 
glaber; discus annularis glaber; bacca alba, calyce purpureo suffulta. 
La Palma, Prov. San Jose", alt. 1,500 meters, Werckle (Herb. Nac. 
Cost. 11603; type in U. S. Nat. Herb., No. 588,468). La Hondura, 
Prov. San Jose", M. Valeria 749. La Palma, M. Valeria 11*12; Tonduz 
7456. Cerro de La Palma near San Ramon, Prov. Alajuela, Brenes 
3750. Pejivalle, Prov. Cartago, Standley & Valeria 49121. A speci- 
men from Cricamola Valley, Prov. Bocas del Toro, Panama (Cooper 
197) is possibly to be referred to this species. 

Besleria princeps Hanst. Linnaea 34: 317. 1865-66. Chirrite. 
Costa Rica, without special locality, Wendland. Central mountains, 
600-1,600 meters elevation; Pejivalle; region of San Ramon. En- 
demic. Stems densely short-hirsute; petioles hirsute, 1-3.5 cm. 
long; leaf blades elliptic, up to 13 cm. long and 6 cm. wide, acute or 



1154 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

short-acuminate, rounded or obtuse at the base, entire, membranous, 
tuberculate above, the tubercles surmounted by a long hair, soft- 
hirsute beneath, the primary veins 7-10 pairs; common peduncle 
absent, the pedicels solitary, axillary, 2.5-4 cm. long, pilose; calyx 
lobes lanceolate, 15-17 mm. long, long-acuminate, erect, nearly free, 
irregularly sharp-serrate, densely sericeous-villous externally, gla- 
brous within; corolla bright orange, erect, not spurred or gibbous at 
the base, the tube posteriorly about 15 mm. long, anteriorly about 
20 mm. long, 7-8 mm. wide at the middle, externally pilose, bearing 
a dense, hairy ring within near the base, the throat oblique and 
densely pubescent within, the limb bilabiate, widely spreading, 20- 
35 mm. wide, the posterior lobes about 1 cm. long and wide, the 
lateral lobes about 15 mm. long and 20 mm. wide, the anterior about 
12 mm. long and 14 mm. wide, narrowed at the base; filaments and 
anthers densely pilosulous; ovary and style densely villous; disk 
annular, low, glabrous. 

Besleria pycnosuzygia Bonn. Smith, Bot. Gaz. 52: 53. 1911. 
Forests at La Palma, Prov. San Jose", 1,460 meters, Tonduz 12545. 
Known only from La Palma. An epiphytic shrub, the stems un- 
branched, strigillose at the apex; leaves subequal, petiolate, the 
petiole up to 2 cm. long, the blades oblanceolate, up to 14 cm. long 
and 5 cm. wide, acuminate at the apex, attenuate at the base, sub- 
coriaceous, entire, green above, pale beneath, glabrous above, 
minutely and sparsely strigillose beneath; flowers fasciculate, the 
pedicels up to 1 cm. long, strigose, the bracts small, oblong; calyx 
lobes oblong, 7 mm. long, erect, obtuse, entire, puberulous externally, 
pilose within near the middle; corolla scarlet, 21-24 mm. long, erect 
in the calyx, gibbous at the base, long-pilose externally, glandular- 
pilose within toward the base, the tube curved, scarcely ampliate 
or ventricose, slightly contracted in the throat, the limb narrow, 
about 5 mm. wide, regular, the lobes minute; anthers transversely 
oblong, 1 mm. long, 2 mm. broad, connate in pairs, the cells not 
confluent; ovary villous; disk annular, thick, glabrous, slightly 
interrupted dorsally. Because of the non-confluent anther cells the 
proper generic position of this species is doubtful. 

Besleria robusta Bonn. Smith, Bot. Gaz. 25: 156. 1898. Forests 
of Tsaki, Talamanca, Tonduz 9556. Low elevations in the Province 
of Limon. Panama. A shrub; stems 8-10 mm. thick, densely 
tomentose; petioles 2.5-5.5 cm. long, thick; leaf blades obovate, 
up to 32 cm. long and 17.5 cm. wide, subcoriaceous, entire, cuneate 



FLORA OF COSTA RICA 1155 

at the base, subtomentose on the veins beneath and on the margins, 
the primary veins 11-12 pairs; pedicels densely sericeous, 6-10 mm. 
long, elongating in fruit; calyx 10-12 mm. long, the lobes free, oblong- 
ovate, acute, coriaceous, entire, densely sericeous without, glabrous 
within; corolla orange, 9-14 mm. long, ventricose, glabrous except 
for the glandular throat within; ovary glabrous; disk annular. 

Besleria solanoides HBK. B. acutifolia Benth. Parabesleria 
costaricensis Oerst. Centralamer. Gesner. 53. 1858 (Turrialba, 900 
meters, Oersted). B. costaricensis Hanst. Linnaea 34: 330. 1865-66. 
Central mountains at middle elevations; Guanacaste. Honduras to 
Peru. A shrub 1.5-3 meters high; stems strigose upwardly, almost 
glabrous below; petioles 5 cm. long or less; leaf blades elliptic, up to 
15 cm. long and 5.5 cm. wide, cuneate at the base, inconspicuously 
denticulate, strigillose on the veins beneath, otherwise glabrous or al- 
most, the primary veins 6-8 pairs; pedicels few, up to 2 cm. long; calyx 
lobes oblong, 3-4 mm. long, obtuse, free, membranous, not concave, 
faintly nerved, nearly glabrous, ciliolate; corolla orange, 13-15 mm. 
long, erect, not spurred or saccate at the base, ventricose upwardly, 
glabrous externally, lacking a hairy ring within; androecium glabrous; 
ovary and style glabrous; disk entire, glabrous; berry orange. 

Besleria solanoides var. tenera Morton, var. nov. A var. 
typica corolla paullo puberula, ovario piloso differt. Cerro de La 
Carpintera, Prov. Cartago, alt. 1,500-1,850 meters, Standley 35554 
(type in U. S. Nat. Herb., No. 1,227,167). Estrella, Prov. Cartago, 
Cooper 5879. 

Besleria Standleyi Morton, sp. nov. Frutex 1 m. altus, caulibus 
crassis, 4 mm. diam., sericeis; petioli 1.5-2.5 cm. longi, sericei; 
lamina foliorum late elliptica, usque ad 18 cm. longa et 10 cm. 
lata, obtusa, basi obtusa vel cuneata, denticulata, supra viridis 
glabra, subtus pallida, minute sericea, venis primariis ca. 10-jugis; 
pedunculus communis obsoletus, pedicellis numerosis, in axillis 
foliorum dense aggregatis, ca. 10 mm. longis, apice incrassatis, 
brunneo-sericeis; calycis lobi suborbiculares, 4.5 mm. longi, rotundati 
venosi suberosi ciliolati, extus sericei; corolla aurantiaca, 13 mm. 
longa, erecta vel obliqua, ecalcarata, basi saccata, paullo ventricosa, 
tubo 5-6 mm. lato, fere glabro, annulo piloso carente, limbo terminali 
angusto, lobis parvis rotundatis aequalibus; androeceum glabrum; 
ovarium pilosulum; stylus crassus; discus annularis integer, glaber. 
La Palma, Prov. San Jose', alt. 1,600 meters, Standley 33005 (type 
in U. S. Nat. Herb., No. 1,226,074). 



1156 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Besleria trichostegia Bonn. Smith, Bot. Gaz. 61: 380. 1916. 
Tsaki, Prov. Limon, 200 meters, Tonduz 9558. Endemic. Stems 
slender, sparingly hirsute; petioles hirsute, up to 2 cm. long; leaf blades 
oblanceolate, up to 17 cm. long and 6.5 cm. wide, narrowed at the 
base, entire, ciliate, hirsute beneath, especially on the veins, the 
primary veins 7-10 pairs; pedicels few, 8-13 mm. long, hirsute, very 
slender; calyx 5-6 mm. long, the lobes free, linear-lanceolate, about 
7 mm. wide at the base, entire, long-pilose; corolla red, 11-13 
mm. long, pilose without, glabrous within at the insertion of the 
filaments, sparsely pilose within the throat; ovary pilose; disk 
semiannular. 

Besleria triflora (Oerst.) Hanst. Linnaea 34: 329. 1865-66. Para- 
besleria triflora Oerst. Centralamer. Gesner. 53. pi. 6. 1858. B. triflora 
var. subcorymbosa Bonn. Smith, Bot. Gaz. 61 : 380. 1916 (La Palma, 
Prov. San Jose", 1,460 meters, Tonduz 7453). Mountains of Naranjo, 
1,200-1,500 meters, Oersted. Central mountains, 1,300-2,400 meters; 
region of San Ramon. A shrub; stems sparingly strigose; petioles 
1.5-3 cm. long, strigose; leaf blades elliptic, up to 15 cm. long and 
7 cm. wide, usually much smaller, entire, membranous, cuneate at 
the base, nearly glabrous at maturity, the primary veins 5-7 pairs; 
peduncles variable in length, commonly shorter than the petioles; 
pedicels umbellate, subcorymbose, usually longer than the peduncle; 
calyx white at maturity, 5-6 mm. long, the lobes free, suborbicular, 
ciliolate, strigillose externally, the margins subscarious; corolla 
orange-red, about 15 mm. long, puberulous without, the limb about 
7 mm. broad, scarcely wider than the corolla tube; ovary hairy; 
disk annular; berry white. 

Besleria Wendlandiana Hanst. Linnaea 34 : 318. 1865-66. Costa 
Rica, without definite locality, Wendland 568. Central mountains, 
900-2,400 meters elevation; Pejivalle; region of San Ramon. Chiri- 
qui region, Panama. Stems densely lanate-tomentulose; petioles 
tomentulose, up to 2 cm. long; leaf blades elliptic, up to 20 cm. long 
and 9 cm. wide, cuneate at the base, serrate, lanate-puberulous on 
the veins beneath, the primary veins about 13 pairs; common 
peduncle elongate, up to 9 cm. long, the pedicels numerous, about 
1 cm. long; calyx green, the lobes ovate-lanceolate, 8-12 mm. long 
in anthesis, up to 16 mm. long in fruit, callose-serrulate, sparsely 
pilosulous; corolla yellow, 18-23 mm. long, horizontal, spurred at 
the base, the tube cylindric, not ventricose, externally pilosulous, 
the limb terminal, 10 mm. wide; ovary sericeous- ^ ; *V -allosulous, 
thicker posteriorly. 



FLORA OF COSTA RICA 1157 

CAM PANE A Dene. 

Suffrutescent herbs, shrubs, or short-climbing vines; leaves 
opposite, equal, petiolate, crenate-serrate; common peduncles 
elongate, axillary, pendent, one-several-flowered; calyx large, the 
tube costate, the lobes spreading, entire, pubescent; corolla large, 
open-campanulate, erect in the calyx, not spurred, the tube not 
ventricose, hairy externally, not contracted in the throat, yellowish 
green, the limb broad, subregular, the lobes spreading, purple-spotted 
within; filaments pilosulous; anthers connate, the cells divergent, 
oblong, not confluent; ovary over half inferior; style thick, exserted; 
stigma stomatomorphic; disk glands 5, free, large, glabrous; fruit 
capsular, opening by 2 valves. One additional species is found in 
Central America. 

Leaf blades soft-tomentose on both sides C. Humboldtii. 

Leaf blades pilosulous on both sides or glabrous above. .C. Oerstedii. 

Campanea Humboldtii (Klotzsch) Oerst. Centralamer. Gesner. 
34. 1858. Rhytidophyllum Humboldtii Klotzsch in Otto & Dietr. 
Allgem. Gartenz. 20: 18. 1852. Veragua, Panama. Warscewicz. 
Central mountains, 1,100-2,100 meters; region of San Ramon. 
Shrubs or shrubby vines to 2 meters or more; stems and peti- 
oles softly brown-tomentose; leaf blades oblong-lanceolate, oblique, 
up to 19 cm. long and 8 cm. wide, acuminate, brown-tomentose 
on both sides, at least when young; peduncles 10-16 cm. long, 1^4- 
flowered, the flowers nodding; calyx lobes ovate, 15 mm. long or 
more, 5-nerved, pubescent on both sides; corolla 4-5 cm. long, 1.7-2 
cm. wide, densely brown-tomentose externally, the limb 3-4 cm. wide. 

Campanea Oerstedii (Klotzsch) Oerst. Centralamer. Gesner. 34. 
pi. 1, 2. 1858. Rhytidophyllum Oerstedii Kl. in Otto & Dietr. Allgem. 
Gartenz. 20: 17. 1852. Between Candelaria and Cartago, 1,800-2,100 
meters, Oersted. Common in the central mountains, 1,300-1,800 
meters; Zarcero. Endemic. Similar to the preceding but always 
herbaceous, frequently epiphytic and pendent, not over 1 meter long, 
the stems often more coarsely hirsute, the leaf blades scabrous- 
pilosulous above, pilosulous beneath chiefly on the veins and veinlets, 
often reddish beneath; calyx lobes narrower, commonly oblong- 
lanceolate, usually 3-nerved. This species has often been confused 
with Campanea grandiflora (HBK.) Dene, of Colombia. 

Campanea Oerstedii var. laevis Morton, var. nov. A var. 
typica foliis supra semper glaberrimis differt. La Palma, Prov. San 



1158 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Jose", 1,450-1,550 meters, May 6-8, 1906, Maxon 448 (type in U. S. 
Nat. Herb., No. 1,336,367). 

CENTROSOLENIA Benth. 

Epiphytic, scandent shrubs; flowers in sessile or short-peduncu- 
late, axillary umbels, the bracts linear or subulate, the pedicels short, 
numerous; calyx lobes oblique, oblong-linear to subulate, acuminate, 
free, entire, or serrulate near the apex, hirsute, unequal, the posterior 
one shorter, saccate at the base; corolla horizontal in the calyx, 
spurred at the base, the tube ampliate upwardly or abruptly ventri- 
cose, not contracted in the throat, the limb spreading, subbilabiate; 
filaments inserted high in the corolla tube, contorted, glabrous; 
anthers connate, quadrate, the cells discrete or divergent, glabrous 
or bearded, dehiscent only part of their length; ovary villous; disk 
glands 1 or 2; lamellae of the placentae ovuliferous on the inner 
surface only. One or two additional species, represented by incom- 
plete material, occur in Costa Rica. 

Anthers bearded at the apex; leaf blades attenuate and decurrent 
into the petioles C. decurrens. 

Anthers glabrous; leaf blades rounded at the base, not decurrent, 
very long-petiolate C. longipetiolata. 

Centrosolenia decurrens Morton, sp. nov. Frutex scandens, 
caulibus elongatis vel raro abbreviatis, novellis rubro-hirsutis; folia 
petiolata, petiolo brevi vel usque ad 11 cm. longo, sursum alato; 
lamina foliorum oblanceolata, usque ad 29 cm. longa et 10.5 cm. lata, 
acuminata, basi attenuata et longe in petiolum decurrens, chartacea, 
denticulata, supra strigosa, subtus pallida strigosa, in venis rubro- 
hirsuta, venis primariis ca. 8-jugis; flores umbellati, pedunculo 
communi subnullo, bracteis subulatis, ca. 1 cm. longis, pedicellis 
numerosis 1-1.5 cm. (raro 3.5 cm.) longis, rubro-hirsutulis; calycis 
lobi virides, subulati, 12-17 mm. longi, 1 mm. lati, integri, rubro- 
hirsutuli, paullo inaequales, posticus minor, basi saccatus; corolla 
ochroleuca, rubro-tincta, ca. 4 cm. longa, basi calcarata (2.5 mm.), 
in calyce horizontalis, tubo extus rubro-pilosulo, supra basin angus- 
tato, in medio deflexo et abrupte ventricoso, turn angustato, faucem 
versus paullo ampliato, limbo patente, subbilabiato, lobis subintegris; 
filamenta gracilia, glabra, contorta, alte in corollae tubo affixa; 
antherae connatae, quadratae, loculis oblongis, ca. 1.2 mm. longis, 
divergentibus, apice perspicue barbatis; ovarium rubro-villosum; 
stylus pilosulus; disci glandula postica oblonga glabra; placentae 
lamellae intus solum ovuliferae. Finca Montecristo, on Rio Reven- 



FLORA OF COSTA RICA 1159 

tazon, below El Cairo, Prov. Limon, alt. 25 meters, Feb. 18-19, 1926, 
Standley & Valeria 48589 (type in U. S. Nat. Herb., No. 1,305,619). 
Same locality, Standley & Valeria 48416. La Conception, Llanuras 
de Santa Clara, Prov. Limon, Donnell Smith 6732. Vicinity of 
Guapiles, Prov. Limon, alt. 300-500 meters, Standley 37022, 37238, 
37501, 37553. The barbate anthers ally this with Centrosolenia 
glabra Benth. The Costa Rican record for Centrosolenia congesta 
(Oerst.) Morton, comb. nov. (Trichodrymonia congesta Oerst. Dansk. 
Vid. Selsk. Skrivt. V. 5: 112. 1861), a species of Mexico, is based on 
specimens of C. decurrens. 

Centrosolenia longipetiolata (Bonn. Smith) Morton, comb, 
nov. Episcia longipetiolata Bonn. Smith, Bot. Gaz. 25: 152. 1898. 
Known only from the type, Carillo, alt. 300 meters, Pittier 2493. 
Leaves long-petiolate (about 25 cm.), the blades broadly elliptic, 
22-25 cm. long, 12-16 cm. wide, short-cuspidate, rounded at the 
base, membranous, inconspicuously denticulate, glabrous on both 
sides, the primary veins 14-15 pairs; inflorescence congested, the 
common peduncle short, bracteate, the bracts subulate, 18-25 mm. 
long, pilosulous, the pedicels about 6 mm. long, numerous; calyx 
lobes narrowly linear, about 15 mm. long, 1 mm. wide, entire, pilosu- 
lous, unequal, the posterior shorter; corolla tubular, about 3 cm. 
long, spurred at the base, horizontal in the calyx, the tube not ven- 
tricose, about 2 mm. wide, slightly enlarged in the throat, the limb 
about 1 cm. wide, the lobes fimbriate; anthers subquadrate, 1 mm. 
long, not bearded. 

CODONANTHE (Mart.) Hanst. 

Codonanthe crassifolia (Focke) Morton, comb. nov. Hypo- 
cyrta crassifolia Focke, Tijdschr. Nat. Wetens. 5: 199. 1852. C. 
macradenia Bonn. Smith, Bot. Gaz. 25: 154. 1898 (Boruca, 460 
meters, Tonduz 6769). C. confusa Sandw.; C. cakarata auct., non 
Hanst. Low elevations, Atlantic and Pacific coasts. Panama to 
northern South America. Epiphytic, herbaceous vines with glabrous, 
shining stems; leaves of a pair equal, short-petiolate (4-5 mm.), 
the blades ovate or oval, up to 3.5 cm. long and 2 cm. wide, fleshy, 
entire, obtuse at both ends, glabrous throughout; flowers solitary, 
axillary, short-pedicellate; calyx lobes 5, free, subulate, 7-9 mm. 
long, entire, puberulous on both sides, oblique and slightly unequal; 
corolla white or yellowish, about 2 cm. long, horizontal in the calyx, 
spurred at the base, puberulous externally, the tube strongly ampliate 
upwardly, not contracted in the throat, the limb spreading, oblique, 



1160 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

slightly irregular; anthers orbicular, the cells discrete, separated by 
the enlarged connective; disk gland posterior, large. 

COLUMNEA L. 

Erect or scandent shrubs; leaves opposite, those of a pair equal or 
strongly unequal, usually short-petiolate; flowers axillary, solitary or 
fasciculate, short- or long-pedicellate, usually ebracteate; calyx 5- 
parted, the lobes nearly free, usually colored, acute or acuminate, 
often toothed or incised; corolla yellow or red, erect in the calyx, not 
spurred at the base, often gibbous at the base, the tube hairy, am- 
pliate upwardly, usually not ventricose, the limb regular or often 
strongly bilabiate; filaments connate, usually not adnate to the 
corolla except at the very base, the anthers persistently connate, 
oblong, dehiscent throughout; disk gland posterior, solitary; fruit 
baccate. A large, polymorphic genus, reaching its best development 
in Costa Rica. 

Corolla limb regular or slightly bilabiate. 

Leaves of a pair equal; calyx lobes entire (Stygnanthe) . 

C. calotricha. 

Leaves of a pair strongly unequal; calyx lobes often pectinate- 
toothed. 

Corolla yellow; flowers fasciculate, subsessile or short-pedicel- 
late; erect shrubs (Collandra). 

Leaf blades glabrous above or nearly so, marked with scarlet 
spots on the lower surface. 

Calyx lobes entire or nearly so C. consanguinea- 

Calyx lobes pectinate-toothed C. florida. 

Leaf blades pilose above, lacking scarlet spots. 

Calyx equaling or exceeding the corolla; corolla sericeous. 

C. purpurata. 

Calyx one-half as long as the corolla or less; corolla pilose. 

C. aureonitens. 

Corolla scarlet; flowers solitary or paired, long-pedicellate; 
epiphytic, pendent shrubs. 

Leaves glabrous above; calyx lobes pectinate-fimbriate (Sten- 

anthus). 

Leaf blades serrate; stems villous at the apex. .C. serrata. 
Leaf blades subentire; stems appressed-pubescent. 



FLORA OF COSTA RICA 1161 

Pedicels exceeding the leaves; teeth of calyx lobes not or 
scarcely longer than the width of the lobe. 

C. grata. 

Pedicels half as long as the leaves or less; teeth of the 
calyx lobes much longer than the width of the lobe. 

C. sanguinolenta. 

Leaves hirsute above; calyx lobes entire, serrate, or with a few 
subulate teeth (Ortholoma). 

Calyx lobes subserrate C. acuminata. 

Calyx lobes entire or with a few subulate teeth. 

Leaves lanceolate, purple beneath , . . .C. ochroleuca. 

Leaves oblanceolate, green beneath . . C. Warszewicziana. 
Corolla limb strongly bilabiate, the lower lobe linear, reflexed. 
Leaves of a pair strongly unequal, purple-spotted beneath; flowers 
fasciculate, short-pediceled, conspicuously bracteate (Crypto- 

columnea) C. praetexta. 

Leaves of a pair equal or nearly so, not purple-spotted beneath; 

flowers usually solitary, pedicellate (Eucolumnea). 
Corolla densely sericeous externally, the tube not much exceeding 

the calyx; filaments puberulous. 

Leaf blades linear; calyx lobes 12-14 mm. long. . . .C. linearis. 
Leaf blades not linear; calyx lobes 2-3 cm. long (except in C. 

mrecunda). 
Corolla 4 cm. long or less. 

Calyx lobes oblong-lanceolate, 3-5 mm. wide at the base. 

C. querceti. 
Calyx lobes linear-subulate, -not over 2 mm. wide. 

C. verecunda. 
Corolla 6-8 cm. long. 

Leaf blades strigillose beneath; epidermis of the stem 

peeling off in scales C. lepidocaula. 

Leaf blades appressed-pilose beneath; epidermis not 

peeling off in scales C. nicaraguensis. 

Corolla pilose externally, the tube much exceeding the calyx; 

filaments glabrate (except in C. magnified). 
Calyx lobes toothed. 
Stems and leaves strigose. 

Calyx lobes deeply pinnatifid C. flaccida. 



1162 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Calyx lobes slightly toothed at the base. .C. Oerstediana. 
Stems and leaves not strigose. 

Stems and leaves tomentose C. tomentulosa. 

Stems and leaves pilose or hirtous. 

Pedicels shorter than the calyx C. hirta. 

Pedicels longer than the calyx C. magnified. 

Calyx lobes entire. 
Calyx lobes broadest above the base. 
Stem hairs spreading. 
Leaf blades suborbicular, 1 cm. long or less. 

C. macrophylla. 

Leaf blades oblong-elliptic, 2-5 cm. long. . . .C. localis. 
Stem hairs subappressed; leaf blades 1.5-2.5 cm. long. 

C. microcalyx. 
Calyx lobes broadest at the base. 

Stems hirsute; leaf blades hirsute on both sides. 

Pedicels 5 mm. long or less C. hirta. 

Pedicels 2-2.5 cm. long C. gloriosa. 

Stems strigose; leaf blades strigose beneath, glabrous 

above. 

Calyx lobes ovate C. Oerstediana. 

Calyx lobes linear-lanceolate. 

Calyx lobes densely white-sericeous C. oxyphylla. 

Calyx lobes strigose near the base C. glabra. 

Columnea acuminata Benth. Ortholoma acuminatum Hanst. 
Colombia. Reported from Costa Rica by Oersted. Near C. War- 
scewicziana, but the calyx lobes serrate, not entire or subulate- 
toothed. 

Columnea aureonitens Hook. Atlantic coastal region. Colom- 
bia. Leaves strongly unequal, the larger ones oblanceolate, up to 21 
cm. long, serrate, densely red-pilose on both sides, not red-spotted; 
flowers fasciculate, pedicellate, the bracts deciduous; calyx lobes 
green, 13 mm. long, pectinate-toothed, brown-hirsute; corolla yellow, 
about 3.5 cm. long, pilose, the limb regular, about 7 mm. wide, the 
lobes suborbicular, erect. The Costa Rican plant may be specifically 
distinct from the Colombian type. 



FLORA OF COSTA RICA 1163 

Columnea calotricha Bonn. Smith, Bot. Gaz. 40: 9. 1905. 
Atirro, Donn. Smith 6728. Guatemala. An erect herb, the stems 
unbranched, red-hirsute; leaves subequal, petiolate (11-14 mm.), the 
blades oblong-elliptic, obtuse, cuneate at the base, remotely crenu- 
late, pilose on both sides, red beneath; pedicels 6-8 mm. long; calyx 
lobes spatulate-oblong, 13-15 mm. long, obtuse, pilose, entire; 
corolla reddish, tubular, 3 cm. long, long-hirsute, the tube not ventri- 
cose, the limb small, about 6 mm. wide, nearly regular, the lobes 
small, erect, subequal. 

Columnea consanguinea Hanst. Linnaea 34: 383. 1865-66. 
Turrialba, Wendland 509. Central mountains, 1,200-1,700 meters; 
region of San Ramon. Endemic. An epiphytic shrub with un- 
branched stems, 1.2 meters high; leaves unequal, the larger ones 
narrowly oblanceolate, up to 25 cm. long, entire, above glabrous, not 
red-spotted, beneath appressed-pilose, red-spotted, the smaller 
ones linear-lanceolate, about 2.5 cm. long; flowers fasciculate, the 
bracts persistent, entire, strigose; calyx lobes green, lanceolate, about 
2 cm. long, entire, strigose; corolla yellow, tubular, about as long 
as the calyx, densely sericeous, the limb narrow, regular, the lobes 
erect. 

Columnea flaccida Seem. Atlantic and Pacific coasts, at low 
elevations. Panama. An epiphytic, branched shrub, with slender, 
strigose stems; leaves subequal, very short-petiolate, the blades 
oblong-lanceolate, long-acuminate, about 3.5 cm. long, subentire, 
subcoriaceous, sparingly strigose on both sides; pedicels shorter than 
the calyx; calyx lobes red, linear-lanceolate, about 13 mm. long, 
deeply pinnatifid, red-hirsute; corolla red, 5-6 cm. long, gibbous at 
the base, pilose externally, the tube 4 cm. long, lateral lobes free for 
about 9 mm., the lower lobe ovate, about 1 cm. long; filaments 
glabrous. 

Columnea florida Morton, Journ. Wash. Acad. Sci. 27: 310. 
1937. Vicinity of El General, Prov. San Jose", 915 meters, Skutch 
2436. Pacific drainage, 500-1,100 meters. Panama. An epiphytic 
shrub with thick, hirsute stems; leaves opposite, strongly unequal, 
the larger ones short-petiolate, oblanceolate, up to 35.5 cm. long and 
10.5 cm. wide, short-acuminate, entire, glabrous above and bearing 2 
red spots, appressed-pilose beneath, red-spotted, the smaller leaves 
narrowly elliptic, about 3 cm. long, subsessile; flowers fasciculate, the 
pedicels up to 1 cm. long, bracteate, the bracts small, deciduous; 
calyx lobes ovate, 1.5-2.3 cm. long, densely brown-hirsute, pectinate- 



1164 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

dentate; corolla yellow, about 2.5 cm. long, cylindric, densely brown- 
hirsute, the limb subregular, about 5 mm. wide, the lobes suborbicular, 
2.5 mm. long. This species has been misidentified as C. sanguined 
Hanst. 

Columnea glabra Oerst. Centralamer. Gesner. 62. 1858. 
Mountains between Candelaria and Cartago, 1,800-2,100 meters, 
Oersted. Central mountains, 900-2,000 meters. Endemic. An 
epiphytic shrub with branched, strigose stems; leaves subequal, very 
short-petiolate, the blades oblanceolate, up to 3.5 cm. long, rounded 
at the apex, attenuate at the base, entire, glabrous above, thin- 
strigose beneath or glabrate; pedicels shorter than the calyx; calyx 
lobes green, linear-lanceolate, about 15 mm. long, obtuse, entire, 
subappressed-pubescent near the base; corolla scarlet, about 7 cm. 
long, the tube about 4 cm. long, pilose externally, not contracted 
in the throat, the lateral lobes deltoid, acute, free for about 9 mm., 
the lower lobe linear-oblong, acute, about 15 mm. long; filaments 
glabrous. 

Columnea gloriosa Sprague, Bot. Mag. 137: pi. 8378. 1911. 
Known only from the type, cultivated from Costa Rica. Epiphytic, 
the stems hirsute; leaves subequal, the blades ovate or ovate-oblong, 
2-3.5 cm. long, obtuse, rounded at the base, hirsute above, the nerves 
impressed, hirsute beneath on the elevated nerves; pedicel 2-2.5 cm. 
long; calyx lobes broadly ovate, 1.5 cm. long, entire, villous; corolla 
scarlet, 6.5-8 cm. long, sparsely villous. 

Columnea grata Morton, nom. nov. Columnea heterophylla 
Hanst. Linnaea 34: 390. 1865-66, non Roxb. Stenanthus heterophyllus 
Oerst. Centralamer. Gesner. 48. pi. 5. 1858. Naranjo, Oersted. 
Pacific and Atlantic coasts, at low elevations. Endemic. A pendent, 
epiphytic shrub with branched, appressed-pubescent stems; leaves 
strongly unequal, the larger ones oblanceolate, 10 cm. long, suben- 
tire, glabrous above, strigose beneath, usually red-spotted, the smaller 
ones stipule-like, not over 1 cm. long; flowers solitary, the pedicels 
pendent, longer than the leaves, bracteate at the base; calyx lobes 
lanceolate, about 3 cm. long, green, hirsute, pectinate-toothed, the 
teeth usually shorter than the width of the lobes; corolla scarlet, 4-5 
cm. long, pilose, the limb 15-17 mm. wide, subregular. 

Columnea hirta Klotzsch & Hanst. Linnaea 34: 403. 1865-66. 
C. hirsuta Kl. ex Oerst. Centralamer. Gesner. 61. 1858, non Swartz. 
Veragua, Panama, Warscewicz. Central mountains. A pendent, 



FLORA OF COSTA RICA 1165 

branched shrub; stems red-villous; leaves equal, short-petiolate (4-5 
mm.), oblong, 2.5-3 cm. long, obtuse, densely pilose on both sides, 
entire; pedicels 7 mm. long or less; calyx lobes green, linear-lanceo- 
late, about 15 mm. long, with 1 or 2 short teeth on either side, long- 
pilose; corolla scarlet, about 7 cm. long, pilose, the tube slender, 
about 4 cm. long, becoming 8 mm. wide in the throat, the galea 
oblong, 3 cm. long, entire, concave, the lateral lobes oblong, free 
for about 1 cm., the lower lobe lanceolate, reflexed, about 1.8 cm. 
long; filaments nearly glabrous. 

Columnea lepidocaula Hanst. Linnaea 34: 411. 1865-66. 
Costa Rica, without locality, Valentini. Central mountains, at 
middle elevations. Endemic. An epiphytic shrub, with coarse 
stems, the epidermis of which peels off in scales; leaves subequal, 
short-petiolate, oblanceolate, 5-9 cm. long, acute, narrowed at the 
base, entire, glabrous above, sparsely strigose beneath; pedicels 
shorter than the calyx; calyx lobes green, oblong-lanceolate, 2-2.5 
cm. long, acute, entire, strigose; corolla 6.5-7.5 cm. long, scar- 
let, the throat yellow, densely white-sericeous externally, the tube 
shorter than the limb, little longer than the calyx; filaments densely 
puberulous. 

Columnea linearis Oerst. Centralamer. Gesner. 59. 1858. At 
1,800-2,100 meters, Oersted. Central mountains, at middle eleva- 
tions; Atlantic coast. Endemic. An epiphytic shrub, the stems 
branched, substrigose; leaves subequal, subsessile, the blades linear, 
4-8 cm. long, acuminate, entire, glabrous above, strigillose beneath; 
pedicels shorter than calyx; calyx lobes red, ovate-lanceolate, 12-14 
mm. long, acuminate, entire, strigillose; corolla scarlet, about 4 cm. 
long, white-sericeous externally; filaments puberulous. 

Columnea localis Morton, nom. nov. Columnea microcalyx var. 
macrophylla Bonn. Smith, Bot. Gaz. 31: 118. 1901, non C. macro- 
phylla Kuntze. Known only from Tucurrique (type, Tonduz 12932). 
An epiphytic shrub, the stems slender, branched, white-pilose; leaves 
green, subequal, short-petiolate (1.5 mm.), the blades oblong-elliptic, 
2-5 cm. long, 1.1-1.7 cm. wide, acute or obtuse, rounded at the base, 
membranous, entire, soft-pilose on both surfaces; pedicels solitary, 
about 14 mm. long, pilose; calyx lobes green, oblanceolate, about 
12 mm. long, acute, narrowed at the base, entire, pilose on both 
surfaces; corolla scarlet, 7 cm. long, gibbous at the base, thin-pilose 
externally, the tube short (2 cm.), the galea broad, the lateral lobes ad- 
nate for most of their length, the lower lobe linear, about 2.2 cm. long. 



1166 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Columnea magnifica Klotzsch & Hanst. ex Oerst. Centralamer. 
Gesner. 60. 1858. ?C. Wendlandiana Hanst. Linnaea 34: 402. 
1865-66 (without definite locality, Wendland). ?C. oblanceolata 
Sprague, Kew Bull. 449. 1908 (without special locality, Hubsch). 
Cerro de Aguacate, Hoffmann. Central mountains, 1,400-2,500 
meters. Panama. Stems hirsute, unbranched; leaves subequal, 
lanceolate or oblanceolate, not over 9 cm. long, pilose on both sides, 
eventually glabrate above, subentire, not red-spotted; flowers 
solitary, the pedicels about 1.5 cm. long, erect; calyx lobes green, 
lanceolate, about 15 mm. long, pilose, remotely serrulate; corolla 
scarlet, up to 7 cm. long, pilose, not narrowed in the throat, strongly 
bilabiate, the galea entire; filaments densely puberulous. 

Columnea microcalyx Hanst. Linnaea 34: 408. 1865-66. Costa 
Rica, without definite locality, Wendland 910. Central mountains, 
at middle elevations; region of San Ramon. Endemic. A pendent, 
epiphytic shrub, the stems branched, substrigose; leaves subequal, 
short-petiolate, the blades lanceolate or ovate-lanceolate, 1.5-2.5 cm. 
long, acute, rounded at the base, entire, subcoriaceous, glabrate or 
thin-strigose on both sides; pedicels shorter than the calyx; calyx 
lobes oblanceolate to obovate, about 12 mm. long, obtuse, narrowed 
toward the base, entire, pilose, usually reddish within; corolla scarlet, 
6-7 cm. long, pilose externally, the tube shorter than the limb; 
filaments glabrous. 

Columnea microphylla Klotzsch & Hanst. ex Oerst. Central- 
amer. Gesner. 63. 1858. Without definite locality, Warsceuricz. 
Central mountains, at middle elevations. Endemic. A small, 
pendent shrub, the stems slender, red-hirtous; leaves subequal, 
short-petiolate, the blades oval or suborbicular, 1 cm. long or less, 
obtuse, entire, pilose on both sides; pedicels shorter than the calyx; 
calyx lobes oblanceolate, narrowed at the base, obtuse at the apex, 
about 1 cm. long, entire, long-pilose; corolla scarlet and yellow, about 
6 cm. long, pilose externally, the tube shorter than the limb; filaments 
glabrous. 

Columnea nervosa (Klotzsch) Hanst. (Pentadenia nervosa Klotzsch 
ex Oerst.) was reported from Costa Rica, but the locality is in Panama. 

Columnea nicaraguensis Oerst. Atlantic coast, at low eleva- 
tions. Nicaragua. Changuinola Valley, Panama. An epiphytic shrub, 
the stems closely strigose, the epidermis not peeling in scales; leaves 
subequal, petiolate (5 mm.), the blades oblong or lanceolate, 7-11 cm. 
long, acute, rounded at the base, entire, strigillose above, appressed- 



FLORA OF COSTA RICA 1167 

pilose beneath; pedicels shorter than the calyx; calyx lobes lanceolate 
or ovate-lanceolate, 2.5-3 cm. long, acuminate, entire, long-pilose 
at the base, strigose upwardly; corolla red, 6-8 cm. long, densely 
sericeous externally, the tube not much longer than the calyx; fila- 
ments puberulous. 

Columnea ochroleuca (Klotzsch) Hanst. (Ortholoma ochroleiLcum 
Klotzsch ex Oerst.) was reported from Costa Rica, but the locality 
is in Panama. 

Columnea Oerstediana Klotzsch ex Oerst. Centralamer. Gesner. 
61. pi. 8. 1858. ?C. tennis Klotzsch ex Oerst. op. cit. 61 (Veragua, 
Panama, Warscewicz). Naran jo, 1,200-1, 500 meters, Oersted. Central 
mountains, at middle elevations. A pendent, epiphytic shrub, the 
stems branched, strigose; leaves subequal, short-petiolate, the blades 
ovate, 1-1.5 cm. long, coriaceous, obtuse, rounded at the base, 
entire, or slightly toothed at the base, glabrous above, thin-strigose 
beneath; pedicels shorter than the calyx; calyx lobes green, ovate, 
about 15 mm. long, acute, entire, sparingly pilose, the tube longer 
than the limb ; filaments glabrous. 

Columnea oxyphylla Hanst. Linnaea 34: 405. 1865-66. Costa 
Rica, without definite locality, Wendland 778. Low elevations on the 
Atlantic coast. Endemic. A shrub (0.9-1.5 meters), pendent from 
trees; stems strigose; leaves subequal, short-petiolate (2 mm.), the 
blades lanceolate, 3-4 cm. long, 8-10 mm. wide, subcoriaceous, long- 
acuminate, entire, glabrous above, thin-strigose beneath; flowers 
solitary, the pedicel about 15 mm. long, white-sericeous; calyx lobes 
green, linear-lanceolate, about 20 mm. long, long-acuminate, entire, 
densely white-sericeous; corolla scarlet, about 7 cm. long, erect in the 
calyx, pilose, the tube about 3 cm. long, not contracted in the throat, 
the galea about 4 cm. long, obtuse, entire, the lateral lobes oblong, 
obtuse, free for about 12 mm., the lower lobe reflexed, 2.5 cm. long; 
filaments glabrous. 

Columnea pendula (Klotzsch) Hanst. (Ortholoma pendulum 
Klotzsch ex Oerst.) was reported from Costa Rica, but the locality 
is in Panama. 

Columnea praetexta Hanst. Linnaea 34: 394. 1865-66. Costa 
Rica, without special locality, Warscewicz 5. Known only from 
the type. Leaves strongly unequal, the larger ones oblanceolate, 
20-25 cm. long, villous on both sides, purple-spotted beneath, the 
smaller ones 2.5-5 cm. long; flowers fasciculate, the pedicels very 



1168 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

short, the bracts large, orbicular, crenulate; calyx lobes lanceolate, 
subentire, villous, about 2.5 cm. long; corolla orange, the tube tomen- 
tose, clavate, the throat narrowed, the limb bilabiate. 

Columnea purpurata Hanst. Linnaea 34: 386. 1865-66. 
Costa Rica, without definite locality, Wendland 548. Common at 
elevations from sea level to 1,500 meters; region of San Ramon; 
Guanacaste. Endemic. A soft-woody shrub with unbranched stem ; 
leaves clustered at the apex, strongly unequal, the larger ones oblan- 
ceolate, up to 30 cm. long, serrate, pilose on both sides, not red- 
spotted, the smaller leaves not over 3 cm. long, long-acuminate; 
flowers fasciculate, subsessile, bracteate, the bracts scarlet, about 
3 cm. long, spinulose-toothed, long-pilose; calyx lobes scarlet, lanceo- 
late, about 3 cm. long, spinulose-toothed, long-pilose; corolla yellow, 
tubular, 3 cm. long, sericeous, the limb narrow, regular, the lobes 
erect, 4 mm. long. 

Columnea querceti Oerst. Centralamer. Gesner. 59. 1858. 
C. erythrocalyx Klotzsch ex Oerst. loc. cit. (without locality, War- 
scewicz). At 1,800-2,100 meters, Oersted. In central mountains, at 
middle elevations. Endemic. An epiphytic shrub, the stems coarse, 
the epidermis peeling off in scales; leaves subequal, the blades oblan- 
ceolate or linear-oblanceolate, 4-10 cm. long, acute, attenuate at the 
base, entire, glabrous above, strigillose beneath; pedicels much 
shorter than the calyx; calyx lobes red, oblong-lanceolate, about 2 cm. 
long, acute, entire, pilose at the base; corolla scarlet, about 4 cm. 
long, densely sericeous externally, the tube shorter than the calyx; 
filaments puberulous. 

Columnea sanguinolenta (Klotzsch) Hanst. Linnaea 34: 389. 
1865-66. Stenanthus sanguinolentus Klotzsch ex Oerst. Centralamer. 
Gesner. 49. 1858. S. squarrosus Klotzsch ex Oerst. loc. cit. (Veragua, 
Panama, Warscewicz). Low elevations, Atlantic and Pacific coastal 
regions. Panama. An epiphytic shrub, the stems unbranched, 
appressed-pubescent; leaves strongly unequal, the larger ones oblan- 
ceolate, up to 12 cm. long, subentire, glabrous above, appressed- 
pilose beneath, often red-spotted beneath, the smaller ones stipuli- 
form, less than 1 cm. long; pedicels erect, solitary, axillary, usually 
less than half as long as the leaves, bracteate at the base; calyx lobes, 
2.5-3 cm. long, green or red, hirsute, deeply pectinate-fimbriate, the 
teeth longer than the width of the lobe; corolla scarlet, about 4 cm. 
long, pilose, the limb sub-regular, about 12 mm. wide. 



FLORA OF COSTA RICA 1169 

Columnea serrata (Klotzsch) Hanst. Stenanthus serratus 
Klotzsch. Veragua, Panama, Warscewicz. Doubtfully reported from 
Costa Rica by Hanstein. Near C. sanguinolenta, but the stems hir- 
sute at the apex and the leaf blades serrate, not red-spotted. 

Columnea tomentulosa Morton, nom. nov. Columnea tomen- 
tosa Oerst. Centralamer. Gesner. 64. 1858, non Roxb. Low eleva- 
tions on the Atlantic coast. Nicaragua. A scandent, branched shrub ; 
stems tomentose; leaves short-petiolate (2 mm.), subequal, the blades 
oval, 2.5 cm. long or less, obtuse, soft-tomentose on both sides, entire; 
flowers solitary, the pedicel 1 cm. long or less, white- tomentose; calyx 
lobes linear, 7-10 mm. long, acuminate, deeply subulate-toothed (the 
teeth 2-3 to a side), pilose externally, glabrous within; corolla scarlet, 
4-4.5 cm. long, gibbous at the base, erect in the calyx, the tube 
slender, pilose, not contracted in the throat (this 6 mm. wide), the 
galea oblong, 15 mm. long, obtuse, entire, the lateral lobes triangular, 
free for about 3.5 mm., the lower linear, reflexed, 8.5 mm. long; 
filaments glabrous, nearly equaling the galea; style pilosulous; stigma 
bilobed. 

Columnea verecunda Morton, sp. nov. Frutex epiphyticus, 
0.3-0.8 m. altus, caulibus crassis, ca. 6 mm. diam., parce ramosis, 
hirsutis, inferne squamulis epidermidis retroversis instructis; petioli 
brevissimi, 3-4 mm. longi, hirsuti; lamina foliorum oblongo-oblanceo- 
lata, usque ad 11 cm. longa et 2.5 cm. lata, acuta, basi rotundata, 
non decurrens, subcoriacea, integra, supra glabrescens, subtus rubra, 
strigosa, venis primariis ca. 6-jugis; flores fasciculati, axillares, pauci, 
pedunculo communi nullo, pedicellis 5-6 mm. longis, bracteis minutis, 
linearibus; calycis lobi liberi, lineari-subulati, 13-15 mm. longi, 2 mm. 
basi lati, integri, rubri, externe sericei, intus glabri, aequales, erecti; 
corolla flava vel pallide rubra, 3-4 cm. longa, in calyce erecta, basi 
parum gibbosa, tubo externe albo-sericeo, sursum gradatim ampliato, 
12-15 mm. longo, quam limbo multo breviore, limbo valde bilabiate, 
galea integra porrecta, lobis lateralibus cum galea alte connatis, lobo 
inferiore reflexo, lineari, ca. 15 mm. longo; filamenta basi in tubum 
postice fissum connata, sursum per paria connata, partibus liberis 
filiformibus, pilosulis, rectis; ovarium villosum; stylus pilosulus; 
stigma stomatomorphum; disci glandula emarginata, crassa. Vara 
Blanca de Sarapiqui, Prov. Alajuela, alt. 1,615 meters, Jan., 1938, 
Skutch 3466 (type in U. S. Nat. Herb., No. 1,643,546). Zarcero, 
Prov. Alajuela, alt. 1,350 meters, A. Smith H67. Region of San 
Ram6n, Prov. Alajuela, Brenes 3708, 4497, 5336, 13584; Soils 497; 
Tonduz 17825. 



1170 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Columnea Warszewicziana (Klotzsch) Hanst. Ortholoma War- 
szewiczianum Klotzsch ex Oerst. Ortholoma vestitum Klotzsch ex 
Oerst. Central mountains, 1,400-2,400 meters; region of San 
Ramon. Panama. A pendent, epiphytic shrub, the stems branched, 
0.6-1.2 meters long, reddish-villous at the apex; leaves strongly 
unequal, the larger ones oblanceolate, up to 12 cm. long, subserrate, 
hirsute on both sides, not red-spotted, the smaller ones stipule-like; 
pedicels up to 4 cm. long, pendent, red-villous; calyx lobes about 
1 cm. long, lanceolate, acuminate, villous, entire or with a few subu- 
late teeth; corolla scarlet, 4-5 cm. long, strongly ventricose, pilose, 
the limb subregular, about 13 mm. wide. 

DIASTEMA Benth. 

Low, delicate herbs, the stems unbranched, hirsute; leaves green, 
opposite, subequal, thin-membranous, coarsely toothed; inflorescence 
axillary or terminal, racemose, bracteate, the bracts opposite; calyx 
tube turbinate, the lobes free, elongate, entire, green; corolla white, 
tubular, erect in the calyx, not spurred, not ventricose, not markedly 
ampliate upwardly, the limb terminal, regular or subbilabiate; 
anther cells discrete, orbicular; ovary half inferior; style slender, 
elongate; stigma subbilobed; disk glands 5, linear. One or two 
additional species are found in Central America. 

Calyx tube and lobes glabrate or sparingly strigose. 

Leaf blades dentate or crenate-dentate D. exiguum. 

Leaf blades serrate or biserrate D. cristatum. 

Calyx tube and lobes spreading-pilose D. micranthum. 

Diastema cristatum Hanst. Linnaea 34: 438. 1865-66. ?Dias- 
temella bracteosa Oerst. Centralamer. Gesner. 26. 1858 (Cerro de 
Aguacate, 360 meters, Oersted). Diastema bracteosum Hanst. loc. 
cit. Cerro de Aguacate, Hoffmann 692. Atlantic coast; El General; 
region of San Ramon. From 400-1,100 meters elevation. Endemic. 
Leaf blades ovate or oblong, pilose above, setulose-pilosulous beneath 
on the veins and veinlets; inflorescence usually terminal, the common 
peduncle elongate, glabrate; calyx lobes linear-lanceolate, 3.5-5 mm. 
long, 1-1.5 mm. wide at the base, ciliate, glabrate or strigose; corolla 
15-20 mm. long, sparsely pilose toward the apex, the tube slender, 
about 4 mm. wide; disk glands long and slender, often equaling the 
ovary, often clavate. 

Diastema exiguum Morton. Pejivalle, 900 meters. Panama. 
Similar to the preceding, but the leaf blades dentate or crenate- 



FLORA OF COSTA RICA 1171 

dentate, glabrate, the calyx lobes larger (5-7 mm. long), and the disk 
glands not clavate. 

Diastema micranthum Donn. Smith, Bot. Gaz. 61: 377. 1916. 
JeYico, Llanuras de Santa Clara, 400 meters, Pittier 7602. La Palma 
de San Jose"; La Hondura. Endemic. Stems and petioles densely 
pilose; leaf blades pilose on both sides, dentate or dentate-serrate, 
never biserrate; peduncle and rachis pilose, often reduced, the 
flowers sometimes appearing as if aggregate in the leaf axils; calyx 
tube densely spreading-pilose, the lobes sparingly pilose, broadly 
lanceolate, spreading, often recurved at the apex; corolla 10-18 
mm. long; disk glands broadly linear, shorter than the ovary, 
not clavate. This is perhaps the species mentioned from Costa 
Rica by Fritsch as D. pictum Regel. 

DRYMONIA Mart. 

Shrubs or vines; leaves opposite, equal, usually long-petiolate; 
flowers usually solitary, rarely fasciculate; calyx lobes large, broad, 
5-parted, free or rarely partially connate, often colored, often toothed 
or fimbriate; corolla infundibuliform, horizontal in the calyx, spurred 
at the base, the tube not ventricose, ampliate upwardly, not con- 
tracted in the throat, the limb broad, spreading, slightly bilabiate, 
the lobes often toothed or fimbriate; filaments short-connate at the 
base, the anthers narrowly oblong, large, usually free, sagittate at 
the base, only partially dehiscent; disk gland solitary, posterior. 

Calyx lobes obtuse or rounded. 

Ovary red-villous; stems and leaves hirsute. Corolla red, long- 
pilose; leaves green beneath D. cdloplectoides. 

Ovary glabrous; stems and leaves strigillose or glabrous. 

Corolla white, minutely puberulous; leaves purple beneath, 
minutely strigillose, crenulate. 

Filaments contorted D. marmorata. 

Filaments straight D. turrialvae. 

Corolla pink, glabrous; leaves green beneath, glabrous, entire. 

D. conchocalyx. 
Calyx lobes acute or acuminate. 

Plants epiphytic vines. Calyx lobes usually serrulate. 
Leaves and calyces puberulous. Corolla yellowish white. 

D. mollis. 
Leaves and calyces strigillose. 



1172 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Leaf blades broadly ovate D. ovata. 

Leaf blades oblong D. spectabilis. 

Plants erect, terrestrial shrubs (epiphytic in D. parviflora). 
Calyx lobes sharply toothed or pectinate. 

Corolla yellow, 2.8 cm. long or less D. parviflora. 

Corolla white, veined with red, about 4 cm. long . . D. fimbriata. 
Calyx lobes entire. 

Corolla pure white, the lobes long-fimbriate; calyx densely 

puberulous D. lanceolata. 

Corolla not pure white, the lobes erose or dentate; calyx gla- 
brous or slightly strigillose. 
Pedicels longer than the calyx; corolla 4.5-6 cm. long, the 

limb about 4 cm. wide D. rubra. 

Pedicels shorter than the calyx; corolla 2.5-3 cm. long, the 
limb 2 cm. wide or less D. Warszewicziana. 

Drymonia alloplectoides Hanst. Linnaea 34: 358. 1865-66. 
Costa Rica, without definite locality, Warscewicz 34. Low elevations 
on the Atlantic coast. Nicaragua. A subscandent, epiphytic shrub, 
0.6-1.2 meters high, the stems brown-hirsute; leaves subequal, short- 
petiolate (0.5-1.5 cm.), the blades ovate to oblong, up to 10 cm. long 
and 5 cm. wide, acute, rounded at the base, serrulate or serrate, 
hirsute on both sides; flowers solitary, the pedicel up to 1.8 cm. long, 
densely hirsute; calyx lobes red, connate at the base, strongly un- 
equal, the 2 anterior ones broadly elliptic, 2.5-2.8 cm. long, the 2 
lateral ones about 2.3 cm. long, the posterior one suborbicular, about 
0.9 cm. long, all obtuse, serrulate, hirsute on both sides, the midrib 
prominent, elevated; corolla red, 5-5.5 cm. long, saccate at the 
posterior base, horizontal in the calyx, densely pilose, the tube 
gradually ampliate, becoming 2 cm. wide, not contracted in the 
throat, the limb slightly bilabiate, 3 cm. wide, glabrous within, not 
spotted, the anterior lobe fimbriate; filaments flattened, connate, 
partially adnate to the corolla, the anthers oblong, 7 mm. long; ovary 
red-villous; gland posterior, glabrous; style pilosulous. 

Drymonia conchocalyx Hanst. Linnaea 34: 360. 1865-66. 
Between Desengano and Cariblanco, Wendland 954. Central moun- 
tains, 1,400-1,700 meters; region of San Ramon. Endemic. An 
epiphytic shrub to 1.8 meters high, the stems glabrous; leaves 
slightly unequal, short-petiolate (up to 2.5 cm.), the blades elliptic 
to oblong, up to 22 cm. long and 9 cm. wide, cuspidate, rounded and 



FLORA OF COSTA RICA 1173 

oblique at the base, entire, green, wholly glabrous; flowers solitary 
or in pairs, not bracteate, the pedicels red, 3-5 cm. long, glabrous; 
calyx red, oblique, irregular, the 2 anterior lobes oblong, about 3 cm. 
long, 1 cm. wide, obtuse, connate anteriorly for 1.5 cm., the lateral 
lobes ovate, obtuse, about 3 cm. long, 2 cm. wide, connate to the 
anterior lobes for 1 cm., the posterior lobe 2 cm. long, acuminate, 
incurved, all the lobes entire, glabrous; corolla tube whitish, spotted 
with lavender inside, the lobes pink, drying purple, about 7 cm. long, 
saccate at the base, horizontal in the calyx, glabrous, the tube 
slightly ampliate, not contracted in the throat, the limb about 4 cm. 
wide, the lobes all fimbriate; filaments flattened, glabrous, contorted, 
adnate toward the base to the corolla, connate in pairs; anthers 
oblong, 6.5 mm. long; ovary strigillose; style glabrous. 

Drymonia fimbriata Morton, Journ. Wash. Acad. Sci. 27: 311. 
1937. Near El General, Prov. San Jose", 880 meters, Skutch 2839. 
Known only from El General. A shrub 1.5 meters high, the stems 
quadrangular, strigillose; petioles up to 4.7 cm. long; leaf blades 
broadly ovate, 22 cm. long, 11 cm. wide, short-acuminate, oblique 
and decurrent at the base, denticulate, nearly glabrous above, sparsely 
strigillose beneath; flowers aggregate in the leaf axils, the pedicels 
strigillose, about 13 mm. long, bracteate at the base, the bracts lance- 
olate, 5 mm. long, entire, puberulous; calyx lobes green, oblong, free, 
subequal, oblique, about 15 mm. long, puberulous on both sides, 
pectinate-fimbriate, the teeth filiform, up to 10 mm. long, puberulous; 
corolla white with red stripes, about 4 cm. long, horizontal in the 
calyx, spurred at the base (4.5 mm.), the tube pilosulous, curved, 
ampliate upwardly, not contracted in the throat, the limb glabrous, 
oblique, bilabiate, the upper and lower lips lacerate-dentate, the 
lateral lobes entire; filaments glabrous; anthers oblong, connate; 
ovary puberulous. 

Drymonia lanceolata (Hanst.) Morton, comb. nov. Episcia 
lanceolate, Hanst. Linnaea 34: 346. 1865-66. Cariblanco, Wendland 
902. Central mountains, at middle elevations. Panama. Ter- 
restrial herbs, 1-1.2 meters high, the stems unbranched, minutely 
strigillose at the apex; petioles up to 6 cm. long; leaf blades elliptic 
or broadly elliptic, up to 20 cm. long and 9 cm. wide, short-acuminate, 
cuneate at the base, membranous, crenulate, sparsely strigose above, 
soon glabrous, puberulous beneath, the primary veins about 6 pairs; 
flowers axillary, solitary, the pedicels 2.4-5 cm. long, longer than the 
calyx; calyx lobes pale green, oblique, ovate-lanceolate, 2.2-4 cm. 



1174 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

long, long-acuminate, entire, densely puberulous externally, unequal, 
the posterior smaller, saccate at the base; corolla white, 3.6-4.5 cm. 
long, oblique in the calyx, short-spurred at the base, the tube curved, 
puberulous externally, broad, not or scarcely contracted in the 
throat, the limb broad, the lobes unequal, long-fimbriate; anthers 
oblong, sagittate, about 5 mm. long; ovary strigillose. 

Drymonia marmorata Hook. f. Bot. Mag. 110: pi 6763. 1884. 
Locality given doubtfully as Guiana, but probably Costa Rica. 
La Palma; Llanuras de Santa Clara; El General. A terrestrial herb 
with unbranched, strigillose stems; leaves long-petiolate (up to 12 
cm.), the blades oblong to orbicular, up to 30 cm. long and 20 cm. 
broad, cuspidate, rounded or cordate at the base, crenulate, green 
and glabrous above, purple and minutely strigillose beneath; flowers 
in short, congested, bracteate racemes, the bracts purple, oblong, 
about 3 cm. long, sparingly strigillose, the pedicels up to 4 cm. long, 
glabrous; calyx lobes red-purple, ovate, about 2 cm. long, obtuse, 
entire, sparingly strigillose; corolla white, about 6 cm. long, saccate 
at the base, horizontal in the calyx, finely puberulous, the tube 
ampliate upwardly, not contracted in the throat, the limb bilabiate, 
the anterior lobe the largest, fimbriate; filaments glabrous, contorted; 
anthers oblong, sagittate; ovary glabrous. 

Drymonia mollis Oerst. Centralamer. Gesner. 37. 1858. 
Ujarras, Oersted. Central mountains; Atlantic coast; region of San 
Ramon. Endemic. An epiphytic vine, the stems densely puberulous 
toward the apex; leaf blades oblong, up to 12.5 cm. long and 5 cm. 
wide, acute, rounded or cuneate at the base, serrulate, puberulous on 
both sides; pedicels solitary, 1-1.5 cm. long; calyx lobes ovate, 2.5-3 
cm. long, serrulate, puberulous on both sides; corolla yellowish white, 
scarcely longer than the calyx, horizontal in the calyx, spurred at the 
base, the tube villosulous, enlarged to the throat, the limb bilabiate, 
the lobes fimbriate. 

Drymonia ovata Hanst. Linnaea 34: 355. 1865-66. Rio 
Virilla, Hoffmann 545. Description adapted from the original. 
Stems reddish, hirto-pubescent, scandent; leaf blades broadly ovate, 
7.5-10 cm. long, 3.7-6.7 cm. wide, subentire or denticulate, scabrid- 
pubescent; pedicel about equaling the petiole; calyx lobes broadly 
ovate, 2.5-3 cm. long, acute, lightly crenate-serrate, densely pubes- 
cent, unequal; corolla yellowish rose, about 3 cm. long, short- tomen- 
tulose, the limb 1.8 cm. wide, the lobes fimbriate-denticulate; anthers 
very long, sagittate. 



FLORA OF COSTA RICA 1175 

Drymonia parviflora Hanst. Linnaea 34: 352. 1865-66. Can- 
delaria, Hoffmann 798. Description adapted from the original. 
Epiphytic, the stems thick, tetragonal, branched, pubescent; leaves 
about 30 cm. long, 7.5-10 cm. wide, the blades elongate-elliptic, 
acuminate, narrowed at the base and long-decurrent into the petiole, 
sinuate-denticulate; flowers fasciculate, numerous in the leaf axils, 
the pedicels hirto-pubescent, shorter than the flowers; calyx lobes 
oblong-lanceolate, 12-14 mm. long, membranous, mucronate-acumi- 
nate, cristate-dentate, the teeth 5-6 on each side; corolla yellow, 
2-2.8 cm. long, horizontal in the calyx, the tube curved, puberulous, 
ampliate in the throat, the limb spreading, 1.6-2 cm. wide, the lobes 
denticulate-fimbriate; ovary pilose. 

Drymonia rubra Morton, sp. nov. Frutex terrestris, 1.2-1.8 m. 
altus, caulibus glabris; folia paullo inaequalia, petiolata (usque ad 
3.5 cm.); lamina foliorum elliptica, usque ad 13 cm. longa et 6 cm. 
lata, acuminata, basi cuneata, membranacea, integra, utrinque 
glabra; flores axillares, solitarii, longe pedicellati, pedicello glabro, 
usque ad 7 cm. longo, quam calyce longiore; calyx obliquus, viridis, 
3.5-4 cm. longus, lobis ovatis subaequalibus, longe et acriter acumina- 
tis, integris, utrinque glabris; corolla rubra, 4.5-6 cm. longa, in calyce 
obliqua, basi saccata, tubo sursum ampliato, non ventricoso, externe 
minute strigilloso, in fauce non contracto, limbo patente, ca. 4 cm. 
lato, subbilabiato, lobis crenato-dentatis; antherae oblongae, 5.5 
mm. longae; ovarium strigillosum. Yerba Buena, Prov. Heredia, 
alt. 2,000 meters, Standley & Valeria 49933 (type in U. S. Nat. Herb., 
No. 1,306,254). Same locality, Standley & Valeria 49188, 49245, 
49670, 49733. Cerro de Las Caricias, Prov. Heredia, alt. 2,000-2,400 
meters, Standley & Vakrio 51959, 51995, 52033. La Palma, Prov. 
San Jose", Pittier 682B; Standley 33109. La Carpintera, Prov. Car- 
tago, Standley 35511, 35630. Viento Fresco, Prov. Alajuela, alt. 
1,600-1,900 meters, Standley & Torres 47872. Fraijanes, Prov. 
Alajuela, Standley & Torres 47674- Los Angeles de San Ramon, Brenes 
3946. La Palma de San Ramon, Brenes 3737. San Rafael de San 
Ramon, Brenes 22018. Santo Domingo de Vara Blanca, M. Valerio 
1547. Zarcero, A. Smith A240, A684. 

Drymonia spectabilis (HBK.) Mart. Llanuras de Santa Clara. 
Changuinola Valley, Panama. Reported from Mexico to Colombia. 
An epiphytic vine, the stems puberulous toward the apex; leaf blades 
oblong, up to 10 cm. long and 4.5 cm. wide, short-acuminate, cuneate 
at the base, serrulate, sparingly strigillose on both sides; pedicels 



1176 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

solitary, 1-1.5 cm. long; calyx lobes ovate-lanceolate, up to 4 cm. 
long, entire or serrulate, acuminate, sparingly strigillose; corolla not 
much longer than the calyx, horizontal in the calyx, spurred at the 
base, minutely puberulous, ampliate toward the throat, the limb 
bilabiate, the anterior lobe erose-fimbriate. The identification of 
the Costa Rican plants with D. spectabilis of Colombia is doubtful. 

Drymonia turrialvae Hanst. Linnaea 34 : 359. 1865-66. Volcan 
de Turrialba, Wendland 517. Central mountains; Atlantic coast. 
Panama. Similar to D. marmorata, but the filaments not contorted ; 
leaves and calyces more densely strigillose. Perhaps not specifically 
different. 

Drymonia Warszewicziana Hanst. Linnaea 34: 352. 1865-66. 
Costa Rica, without special locality, Warscewicz 17. Low elevations, 
Atlantic and Pacific coasts; Guanacaste. Terrestrial shrubs, 1-2 
meters high, the stems sparingly red-pilosulous at the apex; petioles 
1-3.5 cm. long; leaf blades elliptic or ovate-oblong, 7.5-14 cm. long, 
3-5 cm. wide, short-acuminate, cuneate at the base, subentire, gla- 
brate above, very sparsely strigillose beneath; pedicels axillary, 
solitary or paired, 10-16 mm. long, shorter than the calyx; calyx 
lobes green, oblique, oblong-lanceolate or ovate-lanceolate, 2-2.8 cm. 
long, 6-10 mm. wide, long-acuminate, subentire, strigillose, unequal, 
the posterior shorter, saccate at the base; corolla red, pinkish, or 
brown and yellow, infundibuliform, 2.5-3 cm. long, horizontal in 
the calyx, saccate at the base, the tube curved, ampliate toward 
the throat, nearly glabrous, the limb spreading, crenulate, 16-20 
mm. wide; anthers oblong; ovary densely short- tomentose; style 
glabrous. 

EPISCIA Mart. 

Episcia lilacina Hanst. Linnaea 34: 342. 1865-66. Rio Sarapi- 
qui, near San Miguel, in forest, Wendland 782. E. acaulis Bonn. 
Smith, Bot. Gaz. 61: 378. 1916 (banks of Rio de Las Vueltas, Tucu- 
rrique, Prov. Cartago, 635 meters, Tonduz 13167). ?E. Fendleriana 
Kuntze. Low elevations, Atlantic and Pacific coasts. A prostrate, 
stoloniferous, subacaulescent herb, the stolons and short stems 
hirsute; leaves crowded, petiolate (0.8-1.5 cm.), the blades broadly 
elliptic, up to 5 cm. long and 3 cm. wide, obtuse, rounded at the base, 
crenate, bullate on the upper surface, hirsute, beneath pilose, purple; 
flowers solitary, axillary, the pedicels 12-16 mm. long, pilose; calyx 
lobes purple, oblong-spatulate, about 8 mm. long, obtuse, entire, 
slightly unequal, pilose on both sides; corolla about 4.5 cm. long, 



FLORA OF COSTA RICA 1177 

spurred at the base, horizontal in the calyx, sparsely pilosulous, 
the tube not ventricose, gradually enlarged to the throat, the limb 
about 3 cm. wide, bilabiate, the lobes orbicular, entire; filaments 
slender, glabrous; anthers oblong, 2 mm. long; disk gland emarginate; 
lamellae of the placentae ovuliferous on both surfaces. One or two 
additional species are found in Central America. The genus is here 
adopted as limited by Sprague (Kew Bull. 1912: 85-90). 

GESNERIA L. 

Gesneria tenera (Oerst.) Kuntze, Rev. Gen. 2: 473. 1891. 
Chorisanthera tenera Oerst. Centralamer. Gesner. 35. 1858. Described 
from Cerro de Aguacate, Oersted. A wholly dubious species. No 
specimens of Gesneria from Costa Rica have been seen. 

GLOXINIA L'He"r. 

Gloxinia perennis (L.) Fritsch. Cultivated for ornament. 
Native probably of Brazil. An herb with large, long-petiolate, 
cordate-based, glabrate leaves; flowers in an elongate, terminal, 
leafy-bracteate raceme; calyx lobes glabrous, large and broad; corolla 
pale bluish, open-campanulate, densely pilose, horizontal in the calyx, 
the limb regular; disk annular. 

HEPPIELLA Regel 

Heppiella ulmifolia (HBK.) Hanst. Reported doubtfully from 
Costa Rica by Fritsch (Bot. Jahrb. 50: 400. 1913). South America. 
Plants with the habit of Kohleria but with an annular disk; leaf 
blades bullate above, coarsely reticulate beneath; flowers clustered 
in the leaf axils; calyx tube short, the lobes narrow, free; corolla 
red, tubular, not ventricose, the limb narrow, regular; stamens 
not connate or connivent, the anthers oblong, the cells distinct; 
ovary half inferior. 

HYPOCYRTA Mart. 

Hypocyrta nummularia Hanst. Linnaea 34: 381. 1865-66. 
Costa Rica, without special locality, Wendland 1272. Central 
mountains at middle elevations; region of San Ramon. Guatemala. 
Creeping, epiphytic herbs, the stems reddish, red-hirsute; leaves 
clustered at the apices of short, lateral branches, the petiole up to 
2 cm. long, the blades obovate, up to 4 cm. long and 2 cm. wide, 
rounded, attenuate and decurrent at the base, thin-membranous, 
entire, sparsely pilosulous on both sides; flowers solitary, axillary, 
the pedicels slender, about 2 cm. long, pilosulous; calyx lobes green, 
linear-lanceolate, about 8 mm. long and 1.5 mm. broad at the base, 



1178 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

entire, membranous, pilosulous, equal, erect; corolla red, about 
2 cm. long, erect in the calyx, not spurred at the base, puberulous 
externally, the tube about 2 mm. wide at the base, near the apex 
abruptly and conspicuously ventricose, becoming 1 cm. wide, 
strongly contracted in the throat, the limb oblique, yellow, about 3 
mm. wide, the lobes small, erect; anthers connate, quadrate, the cells 
discrete, parallel; disk gland posterior. 

KLUGIA Schlecht. 

Klugia azurea Schlecht. Valley of Desengano. Mexico. An 
erect herb, 30 cm. high or more, the stems slender, little branched, 
puberulous; leaves alternate, short-petiolate (1-2 cm.), the blades 
ovate, up to 15 cm. long and 8 cm. wide, sharply acuminate, rounded 
and oblique at the base, membranous, entire, sparingly pilosulous 
above, glabrate beneath; inflorescence a terminal raceme, the com- 
mon peduncle short, the pedicels and bracts alternate, the bracts 
linear, the pedicels slender, 6-8 mm. long; calyx purple, tubular, 
about 1 cm. long, erect, the lobes connate most of their length, 
saccate at the base, glabrous; corolla blue, tubular, about 3 cm. long, 
erect in the calyx, not spurred or gibbous at the base, glabrous, the 
tube cylindric, not ventricose, the limb spreading, bilabiate, the lobes 
denticulate; anthers reniform, the cells divaricate, confluent at the 
apex; disk annular; placentae ovuliferous on both sides. 

KOELLIKERIA Regel 

Koellikeria argyrostigma (Hook.) Regel. Central mountains; 
Puntarenas; region of San Ramon. Panama to South America. A 
delicate herb, propagating by scaly offsets, the stems unbranched, 
pilose; leaves opposite, petiolate, clustered near the apex of the 
stem, obovate or broadly elliptic, up to 6.5 cm. long, obtuse or 
rounded, broadly cuneate at the base, membranous, crenate, pilose 
on both sides; inflorescence a terminal, elongate raceme, the pedicels 
and minute bracts alternate, the bracts spatulate, about 2 mm. 
long; calyx about 2.5 mm. long, the tube turbinate, the lobes free, 
lanceolate, pilosulous without, glabrous within; corolla white, purple 
in the throat, about 5 mm. long, erect in the calyx, not spurred, 
the limb bilabiate; filaments glabrous; anthers quadrate, the cells 
oblong, distinct; ovary half inferior, the free part pilose; style 
glabrous; stigma bilobed; disk annular, entire, glabrous. 

KOHLERIA Regel 

Herbs or shrubs, the stems and leaves usually hirsute; leaves 
opposite or ternate, usually subequal, thick, serrate or crenate; 



FLORA OF COSTA RICA 1179 

flowers solitary, fasciculate, or umbellate, sometimes falsely racemose; 
calyx lobes thick, erect, equal, pubescent on both surfaces; corolla 
scarlet, erect in the calyx, tubular, usually ventricose at the middle, 
conspicuously red-hirsute, the limb narrow, subregular; filaments 
inserted in the base of the corolla, not contorted; anthers free or con- 
nate, the cells not confluent, longitudinally dehiscent; ovary over 
two-thirds inferior, the lamellae of the placentae ovuliferous on both 
surfaces; style pilosulous, thick; stigma bilobed or stomatomorphic; 
disk of 5 glands, or these united into an entire, very thin annulus. 

Stigma stomatomorphic; disk glands united into an entire or sub- 
entire ring; calyx lobes linear-lanceolate or linear-subulate. 
Flowers umbellate on a common peduncle; filaments pilose at 
the base (Moussonia). 

Pubescence of stems, petioles, peduncles, and pedicels appressed. 

K. strigosa. 

Pubescence hirsute or hirsutulous K . elegans. 

Stigma bilobed; disk glands 5, free or the 2 posterior ones partially 

connate; calyx lobes deltoid-ovate or deltoid. 
Flowers umbellate, a common peduncle present; filaments pilosu- 
lous; 2 posterior disk glands connate (Brachyloma). 

K. maculata, 

Flowers solitary or fasciculate in the axils of leaves or bracts, a 
common peduncle absent; filaments glabrous; all disk glands 
free. 
Corolla limb narrower than the tube, the lobes minute, equal, 

erect (Cryptoloma) K. tubiflora. 

Corolla limb broader than the tube, the lobes spreading or partly 

reflexed, somewhat unequal (Eukohleria). 
Pubescence whitish lanate-tomentose; calyx lobes deltoid, 
shorter than the calyx tube, the margins plane. 

K. longifolia var. petiolaris. 

Pubescence reddish, hirsute; calyx lobes triangular-ovate, 
longer than the calyx tube, the margins thickened, 
recurved K . spicata. 

Kohleria elegans (Dene.) Loes. Moussonia costaricensis 
Klotzsch ex Oerst. Centralamer. Gesner. 33. 1858 (Veragua, Pan- 
ama, Warscewicz). Naranjo; Escasu. Mexico and Guatemala. 
A branched shrub, the stems, peduncles, and pedicels hirto-tomen- 
tose, the hairs spreading; leaf blades pilosulous beneath; flowers 



1180 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

umbellate on a short or elongate common peduncle; calyx lobes 
linear-subulate; corolla tube ampliate upwardly, not ventricose, not 
contracted in the throat, the limb subregular, patent, the lobes crenu- 
late; filaments pilose at the base; stigma stomatomorphic; disk glands 
connate into a ring. 

Kohleria longifolia (Lindl.) Hanst. var. petiolaris (Benth.) 
Morton, comb. nov. Gesneria petiolaris Benth. Bot. Voy. Sulph. 131. 
1844. Cocos Island. Originally described from "Island off the 
coast of Veragua," perhaps Coiba(?). An herb with unbranched, 
closely tomentose stems; leaves ternate, long-petiolate, elliptic, 
obtuse or merely acutish, densely and closely tomentose beneath; 
inflorescence elongate, the pedicels fasciculate in the axils of reduced 
floral leaves, whitish-tomentose; calyx tube and lobes appressed- 
whitish-tomentose, the lobes deltoid, about 2 mm. long; corolla erect, 
slightly ventricose, contracted in the throat, the limb spreading, a 
little irregular, glandular-pilose within; filaments glabrous; stigma 
bilobed; disk glands 5, free. Differs from the typical form of Guate- 
mala in its smaller, obtuse or acutish, rather than acuminate leaf 
blades, these more densely and closely tomentose beneath. 

Kohleria maculata Morton, sp. nov. Herba alta, caulibus non 
ramosis, sulcatis, longe hirsutis, pilis iridescentibus flaccidis multi- 
septatis eglanduliferis; folia opposita aequalia, breviter petiolata, 
petiolo crasso, usque ad 13 mm. longo, dense longeque hirsuto; 
laminae foliorum ellipticae, 9 cm. longae, 4 cm. latae, acuminatae, 
basi rotundatae, regulariter usque ad basin crenatae, crassae, supra 
hirsutae, subtus molliter pilosae, venis primariis 7-jugis; inflorescentia 
axillaris umbellata, usque ad 7 cm. longa, pedunculo communi usque 
ad 2.5 cm. longo, 1.5 mm. diam., dense hirsuto, apice bibracteato, 
bracteis lineari-lanceolatis, sessilibus, usque ad 15 mm. longis, pedi- 
cellis 4-6, usque ad 3 cm. longis, dense rubro-hirsutis; calycis tubus 
late turbinatus, 4 mm. longus, rubro-hirsutus, lobis triangulari- 
ovatis, 6.5 mm. longis, 3.5 mm. latis, acutis integris erectis aequalibus, 
utrinque hirsutis; corolla coccinea tubulosa, 3 cm. longa, in calyce 
erecta, externe dense rubro-hirsuta, tubo basi 5.5 mm. lato, medio 
ventricoso et 9 mm. lato, fauce contracta, 6 mm. lata, limbo angusto, 

1 cm. lato, lobis aequalibus brevibus rotundatis patulis integris, ca. 

2 mm. longis, intus maculatis glabris, in fauce glandulosis; filamenta 
basi corollae tubi affixa, recta, glanduloso-pilosula; antherae liberae 
quadratae, ca. 2 mm. longae et latae, loculis oblongis; ovarium (pars 
libera) conicum, longe hirsutum; stylus pilosulus; stigma bifidum; 



FLORA OF COSTA RICA 1181 

disci glandulae 5, ca. 1 mm. altae, 2 posticae connatae; placentae 
lamellae utrinque ovuliferae. San Jose", Sept. 4, 1936, Estrella Morana 
40 (type in Herb. Field Mus., No. 873,011). 

Kohleria spicata (HBK.) Oerst. K. Schiedeana (DC.) Hanst. 
K. Wageneri Regel. K. tetragona Oerst. Centralamer. Gesner. 27. 
1858 (Aguacaliente, Oersted). Brachyloma strictum Oerst. op. cit. 
29 (Cerro de Aguacate, Oersted). Cryptoloma strictum Hanst. Lin- 
naea 29: 538. 1857-58. Isoloma strictum Hemsl. Biol. Centr. Amer. 
Bot. 2 : 479. 1882. I. tetragonum Hemsl. op. cit. 480. Tropical Atlan- 
tic region; in central mountains, ranging up to 1,500 meters. 
Mexico to South America. An herb with erect, unbranched, reddish- 
hirsute stems up to 1.5 meters high; leaves opposite or ternate, 
elliptic or elliptic-lanceolate, obtuse or cuneate at the base, crenate- 
serrate, thick but flexible, hirsutulous above, tomentose-hirsute 
beneath, often with red veins; flowers solitary or clustered in the leaf 
axils, the inflorescence often racemose by reduction of the floral 
leaves to bracts, the pedicels up to 2.5 cm. long, hirsute; calyx tube 
hirsute, the lobes green, triangular-ovate, hirsute on both sides, the 
margins thickened and recurved; corolla scarlet, up to 2 cm. long, 
erect in the calyx, red-hirsute externally, the tube ventricose, a little 
deflexed, the throat contracted, the limb spreading, slightly bilabiate, 
the lower lobes reflexed, the upper erect, not spotted within; filaments 
glabrous; anthers about 2 mm. long; style pilosulous; stigma bilobed; 
disk of 5 distinct glands. 

Kohleria strigosa Morton, sp. nov. Frutex ramosus, usque 
ad 1 m. altus, caulibus teretibus, dense strigosis; folia opposita, 
subaequalia, petiolo usque ad 2 cm. longo, strigilloso-puberulo; 
laminae foliorum oblique ovatae, usque ad 12 cm. longae et 5 cm. 
latae, acuminatae, basi rotundatae, crassae, serrulatae, supra virides 
pilosulae, subtus puberulae, venis primariis 7-8-jugis; inflorescentia 
cymosa axillaris, 10-12 cm. longa, ca. 7-flora, pedunculo communi 
elongate, 6-8 cm. longo, strigoso, pedicellis usque ad 15 mm. longis; 
calycis tubus subcylindricus, ca. 3.5 mm. longus, 2.5 mm. latus, basi 
attenuatus, externe dense strigilloso-puberulus, lobis erectis, lineari- 
lanceolatis, ca. 4 mm. longis, 1.5 mm. basi latis, integris subaequali- 
bus, utrinque puberulis; corolla coccinea tubulosa, 20-25 mm. longa, 
in calyce erecta, basi paullo ampliata, externe rubro-pilosa, tubo 
ca. 3.5 mm. supra basin lato, non ventricoso, sursum paullo ampliato, 
fauce non contracta, 8-9 mm. lata, limbo ca. 15 mm. lato, lobis 
patulis aequalibus rotundatis apiculatis, intus glabris et eglanduli- 



1182 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

feris; filamenta basi corollae tubi affixa, basi pilosa; antherae liberae 
subquadratae, ca. 1.3 mm. longae, 1 mm. latae, loculis oblongis; 
ovarium (pars libera) conicum hirsutum; stylus pilosulus eglandu- 
losus; stigma stomatomorphum ; disci glandulae omnino in annulum 
integrum tenerum conjunctae; placentae lamellae utrinque ovuliferae. 
Tapesco de Zarcero, Alajuela, 2,000 meters, Austin Smith H428 
(type in Herb. Field Mus., No. 919,559). 

Kohleria tubiflora (Cav.) Hanst. Cryptoloma Hookerianum 
Hanst. ; Gesneria picta Hook.; ?C.rhynchocarpa Benth. Brachyloma 
pilosum Oerst. Centralamer. Gesner. 29. 1858 (Cerro de Aguacate, 
Oersted). Cryptoloma pilosum Hanst. Linnaea 29: 538. 1857-58. 
?C. cordifolium Hanst. Linnaea 29: 538. 1857-58 (without definite 
locality, Hoffmann). Isoloma cordifolium Hemsl. Biol. Centr. 
Amer. Bot. 2: 478. 1882. I. pilosum Hemsl. op. cit. 479. Poas; 
Aguacate; region of San Ramon. Panama. An herb with un- 
branched, hirsute stems up to 1 meter high; leaves opposite, 
short-petiolate, the blades obliquely elliptic, coarsely crenate- 
serrate, pilosulous on both sides; flowers axillary, solitary or rarely 
in 2's or 3's, the floral leaves usually not reduced, the pedicels up to 
1.5 cm. long, densely hirsute; calyx tube hemispheric, hirsute, the 
lobes equal, erect, triangular-ovate, hirsute on both sides, the margins 
not recurved; corolla scarlet, yellowish in the throat, 2-2.5 cm. long, 
erect in the calyx, red-hirsute, the tube ventricose upwardly, not 
deflexed, gradually contracted to the throat, the limb very narrow, 
the lobes small, equal, erect, about 1.5 mm. long, glabrous and 
eglandular within; filaments glabrous; anthers 2.5 mm. long, 2 mm. 
wide, the cells oblong; style pilosulous; stigma bilobed; disk glands 5, 
the 2 posterior ones approximate, free. 

MONOPYLE Moritz 

Perennial herbs with sparingly branched, reddish stems; leaves of 
a pair connected by a stipular line, very unequal, the smaller often 
stipuliform, serrate, thin-membranous; inflorescence paniculate, 
terminal, several-many-flowered, the flowers pedicellate; calyx lobes 
5, entire, equal; corolla white and purple, open-campanulate, not 
spurred at the base, minutely strigillose-pilosulous externally, the 
limb broad, with spreading, subequal lobes; stamens 4, with a rudi- 
mentary fifth, the filaments inserted on the base of the corolla tube, 
incurved upwardly, the anthers connate, 2-celled, the cells a little 
divergent; ovary almost wholly inferior, the lamellae of the placentae 
ovuliferous on both sides; style short and thick, glabrous; stigma 



FLORA OF COSTA RICA 1183 

stomatomorphic, pilose; disk none; fruit capsular, linear-oblong, 

opening by 2 longitudinal slits. The two species below described 

are the only ones known from North America. 

Calyx lobes oblong, red-purple, recurved at the tip; inflorescence 

(especially calyx tube) with numerous dark, reddish brown, 

uncinate, glandular hairs; petioles over 1 cm. long, bearing long, 

brown, flaccid, multiseptate hairs; lower leaf surface glabrous; 

corolla oblique in the calyx M. Maxonii. 

Calyx lobes linear-oblong, green, not recurved; inflorescence with 
hyaline, eglandular, uncinate hairs; petioles commonly not over 
5 mm. long, closely puberulous, lacking long, brown hairs; lower 
leaf surface minutely strigillose; corolla erect in the calyx. 

M. puberula. 

Monopyle Maxonii Morton, sp. nov. Herba perennis 30-100 
cm. alta, caulibus non vel parum ramosis, subflexuosis rubescentibus, 
parce pilosis, pilis longis brunneis multiseptatis, demum deciduis; 
folia opposita valde inaequalia, majora longe petiolata, petiolo 1-3.5 
cm. longo, parce brunneo-piloso; laminae foliorum majorum oblongae, 
usque ad 19 cm. longae et 6.5 cm. latae, acuminatae, basi obliquae 
et obtusae, tenuiter membranaceae, serratae, vix ciliatae, supra 
virides, parce pilosulae, subtus saepe rubescentes, venis sparse pilo- 
sulae, mesophyllo glabro, venis primariis 7-12-jugis; folia minora 
breviter petiolata, reducta, saepe 1-2 cm. longa; inflorescentia 
terminalis, laxe paniculata, 10-21 cm. longa, pedunculo communi 
5-9 cm. longo, rubro, minute glanduloso-pilosulo, pilis basi hyalinis, 
sursum glandulosis cylindricis gracilibus atrorubris apice uncinatis; 
bracteae saepe parvae, late lineares, raro foliosae et usque ad 3.5 cm. 
longae; inflorescentiae rami primarii 1-4-jugi, 3-35 mm. longi, apice 
bibracteolati, simplices vel pluries furcati, pedicellis 2-4, terminalibus 
subumbellatis, inaequalibus, 8-30 mm. longis; calycis tubus cylindri- 
cus, ad anthesin ca. 5 mm. longus et 2.2 mm. latus, basi attenuata 
gibbosus et curvatus, ubique pilis minutis rubris glanduliformibus 
uncinatis vestitus; calycis lobi rubropurpurei, oblongi, ca. 5 mm. 
longi, 2.5 mm. lati, integri eciliati, externe puberuli, intus glabri, 
apice recurvati; corolla basi alba, sursum violacea, apice brunneo- 
purpurea, (1.5) 2-3 cm. longa, late campanulata, in calyce obliqua, 
basi ecalcarata, non gibbosa, externe parce appresso-pilosula, tubo 
basi ca. 4 mm. lato, abrupte ampliato, in fauce usque ad 2 cm. lato, 
limbo 2-3 cm. lato, lobis patentibus rotundatis subaequalibus, ca. 
8 mm. longis, integris eciliatis, intus glabris et eglanduliferis; fila- 
menta basi corollae affixa, glabra, antheris connatis, loculis discretis, 



1184 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

subdivergentibus, rimis brevibus longitudinalibus dehiscentibus; 
ovarium fere omnino inferum, pars libera conica brevis pilosula; 
stylus crassus glaber; stigma stomatomorphum longe pilosum; discus 
nullus; fructus capsularis, rimis 2 longtitudinalibus dehiscens, 
elongato-cylindricus, 16-19 mm. longus, 2.5 mm. latus; calycis lobi 
persistentes, indurati; placentae lamellae utrinque ovuliferae. La 
Palma, Prov. San Jose", 1,500-1,700 meters, July 17-18, 1923, Maxon 
& Harvey 7947 (type in U. S. Nat. Herb., No. 1,181,611). Same 
locality, Tonduz 7457 (Herb. Nac. C. R. 12491, 12441}; Pittier 10179; 
M. Valeria 1426 p.p. Desamparados, Pittier 1158. Yerba Buena, 
Prov. Heredia, Standley & Valeria 49701, 49759. Los Angeles de 
San Ramon, Prov. Alajuela, Brenes 4916, 6106. It is likely that this 
is the plant described as M. macrocarpa var. costaricana Hemsl. 
(Biol. Centr. Amer. Bot. 2: 472. 1882; at 1,200 meters, Endres 82), 
but the brief description of Hemsley ("foliis minoribus concoloribus, 
calycis lobis brevioribus latioribus") does not adequately charac- 
terize it. The typical Monopyle macrocarpa Benth., a native of 
Amazonian Peru, is evidently a quite different plant with linear calyx 
lobes, eglandular inflorescence, and leaves soft-pubescent beneath. 
The present species has been identified as M. paniculata Benth., 
which, as represented by a specimen of the type collection in the 
New York Botanical Garden, differs in its biserrate leaves, strigillose 
on the mesophyll beneath, in its eglandular inflorescence, and in its 
linear calyx lobes, not recurved at the apex. 

Monopyle puberula Morton, sp. nov. Herba perennis, 60-80 
cm. alta, caulibus rubescentibus, ramosis, parce pilosulis, pilis 
brevibus, albis, uncinatis; folia opposita, valde inaequalia, majora 
breviter petiolata, petiolo saepe ca. 5 mm. longo, dense puberulo, 
pilis minutis, albis, uncinatis, numquam longis, brunneis, multi- 
septatis; laminae foliorum majorum oblongae, usque ad 13.5 cm. 
longae et 5.5 cm. latae, acuminatae, basi obtusae et obliquae, usque 
ad basin regulariter serratae, tenuiter membranaceae, supra virides, 
subtus saepe rubescentes, supra pilosulae, subtus in venis puberulae, 
in mesophyllo minute strigillosae; folia minora saepe stipuliformia, 
subsessilia; inflorescentia terminalis, paniculata, interdum foliosa, 
5-15 cm. longa, pedunculo communi 2-4.5 cm. longo, minute pilosulo, 
eglandulifero, bracteis saepe parvis, raro foliosis; rami primarii 
inflorescentiae 1-3-jugi, ca. 6 mm. longi, numquam furcati, apice 
bibracteolati, pedicellis 2 vel 3, subumbellatis inaequalibus, 6-15 mm. 
longis; calycis tubus cylindricus, ad anthesin 4.5 mm. longus, 2 mm. 
latus, basi attenuatus, vix curvatus vel gibbosus, externe dense 



FLORA OF COSTA RICA 1185 

pilosulus, pilis hyalinis eglandulosis uncinatis; calycis lobi lineari- 
oblongi, 6.5 mm. longi, 2 mm. lati, integri virides acuminati eciliati, 
externe breviter pilosuli, intus glabri, apice non recurvati; corolla 
basi alba, sursum violacea, late campanulata, 2-2.8 cm. longa, in 
calyce erecta, externe appresso-pilosula, tubo basi angusto, abrupte 
ampliato, in fauce usque ad 2 cm. lato, limbo ca. 2.8 cm. lato, lobis 
magnis subaequalibus rotundatis patentibus integris eciliatis, intus 
glabris eglanduliferis; filamenta basi corollae affixa, glabra, antheris 
connatis, ca. 1 mm. longis et 1.5 mm. latis, loculis suborbicularibus, 
rimis longitudinalibus dehiscentibus; ovarium fere omnino inferum, 
pars libera brevis conica pilosula; stylus crassus glaber; stigma 
stomatomorphum, pilosum; discus nullus. La Palma, Prov. San 
Jose", 1,500-1,700 meters, July 17-18, 1923, Maxon & Harvey 7990 
(type in U. S. Nat. Herb., No. 1,181,652). Same locality, M. Valeria 
1426 p.p. La Hondura, Prov. San Jose", Standley 36161, 37591, 
37765; M. Valeria 699. Cerro de Las Caricias, Prov. Heredia, 
Standley & Valeria 51898. Carillo, Pittier 1203. La Balsa de San 
Ramon, Prov. Alajuela, Brenes 4419. El Muneco, Prov. Cartago, 
Stork 2683. El Silencio, J. Valeria 136. Without locality, Stork 2836. 
Chicoy, Alta Verapaz, Guatemala, Tuerckheim 7929. Quebrada 
Seca, Alta Verapaz, Johnson 268. 

NAPEANTHUS Gardn. 

Napeanthus apodemus Donn. Smith, Bot. Gaz. 20: 6. 1895. 
Shaded banks of Rio Jimenez, Llanuras de Santa Clara, 200 meters, 
J. D. Smith 5109. Low elevations, provinces of Limon and Guana- 
caste. Low, acaulescent herbs; leaves rosulate, sessile, the blades 
oblong-spatulate, up to 10 cm. long and 3 cm. wide, rounded or 
obtuse, narrowed toward the base, membranous, entire, glabrous; 
inflorescence racemose-paniculate, the common peduncle up to 4 cm. 
long, slender, terete, glabrate, the branching dichotomous, the bracts 
opposite, linear; pedicels 8-16 mm. long; calyx lobes lanceolate, 
about 5 mm. long, acuminate, entire, erect, equal, nerved, puberulous, 
nearly free; corolla white, about as long as the calyx, rotate, the tube 
very short; stamens 4, the fifth one rudimentary, the filaments not 
connate, glabrous, about 1.5 mm. long, the anthers ovate, about 1.5 
mm. long, glabrous, the cells discrete, diverging; disk none; fruit 
capsular, 2-valved. No other species occur in Central America. 

PHINAEA Benth. 

Phinaea lacerata Morton. Region of San Ramon. Panama. 
A delicate herb 9-20 cm. high, reproducing by scaly offsets, the 



1186 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

stems red, unbranched, substrigose; leaves clustered near the apex 
of the stem, opposite, petiolate, the blades elliptic, up to 8 cm. long 
and 4 cm. wide, coarsely biserrate or incised, strigillose beneath; 
common peduncle none, the pedicels solitary, axillary, filiform, 
puberulous; calyx about 3.5 mm. long, the tube very short, the lobes 
lanceolate, free, acuminate, entire, pilosulous on both sides; corolla 
white, unspotted, rotate, 11-15 mm. wide, actinomorphic or nearly 
so, the tube 2 mm. long, the lobes rounded; filaments slender, gla- 
brous, the anterior curved; anthers much shorter than the filaments, 
the cells distinct, dehiscent by a short slit; disk none; ovary half 
inferior, the free part glabrous, globose; style glabrous, curved. 

SOLENOPHORA Benth. 

Solenophora calycosa Bonn. Smith, Bot. Gaz. 25: 152. 1898. 
Slopes of Volcan de Barba, 2,300 meters, Pittier 283. Common in 
the central mountains, 1,800-2,500 meters. Endemic. A shrub, up 
to 4.5 meters high, branched at the base; leaves opposite, long- 
petiolate, the blades ovate, up to 22 cm. long and 12 cm. wide, 
rounded and oblique at the base, doubly serrate, membranous, pilosu- 
lous above, scabrous beneath on the veins; inflorescence axillary, 
one-several-flowered, the common peduncle short, bibracteolate at 
the apex; calyx cylindric, 3 cm. long or more, the tube extended far 
beyond the ovary, hirtous-pilose on both sides, the lobes deltoid, 
not over 5 mm. long, glandular-serrate; corolla tubular, 7-9 cm. 
long, scarlet-orange outside, yellow within, red-spotted on the lobes, 
erect in the calyx, not spurred, pilose externally, glabrous within; 
anther cells slightly divaricate, not confluent, oblong, about 3.5 mm. 
long; ovary entirely inferior, the lamellae of the placentae ovuliferous 
on both surfaces; style pilosulous; stigma stomatomorphic; disk 
glands thick, 2, posterior, connate, hirsutulous, about 3.5 mm. 
long. This is probably the plant reported from Desengano as S. in- 
signis (Mart. & Gal.) Hanst. by Hemsley. 

Solenophora sp. A specimen collected at La Hondura by C. 
Werckle* (Herb. Nac. Cost. 11570} was reported by Donnell Smith 
as S. coccinea Benth. It doubtless represents an undescribed species, 
differing from S. calycosa in its smaller corollas, smaller, nearly 
glabrous calyces, and in having the leaf blades entirely glabrous 
beneath. The material is inadequate for a proper diagnosis. 

TUSSACIA Reichenb. 

Tussacia Friedrichsthaliana Hanst. Puntarenas. Changui- 
nola Valley, Panama. Guatemala. An erect, terrestrial herb, 30- 



FLORA OF COSTA RICA 1187 

40 cm. high, the stems puberulent; leaves of a pair equal, short- 
petiolate, the blades elliptic or broadly elliptic, up to 30 cm. long and 
13 cm. wide, acuminate, decurrent into the petiole at the base, mem- 
branous, dentate, pilosulous above, puberulent beneath, especially 
on the veins; flowers umbellate, the common peduncles solitary, 
axillary, up to 3 cm. long, puberulous, the pedicels numerous, erect, 
puberulous, about 2 cm. long; calyx open-campanulate, pentagonal, 
yellow, about 15 mm. long, 7-8 mm. wide, the lobes united for most 
of their length, puberulous; corolla yellow, tubular, about 2.5 cm. 
long, erect in the calyx, not spurred or gibbous at the base, the tube 
cylindric, glabrous, not ventricose, the limb spreading, subbilabiate; 
anthers free, the cells parallel, discrete, dehiscent throughout; disk 
gland posterior, bilobed. 

LENTIBULARIACEAE. Bladderwort Family 
Herbs, growing in water or on wet soil, rarely epiphytic; leaves 
various, often submersed and finely dissected, often bearing small 
bladders, sometimes aerial and entire; scapes 1-many-flowered, 
naked or bracteate; flowers perfect, very irregular; calyx 2-5-lobate, 
often almost to the base; corolla bilabiate, the tube saccate or 
calcarate; stamens 2, adnate to the base of the corolla tube, the 
filaments compressed, twisted, the anthers 1-celled; ovary 1-celled, 
with a free, central placenta, the ovules usually numerous; fruit 
capsular, bivalvate or irregularly dehiscent. One other genus, 
Pinguicula, is represented elsewhere in Central America. 

UTRICULARIA L. Bladderwort 

With the characters of the family, the scapes being bracteate and 
1-many-flowered, the calyx bilobate. In Pinguicula the scape is 
ebracteate, always 1-flowered, and the calyx is 5-lobate. Numerous 
other species are known from northern Central America, and one or 
two others may be expected in Costa Rica. 

Utricularia Endresii Reichenb. f. Gard. Chron. 582. 1874. 
Type collected by Endres, at an elevation of 600 meters (Endres 
material seen, in Herb. Field Mus.). Frequent in mountains of the 
central region, at 600-2,400 meters; growing among mosses on tree 
trunks. Endemic. Plants large for the genus, sometimes 35 cm. 
high or even taller; basal leaves long-petiolate, lanceolate to lance- 
elliptic or oblanceolate, the blades mostly 5-7 cm. long, subobtuse to 
attenuate, long-attenuate to the petiole; scapes slender, mostly 2-5- 
flowered, bearing a few inconspicuous, subulate bracts; flowers long- 
pedicellate, the bract at the base 3-parted; calyx lobes ovate, 2-3 cm. 



1188 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

long; corolla as much as 4.5 cm. long, bright purple. The bracts are 
purplish green. The roots bear small, white, translucent tubers. 
The plant often is taken to be an orchid, because of its epiphytic 
habit and its large and showy, very handsome, orchid-like flowers. 

Utricularia mixta Barnhart. Ochomogo, 1,500 meters. Cuba; 
Panama to northern South America. Plants floating on quiet water, 
mostly immersed, with elongate branches; leaves much dissected 
into threadlike segments, bearing numerous small bladders; scapes 
erect, usually 10-20 cm. tall, with 20 or fewer pedicellate flowers; 
bracts entire, not peltate, small; calyx 4-5 mm. long; corolla yellow, 
15-20 mm. broad. 

Utricularia obtusa Swartz. Collected in the regions of Cartago 
and San Ramon. Widely distributed in tropical America. Plants 
floating on quiet water and more or less submersed, the branches 
short or elongate; leaves dissected into capillary segments, bearing 
few minute bladders; scapes 1-6-flowered, very slender, usually less 
than 10 cm. tall; bracts entire, attached by the base; pedicels long 
and slender; calyx scarcely 2 mm. long; corolla yellow, 8-12 mm. long. 

Utricularia pusilla Vahl. Collected in the region of Turrialba 
and in Guanacaste, and doubtless widely distributed in the tierra 
caliente. Widely distributed in tropical America. Plants growing 
in wet soil, with short, rootlike branches at the base, but these soon 
disappearing; scapes filiform, wiry, mostly 2-10 cm. high; bracts 
minute, peltate, entire, present at the base of each pedicel and 1 also 
between each 2 pedicels; flowers slender-pedicellate; calyx scarcely 
2 mm. long; corolla yellow, 4-6 mm. long. 

Utricularia verapazensis Morong. Frequent in mountains of 
the central region, descending to the upper part of the Atlantic tierra 
caliente; region of San Ramon; at 900-2,400 meters, growing among 
mosses on tree trunks. Guatemala. Plants similar in appearance 
and structure to U. Endresii, but smaller in all parts; basal leaves 
linear-oblanceolate or oblanceolate, the blades mostly 1 cm. long or 
less, sometimes larger, long-petiolate, obtuse or acute; scapes usually 
5-12 cm. high, mostly 1-3-flowered; calyx about 1 cm. long; corolla 
about 1 cm. long, pale greenish purple. 

ACANTHACEAE. Acanthus Family 1 

By E. C. Leonard 

Herbs, shrubs, or small trees; leaves simple, opposite, exstipulate; 
flowers irregular to nearly regular, perfect; calyx persistent, inferior, 
1 Published by permission of the Secretary of the Smithsonian Institution. 



FLORA OF COSTA RICA 1189 

the segments 5 or occasionally fewer; corolla gamopetalous, the limb 
5-lobed or 2-lipped (rarely 1-lipped); stamens 4, didynamous, or 2 
only; staminodes often present in 2-stamened flowers; anther sacs 
2 or 1, longitudinally dehiscent; ovary 2-celled, the ovules 2-10 in 
each cavity; style filiform, simple; stigmas 1 or 2; fruit a capsule 
(drupaceous in Mendoncia), 2-celled, 2-valved; seeds usually flat, 
borne on retinacula (papilliform in a few genera, but usually hook- 
shaped), the testa smooth or roughened, often mucilaginous when 
moistened. 

Corollas 1-lipped Acanthus. 

Corollas 2-lipped or regular. 
Fruit drupaceous; vines Mendoncia. 

Fruit capsular; herbs, shrubs, small trees, or sometimes vines 

(Thunbergia). 
Seeds borne on papilliform retinacula. 

Peduncles with imbricate, coriaceous bracts Elytraria. 

Peduncles, if present, without imbricate, coriaceous bracts. 

Capsules strongly beaked, 2-4-seeded; seeds relatively 
large; plants often scandent Thunbergia. 

Capsules not beaked, many-seeded; seeds minute; diffuse, 
villous herbs Nelsonia. 

Seeds borne on hooklike retinacula. 

Calyx spathaceous, 2-lobed Spathacanthus. 

Calyx deeply 3-5-parted. 

Calyx segments 3 Louteridium. 

Calyx segments 4 or 5. 
A. Stamens 4. 

Anthers 1-celled Aphelandra. 

Anthers 2-celled or the anterior pair 1-celled. 

Anterior stamens with 1-celled anthers. 

Chamaeranthemum. 
Anterior stamens with 2-celled anthers. 

Calyx segments very dissimilar, the posterior and 
anterior ones much larger than the lateral, 
the anterior ones more or less united. 

Corolla 2-lipped, 5 mm. long, white or violet. 

Lepidagathis. 



1190 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Corolla subregular, 5 cm. long or more, yellow 
(bluish purple when dry) Barleria. 

Calyx segments similar or nearly so. 
Shrubs or small trees. 
Anther lobes calcarate at the base. .Bravaisia. 

Anther lobes rounded at the base. 

Trichanthera. 
Herbs (sometimes becoming suffrutescent). 

Corollas 2-lipped; flowers solitary or fascicled, 
axillary Hygrophila. 

Corollas regular or nearly so; inflorescence 

various. 

Flowers in dense spikes; bracts closely im- 
bricate Blechum. 

Flowers borne in loose, terminal or axillary 
cymes or panicles (rarely in terminal 
heads) ; bracts small, not imbricate. 

Anther sacs mucronate or aristulate at 
the base Dyschoriste. 

Anther sacs muticous at the base . Ruellia. 
A. Stamens 2. 

B. Anthers 1-celled. 
Stamens included. 

Calyx segments 4 Hansteinia. 

Calyx segments 5 Buceragenia. 

Stamens exserted. 

Corollas blue, the lower lip elongate, ligulate. 

Habracanthus. 

Corollas red or yellow, the lower lip scarcely 
distinct from the tube. 

Corolla tube gradually enlarged from base to 
mouth Razisea. 

Corolla tube abruptly enlarged above the ovary. 

Glockeria. 
B. Anthers 2-celled. 

Calyx segments widely dissimilar Barleria. 

Calyx segments similar or nearly so. 



FLORA OF COSTA RICA 1191 

Outer bracts large, conspicuously 4-ranked; flowers 
borne in dense spikes Tetramerium. 

Outer bracts usually small, if large, not conspicu- 
ously 4-ranked; flowers variously arranged. 

C. Staminodes present. 

Leaf blades mottled; cultivated plants. 

Graptophyllum. 
Leaf blades green; wild plants. 

Bracts imbricate, reticulate-nerved. 

Eranthemum. 

Bracts not imbricate, green. 
Corolla limb 2-lipped Odontonema. 

Corolla limb equally or subequally 5- 
lobed. 

Pollen marked with straight, longitu- 
dinal grooves ("Spangenpollen"). 

P sender anthemum. 

Pollen marked with replum-shaped 
grooves ("Rahmenpollen"). 

Chamaeranthemum. 
C. Staminodes none. 

Stems conspicuously 6-angled; calyx sub- 
tended by 2 partially united bracts; 
placenta separating from the mature 
capsules Dicliptera. 

Stems terete or subquadrangular; bracts dis- 
tinct; placenta remaining attached to the 
capsule walls at maturity. 

Anther cells parallel, equal, both inserted 
at the same height upon the filament. 

Carlowrightia. 

Anther cells unequal, inserted at different 
heights. 

Inflorescence headlike, the bracts white- 
hirsute, linear-subulate. 

Chaetochlamys. 

Inflorescence racemose, paniculate, or 
spicate, rarely headlike but the 



1192 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

bracts then broad and obtuse and 
not hirsute. 

Anther cells muticous at the base, or 
the lower cell occasionally acute 
or subcalcarate Jacobinia. 

Anther cells mucronate or calcarate 
at the base. 

Corolla short, the tube ampliate at 
the throat Justicia. 

Corolla elongate, not or scarcely 
ampliate. 

Pollen grains marked by longi- 
tudinal grooves ("Spangen- 
pollen") Streblacanthus. 

Pollen marked by knobs, pores, or 
plates. 

Pollen bearing small knobs 
("Knotchenpollen"). 

Beloperone. 

Pollen marked by pores and 
angular plates ("Facettier- 
ter pollen") . Poikilacanthus. 

ACANTHUS [Tourn.] L. 

Mostly large, herbaceous perennials with broad, pinnatifid leaves; 
flowers borne in spikes, sessile, solitary in the axils of the bracts; 
corollas white, rose, or purplish, 1-lipped, the lip 3-lobed; anthers 
1-celled, ciliate. A genus of about 20 species, found in the temperate 
and subtropical regions of the Old World, mostly in southern 
Europe. 

Acanthus mollis L. Cultivated in gardens for ornament. 
Native of the Old World. Leaves petioled, mostly radical, the blades 
cordate, up to 60 cm. long and 30 cm. wide, sinuately pinnatifid and 
toothed, sparingly pilose; flowers borne in large, terminal spikes; 
bracts ovate, about 3 cm. long and 1.5 cm. wide, acuminate, puberu- 
lent, each margin with 3 or 4 slender, spreading, spine-tipped teeth 
up to 5 mm. long; bractlets linear, 15-17 mm. long, 1.5 mm. wide, 
spine-tipped, puberulent; corolla rose-white, the lip 4 cm. long; 
filaments 15-18 mm. long, indurate, sigmoid, glabrous. 



FLORA OF COSTA RICA 1193 



APHELANDRA R. Br. 

Shrubs or large herbs; leaf blades usually large, oblong or elliptic, 
usually entire; flowers red, yellow, or white, borne in large, terminal, 
bracted spikes; calyx 5-parted, the segments narrow, subequal or 
the posterior one larger; corolla tube straight or incurved, sometimes 
ampliate upwardly, the limb usually bilabiate, the upper lip erect, 
entire or 2-lobed, the lower reflexed-spreading, 3-lobed; stamens 4; 
anthers 1-celled; ovules 2 in each cell. 

Bracts toothed. 

Bracts with a number of small, round, sessile, submarginal glands. 
Bracts 15 mm. long or less, their teeth 1 mm. long or more. 

A. Deppeana. 

Bracts 25 mm. long or more, their teeth about 0.25 mm. long. 

A. Storkii. 
Bracts without glands. 

Bracts hirsute or villous, the hairs up to 1 mm. long. 
Herbs. 
Bracts 5-7 mm. wide, bearing 2 or 3 pairs of teeth. 

A. Seibertii. 
Bracts 1.5 mm. wide, bearing a single pair of teeth. 

A. Tonduzii. 

Shrubs A. tridentata. 

Bracts puberulent. 

Leaf blades oblong to elliptic A. aurantiaca. 

Leaf blades narrowly lanceolate A. repanda. 

Bracts entire. 

Bracts essentially glabrous. 

Bracts obtuse A. micans. 

Bracts acuminate A. dolichantha. 

Bracts manifestly pubescent or puberulent. 
Bracts glandular-pilose, the surface without glands. 

A. gigantiflora. 

Bracts puberulent, with a number of small, round, sessile, sub- 
marginal glands. 

Bracts acute or obtusish A. tetragona. 

Bracts, at least the upper, obtuse or rounded. A. Sinclairiana. 

Aphelandra aurantiaca (Scheidw.) Lindl. Pavoncillo. A. acuti- 
folia Nees. Wet forests. Southern Mexico to northern South 



1194 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

America. Herbaceous or suffrutescent, up to 1 meter high; leaves 
oblong to elliptic, 8-20 cm. long, 2.5-12 cm. wide, acute or acuminate, 
narrowed at the base, entire, glabrous or nearly so; spikes solitary, 
terminal; bracts ovate-lanceolate, green or red, puberulent, up to 
3 cm. long and 1 cm. wide, acuminate, pectinate- toothed; bractlets 
and calyx segments lanceolate, puberulent and ciliolate; corolla red- 
orange or scarlet, puberulent, about 6 cm. long, the upper lip erect, 
acute, entire, the lower lip 3-lobed, the lateral lobes about one-half 
as long as the middle one; capsule 1.5 cm. long, puberulent. 

Aphelandra Deppeana Schlecht. & Cham. A. cristata HBK.; 
A. pectinata Willd.; A. Haenkeana Nees. Woods or open places. 
Southern Mexico to northern South America and West Indies. 
Shrubs 1-4 meters high; stems pubescent; leaf blades ovate-elliptic 
to lance-oblong, 5-20 cm. long, 2-10 cm. wide, acuminate, narrowed 
at the base and decurrent on the petioles, entire or undulate, scabrous 
above, sparingly or densely and softly pubescent beneath; spikes 
terminal, one or several; bracts lance-ovate, 8-15 mm. long, 4-6 mm. 
wide, acuminate, pilosulous, the upper portion bearing one to several 
slender teeth 1 mm. long or more, with a number of small, round, 
sessile, submarginal glands; flowers pale to dark red or crimson, 
pubescent, the upper lip erect, 2-lobed, the lower lip spreading, sub- 
entire or, if 3-lobed, the lateral lobes very small; capsules 15 mm. long, 
obtuse, glabrous, 4-seeded. 

Aphelandra dolichantha Bonn. Smith, Bot. Gaz. 27: 438. 
1899. Suerre, Llanuras de Santa Clara, 300 meters, J. D. Smith 
6689. Wet forests of Costa Rica, Panama, and Colombia. Suffrutes- 
cent, up to 1.2 meters high, the tips of the stems pubescent; leaf 
blades elliptic, up to 22 cm. long and 12 cm. wide, short-acuminate 
(the tip blunt), narrowed at the base and decurrent on the petioles, 
entire, sparingly pubescent, the hairs confined chiefly to the costa 
and lateral veins beneath; spikes terminal or lateral, solitary or several 
in a cluster, up to 12 cm. long, 1.5 cm. in diameter, the bracts closely 
imbricate but spreading with age, the rachis stout, winged, sparingly 
pubescent; bracts ovate-lanceolate, 2.5-4 cm. long, 1-1.3 cm. wide, 
acuminate, nerved, glabrous or the margins puberulous; bractlets 
subulate, about 6 mm. long; calyx segments subulate, 2-3 mm. long; 
corolla white, the tube 5-6 cm. long, 1 mm. in diameter at the base, 
about 3 mm. at the throat, glandular-pubescent without, the upper 
lip oblong, 13 mm. long, 2-lobed at the tip, the lower lip 3-lobed, 
the middle lobe oblong-elliptic, 18-20 mm. long, 6-7 mm. wide, the 



FLORA OF COSTA RICA 1195 

lateral lobes linear, 15-17 mm. long, 4-5 mm. wide; capsules slender- 
clavate, about 15 mm. long, 3 mm. broad, minutely punctate; seeds 
flat, light brown. 

Aphelandra gigantiflora Lindau. A. Padillana Standl. Moist 
woods, often on banks of streams. Guatemala, Costa Rica, and 
Salvador. Shrubs, up to 2.5 meters high, the stems glabrous or 
puberulent; leaf blades oblong-ovate, 15-25 cm. long, 4-9 cm. wide, 
more or less abruptly acuminate, narrowed or rounded at the base 
and decurrent on the petiole, thin, bright green above, paler and 
with a few scattered, short hairs beneath; spikes stout, terminal and 
axillary, 12-30 cm. long, the rachis densely glandular-pilose, the upper 
bracts closely imbricate, the lower loosely so, spreading, the lower- 
most sterile, all broadly ovate, 2-3 cm. long, obtuse or rounded at 
the apex or the upper acutish, entire, green or purplish, densely 
glandular-pilose; bractlets linear-subulate, equaling the calyx, 
glandular-pilose; calyx segments linear-lanceolate, 15 mm. long, 
long-attenuate, delicately striate-nerved, finely glandular-pilose; 
corolla bright red, 6.5-7.5 cm. long, finely glandular-pilose, the tube 
4-5 cm. long, 7-9 mm. in diameter at the throat, the posterior lip 
oblong, 2.5-3 cm. long, 10 mm. wide at the base, obtuse or rounded, 
the lower lip 3-parted, the middle segment obovate, 2.5-4 cm. long, 
15-17 mm. wide, the lateral segments oblong, 15 mm. long, 5 mm. 
wide, acute, sometimes unequally 2-lobed at the apex; capsules 2 cm. 
long, oblong-elliptic, acutish, finely puberulent. 

Aphelandra micans Moritz. Guatemala, Costa Rica, Panama, 
and Venezuela. Shrubs, up to 2 meters high, the stems strigose; 
leaf blades oblong-ovate to oblong-elliptic, up to 35 cm. long and 15 
cm. wide, acuminate to obtuse, narrowed at the base, thin, entire, 
sparingly appressed-pilosulous; petioles up to 4 cm. long; spikes 
terminal, solitary or several in a cluster, rigid, up to 25 cm. long, 1-2 
cm. in diameter, the rachis glabrous; bracts closely imbricate, about 
12 mm. long and 10 mm. wide, firm, ovate, obtuse, glabrous, or 
minutely ciliolate toward the tip, rather strongly nerved, bearing 
dorsally a pair of oval, glandular-alveolate spots about 3 mm. long 
and 1.5 mm. wide; bractlets linear, 12 mm. long and 2 mm. wide, 
carinate, striate, the keel tomentose; calyx segments oblong, 17-18 
mm. long, 3-5 mm. wide, coriaceous, glabrous, striate; corolla orange- 
yellow, about 5 cm. long, the posterior lip erect, about 15 mm. long, 
arching and acuminate at the tip, the lower lip 3-lobed, spreading, 
the middle lobe ovate, 2 cm. long and about 9 mm. wide, acuminate, 



1196 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the lateral lobes about 5 mm. long, rounded, their upper edge adnate 
to the upper lip; capsule about 2 cm. long, glabrous, minutely punc- 
tate; seeds flat, brown, about 5 mm. long and 4 mm. wide, glabrous. 

Aphelandra repanda Nees. A. aurantiaca var. stenophylla 
Standl. Wet forests. Central America to Peru. Suffrutescent, 
about 1 meter high; leaf blades narrowly lanceolate, 20-25 cm. long, 
2.5-3 cm. wide, gradually narrowed at both ends, glabrous, shining, 
coarsely sinuate-repand; petioles 1-3 cm. long; spikes terminal, 
sessile; bracts oblong, 2.5 cm. long, acuminate, puberulent, pectinate- 
dentate; corolla 2.5-3 cm. long, orange-scarlet, the upper lip erect, 
entire, the lower lip 3-lobed, the lobes subequal, the middle one oval, 
the lateral ones oblong-lanceolate. 

Aphelandra Seibertii Leonard. Found at La Colombiana, 
Prov. Limon, and in Talamanca. The type was collected in the 
vicinity of El Valle de Anton, Panama, Seibert 460. Herbs, up to 20 
cm. high, ascending, rooting at the lower nodes, pilose, the hairs up 
to 1 mm. long; leaf blades oblong-elliptic, 2-9 cm. long, 1.5-5 cm. 
wide, obtuse, narrowed at the base, entire or undulate, both surfaces 
thinly pilose, the costa and lateral veins densely so; petioles 1-1.5 cm. 
long, densely pilose; flowers borne in 1 or several terminal spikes 4-8 
cm. long, the peduncles 3-5 cm. long, densely yellowish-pilose, the 
rachis pilose, the bracts obovate or oblanceolate, 13-15 mm. long, 
5-7 mm. wide, purplish at the tip, closely imbricate, appressed 
(spreading at maturity), hirsute, the hairs up to 1 mm. long, with 
2 or 3 narrow, erect-spreading teeth on each side above the middle; 
bractlets lanceolate, 8 mm. long, 1.5 mm. wide, acuminate, carinate, 
conduplicate, subhyaline, delicately nerved, sparingly ciliolate; calyx 
segments subhyaline, striate-nerved, the posterior segment lanceolate, 
6.5 mm. long, 1.5 mm. wide, the anterior pair lanceolate, 6.6 mm. long 
and 1 mm. wide, the middle pair subulate, 5.5 mm. long and 0.5 mm. 
wide, all sparingly ciliolate with minute, capitate hairs; corolla 2 cm. 
long, yellow, the lobes tipped with lavender, finely and sparingly 
pubescent, the tube slender, 3 mm. in diameter at the mouth, the 
limb about 13 mm. broad, the lobes rounded or emarginate; stamens 
slightly exserted; capsule about 1 cm. long, cylindric, glabrous except 
the pilose tip, 4-seeded; seeds obovoid, acute at the base, light 
brown, 3 mm. long and 2.5 mm. broad; with scattered, minute, short, 
thick hairs. 

Aphelandra Sinclairiana Nees. Low, damp woods. Costa 
Rica and Panama. Shrubs or small trees, up to 5 meters high; stems 



FLORA OF COSTA RICA 1197 

hirsute; leaf blades oblong-lanceolate, up to 35 cm. long and 10 cm. 
wide, acuminate, narrowed at the base, sparingly hirsute or the costa 
and lateral veins densely so; petiole 1-2 cm. long; spikes terminal, 
solitary or several in a cluster, up to 20 cm. long, the rachis hirsutu- 
lous; bracts orange-red, oval, about 2 cm. long and 1 cm. wide, silky- 
puberulent, bearing several well defined glands on either side, the 
upper bracts rounded or obtuse, closely imbricate, the lower acute 
and spreading; bractlets lance-ovate, 6 mm. long, 2 mm. wide, acute, 
sometimes bearing one or more minute teeth on either side, puberu- 
lent, striate; calyx segments lanceolate, 9-10 mm. long, acute (tipped 
by a slender awn 0.5 mm. long), a few hairs near the tip, striate; 
corolla rose-red or purplish red, finely pubescent, about 6 cm. long, 
the tube 3 mm. in diameter at the base, 6-7 mm. at the mouth, the 
upper lip erect, 15 mm. long and 8 mm. wide, 2-lobed, the lobes lance- 
ovate, acute, about 6 mm. long, the lower lip spreading, the middle 
lobe 21 mm. long, 7-8 mm. wide, acuminate, carinate, the lateral 
lobes attached by one edge to the upper lip, the free portion triangular- 
ovate, 2.5 mm. long and 2 mm. wide, obtuse; capsules 2 cm. long, 
clavate, puberulent, obtuse at the apex, 6 mm. broad near the tip, 
3 mm. broad at the base; seeds flat, about 3.5 mm. in diameter, 
minutely hispidulous. 

Aphelandra Storkii Leonard, sp. nov. Suffrutex, caulibus 
pilosis vel glabris; lamina foliorum ovata, apice breviter acuminata, 
basi angustata, in petiolum decurrens, pilosa, undulata; spica termi- 
nalis, rhache dense pilosa; bracteae imbricatae, ovato-oblongae, 
acutae, supra puberulentae, subtus pilosae, medio glandulas parvas 
rotundas sessiles submarginales gerentes, medio minute denticulatae; 
bracteolae lineari-lanceolatae, pilosae, striatae; calycis segmenta 
anguste lanceolata, minute pilosa, striata; corolla pilosa, bilabiata, 
labio superiore acuto, apice bidentato, inferiore trilobo, lobo mediali 
acuto, lateralibus minoribus obtusis. 

Suffrutescent; stems terete, more or less pilose toward the tip, 
glabrous below; leaf blades ovate, 15-20 cm. long, 10-11 cm. wide, 
short-acuminate, abruptly narrowed at the base and decurrent on 
the petiole, undulate, sparingly pilose above, densely so beneath, the 
veins prominent, coarsely reticulate toward the margin; petioles 
about 10 cm. long, winged; spikes terminal, 11 cm. long or more, 
about 2 cm. in diameter, the bracts closely imbricate, the rachis 
densely spreading-pilose; bracts oblong-ovate, 25-28 mm. long, 10-12 
mm. wide, acute, red(?), veiny, puberulent above, pilose beneath, 
bearing at the middle on the margins one to several minute, ascending 



1198 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

teeth about 0.25 mm. long, with a number of small, round, sessile, 
submarginal glands near the middle; lateral bractlets linear-lanceo- 
late, 12 mm. long, 2.5 mm. wide near the base, narrowed to a slender 
point, pilose, striate; calyx segments subequal, 13-14 mm. long, 
narrowly lanceolate, the posterior segment 3.5 mm. wide, the anterior 
pair 2.5 mm. wide, and the lateral pair 2 mm. wide, all gradually 
narrowed to a slender point, minutely pilose, striate; corollas red(?), 
pilose, the upper lip erect, acute, 2-toothed at the apex, the lower lip 
3-lobed, probably spreading, its middle lobe acute, much larger than 
the obtuse lateral ones (mature corolla not seen). Livingston, Rio 
Reventazon, July to August, 1920, Rowlee & Stork 690 (type in U. S. 
Nat. Herb., No. 1,081,476). Related to A. Deppeana, but easily 
separated by its much larger bracts with very small teeth. In A. 
Deppeana the bracts are rarely as much as 15 mm. long, whereas in 
this new species they are at least 25 mm. long. 

Aphelandra tetragona (Vahl) Nees. Wet forests. Costa Rica 
to northern South America, also Martinique. A shrub, the tips of 
the branches densely hirtellous; leaf blades elliptic to ovate or broadly 
lanceolate, 8-40 cm. long, 3-11 cm. wide, acuminate, narrowed at 
the base and decurrent on the petiole, thin, entire or undulate, both 
surfaces usually rather sparingly hirtellous; spikes terminal, solitary 
or several in a cluster, 2-20 cm. long, subquadrangular in cross 
section; bracts ovate to lanceolate, 8-10 mm. long, 3-5 mm. wide, 
acute to obtusish at the apex, puberulent, sometimes ciliolate, with 
a number of small, round, sessile, submarginal glands, the rachis 
tomentose or occasionally subglabrous; lateral bracts lanceolate, 
8 mm. long, 2 mm. wide; calyx segments lanceolate to ovate, 7-8 mm. 
long, the posterior segment about twice as wide as the others, all 
acute, striate, chartaceous, ciliolate toward the apex; corolla scarlet, 
puberulent, 4-7 cm. long, the upper lip erect, about 2 cm. long, 
divided from the middle into 2 curved, acuminate lobes, the lower lip 
ringent in fully expanded flowers, the middle lobe lanceolate, about 
22 mm. long, 7 mm. wide, acute, the lateral lobes about 5 mm. long, 
rounded or occasionally notched at the apex, connate to the upper 
lip for about two-thirds their length; capsules about 17 mm. long and 
6 mm. in diameter near the tip, narrowed to 3 mm. at the base, 
glabrous, minutely dotted; seeds flat, reddish brown, about 5 mm. 
long and 4 mm. wide, minutely roughened. 

Aphelandra Tonduzii Leonard, sp. nov. Herba, caulibus 
erectis vel adscendentibus, hirsutis vel strigosis; folia petiolata, 



FLORA OF COSTA RICA 1199 

laminis ovatis, ellipticis vel late lanceolatis, apice acutis, obtusis vel 
rotundatis, basi angustatis, integris, pilosis; spicae axillares, graciles, 
laxae, pedunculatae, pilosae; bracteae lanceolatae, venosae, hirsutae, 
utroque latere minute unidentatae; bracteolae subulatae, subhya- 
linae, striatae, apicem versus glanduloso-pubescentes; corolla pallide 
rosea, intus pilosa, lobis obovatis, rotundatis; stamina inclusa; 
capsulae obtusae, glabrae; semina obovata, plana, brunnea, sparse 
pubescentia. 

Herbs; stems up to 22 cm. high, simple, erect or ascending, 
hirsute or occasionally strigose; leaf blades ovate to elliptic, or 
broadly lanceolate, up to 18 cm. long and 5.5 cm. wide, acute to 
obtuse or rounded at the apex, narrowed at the base, entire, both 
surfaces pilose, the hairs about 1 mm. long; petioles up to 2 cm. long, 
pilose; flowers borne in slender, lax, peduncled, axillary spikes up to 
15 cm. long, the rachis pilose; peduncles up to 16 cm. long, pubescent 
or occasionally strigose; bracts lanceolate, about 7 mm. long and 1.5 
mm. wide, tapering to a slender point, veiny, hirsute (the hairs up to 
1 mm. long), usually with a pair of minute, slender, marginal teeth; 
bractlets subulate, about 5 mm. long and 1.5 mm. wide at the base, 
striate-nerved, subhyaline, finely pubescent toward the tip, some of 
the longer hairs gland- tipped ; calyx segments 4.5 mm. long, the 
posterior one lanceolate, 1 mm. wide, the other 4 subulate, 0.5 mm. 
wide, all striate-nerved, subhyaline and pubescent toward the tip 
with some of the longer hairs glandular; corolla 1.5 cm. long, pale 
pink, pubescent within above the insertion of the stamens, the tube 
slender, glabrous, 2 mm. broad at the base and the mouth, constricted 
below the middle to 1.5 mm., the limb 12 mm. broad, the lobes 
obovate, rounded, the 2 upper ones about 3 mm. long and 2 mm. 
wide, the middle lobe of the lower lip 5 mm. long and 3.5 mm. wide, 
the lateral ones slightly smaller, all sparingly and minutely pubescent 
without; stamens included; capsules 10 mm. long and 3 mm. wide, 
blunt at the apex, glabrous; seeds obovate, flat, brown, about 2.5 mm. 
long, sparingly pubescent with small, short, thick hairs. Forests 
of Las Vueltas, Tucurrique, 900-1,000 meters, January, 1899, Tonduz 
13162 (type in U. S. Nat. Herb., No. 938,550). El Silencio de San 
Ramon, 950 meters, Brenes 3693. Edge of forest in the vicinity of 
El General, Prov. San Jose", 1,070 meters, Skutch 3004- Moist forests 
of Cerro de La Carpintera, Prov. Cartago, 1,500-1,850 meters, 
Standley 34205, 34495. Deep woods of La Carpintera, Stork 1082. 
Dissimilar to all other species of Aphelandra hitherto described from 
Central America, because of its slender spikes and extremely narrow, 



1200 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

pubescent bracts. Its closest relative is, probably, A. Seibertii 
Leonard. 

Aphelandra tridentata Hemsl. Biol. Centr. Amer. Bot. 2: 513. 
1882. Costa Rica, without definite locality, Endres 236. Specimens 
examined from San Ramon and the vicinity of Tilaran, Guanacaste. 
Found in wet or damp forests. Shrubs, up to 2 meters high; young 
stems densely villous; leaves oblong-ovate to elliptic, up to 30 cm. 
long and 8 cm. wide, acuminate, narrowed at the base, decurrent on 
the petiole, sparingly hirsutulous or the costa and lateral nerves 
densely so; spikes solitary and terminal, up to 10 cm. long and 3 cm. 
in diameter; bracts oblong-lanceolate, 3-4 cm. long, 8-10 mm. wide 
(the uppermost often much smaller), acuminate, villous, often with 
1 to several subulate teeth on each side; bractlets linear, 10-13 mm. 
long, about 2 mm. wide at the base, densely and softly hirsute; calyx 
segments as long as the bractlets, 2-3 mm. wide at the base, gradually 
narrowed to a slender tip, striate-nerved, appressed-pilose; corolla 5-6 
cm. long, bright red, villous, the upper lip oblong, entire, about 15 
mm. long and 7 mm. wide, the lower lip elliptic, slightly longer than 
the upper, 3-toothed at the apex, the teeth deltoid, about 0.5 mm. 
long; stamens scarcely exserted; capsules oblong, 16 mm. long, about 
6 mm. broad, blunt at the apex, glabrous, minutely punctate, shining; 
seeds dark brown, flat, about 4 mm. long and 3 mm. wide, minutely 
pilosulous. 

BARLERIA L. 

Shrubs or herbs; leaves often with axillary spines, the blades 
entire; flowers often yellow; corolla with a tube about as long as the 
spreading limb, sometimes longer, the lobes 5, rounded; stamens 4, 
didynamous, 2 perfect and 2 imperfect (staminodes), the anther sacs 
blunt at the base; capsules oblong or club-shaped; seeds compressed, 
ovate or orbicular. Several species other than the one described 
here grow in South America and the West Indies. 

Barleria micans Nees. Barleriopsis glandulosa Oerst. Vid. 
Medd. Kjoebenhavn 1854: 134. 1855 (Cartago, Oersted). Found in 
forests and thickets from Mexico to Colombia. Herbaceous or 
suffruticose, up to 1.5 meters high; leaves ovate to narrowly lanceo- 
late, up to 25 cm. long and 8 cm. wide, long-acuminate, attenuate at 
the base and decurrent on the petiole, sparingly strigose or hirtellous; 
flowers borne in dense, sessile, terminal spikes 3-8 cm. long and about 
3 cm. in diameter; bracts lance-ovate to broadly ovate, 1-1.5 cm. 
long, 5-7 mm. wide, acute or obtuse, strigose and hirsute-ciliate, 



FLORA OF COSTA RICA 1201 

minutely toothed, bluish when dry; lateral bractlets linear, 1.5-2 cm. 
long; calyx segments very unequal, the anterior segment oblong, 
1.5-2 cm. long, 1-1.4 cm. wide, bidentate at the apex, the posterior 
segment slightly longer and narrowed, acuminate at the apex, both 
bordered by spine-tipped teeth up to 4 mm. long, the lateral segments 
narrowly lanceolate and about one-half as long as others; corolla 
5-5.5 cm. long, turning bluish purple when dry, the tube slender, the 
lobes oblanceolate, subequal, spreading; capsules 4-seeded. 

BELOPERONE Nees 

Herbs or shrubs; leaves usually elliptic or ovate, entire; flowers 
fasciculate in the axils of the bracts, or cymose, or sometimes borne 
in spikes or in terminal thyrsi, the bracts small or large; calyx 5-parted, 
the segments narrow; corolla purple or white, the tube slender, 
elongate, scarcely ampliate above, the limb deeply 2-lipped, the 
upper lip narrow, erect, entire or minutely 2-lobed, the lower 3- 
lobed; stamens 2, the anther cells unequal, usually superposed, at 
least the lower one mucronate or calcarate at the base; staminodes 
none; capsules clavate, 4-seeded. A genus limited to tropical 
America. 

Flowers white; capsules glabrous B. urophylla. 

Flowers purple; capsules puberulent. 

Spikes short, dense; bracts lanceolate, about 1 cm. long, obtuse. 

B. Brenesii. 
Spikes slender, lax; bracts subulate, about 4 mm. long. . 

B. variegata. 

Beloperone Brenesii Leonard, sp. nov. Herbae erectae, cauli- 
bus quadrangularibus, angulis pilosis; lamina foliorum ovata, apice 
breviter acuminata, basi angustata, pilosa; petioli pilosi; bracteae 
lanceolatae, obtusae, pilosae, ciliatae; bracteolae anguste lanceo- 
latae, asymmetricae, ciliatae, pilosae; calycis segmenta lineari- 
subulata, pilosa; corolla roseo-purpurea, pubescens, bilabiata, labio 
superiore acuto, subapiculato, inferiore trilobate, lobis oblongis; 
antherae loculi superpositi, basi calcarati; capsulae minute pubes- 
centes, pilis retrorsis. 

Erect herbs, up to 40 cm. high or more; stems quadrangular, the 
angles densely brownish-pilose, the hairs up to 1 mm. long, or the 
basal portions sub terete and glabrescent; leaf blades ovate, up to 
5 cm. long and 3 cm. wide, short-acuminate (the tip blunt), narrowed 
at the base, both surfaces pilose; petioles up to 2 cm. long, pilose; 



1202 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

flowers borne in dense, terminal spikes or narrow thyrsi up to 10 cm. 
(usually 3-4 cm.) long; rachises and peduncles (1-2 cm. long) pilose; 
bracts lanceolate, about 1 cm. long and 2.5 mm. wide, obtuse, pilose 
and ciliate (sometimes sparingly so), the hairs up to 2 mm. long; 
bractlets narrowly lanceolate, asymmetric, 9 mm. long, 1.5 mm. 
wide, ciliate and pilose; calyx segments linear-subulate, 5 mm. long, 
0.75 mm. wide, sparingly pilose; corollas 2.5 cm. long, pubescent, 
rose-purple, the tube slender, 1.5 mm. in diameter at the base, 3-4 
mm. at the mouth, the lips about 7 mm. long, the upper one acute or 
subapiculate, the lobes of the lower lip oblong, about 3 mm. long 
and 1.5 mm. wide; anther lobes superposed, both calcarate at the 
base, the spur of the lower one 0.75 mm. long, that of the upper one 
about half as long; capsules clavate, 1 cm. long, puberal ent, the hairs 
retrorse. Los Angeles de San Ramon, 1,050 meters, November 20, 
1923, Brenes 3948 (type in Herb. Field Mus., No. 851,797). Catara- 
tas de San Ramon, Brenes 20518. Los Angeles to Piedades de San 
Ramon, Brenes 4767. Without definite locality, Brenes 13420. 
Santo Domingo de Vara Blanca, Manuel Valeria 1549, 1553. Well 
marked by the densely pilose angles of the stems and by the relatively 
broad, short, crowded spikes or thyrsi. 

Beloperone urophylla Lindau in Pittier, Prim. 2: 316. 1900. 
Forests of Boruca, 450 meters, Tonduz 4077. Forests of Costa Rica 
and Panama. Suffrutescent; stems terete, glabrous or finely pubes- 
cent in 2 lines; leaf blades ovate to oblong-ovate, up to 17 cm. long 
and 7 cm. wide, acuminate and subfalcate at the apex, the tip blunt, 
narrowed at the base, glabrous or the veins beneath sparingly pubes- 
cent, the cystoliths numerous and conspicuous; petioles 8-12 mm. 
long, pubescent above with curved hairs; flowers borne in 1 to several 
terminal or subterminal, dense spikes up to 8 cm. long and about 4 
mm. in diameter, imbedded in furrows of the rachis, the peduncles 
up to 3 cm. long, pubescent in 2 lines, the rachis glabrous or puberu- 
lent; bracts and bractlets linear-subulate, up to 5 mm. long, sparingly 
and minutely pubescent; calyx segments subulate, 4 mm. long, 
minutely and sparingly pubescent; corolla 3-3.5 cm. long, finely 
pubescent, white, the tube slender, 2 mm. in diameter, narrowed 
above the ovary, thence enlarged to 4 mm. at the mouth, the lips 
about 12 mm. long, the upper one minutely bidentate at the apex, 
the lobes of the lower lip about 7 mm. long, rounded; anther lobes 
superposed, the connective about 1 mm. long; capsules 13-15 mm. 
long, clavate, glabrous; seeds flat, brown, glabrous, about 3 mm. in 
diameter. 



FLORA OF COSTA RICA 1203 

Beloperone variegata Lindau in Pittier, Prim. 2: 315. 1900. 
Streblacanthus chirripensis Lindau, Bull. Herb. Boiss. II. 4: 404. 
1904 (near Hacienda de Chirripo, 200 meters, shaded places, Pittier 
16046). Forests of Cerro del Volcan, near Boruca, 1,200 meters, 
Tonduz 6735. Forests. Endemic. An herb up to 20 cm. high; 
stems ascending, rooting at the lower nodes, terete, densely brownish- 
pubescent, the hairs retrorsely curved; leaves few, the blades ovate 
to oblong, up to 13 cm. long and 7 cm. wide, obtuse or rounded at 
both ends, glabrous above, the costa and veins pubescent beneath, 
the cystoliths numerous and conspicuous; petioles 3-15 mm. long, 
pubescent; flowers borne in terminal or subterminal, slender, simple 
or forked spikes up to 12 cm. long, the rachis and peduncles (up to 
10 cm. long) densely and softly pubescent, the internodes 10-15 mm. 
long at the base of spike, successively shorter toward the tip; bracts 
narrow-subulate, about 4 mm. long, pubescent; bractlets similar but 
smaller; calyx segments 5-6 mm. long, about 0.5 mm. wide, pubes- 
cent; corolla 12 mm. long, light purple, pubescent, the tube slender, 
1 mm. in diameter at the base, 2 mm. at the mouth, the upper lip 
5-6 mm. long, 3 mm. wide, minutely bidentate at the apex, the lower 
lip 5 mm. long, the middle lobe about 2.5 mm. in diameter, the lateral 
lobes 2.5 mm. long and 2 mm. wide; anther lobes superposed, the 
lower calcarate; capsule clavate, 10-11 mm. long, puberulent; seeds 
flat, glabrous, 2.5 mm. in diameter. 

The description of Ecbolium Chamaeranthemum Kuntze (Rev. 
Gen. 2: 487. 1891; type from Limon, Kuntze) suggests this in many 
respects. Should the two species prove identical, the name variegata 
would be replaced by Chamaeranthemum. I do not feel that this 
transfer is justifiable at present, since I have been unable thus far 
to examine any of Kuntze's type material. 

BLECHUM P. Br. 

Perennial herbs; leaves petioled, repand-dentate, crenate or 
entire; flowers borne in dense, terminal spikes, the bracts imbricate; 
calyx 5-parted, the segments slightly unequal, linear-subulate; 
corollas whitish, the tube slender, the limb almost equally 5-lobed; 
stamens 4, didynamous, the anther sacs parallel ; ovules few in each 
cavity; capsules broadly oblong, with a short, narrowed base. 
Other species occur in Central America. 

Corolla 12-13 mm. long; bracts grayish-puberulent.B. pyramidatum. 
Corolla 20-25 mm. long; bracts sparingly puberulent and pilose. 

B. costaricense. 



1204 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Blechum costaricense Oerst. Vid. Medd. Kjoebenhavn 1854: 
168. 1855. Cerro de Aguacate, Oersted. Wooded regions of the 
Provincia de San Jose 1 . Herbaceous, or suffrutescent at base, stems 
pilose or the lower portions glabrescent; leaf blades ovate to oblong, 
up to 12 cm. long and 4 cm. wide, acuminate, the tip blunt, narrowed 
at the base and decurrent on the petiole, entire or shallowly sinuate- 
dentate, both surfaces pilose with stiff hairs up to 1.5 mm. long, or 
the costa and lateral veins of the lower surface minutely strigose, 
the cystoliths prominent; petioles slender, up to 3 cm. long; spikes 
ovoid, up to about 3 cm. long, 2 cm. in diameter, the rachis finely 
pubescent; bracts ovate, 1.5 cm. long and 1 cm. wide, acuminate at 
the apex, rounded at the base, entire, sinuate, or the lower ones 
shallowly dentate, sparingly pilose within, puberulent without, the 
hairs appressed and confined chiefly to the costa and veins, the 
margins ciliate with spreading, white hairs up to 2 mm. long; bract- 
lets lanceolate, 11 mm. long, 2 mm. wide, obtuse, conspicuously 
white-ciliate; calyx 5 mm. long, the segments 0.5 mm. wide at the 
base, ciliate, with a tuft of longer hairs at the tip; corolla pale 
lavender, 20-25 mm. long, finely pubescent, the lower portion of 
the tube slender, narrowly campanulate above, about 5 mm. broad 
at the throat, the limb 2 cm. broad when fully expanded, the lobes 
suborbicular, rounded or shallowly emarginate; capsules ovoid, about 
6 mm. long and 5 mm. broad, puberulent, 4-seeded. 

Blechum pyramidatum (Lam.) Urb. Sornia. Banks, fields, 
damp woods, thickets, and sometimes a weed in cultivated land. 
General in tropics of America and the Old World. Stems erect or 
ascending, 20-70 cm. high, the branches slender, more or less- 
puberulent; leaf blades ovate, 2-7 cm. long, 1-5 cm. wide, acute 
or obtuse, obtuse or narrowed at the base, glabrous or sparingly 
pilose, thin; spikes 4-sided, 3-6 cm. long; bracts ovate, pinnately 
veined, 1-2.5 cm. long, about 1 cm. wide, acutish at the apex, rounded 
at the base, gray-puberulent and pilosulous, ciliate; corolla 12-13 mm. 
long, pubescent; capsule broadly oblong, about 6 mm. long, puberu- 
lent. Much used in the vicinity of Tilaran as a remedy for amebic 
and other dysentery (Standley). 

BRAVAISIA DC. 

Shrubs or small trees; leaves petioled, entire; inflorescence a 
terminal, subcorymbose panicle, each flower subtended by a pair 
of bracts; calyx segments 5, subcoriaceous; corolla white or purplish, 
the tube rather abruptly expanded above the ovary, the lobes 5, 



FLORA OF COSTA RICA 1205 

subequal, rounded, spreading; stamens 4, the anther cells contiguous, 
spurred at the base; ovules 2-4 in each cell. Several species of this 
genus are found in Central America. 

Bravaisia integerrima (Spreng.) Standl. Palo de agua, Mangle 
bianco. Onychacanthus speciosus Oerst. Vid. Medd. Kjoebenhavn 
1854: 131. 1855 (near Puntarenas, Oersted). Found in wet, wooded 
regions. Ranging from the Mexican states Tamaulipas and Guerrero 
to Trinidad, Venezuela, Colombia, and Peru. A dense tree, up to 
18 meters high, the branches subquadrangular, glabrous or the young 
tips finely pubescent; petioles 3-5 cm. long, connate at the base; 
leaf blades oblong-elliptic, 8-28 cm. long, 3-10 cm. wide, acuminate 
or acute, narrowed at the base, subcoriaceous, glabrous, or the 
costa and lateral veins pilosulous below; panicle leafy, subcorymbose, 
about 12 cm. long and 10-30 cm. in diameter; bractlets subtending 
the flowers ovate, 2-3 mm. long, subcarinate, puberulent and cilio- 
late; calyx 4-5 mm. long, the segments ovate, 3-4 mm. long, 2-3 mm. 
wide, obtuse or rounded, puberulent and ciliolate; flowers white 
(occasionally yellow), with a purple spot at the throat, glabrous, 
the tube about 8 mm. broad at the throat, the limb nearly regular, 
about 2 cm. broad, the lobes emarginate; capsules oblong, 10-12 mm. 
long, obtuse at the apex, rounded at the base, 4-seeded. 

BUCERAGENIA Greenm. 

Herbs; leaves petiolate, usually ovate or oblong-elliptic; flowers 
borne in a leafy panicle, the branches lax, the flowers single or several 
in a cluster; bracts and bractlets minute; calyx segments 5, linear, 
subequal ; corolla tubular, the limb 2-lipped, the posterior lip incurved, 
emarginate or 2-lobed, the anterior lip erect or scarcely spreading, 
3-lobed; stamens 2, inserted at the middle of the corolla tube, in- 
cluded; anthers 1-celled, oblong, obtuse; staminodes 2; capsule 
.oblong, narrowed below to a slender stipe; seeds 4, flattened, sub- 
orbicular, roughened. The name is of Greek origin and refers to 
the Spanish name Cuernavaca, the Mexican locality of the type 
species. Several species occur in Mexico. 

Buceragenia glandulosa Leonard, sp. nov. Herba vel suf- 
frutex, caulibus glabris vel sparse bifariam pilosulis; folia petiolata 
vel suprema sessilia vel subsessilia, laminis ovatis ad oblongis, apice 
acuminatis, basi cuneatis vel rotundatis vel supremis subcordatis, 
in petiolum decurrentibus; spicae terminales et axillares, floribus 
solitariis vel fasciculatis; rachis glanduloso-pilosula; bracteae et 
bracteolae anguste triangulares, glabrae; calycis segmenta lanceolata, 



1206 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

subglabra; corolla minuta, tubo ovoideo, lobis orbicularibus; capsulae 
clavatae, glabrae; semina plana, marginibus asperis. 

Herbaceous or suffrutescent, up to 2 meters high; stems glabrous 
or sparingly pilosulous in 2 lines, with numerous subpunctiform 
cystoliths; leaves petioled or the uppermost sessile or subsessile, the 
blades ovate to oblong, up to 15 cm. long and 7 cm. wide, acuminate 
(the tip blunt), acute to obtuse or rounded at the base (the upper- 
most subcordate) and decurrent on the petiole, thin, glabrous, the 
costa red-purple, the cystoliths numerous, slender; petioles up to 
6 cm. long; flowers solitary or fascicled, borne in terminal and axillary 
spikes, these forming a large, terminal panicle, the internodes about 
2 cm. long at the base of the spike, thence successively shorter toward 
the tip, the uppermost fascicles confluent, the rachis glandular- 
pilosulous; pedicels up to 1 mm. long; bracts narrowly triangular, 
about 2 mm. long, glabrous; bractlets similar but smaller; calyx 
segments lanceolate, 3.5 mm. long, 1 mm. wide at the base, acumi- 
nate, essentially glabrous; corollas 2 mm. long, glabrous, the tube 
ovoid, the lobes orbicular, about 0.25 mm. in diameter; capsules 
about 14 mm. long, glabrous, the stipe slender; seeds flat, the margins 
roughened. Tres Rios, near San Jose", 1,400 meters, January 17, 
1916, Holway 440 (type in U. S. Nat. Herb., No. 862,612). Wet 
forest of Cerro de La Carpintera, Prov. Cartago, 1,500-1,850 meters, 
Standley 35503. Wet forests of El Mufieco, south of Navarro, Prov. 
Cartago, 1,400 meters, Standley 33686. Although this plant easily 
may be mistaken for P sender anihemum cuspidatum (Nees) Radlk., 
it can be recognized by its fascicled flowers, minute corollas, and 
glandular-pilosulous rachis. 

CARLOWRIGHTIA Gray 

Plants herbaceous or suffrutescent; leaves entire; flowers borne 
in loose spikes or racemes, these often forming large, terminal 
panicles; calyx segments 5; corolla tube slender, scarcely dilated, 
the limb almost equally 4-cleft; stamens 2; anther cells equal, inserted 
at the same height; staminodes none; capsule stipitate, 4-seeded. 
A genus of about 20 species, chiefly of Mexico. 

Carlowrightia costaricana Leonard, sp. nov. Suffrutex, cauli- 
bus teretibus puberulentis; lamina foliorum elliptico-ovata, obtusa 
(supremae mucronatae), basi obtusa vel rotundata, supra puberu- 
lenta vel subglabra; petioli tenues puberulenti; inflorescentia pani- 
culata, spicis numerosis laxis curvatis, rhachibus minute puberulentis; 
bracteae et bracteolae subulatae, puberulentae; calycis segmenta 



FLORA OF COSTA RICA 1207 

triangularia, griseo-puberulenta; corolla pubescens; capsulae glabrae; 
semina albida, minute rugosa. 

Erect, suffrutescent, about 60 cm. high; stems terete, puberulent; 
leaf blades elliptic-ovate, up to 4.5 cm. long and 2.5 cm. wide, acute 
(the uppermost mucronate), rounded or obtuse at the base, puberu- 
lent or subglabrous above; petioles slender, up to 1 cm. long, puberu- 
lent; flowers borne in numerous lax, curved spikes forming a terminal 
panicle up to 20 cm. long, the rachises minutely puberulent or sub- 
glabrous, the lower internodes about 8 mm. long, the others succes- 
sively shorter toward the tip of the spikes; bracts and bractlets 
subulate, up to 3 mm. long, puberulent; calyx segments triangular, 
1.5 mm. long, 0.5 mm. wide at the base, minutely grayish-puberulent; 
corolla about 7 mm. long, pubescent, purplish (?), the limb about 
9 mm. broad; capsules up to 11 mm. long and 4 mm. wide, pointed, 
glabrous; seeds whitish, minutely roughened. Dry forests in the 
vicinity of Libano, Prov. Guanacaste, 260-360 meters, January 15, 
1926, Standley & Valeria 44890 (type in U. S. Nat. Herb., No. 
1,253,830). In general appearance this species is closest to C. glabrata 
Fernald, of central Mexico, but differs in the nature of its pubescence 
and in its smaller leaf blades and larger capsules. The rachises of 
C. glabrata are pilose, instead of minutely puberulent. 

CHAETOCHLAMYS Lindau 

Flowers usually borne in congested, terminal and axillary spikes 
or heads; bracts and bractlets narrow; calyx segments 5, narrow; 
corolla tube but slightly enlarged at the throat, the limb 2-lipped, 
the upper lip 2-lobed, the lower 3-lobed; stamens 2; anther lobes 
affixed at different heights, the lower calcarate; staminodes none; 
stigma capitate; capsules 4-seeded. Several species are found in 
South America. 

Chaetochlamys panamensis Lindau. Forests and thickets of 
Costa Rica and Panama. Herbs, up to 1 meter high, erect or decum- 
bent; stems terete, pubescent in 2 lines (the hairs retrorsely curved) 
or glabrous below; leaf blades oblong-lanceolate, up to 10 cm. long 
and 3 cm. wide, firm, acuminate at both ends, glabrous or sparingly 
hirtellous beneath, the cystoliths prominent; petioles up to 10 mm. 
long, minutely strigose or glabrous; flowers borne in terminal and 
axillary heads; bracts linear, about 12 mm. long and 1 mm. wide, 
pilose, the hairs white, up to 1.5 mm. long; bractlets similar but 
narrower; calyx segments lanceolate, about 1 cm. long, 2 mm. wide, 
the costa and margins pilose; corolla pubescent, rose-purple, the 



1208 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

tube 15 mm. long, 3 mm. in diameter, the upper lip 8 mm. long and 
4 mm. wide, the lobes rounded, 2 mm. in diameter, the lower lip 
11 mm. long, the lateral lobes 6 mm. long and 7 mm. wide, the 
middle one 8 mm. long and 10 mm. wide; capsules 10-12 mm. long, 
about 4 mm. in diameter, narrowed and solid at the base, glabrous. 

CHAMAERANTHEMUM Nees 

Herbaceous or suffrutescent; inflorescence terminal, paniculate 
or spicate, the flowers and bracts small; calyx segments narrow, 
equal; corolla hypocrateriform or infundibular, the tube slender, 
the lobes of the limb subequal; stamens 4, included, the anthers 
of the posterior stamens 1-celled or sterile; capsules 4-seeded, reduced 
below to a slender, sterile stipe; seeds discoid. Two other species 
are found in Central America. Several grow in northern South 
America. 

Leaf blades hirsute C. Durandii. 

Leaf blades glabrous C. Tonduzii. 

Chamaeranthemum Durandii Leonard, sp. nov. Suffrutex, 
caulibus subquadrangularibus, hirsutis; lamina foliorum oblonga, 
acuminata, basi angustata, hirsuta; petioli breves, hirsuti; spicae 
terminales, laxae, rhache hirsuta; bracteae et bracteolae subulatae, 
puberulentae; calycis segmenta subulata, glanduloso-puberulenta; 
corolla puberulenta, lobis rotundatis; stamina postica sterilia. 

Suffrutescent herbs 20 cm. high or more; stems subquadrangular, 
hirsute, the hairs 0.5 mm. long, arranged in 2 lines, or more or less 
scattered; leaf blades oblong, acuminate (the tip blunt), narrowed 
at the base, hirsute, the hairs up to 1 mm. long; petioles about 3 cm. 
long, hirsute; flowers borne in lax, terminal spikes about 5 cm. long, 
the rachis hirsute; bracts and bractlets subulate, up to 1 mm. long, 
sparingly puberulent; calyx segments subulate, 3 mm. long, 0.5 mm. 
wide at the base, glandular-puberulent; corollas 4.5 mm. long 
(immature), finely pubescent, the lobes about 2 mm. long and 1.5 
mm. wide, rounded; posterior stamens sterile. Thickets along Rio 
Ceibo, Buenos Aires, 200 meters, January, 1892, Tonduz 6702 in 
part (type in U. S. Nat. Herb., No. 59,938) . Distinct from C. Tonduzii 
in its hirsute foliage and inflorescence. 

Chamaeranthemum Tonduzii Lindau in Pitt. Prim. 2: 303. 
1900. Forests of Te>raba, Pittier 3984. Herbs; stems erect, up to 
50 cm. high, simple or sparingly branched, pilosulous in 2 lines at 
the tips, otherwise glabrous, terete; leaf blades oblong-lanceolate, 



FLORA OF COSTA RICA 1209 

3-9 cm. long, 1-2.5 cm. wide, acuminate, the tip blunt, rounded or 
narrowed at the base, both surfaces glabrous and bearing numerous 
cystoliths; petioles 1-3 mm. long; inflorescence a sparingly branched 
panicle of lax spikes 2-9 cm. long, the flowers secund, the rachis 
sparingly puberulent; bracts and bractlets subulate (or the lower- 
most bracts linear-lanceolate), 2-8 mm. long, glabrous or sparingly 
puberulent; calyx segments subulate, about 3 mm. long, glandular- 
puberulent; corolla about 10 mm. long, minutely pubescent, the 
limb about 1 cm. broad, the lobes rounded; anthers of the posterior 
pair of stamens 1-celled or sterile, those of the anterior pair 2-celled; 
capsules about 15 mm. long, the upper portion 2-3 mm. broad, finely 
pubescent. 

DICLIPTERA Juss. 

Herbs; leaves entire, usually ovate, petioled; inflorescence spicate, 
cymose, or paniculate, the flowers 1-several, subtended by an involucre 
of 2-4 conspicuous bracts, the cymes often contracted; calyx 5-parted, 
hyaline; corolla narrow, slightly ampliate, the limb 2-lipped; stamens 
2, the anther sacs often unequal, the longer ones sometimes calcarate 
at the base; staminodes none; capsules ovate or suborbicular, the 
placenta separating elastically from the walls; seeds 2 or 4. A large 
genus, found in tropical and subtropical regions in many parts of 
the world. 

Cymes sessile or subsessile (the lowermost sometimes short-pedun- 

cled). 

Bracts subtending the flowers small, the larger bract of the pair 
subtending the middle flower of each cyme up to 9 mm. long 

and 4.5 mm. wide D. imbricata. 

Bracts subtending the flowers larger, the larger bract of the pair 
subtending the middle flower of each cyme 10 mm. long and 
5 mm. wide or more. 
Flower bracts unguiculate, spine- tipped; corolla 14 mm. long 

or less, rose-purple D. unguiculata. 

Flower bracts not conspicuously spine- tipped or unguiculate; 

corolla 2.5-3.5 cm. long, brick-red D. pallida. 

Cymes peduncled. 

Inflorescence strongly glandular-pubescent D. Skutchii. 

Inflorescence eglandular (the corolla more or less glandular in 

D. iopus). 

Corolla 14 mm. long; bracts subtending the flowers elliptic or 
obovate D. podocephala. 



1210 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Corolla 3 cm. long or more; bracts subtending the flowers oblong. 
Flowers bracts rounded and apiculate; pedicels essentially 

glabrous D. trifurca. 

Flower bracts acute to obtusish; pedicels subtomentose. 

D. iopus. 

Dicliptera imbricata Leonard, sp. nov. Herbae, caulibus 
ramosis, hexagonis, angulis puberulentis, pilis curvatis; lamina 
foliorum oblongo-ovata, acuminata, basi angustata, glabra vel in 
venis puberulenta, pilis curvatis; petioli graciles, puberulenti; in- 
florescentiae spiciformes, terminales et axillares, compactae, cymis 
subsessilibus, verticillatis, bracteis viridibus lanceolatis suffultis; 
bracteae floriferae inaequales, obovatae, trinerviae, basin versus 
albidae, majores acutiusculae, minores rotundatae; bracteolae 
lineares, uninerviae, acuminatae, ciliatae, albidae; corolla purpurea; 
capsulae apice tenuiter pubescentes. 

Herbs up to 1 meter high; stems branched, the tips hexagonal, 
puberulent on the angles with curved hairs, the basal portions 
sub terete and subglabrous; leaf blades oblong-ovate, up to 9 cm. 
long and 4 cm. wide, acuminate (the tip blunt), narrowed at the 
base, rather thin, glabrous, or the costa and veins puberulent with 
curved hairs; petioles slender, up to 5 cm. long, finely pubescent with 
curved hairs; flowers crowded in terminal, spikelike racemes up to 
3 cm. long and 1.5 cm. in diameter, each node producing several 
short-peduncled, fanlike cymes of 3-5 flowers each, or the lowermost 
cymes in the axils of the leaves, each cyme subtended by a pair of 
lanceolate, green bracts about 12 mm. long and 1-3 mm. wide, 
extending conspicuously beyond the calyces, the bracts subtending 
the flowers unequal, obovate, whitish toward the base, the larger 
of the pair subtending the middle flower about 9 mm. long and 4.5 
mm. wide, acutish, strongly 3-nerved, the smaller one 7 mm. long 
and 3 mm. wide, rounded, those subtending the outer flowers similar 
but successively smaller, all strongly puberulent and ciliate, the hairs 
0.5 mm. long; bractlets linear, the outer pair 7 mm. long and 1 mm. 
wide, the inner 5 mm. long, 0.75 mm. wide, all acuminate, 1-nerved, 
ciliate, whitish; calyx 3.5 mm. long, subhyaline, the segments subu- 
late, 0.5 mm. wide at the base, ciliolate; corollas 15 mm. long(?), 
purple; capsules about 5 mm. long, finely pubescent toward the tip. 
San Pedro de San Ramon, January 31, 1936, Brenes 21440 (type 
in Herb. Field. Mus., No. 866,336). Calera de San Ramon, Brenes 
6533. Low forests in the vicinity of Capulin, Rio Grande de Tarcoles, 
Prov. Alajuela, Standley 40107. Moist forests of La Tejona, north 



FLORA OF COSTA RICA 1211 

of Tilaran, Guanacaste, 600-700 meters, Standley & Valeria 45868. 
Santiago de San Ramon, Brenes 1 7006. This differs markedly from 
other species of Dicliptera found in Costa Rica in its close, terminal, 
spikelike inflorescences, small, strongly ciliate bracts, and elongate, 
leafy bracts subtending the cymes. 

Dicliptera iopus Lindau in Pittier, Prim. 2: 305. 1900. Forests 
of Rancho Flores, 2,045 meters, Tonduz 2136. Wet forests of Costa 
Rica and Panama, and probably elsewhere in Central America. 
An erect, branched herb, up to 60 cm. high, the branches subhexa- 
gonal, pilosulous with small, curved hairs, or glabrous, the hairs 
chiefly on the angles; leaf blades oblong-ovate, up to 13 cm. long and 
9 cm. wide but usually smaller, acuminate, the tip blunt, narrowed 
at the base, glabrous or pilosulous, the hairs about 0.5 mm. long, 
mainly on the costa and lateral veins; petioles slender, up to 2, rarely 
4 cm. long, subtomentose; flowers borne in peduncled cymes usually 
of 3 flowers each, the peduncles about 3 cm. long although occasion- 
ally as much as 9 cm., pilosulous, the pedicel of the middle flower 
of each cyme 8-10 mm. long, those of the lateral flowers much 
shorter, all subtomentose; bracts subtending the cymes resembling 
small leaves; bracts subtending the flowers linear-oblong, acute to 
obtusish, pubescent toward the tip, unequal, the larger about 15 mm. 
long and 4 mm. wide, the others 10 mm. long and 3 mm. wide; 
bractlets 4, oblanceolate, acute, pubescent toward the tip, the larger 
pair 10 mm. long and 2.5 mm. wide, the smaller 8 mm. long and 1.5 
mm. wide; calyx 8 mm. long, minutely pubescent, the segments 
linear, gradually narrowed to a slender tip; corolla rose-purple, 
about 3 cm. long, inflated at the middle, pilose (some of the hairs 
glandular), the mouth 4-6 mm. in diameter, the posterior lip sub- 
entire, about 9 mm. long and 4.5 mm. wide, the lower one 3-lobed, 
the lobes about 2 mm. long and 1.5 mm. wide. 

Dicliptera pallida Leonard, sp. nov. Herbae erectae, ramosae, 
caulibus hexagonis, subtomentosis; lamina foliorum ovata, breviter 
acuminata, basi acuta, hirsuta; petioli subtomentosi; cymae breviter 
pedunculatae, axillares, bracteis foliosis suffultae; bracteae floriferae 
inaequales acuminatae, hirtellae, basin versus albidae, majores 
oblanceolatae ad ellipticae, minores lineares; bracteolae anguste 
lanceolatae, basin versus albidae; calycis segmenta alba, ciliolata; 
corolla ferruginea, pubescens, labio superiore emarginato, inferiore 
minute trilobo; capsulae pubescentes. 

An herb, up to about 1 meter high, erect, branched; stems hexa- 
gonal, subtomentose or the lower portions glabrescent, the hairs 



1212 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

minute and retrorsely curved; leaf blades ovate, up to 10 cm. long 
and 5 cm. wide, short-acuminate, acute at the base, both surfaces 
hirsute, the hairs up to 1 mm. long; petioles up to 3 cm. long, sub- 
tomentose; flowers borne in short-peduncled, axillary cymes, the 
peduncles up to 1 cm. long, white- tomentose, the flowers of the upper 
cymes sessile, those of the lower ones short-pediceled, the pedicel 
of the middle flower longer than those of the lateral ones, all white- 
tomentose; bracts subtending the cymes leaflike, 2-3 cm. long, those 
subtending the flowers (usually 1 fertile flower flanked by 1 or more 
abortive ones) unequal, the larger of the pair subtending the middle 
flower of each cyme oblanceolate or elliptic, 1.7 cm. long, 6 mm. wide, 
the smaller one linear, 11 mm. long and 2 mm. wide, both acuminate, 
narrowed to a slender point, whitish toward the base, hirtellous, 
the hairs spreading, up to 1 mm. long, the bracts of the lateral 
flowers of each cyme similar but successively smaller; bractlets un- 
equal, the outer pair narrowly lanceolate, 12 mm. long and 1.25 mm. 
wide, ciliate, the inner pair similar but smaller; calyx segments white, 
lanceolate, 3 mm. long, 0.75-1.25 mm. wide, minutely ciliolate; 
corollas brick-red, 2.5-3.5 cm. long, pubescent, the tube slender, 
2 mm. in diameter at the base, 4 mm. at the mouth, the lips about 
8 mm. long, the upper emarginate, the lower minutely 3-lobed; 
stamens barely exserted; capsules 4-5 mm. long, pubescent. Wet 
thicket in the vicinity of Pejivalle, Prov. Cartago, 900 meters, 
February 7-8, 1926, Standley & Valeria 46792 (type in U. S. Nat. 
Herb., No. 1,251,470). Valle de Los Arcangeles, central Costa Rica, 
1,700 meters, Pittier 12305. Thickets of Las Vueltas, Tucurrique, 
635 meters, Tonduz 12935 (Donnell Smith 7570). Shady river bank, 
vicinity of El General, Prov. San Jose", 880 meters, Skutch 2297. 
The name pallida refers to the whitish bractlets and basal portions 
of the bracts. 

Dicliptera podocephala Bonn. Smith, Bot. Gaz. 48: 299. 1909. 
Damp meadows and river banks. Type from Atirro, Prov. Cartago, 
Donnell Smith 6685. Suffrutescent, up to 1.5 meters high, diffuse, 
the branches hexagonal, sparingly pubescent; leaf blades ovate- 
lanceolate, up to 12.5 cm. long and 4.5 cm. wide, acuminate, blunt 
and sometimes curved, narrowed at the base, thin, glabrous or the 
veins with a few minute hairs; petioles up to 2.5 cm. long; flowers 
borne in peduncled capitula, the peduncles up to 9.5 cm. long, solitary 
or several in each axil, sparingly and minutely appressed-pubescent, 
the hairs mostly in 2 lines; bracts subtending the capitula narrowly 
lanceolate, about 5 mm. long, glabrous, sometimes ciliolate, often 



FLORA OF COSTA RICA 1213 

semicircularly curved ; fertile flowers usually 5 or 7 in each capitulum 
(sometimes accompanied by abortive ones), the bracts subtending the 
flowers elliptic to obovate, one of each pair slightly larger than the 
other, the middle pair the largest, the others diminishing in size out- 
wardly, the largest 11-14 mm. long, 8-9 mm. wide, all rounded or 
obtuse and minutely apiculate at the apex, narrowed at the base, 
pergamentaceous, sparingly puberulent and ciliolate; bractlets 4, 
linear-lanceolate, 8 mm. long, puberulent; calyx segments resembling 
the bractlets but slightly smaller; corolla 14 mm. long, pubescent; 
capsule ovoid, 5 mm. long, pubescent; seeds puberulent, 2.5 mm. in 
diameter. 

Dicliptera Skutchii Leonard, sp. nov. Herba diffusa ramosa, 
caulibus hexagonis, angulis puberulentis, pilis albis curvatis; lamina 
foliorum ovata ad elliptica, acuminata, basi angustata, membranacea, 
glabra vel in venis parce puberulenta; petioli graciles, subglabri; 
pedunculi terminales et axillares, apice triflori; inflorescentiae brac- 
teae foliosae; bracteae floriferae inaequales, lineari-oblongae, glandu- 
loso-hirtellae, longiores apice acutiusculae, breviores obtusiusculae; 
bracteolae lanceolatae, acutae, glanduloso-hirtellae; calycis segmenta 
anguste triangularia, puberulenta; corolla purpurea, tenuiter pubes- 
cens, labio superiore acuto, apice emarginato, inferiore trilobo, lobis 
suborbicularibus; ovarium glabrum. 

A sprawling herb, up to 1 meter high or more; stems branched, 
puberulent, the hairs white, curved, distributed in 6 lines; leaf blades 
ovate to elliptic, up to 16 cm. long and 8 cm. wide, acuminate, 
narrowed at the base, thin, glabrous, or the costa and veins bearing a 
few minute hairs; petioles slender, up to 6.5 cm. long, essentially 
glabrous; flowers in 3's, at the tips of axillary and terminal peduncles, 
these up to 8 cm. long, puberulent with white, curved hairs; pedicels 
1-2 mm. long, hirsutulous, the hairs straight or curved; bracts sub- 
tending the pedicels leaflike, about as long as the peduncles; bracts 
subtending the flower clusters leaflike, usually exceeding the calyx; 
bracts subtending the flowers unequal, linear-oblong, one 15 mm. 
long and 3.5 mm. wide, acutish, the other 10 mm. long and 3 mm. 
wide, obtusish, both glandular-hirtellous; bractlets lanceolate, the 
outer pair 9 mm. long and 2.25 mm. wide, the inner pair 5 mm. long 
and 1.5 mm. wide, all 4 acute and glandular-hirtellous; calyx about 
9 mm. long, essentially glabrous without, the segments narrowly 
triangular, 5-6 mm. long, 1.5-2 mm. wide at the base, puberulent 
within, the hairs appressed upwardly; corollas purple, up to 5 cm. 
long, sparingly and finely pubescent, the lower half of the tube 



1214 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

slender, 2 mm. in diameter at the base, the upper half enlarged to 
about 10 mm., then reduced to 6-7 mm. at the mouth, the lips about 
1 cm. long, one acute, minutely notched at the tip, the other 3-lobed, 
the lobes suborbicular, about 0.75 mm. in diameter; ovary glabrous; 
mature capsules not seen. -Woods in the vicinity of El General, 
Prov. San Jose", 1,190 meters, December, 1936, Skutch 3090 (type in 
U. S. Nat. Herb., No. 1,643,133). From its closest relative, D. iopus, 
the present species can be separated by its glandular-hirsutulous 
bracts and the coarser pubescence of its stems. 

Dicliptera trifurca Oerst. Vid. Medd. Kjoebenhavn 1854: 172. 
1855. Between Cartago and Candelaria, 1,800-2,100 meters, Oersted. 
Found chiefly in wet forests in the provinces of Cartago and San 
Jose". Endemic. Branching herbs, up to 2 meters high; stems 6- 
angled, glabrous or the angles minutely strigose; leaf blades oblong- 
ovate or elliptic, up to 19 cm. long and 7 cm. wide, acuminate, acute 
at the base, thin, glabrous or the costa and veins sparingly and 
minutely strigose, both surfaces with minute cystoliths; petioles 
slender, minutely and sparingly strigose; flowers in 3's, borne in 
axillary, peduncled cymes; peduncles up to 16 cm. long, glabrous or 
with a few minute hairs; pedicels up to 5 mm. long, glabrous or with 
a few small hairs on the angles; bracts subtending the cymes leaflike, 
equaling or exceeding the calyx, deciduous; bracts subtending the 
flowers oblong, unequal, one about 9 mm. long, the other about 6 mm. 
long, both 2.5 mm. wide, rounded and apiculate at the apex, essen- 
tially glabrous; bractlets oblong-lanceolate, 6.5 mm. long, 2 mm. 
wide, acute, 1-nerved, minutely puberulent toward the tip; calyx 7 
mm. long, the segments 4 mm. long, about 1 mm. wide at the base, 
gradually narrowed to a slender point, finely pubescent; corolla about 
3 cm. long, pubescent, bright rose-purple, the tube about 12 mm. long, 
slightly swollen below the mouth, about 5 mm. in diameter at its 
widest part, the lips subequal, the upper one entire, the lower 3- 
lobed, the lobes suborbicular, about 0.75 mm. in diameter; capsules 
17 mm. long and 5 mm. broad, pointed at the apex, stipitate at the 
base, retrorsely pubescent; seeds flat, suborbicular, about 4.5 mm. 
in diameter, with a few papilliform hairs on the margins. 

Dicliptera unguiculata Nees. Sornia, Olotillo. Common in moist 
thickets, Mexico to northern South America. Erect or ascending 
herbs, up to about 1 meter high; stems branched, terete and glabrous 
below, hexagonal and pubescent toward the tips, the hairs spreading 
or retrorsely curved, 0.5 mm. long; leaves soon deciduous; leaf blades 



FLORA OF COSTA RICA 1215 

ovate, up to 8.5 cm. long and 5 cm. wide, acuminate, the tip blunt, 
narrowed at the base, thin, both surfaces of the younger leaves 
densely pilosulous but sparingly so at maturity; petioles up to 4 cm. 
long, pilosulous or subglabrous; flowers crowded in sessile, axillary 
umbels or in crowded, terminal and axillary, spikelike racemes up 
to 8 cm. long and about 2 cm. in diameter; bracts spatulate, unequal, 
the upper suborbicular to ovate, 1-1.5 cm. long, 5-7 mm. wide, the 
lower about half the length of the upper, both aristate (the awn 2-3 
mm. long), hirsutulous, more or less densely ciliate with hairs up to 
1 mm. long, 3-nerved; bractlets linear-lanceolate, 4 mm. long, 1- 
nerved, ciliolate; calyx 3 mm. long, the segments narrowly lanceolate, 
minutely ciliolate, 1-nerved; corolla 12-14 mm. long, rose-purple, 
pubescent, the tube slender, the lips subequal, about 7 mm. long, 
the upper one with 3 minute lobes, the lower emarginate; capsules 
clavate, 4 mm. long, retrorsely hirsutulous; seeds brownish, flat, 
suborbicular, 1-1.5 mm. in diameter, with minute, retrorsely barbed 
hairs. Standley states that this plant is a common remedy for in- 
testinal disorders. According to Raul Goto Fernandez, it has been 
employed successfully in Costa Rica in treating amebic dysentery; 
see Revista de Agr. 10: 167. 1938. 

DYSCHORISTE Nees 

Herbs or woody plants; leaf blades usually ovate to lanceolate, 
entire, petioled; flowers solitary or clustered in the leaf axils, some- 
times terminal; calyx segments 5, very slender; corolla blue, purple, 
or sometimes white, slightly 2-lipped. Widely distributed in tropical 
and subtropical regions of the world. Several species occur in Central 
America. 

Dyschoriste Valeriana Leonard, sp. nov. Herbae robustae, 
perennes, caulibus erectis vel adscendentibus, simplicibus vel parce 
ramosis, quadrangularibus, glabris vel parce pilosulis; lamina foliorum 
ovata ad oblongo-ovata, rotundata vel obtusa, vel interdum acutius- 
cula, basi angustata et in petiolum decurrens, membranacea, ciliolata, 
glabra vel subglabra, novellae minute et sparse pubescentes; petioli 
glabri vel subglabri; cymae axillares; bracteae foliosae; calycis 
segmenta subulata, subcarinata, ciliata; corolla pallide purpurea, 
limbo subbilabiato, lobis ovalibus, rotundatis; capsulae lanceolatae, 
glabrae. 

A robust perennial; stems simple or sparingly branched, erect 
or ascending, sometimes rooting at the basal nodes, up to 50 cm. 
high or more, quadrangular, glabrous or sparingly pilosulous; leaf 



1216 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

blades ovate to oblong-ovate, 2-7 cm. long, 1-3.5 cm. wide, rounded, 
obtuse, or sometimes acutish, narrowed at the base and decurrent 
on the petiole, entire or undulate, thin, ciliolate, the young leaves 
minutely and sparingly pubescent, the older ones glabrous or the 
costa and veins with a few small, white hairs, the cystoliths slender; 
petioles slender, 5-15 mm. long, glabrous or sparingly pubescent; 
flowers borne in cymose clusters at the nodes and subtended by 
leaflike bracts 10-15 mm. long and 5-10 mm. wide; calyx 10-11 mm. 
long, the tube 4 mm. long, glabrous, the segments subulate, 0.75 
mm. wide at the base, subcarinate, ciliate; corollas pale purple, 8-10 
mm. long, the lobes sparingly puberulent, the tube 2 mm. in diameter 
at the base, 1 mm. above the ovary, 2.5 mm. at the throat, the limb 
subbilabiate, the lobes oval, about 3 mm. long, 2-2.5 mm. wide, 
rounded at the apex, the filaments of the shorter stamens 1 mm. 
long, those of the longer ones 2.25 mm. long, all glabrous, the anthers 
slightly less than 1.5 mm. long, minutely calcarate at the base; 
capsules lanceolate, 8 mm. long, glabrous, 4-seeded. Vicinity of 
Tilaran, Guanacaste, alt. 500-650 meters, January 10-31, 1926, 
Standley & J. Valeria 46613 (type in U. S. Nat. Herb., No. 1,254,741), 
44522, 44965. Easily recognized by its relatively large, nearly 
glabrous leaves and small flowers. 

ELYTRARIA Michx. 

Caulescent or acaulescent herbs; leaves basal or crowded at the 
ends of the branches; flowers borne in peduncled spikes, both peduncle 
and spike bearing coriaceous bracts; calyx segments narrow, thin, 
entire, or the posterior segment bidentate at the apex; corolla small, 
blue or white. The genus is composed of about a dozen species, 
mostly native to tropical or subtropical America. 

Elytraria imbricata (Vahl) Pers. E. tridentata Vahl. Old 
fields, grassy banks, bushy slopes, and other waste places. Probably 
rather common in Costa Rica. Ranging from Arizona and Texas 
to northern and western South America. Introduced into the 
Philippine Islands. Usually caulescent, up to 30 cm. high, the leaves 
crowded at the tips of slender, glabrous or sparingly pilosulous stems; 
leaf blades oblong to obovate, up to 18 cm. long and 6 cm. wide, 
blunt or acutish at the apex, narrowed at the base to a slender, 
winged petiole; scapes numerous, axillary, up to 24 cm. long, simple 
or branched, covered by closely appressed, sheathing, ovate or subu- 
late bracts; spikes up to 6 cm. long, the bracts elliptic, 3-6 mm. long, 
ending in an awn and bearing near the tip a pair of triangular or 



FLORA OF COSTA RICA 1217 

rhombic, scarious teeth; bractlets 3 mm. long; corolla blue, 5-8 mm. 
long; capsule oblong, glabrous. Extremely variable in habit of 
growth. Surprisingly enough, the scarious teeth of the flower 
bracts, one of the main characters on which this species is based, 
are sometimes much reduced or even entirely lacking. In Salvador, 
a decoction of the plant is said to be used to alleviate dysentery. 

ERANTHEMUM L. 

Shrubs or suffrutescent plants; flowers borne in spikes or panicles, 
the bracts reticulate- veined, imbricate; calyx 5-parted the lobes 
often scarious; corolla tube slender, incurved, the limb oblique, the 
lobes spreading and subequal; stamens 2, included; staminodes 2; 
capsules ovate or oblong, stipitate; seeds 4. A genus consisting of 
about 20 species, which are native to eastern India and the Malay 
Archipelago. 

Eranthemum nervosum (Vahl) R. Br. Native to eastern 
India, but found occasionally both as a cultivated plant and as an 
escape in tropical regions of America. A shrub, up to 2 meters high; 
stems quadrangular, glabrous or puberulent; leaf blades oblong-ovate 
to elliptic, usually 9-12 cm. long (or the lowermost as much as 20 cm. 
long), 4-5 cm. wide, acuminate, rounded or abruptly narrowed at 
the base and decurrent on the petiole, entire, glabrous beneath, 
except the costa and lateral nerves, these prominent and minutely 
puberulent, both surfaces bearing numerous inconspicuous cysto- 
liths; petioles 1-1.5 cm. long; flowers borne in numerous axillary 
spikes up to 6 cm. long or more, the bracts closely imbricate, elliptic, 
1-1.5 cm. long, whitish, strongly veined; corolla about 2 cm. long, 
blue, 5-lobed, the tube slender, the limb about 1.5 cm. broad. 

GLOCKERIA Nees 

Herbaceous or shrubby plants; flowers usually borne in terminal 
and axillary, thyrsiform panicles; bracts small, subulate; calyx 
segments 5, narrow, equal; corollas red or yellow, tubular, narrow, 
usually constricted just above the ovary, thence abruptly and some- 
what obliquely enlarged and oblong, often slightly constricted at the 
mouth, the upper lip narrow, erect, entire, the lower scarcely distinct 
from the tube, minutely 3-lobed; stamens 2, attached at the top of 
the tube, exserted, the anthers 1-celled, obtuse; capsules clavate, 
glabrous, 4-seeded; seeds compressed, roughened. Several species 
other than the four described here occur in Mexico and South 
America. 



1218 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Leaf blades rounded or subcordate at the base. 

Corolla 2.5 cm. long; leaf blades densely pilose G. ventricosa. 

Corolla 1.2 cm. long; leaf blades glabrous or sparingly pilose. 

G. sessilifolia. 
Leaf blades narrowed and decurrent on the petiole at the base. 

Inflorescence open, its branches relatively few, strict, erect or 
ascending G. stricta. 

Inflorescence compact, its branches very numerous, spreading. 

G. blepharorhachis. 

Glockeria blepharorhachis (Lindau) Leonard, comb. nov. 
Kolobochilus blepharorhachis Lindau in Pittier, Prim. 2: 309. 1900. 
Type from Costa Rica or Panama, Warscewicz. Wet forests. En- 
demic. Herbaceous or sufiruticose, up to 1.5 meters high; stems 
terete, glabrous below, the upper portions subtomentose with brown- 
ish hairs about 1 mm. long; leaf blades ovate, elliptic, or oblong, up 
to 22 cm. long and 9.5 cm. wide, acuminate, narrowed at the base, 
entire or crenate, the younger ones pilose, the older ones glabrous 
except as to costa and veins; petioles up to 5 cm. long, pilose; flowers 
numerous, borne in a compact, elongate thyrse up to 30 cm. long or 
more and 3-6 cm. in diameter, the branches slender and fascicled, 
spreading, the rachis densely pilose or subtomentose (hairs brownish, 
about 1 mm. long), the peduncles pubescent with hairs 0.5 mm. long; 
pedicels slender, up to 4 mm. long, minutely pubescent; bracts lanceo- 
late, the lowermost leaflike, the others diminishing successively 
toward the tip; bractlets similar but smaller; calyx 8-10 mm. long, 
puberulent, the minute hairs intermixed with longer (0.5 mm.), 
glandular ones, the segments narrowly linear, 0.5-1 mm. wide; 
corollas red to orange-yellow, 15 mm. long, glabrous without, pilosu- 
lous within, the lower portion of the tube about 2 mm. in diameter, 
enlarged to 6 mm. above the ovary, the mouth scarcely contracted, 
the upper lip erect, 5-7 mm. long, 3-4 mm. wide, entire, the lower lip 
scarcely distinct from the tube, the lobes 1-2 mm. long; stamens 
barely exserted; capsules 15 mm. long, 3 mm. wide; seeds flat, 2 mm. 
in diameter, roughened. 

Glockeria sessilifolia Oerst. Vid. Medd. Kjoebenhavn 1854: 
141. 1855. Volcan de Barba, 1,800 meters, Oersted. Found on the 
wooded, southern slopes of Poas and Barba. Herbaceous or suf- 
frutescent, up to 2 meters high; stems branching, glabrous below, 
the tips quadrangular, glabrous or puberulous in 2 lines; leaf blades 
ovate, up to 6 cm. long or more and 5.5 cm. wide, obtuse or short- 
acuminate, the tip blunt, rounded or subcordate at the base, gla- 



FLORA OF COSTA RICA 1219 

brous or sparingly pilose; petioles up to 4 mm. long; flowers borne in a 
lax, terminal panicle up to 15 cm. long and 12 cm. in diameter, some- 
times with smaller additional axillary panicles, the branches opposite, 
spreading, slender, glabrous or sparingly pubescent; pedicels about 
1 mm. long; bracts narrowly triangular, up to 1.5 mm. long; calyx 
segments linear-subulate, 4 mm. long, sparingly glandular-pubescent, 
the hairs about 0.5 mm. long; corolla 1.2 cm. long, glabrous, red and 
yellow, the narrow basal portion of the tube 2.5 mm. long and 1 mm. 
in diameter, the upper portion enlarged to 3-4 mm., thence gradually 
narrowed to 2.5 mm. at the mouth, the upper lip spatulate, acute, 
1 mm. wide at the base, 0.5 mm. wide near the tip, the lower lip short, 
recurved, about 3 mm. long and broad, the lobes 0.5 mm. long, 1 mm. 
wide, rounded; stamens barely exserted; capsule 12-13 mm. long; 
seeds flat, hispidulous. 

Glockeria stricta Leonard, sp. nov. Herba erecta, caulibus 
superne pubescentibus, pilis flavescentibus, retrorsis, inferne glabris; 
lamina foliorum oblongo-elliptica, breviter acuminata, basi angustata, 
membranacea, glabra, costa et venis strigosis exceptis; petioli graciles, 
canalibus appresso-pubescentibus; inflorescentiae racemosae, racemis 
terminalibus et axillaribus, simplicibus vel ramosis, vel saepe pani- 
culatae; rhachis puberulenta, pilis curvatis, interdum glanduliferis et 
patulis; bracteae anguste triangulares vel lanceolatae, puberulentae; 
calycis segmenta anguste linearia vel subulata, puberulenta, interdum 
pilis patulis glanduliferis instructa; corolla rubra et lutea, labio 
superiore ovato, obtusiusculo, inferiore trilobo; capsulae glabrae vel 
apice parce pubescentes, stipitatae; semina plana, muricata. 

An erect herb, up to 1.5 meters high; stems glabrous below, 
puberulent above with yellowish, retrorsely curved hairs, these 
scattered or arranged in 2 lines; leaf blades oblong-elliptic, up to 12 
cm. long and 5 cm. wide, short-acuminate (the tip blunt), narrowed 
at the base, thin, glabrous except the costa, veins, and margins, these 
minutely strigose; petioles slender, up to 3 cm. long, glabrous below, 
the channels appressed-pubescent; flowers borne in terminal and 
axillary racemes, these erect or ascending, simple or often branched 
and forming a large, terminal panicle, the flowers approximate or 
distant, the internodes up to 1.5 cm. long, the rachis puberulent, or 
the minute hairs sometimes intermixed with long (0.5-1 mm.), 
glandular, spreading ones, the pedicels slender, 2-3 mm. long, puberu- 
lent; bracts narrowly triangular or lanceolate, 2-3.5 mm. long, about 
1 mm. wide, puberulent; calyx segments narrowly linear or subulate, 
5-7 mm. long, 0.5-0.75 mm. wide, puberulent, often with additional 



1220 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

scattered, spreading, glandular hairs up to 1 mm. long; corolla red 
and yellow, 1.5 cm. long, the narrow basal portion of the tube 3 mm. 
long and 1.5 mm. in diameter, thence abruptly and somewhat 
obliquely enlarged to 5 mm., then gradually narrowed to 4 mm. at 
the mouth, the upper portion of the tube sparingly pubescent, the 
upper lip ovate, 3 mm. long, 2.5 mm. wide at the base, obtusish, the 
lobes of the lower lip 0.5 mm. long and 0.75 mm. wide; capsules 13 
mm. long, 2.5 mm. wide, glabrous or sparingly pubescent toward the 
tip, 4-seeded, stipitate; seeds flat, muricate. Moist forest, vicinity 
of Santa Maria de Dota, Prov. San Jose", 1,500-1,800 meters, Decem- 
ber 26, 1925-January 3, 1926, Standley & Valeria 43311 (type in 
U. S. Nat. Herb., No. 1,253,199), 43225, 43301. Moist forests of 
Cerro de La Carpintera, Prov. Cartago, 1,500-1,850 meters, Standley 
34212. San Antonio, Brenes 3766. Vara Blanca de Sarapiqui, north 
slope of Central Cordillera, between Pods and Barba volcanoes, 
Skutch 3577. Wet forests of Viento Fresco, Prov. Alajuela, 1,600- 
1,900 meters, Standley & Torres 47769, 47848, 47858. Well marked 
by its strict inflorescence and glandular-pilose rachis and calyx. 

Glockeria ventricosa Bonn. Smith, Bot. Gaz. 27: 439. 1899. 
Alto del Roble, Volcan de Barba, 2,600 meters, Pittier 3519. Slopes 
of Alto del Roble, Volcan de Turrialba, Cerro Gallito, and Vara 
Blanca de Sarapiqui. Endemic. Shrubby, the branches terete, sub- 
tomentose, the hairs about 1 mm. long; leaf blades ovate to oblong- 
ovate, up to 17 cm. long and 9.5 cm. wide, acuminate, rounded or 
subcordate at the base, both surfaces densely pilose; petioles up to 
8 cm. long, pilose; flowers numerous, borne in a compact, elongate 
thyrse up to 50 cm. long, the rachis and branches puberulent; pedicels 
slender, up to 2.5 cm. long, puberulent, some of the hairs glandular; 
bracts oblong to lanceolate, acutish to acuminate, up to 13 mm. long 
and 4 mm. wide, subglabrous; calyx 7 mm. long, becoming 13 mm. 
long at fruiting stage, glandular-pubescent, the segments linear; 
corolla 2.5 cm. long, orange, sparingly pilose, the tube about 4 mm. 
in diameter at the base, contracted to 3 mm. above the ovary, thence 
abruptly enlarged to 5 mm., incurved, scarcely constricted at the 
mouth, the posterior lip elliptic, 8-10 mm. long, 4-5 mm. wide, 
entire, the lobes of the lower lip 1.5 mm. long; stamens exserted; 
capsules 15-17 mm. long. 

GRAPTOPHYLLUM Nees 

Shrubs; leaf blades usually entire, colored; flowers clustered 
either in a terminal thyrse or in the axils of the leaves; calyx segments 



FLORA OF COSTA RICA 1221 

5; corollas reddish purple, ringent, the tube inflated above, the upper 
lip with 2 short, recurved lobes, the lower lip 3-parted; stamens 2. 
The genus consists of 5 or more species, all oriental. The name is 
from the Greek, referring to the variegated foliage. 

Graptophyllum picturn (L.) Griff. Cultivated in gardens of 
tropical America. Shrubs, up to 3 meters high; stems glabrous; 
leaf blades ovate to elliptic, up to 20 cm. long and 9 cm. wide, acu- 
minate, narrowed at the base, glabrous, mottled with yellow along 
the midrib; flowers borne in terminal racemes, the rachis glabrous 
or sparingly puberulent; corollas up to 8 cm. long, purplish red, 
glabrous without, glandular-puberulent within, the mouth about 
1.5 cm. broad; stamens 2, exserted; staminodes 2. 

HABRACANTHUS Nees 

Herbaceous or shrubby plants; leaves usually thin, ovate to 
lanceolate; flowers borne in terminal panicles; bracts and bractlets 
minute; calyx deeply 5-parted, the segments equal; corolla tubular, 
ringent, the lips elongate, ligulate, the upper one entire, the lower 
one minutely 3-lobed at the apex; stamens 2, attached at the top 
of the tube, exserted; anthers 1-celled, attached to the filament below 
the middle; capsules 4-seeded, the basal half slender, solid. Repre- 
sented in Central America and Mexico by a single species. Other 
species have been described from South America. 

Habracanthus silvaticus Nees. Wet forests and clearings of 
the central cordillera. Originally described from Chiapas, Mexico. 
Herbs or slender shrubs, up to 2 meters high, the branches terete, 
glabrous, or puberulent in 2 lines with retrorsely curved hairs; leaf 
blades oblong-ovate to lanceolate, 5-18 cm. long, 1.5-6 cm. wide, 
acuminate, the tip rounded or obtuse (sometimes slightly falcate), 
narrowed at the base, thin, entire or undulate, glabrous or the costa 
and lateral veins pubescent with minute, curved hairs, both surfaces 
usually bearing minute cystoliths; petioles up to 7 cm. long (usually 
shorter), glabrous beneath, the channeled upper portion puberulent 
with curved hairs; flowers borne in loose, terminal thyrses usually 
5-8 cm. long and 4-5 cm. broad, the lower portion of the thyrse 2-3 
times dichotomously branched, the entire inflorescence essentially 
glabrous; bracts subulate, the lowermost 3-6 mm. long, about 1 mm. 
wide at the base, sometimes bearing several minute hairs, the others 
similar but successively smaller toward the tip of the flower cluster; 
peduncles 2-3 cm. long; branches of the thyrse widely spreading; 
pedicels 5 mm. long or less; rachis and calyx whitish or lavender, 



1222 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the calyx lobes subulate, 3-4 mm. long; corolla 1-1.5 cm. long, pale 
purple or lavender, the tube slender, 1 mm. in diameter at the base, 
1.5 mm. at the mouth, the lips 1 cm. long, delicately nerved, the upper 
oblong, 2 mm. wide, rounded at the tip, the lower elliptic, 5 mm. 
wide, the lobes about 0.5 mm. long, the middle of these 1 mm. 
wide, the lateral about 0.5 mm. wide; filaments about as long as the 
lips of the corolla; capsules 1.5 cm. long, 2.5 mm. in diameter; seeds 
flat, brownish, about 2 mm. in diameter, minutely roughened and 
puberulent. 

HANSTEINIA Oerst. 

Herbaceous or suffrutescent plants; flowers borne in lax, terminal, 
dichotomously branched panicles and in axillary spikes; calyx 
segments 4, the superior ones narrowly linear, the others lanceolate, 
subequal; corolla tubular, the limb short, the upper lip narrow, 
entire, the lower 3-lobed; stamens 2, short, included; anthers 1-celled; 
capsule 4-seeded. 

Hansteinia gracilis Oerst. Vid. Medd. Kjoebenhavn 1854: 143. 
1855. Dense, damp woods on Mt. Jaris, Oersted. The following 
description is adapted from the original. Suffrutescent, 60 cm. high; 
stems erect, slender, simple or sparingly branched, sub tetragonal, 
pubescent; leaf blades elliptic-ovate, up to 20 cm. long and 8 cm. 
wide, acuminate, the tip blunt, narrowed at the base and decurrent 
on the petiole, subrepand-crenate, glabrous except on the costa and 
veins, these minutely strigose; flowers fascicled at the tips of the 
branches of a terminal panicle (up to 15 cm. long), the rachises 
glandular, ferruginous-pubescent; calyx segments red, 6-8 mm. long, 
or reaching 1.5 cm. at the fruiting stage, glandular- villous; corolla 
15-17 mm. long, red, tomentose; capsule 8-10 mm. long, puberulent. 

HYGROPHILA R. Br. 

Herbs; leaves entire, often with cystoliths; flowers sessile, fascicled 
or solitary, borne in the axils of the leaves; calyx 5-parted, the 
segments narrow, nearly similar; corolla tube subcylindric, the limb 
strongly 2-lipped, the upper lip 2-lobed, the lower 3-lobed, the lobes 
sinistrorsely contorted; stamens 4, didynamous; anthers 2-celled, 
their lobes unappendaged. Many species occur in tropical regions 
of both hemispheres. 

Hygrophila guianensis Nees. H. conferta Nees. Marshy 
places, usually along streams. Mexico to northern South America 
and West Indies. A branched herb, up to 50 cm. high, sparingly 
hirtellous or glabrate, the stem obtusely quadrangular; leaf blades 



FLORA OF COSTA RICA 1223 

lanceolate, 5-15 cm. long, 0.5-3 cm. wide, acuminate at both ends, 
entire; flowers fascicled; calyx segments lanceolate, about 5 mm. 
long, white-margined, pilose; corolla white or purplish, puberulent, 
the tube 5 mm. long, the lips about 2.5 mm. long; capsules narrowly 
oblong, 1.2-1.4 cm. long, glabrous, 16-18-seeded. 

JACOBINIA Moric. 

Herbs or shrubs; leaves usually oblong to ovate, entire (rarely 
crenate), petioled; flowers sessile or short-pedicellate in the axils 
of the bracts, solitary, cymose, spicate, or paniculate; calyx 5-parted, 
the segments narrow; corolla usually red or yellow, the tube usually 
slender and elongate, straight or curved, the limb 2-lipped, the upper 
lip narrow, erect, entire or minutely 2-lobed, the lower lip 3-lobed; 
stamens 2, the anther cells more or less unequally attached, muticous 
to acute or subcalcarate at the base; staminodes none; capsules 
clavate, 4-seeded. A rather large genus of shrubs or subshrubs, 
distributed throughout tropical America. Often cultivated in gardens 
or, in temperate regions, in greenhouses. 

Flowers borne in a large, dense, terminal thyrse J. umbrosa. 

Flowers borne in terminal or axillary cymes, panicles, or spikes. 
Leaves crenate; branches of the inflorescence puberulent. 

J. crenata. 

Leaves entire; branches of the inflorescence glabrous or subgla- 
brous. 

Corollas 5-6 cm. long J. macrantha. 

Corollas 3-3.5 cm. long J. tinctoria. 

Jacobinia crenata Leonard, sp. nov. Frutex, caulibus teretibus, 
glabris; lamina foliorum ovato-oblonga ad oblonga, acuminata, basi 
angustata, membranacea, glabra; petioli glabri; spicae breves, laxae, 
paniculatae, rhachibus puberulentis; bracteae oblongo-ovatae, apice 
rotundatae, sparse glanduloso-pubescentes, vel infimae foliosae et 
glabrae; calycis segmenta linearia, sparse puberulenta, minute cilio- 
lata, interdum pilis raris longioribus glanduliferis instructa; corolla 
rubra et flava, tenuiter pubescens, labiis angustis; antherae loculi 
inaequaliter affixi, inferior basi subcalcaratus; capsulae clavatae, 
glabrae vel apice pilos paucos glanduliferos gerentes. 

A shrub, up to 2 meters high; stems terete, glabrous; leaf blades 
ovate-oblong to oblong, up to 24 cm. long and 10 cm. wide, acuminate 
(the tip blunt), narrowed at the base, thin, crenate, glabrous, with 
numerous minute cystoliths; petioles up to 8 cm. long, glabrous; 



1224 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

flowers borne in lax spikes up to 8 cm. long, these forming large, 
terminal panicles up to 30 cm. long, the branches of the panicles 
and the rachises puberulent; bracts oblong-ovate, 5 mm. long and 
3 mm. wide, rounded at the apex, sparingly glandular-pubescent, or 
the lowermost larger, leaflike, and glabrous, all but the uppermost 
soon deciduous; bractlets similar but smaller; calyx segments linear, 
about 7 mm. long, less than 1 mm. wide, sparingly puberulent, 
minutely ciliolate, sometimes with scattered, glandular hairs up to 
0.25 mm. long; corollas 4.5-6 cm. long, finely pubescent, red and 
yellow, the tube slender, 1-3.5 mm. in diameter at the base, narrowed 
above the ovary, 4-6 mm. at the mouth, the lips narrow, 6 mm. wide 
or less, up to 2 cm. long, gradually narrowed from base to tip, the 
lobes about 0.5 mm. long, rounded; anther lobes unequally attached, 
about 3 mm. long, the lower subcalcarate; capsules 1.5 cm. long, 
clavate, glabrous or with a few glandular hairs near the tip. La 
Conception, Llanuras de Santa Clara, 250 meters, February, 1896, 
Donnell Smith 6691 (type in U. S. Nat. Herb., No. 355,229). Buena 
Vista, road to San Carlos Valley, 600 meters, Cook & Doyle 148. 
Wet forests in the vicinity of Guapiles, Prov. Limon, 300-500 meters, 
Standley 37286, 37354. Wet forest, Pejivalle, Prov. Cartago, 900 
meters, Standley & Valeria 46991, 47107. Cataratas de San Ramon, 
Brenes 4422, 13412. Forests between Rio Chirripo and Rio Zent, 
2,500 meters, Pittier 16052. Rio de Las Vueltas, 635 meters, Tonduz 
12887. Rio Hondo, 50 meters, Pittier 16642. Zarcero, A. Smith 
A222, H39. Easily distinguished among Costa Rican species by 
its large, crenate leaves, its large, terminal panicles of short spikes, 
and its rounded bracts. 

Jacobinia macrantha (Benth.) Hemsl. Pata de gallo, Saca- 
tinta, Azul. Justicia macrantha Benth. Cyrtanthera macrantha 
Nees. Cyrtantherella macrantha Oerst. Moist forests of Central 
America. Slender shrubs, up to 4 meters high, often resting on 
other plants; stems subtetragonal, pilose at the nodes or glabrous, 
rarely densely pilose; leaf blades ovate to oblong, up to 20 cm. long 
and 6 cm. wide, acuminate (the tip blunt), narrowed at the base, 
thin, glabrous; petioles up to 2 cm. long, glabrous or pilose above; 
flowers borne in axillary cymes; peduncles (up to 4 cm. long) and 
pedicels glabrous; bracts and bractlets subulate, up to 2 mm. long, 
glabrous; calyx 6-7 mm. long, the segments lance-subulate, ciliolate; 
corollas pale yellowish or orange-red, up to 6.5 cm. long, glabrous, 
the tube slender, 1.5 mm. in diameter at the base, 5 mm. at the mouth, 
the lips about 2.5 cm. long, the posterior lip acute and bidentate, 



FLORA OF COSTA RICA 1225 

provided with 2 medial wings enclosing a portion of the style, the 
anterior lip spreading, about 1 cm. wide, the lobes about 2 mm. long, 
rounded; stamens exserted from the mouth of corolla, the anthers 
curved, converging at the tip and diverging at the base. The 
flowers are said to resemble the open mouth of a serpent. 

Jacobinia tinctoria (Oerst.) Hemsl. Biol. Centr. Amer. Bot. 
2: 522. 1882. Azul, Sacatinta, Azul de mata. Sericographis tinctoria 
Oerst. Vid. Medd. Kjoebenhavn 1854: 150. 1855. Cartago, Oersted. 
Dry thickets of Central America. Glabrous shrubs, up to 2 meters 
high; stems terete; leaf blades lanceolate to oblong-lanceolate, up to 
8 cm. long and 2.5 cm. wide, acuminate (the tip blunt), narrowed 
at the base; petioles up to 1 cm. long; flowers secund, borne in lax, 
curved, glabrous, axillary spikes up to 5 cm. long; bracts and bractlets 
triangular-subulate, up to 2 mm. long and 1 mm. wide at the base; 
calyx segments triangular-subulate, 1.5-2.5 mm. long, about 1 mm. 
wide at the base, ciliolate; corolla 3-3.5 cm. long, red, the tube slender, 
1.5 mm. in diameter at the base, 4 mm. at the mouth, the lips oblong, 
1.5 cm. long, 5 mm. wide, the upper acute, minutely bidentate, the 
lobes of the lower lip about 1 mm. long, rounded, the middle one 
2 mm. wide, the lateral ones 1.5 mm. wide; anther lobes 3 mm. long, 
unequally and obliquely attached, the upper acute, the lower sub- 
calcarate at the base. The leaves, treated with hot water, give a 
blue solution used by laundresses to give a blue tint to white clothes. 
This color changes to red with acids like litmus (Standley). Pittier 
reports the Brunka name as Tebi-ka. 

Jacobinia umbrosa (Benth.) Blake. Pav6n amarillo, Pavdn, 
Pavoncillo. Synonyms of J. umbrosa are Justicia aurea Schlecht. 
and Jacobinia aurea Hemsl., non Hiern. Damp woods, southern 
Mexico and Central America. Often cultivated in gardens. A shrub 
or small tree, up to 5 meters high or more; branches puberulent or 
glabrous; leaf blades lance-oblong to broadly ovate, 10-45 cm. long, 
up to 12 cm. wide or more, acute or acuminate, abruptly decurrent 
at the base, glabrous or puberulent; petioles 5-8 cm. long; inflo- 
rescence a many-flowered thyrse 7-30 cm. long, the rachises pu- 
berulent; bracts linear-lanceolate, 1-2 cm. long, about 2 mm. wide, 
1-nerved, puberulent; calyx segments 5, lanceolate, 7 mm. long, 1.5 
mm. wide, puberulent, 3-nerved; corolla 4.5-5 cm. long, yellow or 
reddish yellow, puberulent or pilosulous, the lips 2.5 cm. long, the 
upper one narrow, erect, entire, the lower 3-lobed, the lobes 2 mm. 
long, rounded; anthers 3 mm. long, the cells slightly unequal, ab- 



1226 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ruptly acute at the base, the tip minute; capsules 16-17 mm. long, 
clavate, puberulent. 

JUSTICIA L. 

Herbs or shrubs; leaves petiolate, usually ovate, entire; flowers 
spicate, paniculate, or solitary; bracts small, or occasionally large 
and imbricate; calyx segments narrow, nearly equal; corollas usually 
purple, pink, or white, sometimes with dark purple markings, the 
tube usually short, the throat ampliate, the upper lip 2-lobed, the 
lower 3-lobed; stamens 2; anther cells 2, unequally attached, acute or 
calcarate; staminodes none; ovules 2 in each cavity; capsules oblong 
to obovate. A large, cosmopolitan genus. 

Leaves of a pair unequal J. trichotoma. 

Leaves of a pair equal. 

Calyx segments 4; leaf blades metallic blue beneath. . J. metallica. 
Calyx segments 5. 

Calyx segments unequal, the posterior one much smaller than 
the others. 

Corolla 2.5 cm. long or more J. Oerstedii. 

Corolla 1 cm. long or less. 
Bracts oblong or oblong-lanceolate. 
Leaves firm ; stem densely yellowish-pilose . . . J. Skutchii. 
Leaves thin; stems finely pubescent in 2 lines. 

J. angustibracteata. 
Bracts elliptic or ovate. 

Stems pilose in 2 lines J. Pittieri. 

Stems not pilose in 2 lines, the hairs more or less evenly 

distributed. 
Hairs of the stem spreading, up to 1 mm. long. 

J. candelariae. 

Hairs of the stem curved, minute J. Valerii. 

Calyx segments equal (the posterior one slightly smaller than 
the others in J. pectoralis). 

Bracts and bractlets narrowly spatulate J. costaricana. 

Bracts and bractlets various, not narrowly spatulate. 

Corollas 25-30 mm. long J. carthaginensis. 

Corollas 20 mm. long or less. 

Bracts subulate, minute, 3 mm. long or less. 

Corollas 8-15 mm. long J. pectoralis. 



FLORA OF COSTA RICA 1227 

Corollas about 6 mm. long J. comata. 

Bracts lanceolate to ovate, mostly more than 4 mm. long. 
Leaf blades pilose, the hairs up to 3 mm. long. 

J. Tonduzii. 
Leaf blades glabrous or minutely and sparingly pilose 

beneath. 
Leaf blades rather small, 7 cm. long or less. 

J. parvibracteata. 
Leaf blades large, 20 cm. long or more. 

Bracts ovate, ciliate J. refractifolia. 

Bracts lanceolate, eciliate J. refulgens. 

Justicia angustibracteata Leonard, nom. nov. Rhytiglossa 
microphylla Oerst. Vid. Medd. Kjoebenhavn 1854: 159. 1855, non 
Justicia microphylla Lam. 1791. Mountains of Candelaria, 1,800- 
2,100 meters, Oersted. Wet forests of the upper mountain slopes. 
Endemic. Herbs, up to 60 cm. high; stems slender, branched, 
ascending, subquadrangular, finely pubescent in 2 lines; leaf blades 
oblong-ovate to lanceolate, up to 11 cm. long and 3 cm. wide, obtuse 
to acuminate, the tip blunt, narrowed at the base, thin, sparingly 
hirtellous above, more densely so beneath; petioles up to 1 cm. long, 
hirtellous; flowers borne in terminal, peduncled, solitary or paired 
spikes (one much smaller than the other) up to 8 cm. long and 1 cm. 
in diameter, the rachis and peduncle densely hirtellous; bracts oblong 
to oblanceolate, up to 1 cm. long and 2 mm. wide, acute to obtusish 
at the apex, hirtellous and ciliate; bractlets up to 1 cm. long and 0.5 
mm. wide, hirtellous and ciliate; calyx 6-7 mm. long, the segments 5, 
subulate, ciliate, the posterior segment much smaller than the others; 
corollas 1 cm. long, 1.5 mm. wide at the base, 2.5 mm. at the 
mouth, puberulent, white(?); capsules clavate, 8 mm. long, sparingly 
puberulent. 

Justicia candelariae (Oerst.) Leonard, Carnegie Inst. Wash. 
Publ. 461: 231. 1936. Rhytiglossa candelariae Oerst. Vid. Medd. 
Kjoebenhavn 1854: 158. 1855. Dianthera candelariae Hemsl. Biol. 
Centr. Amer. Bot. 2: 517. 1882. Mountains of Candelaria, 1,800- 
2,100 meters, Oersted. Wet, wooded mountain slopes (usually above 
600 meters). Herbs, up to 45 cm. high; stems branching, decumbent, 
terete, hirsute, the hairs about 1 mm. long; leaf blades ovate to oblong, 
up to 7 cm. long and 3 cm. wide, but usually much smaller, obtuse to 
acute, with blunt tip, narrowed at the base, hirsute, paler beneath; 



1228 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

petioles up to 1 cm. long, pilose; flowers borne in terminal, solitary or 
paired, bracted spikes (one spike much smaller than the other) up to 
4 cm. long and 1 cm. in diameter, the rachis pilose, the bracts im- 
bricate, elliptic, up to 1 cm. long and 6 mm. wide, rounded and 
mucronulate at the apex, rounded or narrowed at the base, sub- 
membranaceous, nitid, subreticulate, hirsute and ciliate; bractlets 
lanceolate, about 8 mm. long and 2 mm. wide, in other respects 
similar to the bracts; lateral and anterior calyx segments linear, 6 
mm. long and 0.5 mm. wide, the posterior one subulate, 4 mm. long, 
all ciliate; corollas 8 mm. long, white or tinged with violet, puberulent 
toward the tip, 2 mm. in diameter at the base, 2.5 mm. at the mouth, 
the upper lip 4 mm. long, the lower 5 mm.; capsules 8 mm. long, 
clavate, puberulent. In the publication of the original transfer the 
author was erroneously given as Hemsley. 

JUvSticia carthaginensis Jacq. Thickets in both moist and 
dry situations. Found throughout the West Indies and tropical 
continental America. Herbaceous or suffrutescent, up to 1.5 meters 
high; stems erect, branched, pilosulous in 2 lines or glabrous; leaf 
blades ovate or elliptic-ovate, 5-12 cm. long, acuminate, narrowed or 
rounded at the base and decurrent on the petiole, glabrous or pilosu- 
lous; petioles 0.5-2 cm. long; flowers borne in dense, terminal, bracted 
spikes 3-7 cm. long; bracts oblong, 15 mm. long or less, about 5 mm. 
wide, acute or obtuse, the bractlets similar but narrower, all pilosu- 
lous, some of the hairs glandular; calyx segments 5, linear-lanceolate, 
10-11 mm. long, white-margined, ciliolate toward the tip; corollas 
purple, 2.5-3 cm. long, sparingly glandular-pubescent, the tube about 
1.5 cm. long, the upper lip straight, about as long as the tube, notched 
at the apex, the lower lip spreading, slightly longer than the upper, 
3-lobed, the lobes elliptic, about 1 cm. long, rounded; capsules 1.5-2 
cm. long, finely puberulent. 

Justicia cornata (L.) Lam. Damp thickets and woods, usually 
on banks of streams. West Indies and tropical continental America. 
An herb, up to 20 cm. high; stems weak, ascending or nearly erect, 
usually branched, often rooting at the lower nodes, glabrous or spar- 
ingly pilosulous; leaves sessile or petioled, the blades lanceolate to 
oblong-ovate or oblong, 3-15 cm. long, acuminate or acute, rounded 
or narrowed at the base; petioles, when present, up to 2 cm. long; 
flowers borne in axillary and terminal, peduncled panicles up to 22 
cm. long, the branches nearly filiform, 2-7 cm. long, often forming 
verticillasters, more or less glandular-pubescent; bracts and bractlets 



FLORA OF COSTA RICA 1229 

subulate, 3 mm. long or less; calyx segments 5, subulate, 1.5-2 mm. 
long; corollas white or purplish, about 6 mm. long, the tube relatively 
broad, 2 mm. long, the upper lip erect, 3 mm. long, emarginate, the 
lower lip 4-5 mm. long, 3-lobed, the lobes rounded, 1 mm. long; 
capsules 4-5 mm. long, stipitate, puberulent, sometimes glabrous 
when mature. 

Justicia costaricana Leonard, sp. nov. Suffrutescens, caulibus 
parce ramosis, subteretibus, glabris vel apice bifariam pubescentibus; 
lamina foliorum oblongo-ovata, acuminata, basi angustata, glabra, 
perspicue cystolithigera; petioli glabri; spicae axillares, solitariae 
vel geminae, simplices vel interdum basi ramosae, rhachibus et 
pedunculis glanduloso-puberulentis; bracteae spathulatae, unguicu- 
latae, parce glanduloso-puberulentae; bracteolae anguste lineari- 
spathulatae, glanduloso-puberulentae, saepe patentes; calycis seg- 
menta lanceolato-subulata, puberulenta; corolla virescenti-flava, 
fauce atrorubra, tubo basi ventrali bisaccato; antherae loculi oblique 
et inaequaliter affixi, inferior breviter calcaratus. 

Suffrutescent, up to 1.3 meters high; stems sparingly branched, 
sub terete, glabrous or the tips pubescent in 2 lines; leaf blades oblong- 
ovate, up to 13 cm. long, and 4.5 cm. wide, acuminate (the tip blunt), 
narrowed at the base, glabrous, the cystoliths numerous and con- 
spicuous; petioles up to 2 cm. long, glabrous; flowers borne in axillary 
spikes up to 3 cm. long, or occasionally longer, these solitary, paired, 
simple or sometimes branched at the base, the rachises and peduncles 
(5-10 mm. long) glandular-puberulent (sometimes in 2 lines) ; bracts 
spatulate, 1 cm. long, 3-5 mm. wide, unguiculate, sparingly glandular- 
puberulent; bractlets linear-spatulate, 1 cm. long, 0.5 mm. wide, 
glandular-puberulent; bracts and bractlets usually spreading; calyx 
segments 5, lance-subulate, 5.5 mm. long, 1.5 mm. wide at the base, 
puberulent; corollas 10 mm. long, greenish white, tinged with dark 
red in the throat, the tube short, 1.5 mm. in diameter at the base, 
enlarged to 3 mm. and narrowed again above the ovary, about 4 mm. 
at the mouth, the ventral basal portion of the tube with 2 saccate 
projections, the lips 4.5 mm. long, the upper one about 5 mm. wide 
at the base, bidentate at the apex, the lobes of the lower lip rounded, 
the middle one 3 mm. wide, the lateral 2 mm. wide; anthers ob- 
liquely and unequally attached, the upper apiculate at the base, the 
lower short-spurred (0.5 mm. long). Moist forests of El Muneco, 
Rio Navarro, Prov. Cartago, 1,400-1,500 meters, March 6-7, 1926, 
Standley & Torres 51257 (type in U. S. Nat. Herb., No. 1,306,691), 
50960, 50976, 51237; Standley 33429, 33453. Cataratas de San 



1230 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Ramon, Brenes 13488. Wet forests of La Hondura, Prov. San Jose", 
1,300-1,700 meters, Standley 36539, 37735, 37956. La Brisa de 
Zarcero, A. Smith H396. Easily recognized by its spreading, spatu- 
late bracts and greenish white, red-throated, saccate corollas. 

Justicia metallica Lindau in Pittier, Prim. 2: 311. 1900. 
Forests of El General, 600 meters, Pittier 3497. Found in the forests 
of El General, Terraba, and Canas Gordas. Also in the Changuinola 
Valley, Panama. Herbs, up to 40 cm. high; stems ascending, rooting 
at the lower nodes, subquadrangular, pilosulous in 2 lines, purplish; 
leaf blades ovate, up to 8 cm. long and 4 cm. wide, more or less 
acuminate (the tip blunt), rounded at the base and slightly decurrent 
on the petiole, glabrous or the costa and veins hirsute with curved 
hairs, the lower surface of a shining, metallic purple; petioles up to 
7 mm. long, hirsutulous above; flowers secund in terminal or axillary, 
glandular-pilosulous panicles composed of lax spikes; bracts and 
bractlets minute; calyx segments 4 and 4 mm. long, 1 mm. wide; 
corollas purple(?), finely pubescent, the tube slender, 8 mm. long, 
the upper lip 4.5 mm. long, minutely 2-dentate at the apex, the lower 
lip 4 mm. long, the lobes small, obtuse; capsules 8 mm. long, glabrous, 
slender-stipitate. 

Justicia Oerstedii Leonard, nom. nov. Chiloglossa glabra Oerst. 
Vid. Medd. Kjoebenhavn 1854: 161. 1855. Dianthera glabra Hemsl. 
Biol. Centr. Amer. Bot. 2: 518. 1882. Justicia glabra Lindau in 
Pittier, Prim. 2: 312. 1900, non Koen. ex Roxb., 1820. Candelaria, 
1,500 meters, Oersted. Found in damp thickets and forests, usually 
above 500 meters; also in the Chiriqui region of Panama. Slender, 
suffrutescent plants, rarely over 2 meters high, erect or suberect, the 
branches terete, retrorsely hirsutulous in 2 lines; leaf blades oblong 
to lanceolate, up to 7 cm. long and 3 cm. wide (occasionally larger), 
obtuse to acuminate, narrowed at the base, glabrous; petioles up to 
1.5 cm. long but usually much shorter; flowers borne in lax, terminal 
panicles, the branches glandular-puberulent; bracts and bractlets 
minute, subulate; calyx up to 6 mm. long, the segments linear, 
puberulent, the posterior one much smaller than the others; corolla 
bright red, up to 3 cm. long, finely and sparingly pubescent, the tube 
narrow, 1 mm. in diameter at the base, gradually enlarged to 5 mm. 
at the mouth, the upper lip oblong, 1 cm. long, 5 mm. wide, minutely 
bidentate at the apex, the lower lip slightly longer than the upper, 
the middle lobe 2.5 mm. wide, the lateral one 1.5 mm. wide, all 
3 mm. long and rounded at the apex; stamens barely reaching the 



FLORA OF COSTA RICA 1231 

tip of the lips, the connective about 1 mm. broad; anther cells 
attached one slightly above the other, the lower one rounded, 
acutish at the base; ovary glabrous. 

Justicia parvibracteata Leonard, sp. nov. Herbae erectae vel 
adscendentes, caulibus teretibus, bifariam retrorso-pubescentibus; 
lamina foliorum oblongo-lanceolata, acuminata, basi angustata, 
glabra; petioli glabri; inflorescentia spicata vel paniculata, rhachibus 
et pedunculis patente pubescentibus, interdum glanduliferis, floribus 
2-4 in axillis bractearum; bracteae oblongae, obtusae, basi angus- 
tatae, chartaceae, glabrae, venosae; bracteolae lanceolatae, glabrae 
vel parce pubescentes; calycis segmenta lanceolata, minute pubes- 
centia; corolla alba et purpurea, parce pubescens, tubo subcylindrico, 
labiis subaequalibus, superiore oblongo-ovato, acuto et minute 
bidentato, inferiore trilobo, lobis rotundatis; antherae loculi subae- 
qualiter affixi, inferior basi acutiusculus. 

Erect or ascending herbs up to 1 meter high; stems terete, 
retrorsely pubescent in 2 lines; leaf blades oblong-lanceolate, up to 
7 cm. long and 2.5 cm. wide, acuminate (the tip blunt), narrowed 
at the base, glabrous, with cystoliths; petioles up to 5 mm. long, 
glabrous; inflorescence spicate or subpaniculate, up to 10 cm. long, 
the rachises and peduncles (up to 3 cm. long) spreading-pubescent, 
the hairs about 0.5 mm. long, some of them gland-tipped, the inter- 
nodes about 5 mm. long; flowers 2-4 in the axils of the bracts; pedicels 
1 mm. long; bracts oblong, 4 mm. long, 1 mm. wide or slightly wider, 
obtuse at the apex, narrowed at the base, firm, glabrous, the veins 
prominent; bractlets lanceolate, 1-2.5 mm. long, 0.5 mm. wide, 
glabrous or sparingly pubescent, the hairs barely 0.5 mm. long, some 
of them gland-tipped; calyx about 3 mm. long, minutely pubescent, 
the segments 5, lanceolate, about 0.5 mm. wide, subequal; corollas 
7 mm. long, subcylindric, 1.5 mm. in diameter, barely expanded at 
the mouth, white and purple, sparingly pubescent, the lips subequal, 
2.5 mm. long, the upper lip oblong-ovate, 1.5 mm. wide, acute and 
minutely bidentate at the apex, the lower lip 3 mm. wide, with 
3 rounded lobes 1.25 mm. in diameter; anther cells subequally at- 
tached, the lowermost slightly larger than the upper and acutish 
at the base; capsules clavate, 7 mm. long, puberulent, the hairs of 
the stipe retrorse. Wet forests in the vicinity of Pejivalle, Prov. 
Cartago, 900 meters, February, 1926, Standky & Valeria 47001 (type 
in U. S. Nat. Herb., No. 1,254,955), 46926. Closely related to 
J. Pittieri, but with comparatively narrow bracts, strictly glabrous 



1232 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

leaf blades, and with glandular, spreading hairs, evenly distributed 
on the rachis and peduncle. 

Justicia pectoralis Jacq. Waste and cultivated regions in 
continental tropical America and West Indies. An herb, up to 60 cm. 
high; stems erect, decumbent, or ascending, sparingly branched, 
often rooting at the lower nodes, short-pilose in 2 lines below, 
puberulent and more or less glandular above; leaf blades lanceolate 
to ovate-lanceolate, 2-10 cm. long, acuminate, acute or obtuse at 
the base, glabrous; petioles 2-12 mm. long; inflorescence a terminal, 
few-several-branched panicle, the branches slender, alternate, 2-12 
cm. long, more or less glandular, the flowers subdistant; bracts and 
bractlets subulate, up to 3 mm. long; calyx segments 5, subulate, 
about 2 mm. long, the posterior segment slightly shorter than the 
others; corollas pink, puberulent, 8-15 mm. long, the tube 5 mm. long, 
the upper lip straight, about 4 mm. long, minutely 2-lobed at the 
tip, the lower lip slightly longer, 7 mm. broad, the lobes rounded, 
1.5-2 mm. long, purple, white-striped toward the throat. 

Justicia Pittieri Lindau in Pittier, Prim. 2: 312. 1900. Known 
only from the type, collected at Tskuinak, upper Zhorquin, Tala- 
manca, Pittier 864.2. An ascending herb, rooting at the lower nodes; 
stems terete, pilose in 2 lines; leaf blades oblong-lanceolate, 7-10 cm. 
long, 2-2.5 cm. wide, acuminate, the tip itself blunt, narrowed at 
the base, glabrous or sparingly pilose beneath, the cystoliths numer- 
ous and conspicuous; petioles 5-8 mm. long, pubescent above, the 
hairs usually curved; spikes simple, terminal, about 5 cm. long, the 
rachis and peduncle (2 cm. long) pubescent in 2 lines; flowers fascicled 
in the axils of the bracts, 4-8 at each node, the internodes of the 
spikes about 5 mm. long; bracts ovate or the lowermost oblong, 
about 5 mm. long and 2 mm. wide, obtuse at the apex, narrowed 
and stipitate at the base, ciliate, pilosulous, some of the hairs gland- 
ular; bractlets lanceolate, 2.5 mm. long and 1 mm. wide, puberulent; 
calyx 3-4 mm. long, the anterior segment minute and setulose, the 
other 4 narrowly lanceolate, 0.5 mm. wide at the base, sparingly and 
minutely pubescent; corollas sparingly pubescent, purple(?), the tube 
5 mm. long and about 1.5 mm. in diameter, the lips 3 mm. long, 
the upper one 1.5 mm. wide, minutely bidentate at the apex, the 
3 lobes of the lower lip about 1 mm. in diameter, obtuse. 

Justicia refractifolia (Kuntze) Leonard, comb. nov. Ecbolium 
refractifolium Kuntze, Rev. Gen. 2: 487. 1891. Justicia pseudopolysta- 
chya Cuf. Archivio Bot. 10: 49. 1934 (Finca Waldeck, Rio Madre de 



FLORA OF COSTA RICA 1233 

Dios and Rio Barbilla, 20-40 meters, Cufodontis 349 and 652). Costa 
Rica, at 300 meters, Kuntze. Near sea level, in wet forests of Costa 
Rica and Panama. Herbs, up to 1 meter high; stems simple, sub- 
terete, strigose (the hairs about 0.5 mm. long), or the upper portions 
hirsute with somewhat longer, spreading hairs; leaf blades oblong- 
lanceolate, up to 25 cm. long and 10 cm. wide, acute to acuminate 
(the tip itself usually blunt), narrowed at the base, not at all decur- 
rent, glabrous above, more or less pilose beneath; petioles about 5 mm. 
long, strigillose; spikes axillary, solitary or paired (one much smaller 
than the other), up to about 10 cm. long and 1 cm. in diameter; 
bracts ovate, up to 10 mm. long and 6 mm. wide, acute at the apex, 
obtusish at the base, glabrous above, sparingly puberulent beneath, 
ciliate, drying dark brown, the veins prominent, the sterile basal 
bracts similar but narrower; bractlets lanceolate-falcate, 8 mm. long 
and about 2 mm. wide, in other respects similar to the bracts; calyx 
segments 5, subulate, up to 7 mm. long, ciliolate; corollas 10-12 mm. 
long, purple, finely pubescent; lower anther cells spurred; capsules 
clavate, 8 mm. long, puberulent, 4-seeded; seeds muricate. 

Justicia refulgens Leonard, sp. nov. Frutex, caulibus sub- 
quadrangularibus, glabris vel apice pilosis; lamina foliorum oblonga, 
acuminata, basi angustata, chartacea, laete viridis, nitens, subtus 
pallidior, glabra, costa et venis subtus tenuiter et sparse pilosis 
exceptis; petioli pubescentes; spicae axillares, trifurcatae, peduncu- 
latae, pedunculis glabris vel appresso-pilosis, rhachibus et pedunculis 
secundariis brunneo-tomentosis; spicae bracteae lineares, floriferae 
lanceolatae, appresso-pubescentes vel subtomentosae; calycis seg- 
menta linearia, acuminata, tomentosa; corolla tenuiter pubescens, 
alba, labiis lilacino-tinctis; antherae loculi inaequaliter affixi, inferior 
minute calcaratus; capsulae clavatae, glabrae; semina plana, brunnea, 
papillosa. 

Shrubs, up to 1 meter high; stems subquadrangular, glabrous or 
the tips appressed-pilose; leaf blades oblong, up to 25 cm. long and 
6 cm. wide, acuminate, narrowed at the base, rather firm, bright 
shining green, darker above than beneath, glabrous except the costa 
and veins beneath, these finely and sparingly pilose; petioles up to 
4 cm. long, pubescent; flowers borne in trifurcate, axillary, peduncled 
spikes up to 6 cm. long, the peduncles up to 4 cm. long, subquad- 
rangular, glabrous or appressed-pilose, the rachises and secondary 
peduncles (1-1.5 cm. long) brownish-tomentose, the lower internodes 
about 1 cm. long, the others successively shorter toward the tip; 
bracts subtending the spikes linear, about 2 cm. long and 1.5 cm. 



1234 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

wide, those subtending the flowers lanceolate, 6 mm. long, 1.5-2.5 
mm. wide at the base, appressed-pubescent or subtomentose; bract- 
lets similar but narrower; calyx segments 5, linear, 8 mm. long, 1.5 
mm. wide, acuminate, tomentose; corollas about 18 mm. long, 6 mm. 
in diameter at the mouth, finely pubescent, white, the lips tinted 
lavender, 8 mm. long; anther lobes unequally attached, the lower 
with a minute spur about 0.25 mm. long at the base; capsules clavate, 
2.5 cm. long, 5 mm. wide, glabrous; seeds flat, brown, 4 mm. long, 
3.5 mm. wide, papillose. Between Rio Ladrillo and Los Siguas 
Camp, southern slope of Cerro de La Horqueta, Chiriqui, Panama, 
1,200-1,700 meters, March 18, 1911, Pittier 3272 (type in U. S. Nat. 
Herb., No. 677,663). Rain forests of Bajo Chorro, Boquete District, 
Chiriqui, Panama, Davidson 77. Well marked by its firm, large, 
shining, green leaves and its trifurcate, tomentose spikes. Because 
of the similarity of the Chiriqui Province of Panama to the adjacent 
region of Costa Rica, it is reasonable to expect an extension of the 
range of this species to Costa Rica. 

Justicia Skutchii Leonard, sp. nov. Frutex gracilis, caulibus 
dense flavescenti-pilosis; lamina foliorum oblongo-ovata, acuta, brevi- 
ter acuminata, vel interdum obtusa, basi rotundata vel subcordata, 
chartacea, dense flavescenti-pilosa, costa et venis perspicuis; petioli 
dense pilosi; spicae terminales et axillares, paniculatae, floribus 
fasciculatis, sessilibus, rhachibus et pedunculis dense flavescenti- 
pubescentibus, pilis nonnullis glanduliferis; bracteae anguste ob- 
longae, apice obtusae, basi angustatae, supra glabrae, subtus parce 
pubescentes, venosae; bracteolae lineares, parce hirsutulae; calycis 
segmentum anticum setulosum, cetera anguste lanceolata, omnia 
minute pubescentia; corolla tenuiter pubescens, alba et purpurea, 
labio superiore anguste ovato, acuto, inferiore trilobo, lobis rotun- 
datis; antherae loculi subaequaliter affixi, connectivo lato; capsulae 
clavatae, pubescentes, 4-spermae; semina papillosa. 

Slender shrubs, up to 1 meter high; stems densely yellowish- 
pilose, the hairs up to 2 mm. long; leaf blades oblong-ovate, up to 
10 cm. long and 4 cm. wide, acute, short-acuminate, or occasionally 
obtuse at the apex, rounded or subcordate at the base, firm, both 
surfaces densely yellowish-pilose, the costa and veins prominent 
beneath, the cystoliths conspicuous; petioles about 2 mm. long, 
densely yellowish-pilose; spikes terminal and axillary, up to 12 cm. 
long, forming a large, terminal panicle; flowers borne in fascicles of 
3-8 (or the uppermost solitary), sessile in the axils of the bracts, the 
internodes about 5 mm. long, the rachises and peduncles (up to 3 cm. 



FLORA OF COSTA RICA 1235 

long) densely yellowish-pubescent, the hairs about 0.25 mm. long, 
some of them glandular; bracts narrowly oblong, 6 mm. long and 2 
mm. wide, blunt at the apex, narrowed at the base, sparingly hirsute 
beneath with yellowish hairs 0.5 mm. long, glabrous above, the costa 
and veins prominent; bractlets linear, about 3 mm. long, 0.5 mm. 
wide, sparingly hirsutulous; anterior calyx segment setulose, about 
2.5 mm. long, the other 4 narrowly lanceolate, 4.5 mm. long, 0.75 
mm. wide at the base, minutely pubescent; corollas 8 mm. long, white 
and purple, finely and rather sparingly pubescent, the tube 1.5 mm. 
in diameter at the base, 2.5 mm. at the mouth, the lips 4 mm. long, 
the upper one narrowly ovate, acute, 2 mm. wide near the base, the 
lower 4 mm. wide, its lobes rounded, barely 0.5 mm. long, 1.2-2 mm. 
wide; anther lobes subequally attached, the connective broad; cap- 
sules 8 mm. long, clavate, finely pubescent, 4-seeded, the seeds 
papillose. On rocks beside a forest stream, vicinity of El General, 
Prov. San Jose*, 915 meters, December, 1936, Skutch 3012 (type in 
U. S. Nat. Herb., No. 1,643,087). Possibly related to J. Tonduzii, 
but readily distinguishable by its larger size, dense and yellowish 
pubescence, larger and more densely flowered spikes, and thicker 
leaf blades. 

Justicia Tonduzii Lindau in Pittier, Prim. 2: 313. 1900. Type 
from the forests of Rio Naranjo, Tonduz 7976. Found also in the 
forests of Victoria Farm, Plains of Zent, Pittier 16073, and at La 
Colombiana, Prov. Limon, Standley 36928. Endemic. An erect 
herb, up to 30 cm. high; stems terete, the pubescence a mixture of 
small, curved hairs, arranged in 2 lines, and long, spreading ones 
(up to 3 mm. long) evenly distributed; leaf blades ovate, up to 9 cm. 
long and 4 cm. wide, short-acuminate (the tip blunt), narrowed or 
obtuse at the base, both surfaces pilose, the hairs up to 3 mm. long, 
or subglabrous at maturity; petioles about 5 mm. long, the pubescence 
a mixture of long, spreading hairs and short, curved ones; spikes 
terminal or axillary, slender, up to 7 cm. long, the flowers borne in 
fascicles of 3 or 4 in the axils of the bracts, the internodes up to 6 mm. 
long, the rachis and peduncle (up to 3 cm. long) pubescent with a 
mixture of short, curved and long, spreading hairs, some of them 
glandular; bracts ovate, stipitate, up to 5 mm. long and 2 mm. wide, 
the margins and costa with long, spreading hairs; bractlets narrowly 
oblong, up to 2 mm. long and 0.5 mm. wide, pilose; calyx 2.5 mm. 
long, the segments 5, subulate, pubescent, some hairs glandular; 
corolla 7-8 mm. long, purplish, sparingly and finely pubescent, the 
lips about 4 mm. long, the upper one minutely 2-lobed at the tip, 



1236 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the lobes of the lower lip rounded; anther lobes subequally attached, 
the lower smaller than the upper; capsules clavate, about 5 mm. long, 
pubescent. 

Justicia trichotoma (Kuntze) Leonard, comb. nov. Ecbolium 
trichotomum Kuntze, Rev. Gen. 2: 488. 1891. J. asymetrica Lindau 
in Pittier, Prim. 2: 310. 1900 (Chilamate, bank of Rio Sarapiqui, Biol- 
ley 7383). Limon, Kuntze. At low altitudes in wet forests and 
thickets. Honduras and Panama. Herbs or small shrubs, up to 1 
meter high; branches subquadrangular, pubescent, the hairs small, 
curved, the lower portions of the stems becoming subterete and 
glabrous; leaf blades ovate to ovate-lanceolate, one blade of each 
pair conspicuously smaller than the other, the larger blades up to 
10 cm. long and 4 cm. wide, the smaller up to 5 cm. long and 2 cm. 
wide, all long-acuminate (the tip itself blunt), narrowed at the base, 
the costa and veins pubescent, the hairs minute, curved, the inter- 
costal spaces glabrous or sparingly pilose; petioles up to 2 cm. long, 
slender, puberulent; flowers borne in small, few-flowered, axillary 
cymes, the peduncles up to 4 cm. long, puberulent, the flowers sessile 
on the branches; bracts small, linear; calyx segments 7-8 mm. long, 
less than 1 mm. wide; corollas pale purple, pubescent, the tube sub- 
cylindric, 10 mm. long and about 1.5 mm. in diameter, the upper lip 

5 mm. long and 4 mm. wide, minutely bifid at the apex, the lower lip 

6 mm. long, its 3 lobes 5 mm. long, the middle one 2.5 mm. wide, 
the lateral ones 1.5 mm. wide, all obtuse; capsules 14 mm. long, about 
2.5 mm. wide, puberulent, 4-seeded, stipitate; seeds 1.5 mm. in 
diameter, verruculose. 

Justicia Valerii Leonard, sp. nov. Herbae decumbentes, cauli- 
bus gracilibus, subteretibus, tenuiter retrorse pubescentibus; lamina 
foliorum ovata ad oblonga, acuta vel acuminata, basi angustata, 
supra glabra, subtus minute puberulenta vel subglabra, perspicue 
cystolithigera; petioli appresso-pubescentes; spicae terminales, soli- 
tariae vel gemmae, pedunculis et rhachibus pubescentibus, pilis 
parvis, recurvatis, albidis, bracteis inferioribus lanceolatis, acutis vel 
acutiusculis, ceteris ellipticis, obtusis, vel rotundatis et obtuse apicu- 
latis, basi abrupte angustatis, pallide viridibus, viridi-reticulatis, 
imbricatis, subglabris, ciliatis; bracteolae oblanceolatae, acutae, 
ciliatae; calycis segmenta tenuiter pubescentia, posticum minutum, 
setaceum, cetera lineari-subulata; corolla tenuiter pubescens, alba 
vel purpurascens, intus purpureo-maculata, labio superiore angusto, 
erecto, apice minute bidentato, inferiore patente, trilobo, lobis 



FLORA OF COSTA RICA 1237 

rotundatis; antherae loculi subaequaliter affixi, connective lato; 
capsulae clavatae, puberulentae. 

Decumbent herbs, up to 30 cm. high; stems slender, subterete, 
puberulent or finely retrorse-pubescent; leaf blades ovate to oblong, 
up to 7 cm. long and 3 cm. wide, but usually smaller, acute to acumi- 
nate (the tip blunt), narrowed at the base, glabrous and dark gray- 
green, or with a blue-green, metallic tint above, paler and minutely 
pubescent or subglabrous beneath, the cystoliths conspicuous; peti- 
oles up to 1 cm. long, minutely appressed-pubescent; flowers borne 
in terminal, solitary or paired, bracted spikes up to 5 cm. long and 
1 cm. in diameter; peduncles (up to 3.5 cm. long) and rachises finely 
pubescent, the hairs white and recurved, the lower pair of bracts 
lanceolate, up to 1 cm. long and 3 mm. wide, acute or obtusish, the 
others elliptic, 6-7 mm. long and about 4 mm. wide, obtuse or 
rounded at the apex, bluntly apiculate, rather abruptly narrowed at 
the base, all pale green with dark green, coarse reticulation, imbricate, 
subglabrous, ciliate; bractlets oblanceolate, 6 mm. long, 1.5 mm. wide, 
acute, ciliate; calyx sparingly and finely pubescent, the posterior 
segment setaceous, 3 mm. long, the other 4 linear-subulate, 5 mm. 
long and 0.5 mm. wide; corollas finely pubescent, white or purplish 
white with purple dots within, 9 mm. long, 2 mm. in diameter at the 
base, somewhat constricted above the ovary, 2.5 mm. wide at the 
mouth, the lips about 4 mm. long, the upper one narrow, erect, 
minutely bidentate at the apex, the lower one spreading, 4.6 mm. 
wide across the 3 rounded lobes; anther lobes subequally attached, 
the connective broad; capsules clavate, 7 mm. long, 2 mm. broad, 
puberulent. Moist forest of Quebrada Serena, southeast of Tilaran, 
Guanacaste, 700 meters, January 27, 1926, Standley & Valeria 46147 
(type in U. S. Nat. Herb., No. 1,254,478), 46154, 46178, 46212. 
Wooded stream bank at Dulce Nombre, Prov. Cartago, 1,400 meters, 
Standley 35945. Wet forests of El Muneco, south of Navarro, Prov. 
Cartago, 1,400 meters, Standley 33454. Moist forests of El Silencio, 
Guanacaste, near Tilaran, 750 meters, Standley & Valeria 44731. 
El Silencio de San Ramon, Brenes 17100. El Rosario de Orosi, 1,120 
meters, Pittier 16614. Wet forest in the vicinity of Fraijanes, 1,500- 
1,700 meters, Standley & Torres 47522. La Palma de San Ramon, 
Brenes 15061. La Pifia de Zarcero, A. Smith H305. Wet forest of 
Los Ayotes, near Tilaran, Guanacaste, 600-700 meters, Standley 
& Valeria 45543. Moist forest of Naranjos Agrios, Guanacaste, 
600-700 meters, Standley & Valeria 46456. Piedades Norte de San 
Ramon, Brenes 6717. In a clearing at Vara Blanca de Sarapiqui, 



1238 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

north slope of Central Cordillera, Skutch 3448. Zarcero, A. Smith 
H52. Without definite locality, Brenes 3698. Closely related to J. 
candelariae (Oerst.) Leonard, but easily separated from that by the 
pubescence of the stems. In J. candelariae the hairs are rather 
coarse and spreading, but in this new species they are fine and 
retrorsely curved. 

LEPIDAGATHIS Willd. 

Herbs or low shrubs; leaves entire; flowers borne in dense spikes; 
calyx unequally 5-parted; corolla tube slender, subcylindric, the limb 
2-lipped, the upper lip 2-cleft or emarginate, the lower 3-lobed; 
stamens 4, didynamous, the filaments short, the anther cells blunt; 
style entire; capsules oblong, pointed, 4-seeded (sometimes fewer); 
seeds flat, nearly orbicular. There are about 60 species, mostly of 
the Old World tropics. 

Lepidagathis alopecuroidea (Vahl) Griseb. Wet, shaded banks 
in woods, thickets, and fields. Central America to northern South 
America and West Indies. An herb; stems up to 50 cm. long, usually 
branched, erect, decumbent, or ascending, sometimes rooting at the 
lower nodes, short-pilose; leaf blades thin, ovate to elliptic, 3-8 cm. 
long, 2-3 cm. wide, acute or obtuse, narrowed at the base, repand 
or entire; petioles up to 2 cm. long; spikes usually terminal, 3-8 cm. 
long, 1-1.5 cm. in diameter, many-flowered; bracts lanceolate to 
oblong, about 6 mm. long, venose, ciliate; calyx segments unequal, 
4-6 mm. long, the anterior pair lanceolate, connate at the base, the 
lateral pair linear, the posterior segment obovate; corollas white or 
violet, about as long as the calyx; stamens free at the base; anther 
sacs parallel, slightly unequal; capsules sessile, glabrous, about 
4 mm. long. 

LOUTERIDIUM S. Wats. 

Shrubs; leaves petiolate, crenate or entire; flowers borne in narrow, 
nearly naked, terminal panicles or racemes; bracts small; calyx 
segments 3; corolla large, red or yellowish, the tube abruptly ex- 
panded above the ovary, broad and hood-shaped above, the lobes 5, 
short, subequal; stamens 2 or 4, exserted, the anther cells parallel; 
capsules sessile, the locules 6-8-seeded. Several species are found 
in Mexico and Central America. 

Louteridium costaricense Radlk. & Bonn. Smith, Bot. Gaz. 
37: 422. 1904. Known only from the type, collected in the forests 
of Las Vueltas, Tucurrique, Tonduz 8123. Glabrous shrubs; leaf 
blades oblong-elliptic, 28-31 cm. long, abruptly acuminate, narrowed 









FLORA OF COSTA RICA 1239 



at the base and decurrent on the petiole, membranous, entire; 
petioles up to 6 cm. long; flowers borne in a narrow panicle, the 
peduncle about 15 cm. long; bracts and bractlets ovate-lanceolate, 
6-9 mm. long; pedicels 2-3.5 cm. long; calyx segments oblong, 2-4 
cm. long, acute, reticulate-veined; corolla yellow; stamens 2. 

MENDONCIA Veil. 

Twining, herbaceous or suffrutescent vines; leaf blades usually 
ovate, elliptic, or oblong, petioled; flowers pediceled, one to several 
in the axils of the leaves, each flower subtended by a pair of con- 
spicuous bracts; corolla usually red or white, 5-lobed, the tube narrow; 
stamens 4, paired; anther lobes diverging, bearded; calyx usually 
reduced to a minute ring; fruit drupaceous, usually dry and com- 
pressed at the tip. The genus was named for Cardinal Mendonca, 
Patriarch of Lisbon. Although numerous in tropical South America, 
only five species are known at present in Central America. 

Corolla 1.5 cm. long M. Brenesii. 

Corolla 3 cm. long or more. 

Stems strigose or glabrous; bracts less than twice as long as wide. 
Stems narrowly winged; bracts oblong, retuse at the apex. 

M. retusa. 
Stems quadrangular but not winged; bracts obovate, rounded 

or obtuse at the apex M . costaricana. 

Stems pilose; bracts more than twice as long as wide. 
Bracts oblong-lanceolate, acuminate at the apex . . . M. Lindavii. 
Bracts oblong-elliptic, rounded at the apex M. Tonduzii. 

Mendoncia Brenesii Standl. & Leonard, sp. nov. Frutex 
volubilis, caulibus teretibus, glabris vel sparse et minute strigosis; 
lamina foliorum ovata, abrupte acuminata, mucronata, basi rotun- 
data vel angustata, chartacea, glabra vel novella minute et sparse 
strigosa; petioli glabri; flores fasciculati; bracteae ovatae, obtusae, 
sparse et minute strigosae; corolla alba, parva, glabra, lobis obcor- 
datis; stamina inclusa. 

Scandent, the stems subterete, grooved, glabrous or sparingly 
and minutely strigose; leaf blades ovate, up to 11 cm. long and 
6 cm. wide, abruptly acuminate and tipped by a mucro about 2 mm. 
long, rounded or narrowed at the base, firm, glabrous or the upper- 
most minutely and sparingly strigose; petioles up to 3 cm. long, 
glabrous; flowers several, radiating from short, flattened, rounded, 
axillary spurs; pedicels about 1 cm. long, sparingly and minutely 



1240 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

strigose; bracts ovate, 10-14 mm. long, 6-7 mm. wide, obtuse, 
sparingly and minutely strigose; corollas 1.5 cm. long, white, glabrous, 
the tube about 8 mm. long and 3 mm. in diameter, slightly expanded 
at the base, the lobes obcordate, about 7 mm. in diameter; stamens 
included, the filaments 1.5 mm. long; fruit not seen. La Palma de 
San Ramon, August 26, 192S,Brenes 6269 (type in Herb. Field Mus., 
No. 855,431), 4406, 5631. This is quite unlike any other Mendoncia 
found in Central America. It has axillary umbels of small, white 
flowers. 

Mendoncia costaricana Oerst. Vid. Medd. Kjoebenhavn 
1854: 113. 1855. Cerro de Aguacate, Oersted; collected by Oersted 
also at Quebrada Honda, 540 meters, and south of Cartago. 
Found occasionally in wet forests. Also seen in Panama. Large, 
herbaceous vines; stems obscurely quadrangular, slightly grooved, 
strigose; leaf blades ovate-elliptic 5-10 cm. long, 1.5-5 cm. wide, 
acuminate and sometimes falcate at the apex, tipped by a mucro 
1-2 mm. long, rounded or narrowed at the base and slightly decurrent 
on the petiole, thin, hirtellous and barely scabrous above (the hairs 
arising from star-shaped bases), softly pubescent beneath; petioles 
15-19 mm. long, strigose; flowers 1 or 2 in the axils of the upper 
leaves; peduncles 1-1.5 cm. long, strigose; bracts obovate, 12-15 
mm. long, 10-11 mm. wide, obtuse or rounded and apiculate at the 
apex, subcordate at the base, strigose; corollas white, the throat 
with dark purple spots, glabrous, 3 cm. long, the tube 8 mm. broad 
at the throat, the lobes orbicular, 4-5 mm. wide, emarginate; ovary 
glabrous. 

Mendoncia Lindavii Rusby. Occasional in wet forests of the 
lower mountain slopes (up to 900 meters). Its range extends from 
British Honduras and Guatemala as far south as Bolivia. A large, 
densely ferruginous-pilose vine; leaf blades ovate to broadly oval, 
5-10 cm. long, 2-5.5 cm. wide, acuminate and sometimes mucronate 
at the apex, narrowed or rounded at the base; petioles up to 1.5 cm. 
long; flowers single or in pairs; pedicels 2-3 cm. long; bracts oblong- 
lanceolate, subfalcate, 3-4 cm. long, 1 cm. wide, acuminate and 
usually mucronate at the apex, partly connate; corollas red, pink, 
or white, 4-5 cm. long, tubular, glabrous, the lobes rounded, 4-5 mm. 
long, entire; fruit ovoid, compressed, keeled, about 2 cm. long and 
1 cm. wide, brown-puberulent. 

Mendoncia retusa Turrill. Common in moist forests. Also 
in other parts of Central America. Stem conspicuously quadrangular, 



FLORA OF COSTA RICA 1241 

narrowly winged, glabrous or sparingly pilose toward the tip; leaf 
blades elliptic, 5-12 cm. long, 3-7 cm. wide, abruptly acuminate 
and mucronulate at the apex, rounded or narrowed at the base, 
entire, glabrous, or the costa and lateral veins sparingly pilose; 
flowers solitary, the pedicels 2-3 cm. long; bracts oblong, 1-2.5 cm. 
long, 1.5 cm. wide, rounded at both ends, retuse and apiculate at 
the apex, glabrous; corolla white with a purple throat, glabrous, 
4-5 cm. long, the lobes broadly obovate, spreading, 5-10 mm. 
long; fruit black, minutely puberulent or glabrescent, ovoid, about 
1.5 cm. long. 

Mendoncia Tonduzii Turrill, Kew Bull. 413. 1919. Tucurrique, 
Tonduz 12946. Rarely found except at the type locality. Endemic. 
A suffrutescent vine; stems quadrangular, hirsute; leaf blades elliptic, 
up to 13 cm. long and 8 cm. wide, acuminate, narrowed at the base, 
pilose and subscabrous above (the hairs arising from star-shaped 
bases), rather densely and softly pilose with yellowish, curved hairs 
below; petioles 2-3 cm. long, hirsute; flowers solitary or in pairs 
in the axils of the upper leaves; pedicels 2-3 cm. long, densely 
yellowish-hirsute; bracts oblong-elliptic, up to 3.5 cm. long and 
1.5 cm. wide, rounded at both ends, apiculate at the apex, the costa 
prominent, densely and softly yellowish-hirsute without, glabrous 
within; corolla about 5 cm. long, white, glabrous, about 8 mm. 
broad at the oblique throat, the lobes ovate, spreading, about 8 mm. 
long and 5-6 mm. wide; fruit oblong, slightly compressed, 2 cm. long, 
12-14 mm. wide, doubly keeled when dry, puberulent. 

NELSONIA R. Br. 

A monotypic genus, ranging from Mexico to northern South 
America and the West Indies; also in the Old World tropics. 

Nelsonia brunelloides (Lam.) Kuntze. Common in dry 
thickets or forests. A diffuse, softly villous herb; leaves entire; 
flowers borne in terminal and axillary, bracted spikes; calyx 4-lobed, 
the posterior segment 2-toothed or 2-cleft; corolla blue or purple, 
the tube slender, the limb 2-lipped, the upper lip 2-cleft, the lower 
3-lobed ; stamens 2, the filaments short, the anther sacs mucronulate 
at the base; ovules several in each cavity; stigma 2-lobed; capsules 
oblong, beaked; seeds pubescent with peculiar, anchor-shaped hairs. 

ODONTONEMA Nees 

Herbs or shrubs; leaves usually large, entire; flowers borne in 
narrow, terminal panicles; calyx 5-parted, the segments small and 



1242 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

narrow; corollas white, red, or yellow, the tube slender, the throat 
only slightly enlarged, the limb 2-lipped, nearly regular; stamens 2, 
the anther sacs blunt at the base; staminodes 2; capsules oblong, 
stipitate; seeds 4 or fewer. A genus of 30 or more species, confined 
chiefly to tropical America. 

Rachis tomentose or subtomentose 0. callistachyum. 

Rachis glabrous or sparingly puberulent. 

Inflorescence gradually diminished to a flagellate, flowerless tip. 

0. flagellum. 

Inflorescence flowering to the tip 0. strictum. 

Odontonema callistachyum (Schlecht. & Cham.) Kuntze. 
Thickets, southern Mexico and Central America. Suffrutescent, 
2-4.5 meters high; leaves lance-oblong to elliptic-ovate, 10-30 cm. 
long, acuminate, the uppermost sessile, the lower petiolate; inflores- 
cence an interrupted, racemiform or paniculate thyrse, the flowers 
borne in fascicles, these subtended by triangular, cuspidate bracts 
about 4 mm. long, the rachis and pedicels tomentose or subtomentose; 
calyx segments subulate, about 2 mm. long, ciliolate; corolla pink 
or red, glabrous, 2-3 cm. long, 5 mm. broad at the throat, the lips 
4-5 mm. long, the upper 2-lobed (the lobes about 3 mm. long), 
the lower 3-lobed nearly to the base, all the lobes rounded and 
ciliolate near the tip; staminodes 0.5-1 mm. long; capsules 2 cm. long, 
narrow and solid from middle to base, glabrous. 

Odontonema flagellum (Oerst.) Kuntze, Rev. Gen. 2: 494. 
1891. Thyrsacanthus flagellum Oerst. Vid. Medd. Kjoebenhavn 
1854: 146. 1855. Turrialba, Oersted. Forests. Endemic. Suffrutes- 
cent, up to 2 meters high; stems glabrous; leaf blades oblong-elliptic 
to lanceolate, up to 30 cm. long and 11 cm. wide, acuminate, ending 
in a blunt and sometimes curved tip, narrowed at the base to a short, 
winged petiole, glabrous or minutely and inconspicuously puberulent 
beneath; inflorescence a slender raceme, simple, or branched at the 
base, and ending in a slender, flagellate, sterile tip; flowers 6 or more 
(sometimes fewer toward the tip), crowded in opposite clusters 
usually about 1 cm. distant on the rachis and subtended by triangular 
bracts 3.5 mm. long and 1.5 mm. wide at the base; bractlets similar 
but smaller; pedicels red, slender, up to 7 mm. long, these and the 
rachis and bracts minutely and sparingly hirsutulous; calyx 3-4.5 
mm. long, red, essentially glabrous, minutely ciliolate, the segments 
narrowly triangular, 2-3 mm. long, 0.5-0.75 mm. wide at the base; 
corolla 2-2.5 cm. long, bright red, glabrous, the lower portion of the 



FLORA OF COSTA RICA 1243 

tube slender, about 1.5 mm. in diameter, to about 5 mm. above the 
base, thence rather abruptly enlarged to 4-5 mm., the lobes rounded 
or obtuse, minutely ciliolate; staminodes truncate, about 0.75 mm. 
long; capsules 2 cm. long, glabrous, long-stipitate. 

Odontonema strictum (Nees) Kuntze. Forests of Central 
America. Suffrutescent, 1-2 meters high, glabrous; leaves up to 
35 cm. long and 12 cm. wide, acuminate, narrowed at the base to a 
short petiole; inflorescence a simple or branched raceme up to 30 cm. 
long or more, the rachis minutely puberulent; flowers borne in pseudo- 
whorls, the pedicels short, subtended by a keeled, acuminate outer 
bract (1 mm. wide at the base) and several minute, subulate inner 
bracts; calyx 3 mm. long, the segments subulate, purplish red; 
corolla rich scarlet, tubular, or slightly funnelform, the lips about 
6 mm. long, the upper one 2-lobed at the tip, the lower 3-lobed nearly 
to the base, the lobes ovate, obtuse, minutely ciliolate; fertile stamens 
reaching the notch in the upper lip; staminodes about 0.5 mm. long, 
subulate; style shorter than the corolla tube. 

POIKILACANTHUS Lindau 

Herbs or shrubs; flowers borne in small, terminal or axillary, 
bracted spikes or heads; calyx 5-parted; corolla slender, 2-lipped; 
stamens 2, the anthers unequally attached, the lobes rounded or 
ending in a short spur; staminodes none; pollen elliptic, marked 
with hexagonal plates ("facettierter Pollen"). Several species are 
found in South America. 

Poikilacanthus macranthus Lindau. Wet forests of Costa 
Rica, Guatemala, and Nicaragua. Shrubs up to 2 meters high; 
stems terete, glabrate or the tips yellowish-subtomentose; leaf 
blades oblong-ovate, 2.5-9 cm. long, 1.5-4.5 cm. wide, acuminate, 
the tip blunt, narrowed at the base, subglabrous above, the nerves 
and margins of the lower surface pilosulous, the hairs curved, 
appressed or ascending; petioles 7-15 mm. long, subtomentose; 
flowers borne in short, terminal, sessile heads, each head with 5-6 
flowers; bracts oblong, up to 9 mm. long and 2 mm. wide, hirsutulous; 
bractlets similar but smaller; calyx segments lanceolate, about 5 mm. 
long and 1.5-2 mm. wide, ciliate; corolla light pink or lavender, 
sparingly pubescent (the hairs about 0.5 mm. long, some of them 
gland-tipped), the tube 3.5 cm. long, 3 mm. in diameter at the base, 
gradually enlarged to 6 mm. at the mouth, the upper lip 3 cm. long, 
10 mm. wide at the base, bilobed at the tip, the lobes 1 mm. long, 
the lower lip 24 mm. long, 5 mm. wide at the base, 11 mm. at the 



1244 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

middle, the middle lobe 1.5-2 mm. long, obtusish, the lateral lobes 
1.5 mm. long, 3 mm. wide, acutish; stamens reaching to the tips of 
the lips; anther lobes obtuse, unequally attached; style about 7 cm. 
long; capsules about 1.5 cm. long, pilosulous, the basal half about 
4 mm. broad, solid, the upper half ovate, 4-seeded; seeds flat, black 
to light brown, minutely roughened. Distinct from Beloperonides 
macrantha Oerst., erroneously treated as a synonym by Lindau in 
Pittier, Prim. 2: 307. 1900. 

PSEUDERANTHEMUM Radlk. 

Herbs or small shrubs; leaves usually ovate; flowers borne in 
terminal or axillary, bracteate spikes, the bracts and bractlets small ; 
calyx segments 4 or 5, subulate; corolla white, blue, or pink, the 
tube slender, the limb spreading, the 5 lobes subequal; stamens 2, 
included; staminodes 2, minute; anther cells 2, muticous or acute 
at the base; capsules long-stipitate; seeds 2 or 4. A genus frequently 
found in tropical regions throughout the world. Several species 
other than those here described have been found in Central America. 

Calyx puberulent, some of the hairs gland-tipped. 

Flowers crowded in opposite fascicles or in whorls, and borne in a 
narrow, terminal raceme; leaf blades, or some of them, more 
than 9.5 cm. long P. Pittieri. 

Flowers mostly alternate and solitary, borne in a much branched 
panicle of slender, lax spikes; leaf blades 9.5 cm. long or less. 

P. Standleyi. 
Calyx essentially glabrous, eglandular. 

Leaf blades thin, green, the petioles long and conspicuously winged ; 
flowers sessile or subsessile; native species P. cuspidatum. 

Leaf blades firm, often purple, the petioles short and not conspicu- 
ously winged; pedicels up to 6 mm. long; cultivated species. 

P. atropurpureum. 

Pseuderanthemum atropurpureum (Bull) Bailey. Eran- 
themum atropurpureum Bull. Widely planted in tropical regions of 
America. A shrub, up to 2 meters high; stem glabrous; leaf blades 
ovate, up to 12 cm. long and 5 cm. wide, or occasionally larger, short- 
acuminate or obtuse and apiculate at the apex, narrowed at the base, 
purple, entire, glabrous, both surfaces bearing numerous minute, 
subpunctiform cystoliths; petioles short; flowers numerous, crowded 
in subsessile fascicles forming narrow, terminal, simple or sparingly 
branched racemes, the rachis and pedicels (up to 6 mm. long) usually 



FLORA OF COSTA RICA 1245 

glabrous; bracts oblong, about 5 mm. long, ciliolate; calyx segments 
narrowly triangular, ciliolate; corollas 15-20 mm. long, glabrous, 
purple, or white with purple throat, the limb 15-20 mm. broad, the 
segments ovate, rounded. 

Pseuderanthemum cuspidatum (Nees) Radlk. Eranthemum 
cuspidatum Nees. River banks and borders of woods. Southern 
Mexico, Central America, and northern South America. Herbs, 
30-50 cm. high or more; stems glabrous or sparingly hirtellous, the 
hairs scattered or confined to 2 lines; leaf blades ovate to elliptic, 
variable in size, up to 17 cm. long and 9 cm. wide, acuminate (the 
tip blunt), rounded or narrowed at the base and decurrent on the 
petiole, thin, green, sparingly hirtellous; petioles up to 13 cm. long, 
winged, about 1 cm. broad at the base of the blade; flowers sessile or 
subsessile, solitary or several in a cluster, borne in lax, terminal 
spikes, these sometimes forming a sparingly branched panicle, the 
rachis sparingly and minutely hirtellous, the lowermost pair of bracts 
or those subtending the branches of a paniculate inflorescence sub- 
orbicular, up to 3 cm. long and 2.5 cm. wide, mucronate at the apex, 
the bracts subtending the flowers narrowly triangular, glabrous, or 
ciliate toward the base; bractlets about 1 mm. long; calyx 4-5 mm. 
long, glabrous, ciliolate, the segments 5, subulate; corolla about 3 
cm. long, violet or purple, glabrous, the tube very slender, scarcely 
enlarged to the throat, the limb about 1 cm. broad, the segments 
ovate or elliptic, 8 mm. long; capsules 12-14 mm. long, clavate, 
glabrous, 4-seeded, pointed at the apex, the lower half slender and 
solid; seeds flat, roughened. 

Pseuderanthemum Pittieri Leonard, sp. nov. Herba vel 
suffrutex, caulibus glabris vel apice minute pubescentibus; lamina 
foliorum elliptica, breviter acuminata, basi angustata, glabra vel 
subtus in costa et venis lateralibus sparse puberulenta; petioli puberu- 
lenti; racemi laxi, angusti, pedunculo longo, pilosulo, floribus soli- 
tariis vel laxe fasciculatis, pedicellis et rhachibus glanduloso-puberu- 
lentis; bracteae subulatae, minute pubescentes; calycis segmenta 
subulata, plus minusve glanduloso-pubescentia; corolla minute et 
sparse pubescens, tubo gracili, lobis ovatis. 

Herbaceous or suffrutescent; stems glabrous or minutely pubes- 
cent toward the tip, with minute, subpunctiform cystoliths; leaf 
blades elliptic, up to 14 cm. long and 8 cm. wide, short-acuminate, 
narrowed at the base, glabrous, or the costa and lateral veins beneath 
sparingly puberulent, with slender, inconspicuous cystoliths above; 



1246 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

petioles up to 2 cm. long, puberulent; flowers opposite, usually several 
in a fascicle, borne in loose, narrow, terminal racemes, the peduncles 
up to 12 cm. long, with several pairs of small, sterile bracts, pilosulous, 
the pedicels up to 1.5 mm. long, these and the rachises glandular- 
puberulent; flower bracts up to 3 mm. long, subulate, minutely 
pubescent; bractlets similar but shorter; calyx segments subulate, 
2-3.5 mm. long, minutely puberulent with glandular and eglandular 
hairs; corollas 2 cm. long, minutely and sparingly pubescent, the 
tube of fully expanded flowers about 12 mm. long, 1 mm. in diameter 
at the base, 3 mm. at the mouth, the limb about 2 cm. broad, the 
lobes ovate; capsules not seen. Along Rio Hondo, 50 meters, 
February 15, 1903, Pittier 16641 (type in U. S. Nat. Herb., No. 
1,080,220). Related to P. adenocalyx Lindau, of Guatemala, but 
readily separated from that species by the shape and size of its leaf 
blades and its much larger flowers. In P. adenocalyx the leaf blades 
are oblong-lanceolate and are not more than 7 cm. long and 2 cm. 
wide, and the corolla tube is very slender (0.5 mm. wide at the base 
and 1 mm. at the mouth). The lobes are likewise much smaller in 
that. From the herbarium material it is impossible to determine the 
height of the present plant or the color of the flowers. 

Pseuderanthemum Standleyi Leonard, sp. nov. Herbae, 
caulibus bifariam pilosulis; folia petiolata; lamina ovata ad lanceo- 
lata, acuta vel acuminata, basi angustata vel rotundata, in petiolum 
decurrens, membranacea, supra sparse hispidula, subtus in costa et 
venis hirtella; racemi spiciformes laxi, graciles, in paniculam termi- 
nalem aggregati, floribus solitariis, saepe alternis, rhachibus sparse 
glanduloso-puberulentis; bracteae et bracteolae subulatae, carinatae, 
glabrae vel ciliatae, interdum pilos paucos glanduliferos gerentes; 
pedicelli glanduloso-puberulenti; calycis segmenta subulata, plus 
minusve glanduloso-puberulenta; corolla roseo-purpurea, tenuiter 
et sparse pubescens, tubo gracillimo, lobis ellipticis; capsulae clavatae, 
glabrae; semina plana, pallide brunnea, rugulosa. 

Herbs, up to 1 meter high; stems pilosulous in 2 lines; leaf 
blades ovate to lanceolate, up to 9.5 cm. long and 5 cm. wide, acute 
to acuminate (blunt at the tip), narrowed or rounded at the base 
and decurrent on the petiole, thin, sparingly hispidulous above, the 
costa and veins hirtellous beneath; petioles up to 3 cm. long; flowers 
solitary, mostly alternate, borne in a panicle of slender, lax, spike- 
like racemes, the rachises rather sparingly glandular-puberulent; 
bracts and bractlets subulate, about 2 mm. long, carinate, glabrous 
or ciliolate (sometimes with a few minute, glandular hairs) ; pedicels 



FLORA OF COSTA RICA 1247 

up to 3 mm. long, glandular-puberulent; calyx 5-6 mm. long, puberu- 
lent, some of the hairs gland-tipped, the segments linear-subulate; 
corollas bright rose-purple, 1.5 cm. long, sparingly and minutely 
pubescent, the tube very slender, 0.5-1 mm. in diameter, the upper 
part oblique, enlarged to 1 mm. in diameter, the limb about 1 cm. 
broad, the lobes elliptic, about 3 mm. wide; staminodes slightly 
longer than the filaments of the fertile stamens; capsules 10-15 mm. 
long, clavate, glabrous, 4-seeded; seeds flat, light brown, oblique, 
4 mm. long and 3 mm. wide, roughened. Moist thickets along Rio 
Reventado, north of Cartago, 1,460-1,650 meters, February 26, 
1926, Standley & Valerio 49481 (type in U. S. Nat. Herb., No. 
1,306,060). Vicinity of Santa Maria de Dota, Prov. San Jose", 
1,500-1,800 meters, Standley 41714, 41725, 42538. Oak forest near 
Quebradillas, 7 km. north of Santa Maria de Dota, Prov. San Jose", 
1,800 meters, Standley 42912. Unlike any other Central American 
species because of its relatively small, thin leaves, usually much 
branched panicles, peduncled, alternate, solitary flowers, and slender, 
linear, finely puberulent calyx segments. 

RAZISEA Oerst. 

Shrubs with large, ovate or oblong-elliptic, petioled leaves bearing 
minute cystoliths; flowers borne in a peduncled, terminal thyrse; 
bracts small; calyx segments 5, subequal, narrow; corollas red, the 
tube narrowly infundibuliform, subcurved, gradually enlarged from 
base to throat, the upper lip narrow, acute, minutely bidentate, the 
lower lip broad, blunt, with 3 small, suborbicular lobes at the tip; 
stamens 2, long-exserted ; anthers oblong, 1-celled; capsules slender, 
stipitate, 4-seeded, the seeds roughened. A single species, apparently 
limited to Costa Rica and Panama. 

Razisea spicata Oerst. Vid. Medd. Kjoebenhavn 1854: 142. 
1855. Coral. Kolobochilus leiorhachis Lindau in Pittier, Prim. 2: 
308. 1900 ("Costa Rica et Veragua," Warscewicz). Cerro de 
Aguacate, 450 meters, Oersted. Wet mountain forests of Costa Rica 
and Panama. A shrub, up to 3 meters high; stems quadrangular, 
pilose or subtomentose, becoming glabrous; leaf blades ovate to 
oblong-ovate or elliptic, up to 35 cm. long and 20 cm. wide, acuminate, 
often caudate, narrowed at the base, glabrous, or the veins minutely 
pubescent beneath; petioles up to 3 cm. long, glabrous; inflorescence 
a spiciform, pedunculate, terminal thyrse up to 30 cm. long, the 
rachis minutely pubescent, becoming glabrous, the flowers borne in 
opposite fascicles, these usually from 3-20 mm. distant; pedicels 



1248 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

slender, up to 10 mm. long, glabrous; bracts triangular, about 5 mm. 
long and 2 mm. wide at the base, glabrous; bractlets similar but 
smaller; calyx glabrous, the segments linear, 7-8 mm. long, 1 mm. 
wide at the base; corolla about 4 cm. long, bright red, glabrous, the 
tube subcylindric, gradually enlarged from 2 mm. at the base to 
5-7 mm. at the mouth, the upper lip lance-ovate, 8 mm. long, 3-4 
mm. wide toward the base, ending at the apex in a slender, minutely 
bidentate tip, the lower lip short, subtruncate, 9-12 mm. broad, 
divided at the tip into 3 rounded lobes about 1 mm. in diameter; 
filaments about 4 cm. long, the anthers 5 mm. long; ovary glabrous; 
capsules about 17 mm. long, glabrous; seeds roughened. 

RUELLIA L. 

Perennial herbs or shrubs; leaves petioled, entire or rarely 
dentate; flowers usually large and showy, solitary or clustered in the 
axils, or borne in terminal, cymose panicles; calyx usually 5-parted, 
the segments often narrow; corollas red, yellow, white, or purple 
(usually mauve), funnelform or salverform, sometimes saccate, the 
lower portion of the tube usually narrow, the limb with 5 obtuse, 
spreading lobes; stamens 4, didynamous, the anther sacs blunt at 
the base; stigma lobes unequal; capsules oblong, cylindric, or clavate; 
seeds flat, usually pubescent, mucilaginous when moistened. A 
large, cosmopolitan genus, found in both tropical and temperate 
regions. 

Flowers sessile (or some of them short-peduncled), borne in the axils 
of the leaves or in a terminal head. 

Corolla white or yellow; flowers borne on a terminal, conical rachis. 

R. tetrastichantha. 
Corolla mauve or purplish; flowers axillary. 

Leaf blades large, at least some of them 10 cm. long or more, 

acuminate. 
Corolla 4-5 cm. long, the limb 3 cm. broad; leaves dull. 

R. achimeniflora. 

Corolla 3 cm. long or less, the limb 1-2 cm. broad; leaves 
lustrous. 

Capsules puberulent; corolla 2.5 cm. long R. metallica. 

Capsules glabrous; corolla 3 cm. long R. barbillana. 

Leaf blades small, 5 cm. long or less, obtuse or acutish. 

R. geminiflora. 



FLORA OF COSTA RICA 1249 

Flowers or flower clusters peduncled. 

Ovary and capsule glabrous (the calluses at the tip sometimes 

more or less pubescent). 
Calyx with glandular hairs. 

Corollas large, 5-6 cm. long R. malacosperma. 

Corollas small, 3 cm. long or less. 

Capsules cylindric R- paniculata. 

Capsules clavate, flattened R- inundata. 

Calyx with eglandular hairs, or subglabrous. 

Flowers secund at the ends of long, horizontal or slightly 
ascending peduncles; corollas 3-4 cm. long. 

R. stemonacanthoides. 

Flowers not secund; corollas 2.5 cm. long or less. 

Peduncles 10-22 cm. long; leaf blades up to 21 cm. long. 

R. Pittieri. 

Peduncles up to 9 cm. long; leaf blades up to 10 cm. long. 

R. Tonduzii. 

Ovary and capsule puberulent. 

Flowers fascicled at the ends of the long, sharply angled pedun- 
cles; corollas 2 cm. long . . R- Biolleyi. 
Flowers solitary at the end of the peduncle or borne in dichoto- 

mously branched cymes; corollas 3 cm. long or more. 
Leaf blades obtuse or rounded at the apex, grayish-puberulent. 

R. nudiflora var. puberula. 

Leaf blades acuminate, not gray-puberulent. 

Corollas usually lavender, 4 cm. long or more. 

R. jussieuvides. 

Corollas white, 2.5 cm. long or less. . R- Cooperi. 

Ruellia achimeniflora (Oerst.) Hemsl. Biol. Centr. Amer. Bot. 
2- 503 1882. Arrhostoxylum achimeniflorum Oerst. Vid. Medd. 
Kjoebenhavn 1854: 131. 1855. Cerro de Aguacate, Oersted Wet 
thickets, chiefly in the vicinity of Cartago and San Jose\ 
Herbs, up to 60 cm. high; stems simple or sparingly branched, quad- 
rangular, sharply angled or narrowly winged, minutely pilosulous, 
the hairs confined chiefly to the angles and nodes; leaf blades oblonj 
ovate, 4-14 cm. long, 1-7 cm. wide, acuminate (the tip blunt), acute 
or obtuse at the base and decurrent on the petiole, pilosulous (often 
sparingly so), the hairs confined chiefly to the costa and lateral veins, 



1250 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

these prominent, both surfaces with numerous delicate cystoliths; 
flowers solitary or in 3's, subsessile or borne on short, axillary pedun- 
cles crowded at the tips of the branches; bracts oblong-lanceolate, 
2-3 cm. long, 3-5 mm. wide; bractlets linear-spatulate, 1 cm. long, 
1-2 mm. wide; calyx 9 mm. long, the segments lanceolate, slightly | 
unequal, 6-7 mm. long, 1 mm. wide at the base, ciliolate, the midrib 
prominent; corollas 4-5 cm. long, rose-purple, finely pubescent or 
subtomentose, the tube very slender, curved, 5 mm. broad at the 
mouth, the limb 3 cm. broad, the lobes obcordate; capsules 1.5-1.6 
cm. long, 5 mm. broad, clavate, acutish at the tip, narrowed to a 
solid base 6 mm. long and 2.5 mm. broad, puberulent, 12-seeded; 
seeds brown, about 3 mm. in diameter, glabrous. 

Ruellia barbillana Cufodontis, Archivio Bot. 10: 47. 1934. 
The type was collected near Finca Waldeck on the bank of Rio 
Barbilla, Cufodontis. The following description is adapted from the 
original. Plants up to 40 cm. high; stem quadrangular, bearing 
numerous punctiform cystoliths, the angles retrorsely hirsute; leaf 
blades ovate-lanceolate, up to 10 cm. long and 2.5 cm. wide, long- 
attenuate at the base, abruptly acuminate at the apex, subrepand or 
undulate, glabrous or the costa and nerves appressed-hirsute below, 
both surfaces bearing cystoliths; petioles 1-1.5 cm. long; flowers 
sessile, borne in glomerulate cymes in the axils of the upper leaves; 
bracts and bractlets small ; calyx 5 mm. long, the segments subulate, 
glabrous, with cystoliths; corollas lilac, glabrous, about 3 cm. long, 
2 mm. in diameter at the base, 6 mm. at the mouth, the limb spread- 
ing, the lobes ovate, rounded, about 8 mm. long and 6 mm. wide; 
stamens included; capsules glabrous, 6-seeded. 

Ruellia Biolleyi Lindau in Pittier, Prim. 2: 301. 1900. Junction 
of Rio Puerto Viejo and Rio Sarapiqui, Biolley 7396. Wet forests of 
Costa Rica and Panama. Stem usually erect, quadrangular (the 
angles sharp or narrowly winged), sparingly pilosulous; leaf blades 
oblong-ovate to oblong-elliptic, 10-17 cm. long, 4-6 cm. wide, nar- 
rowed at each end (the tip blunt), membranous, glabrous except the 
costa and veins below, these pubescent; petioles up to 3 cm. long; 
flowers borne in fascicles terminating sharp-angled, axillary peduncles 
up to 20 cm. long; bracts and bractlets small, lanceolate; calyx 4-6 
mm. long, the segments linear-subulate, minutely but rather sparingly 
glandular-pilosulous (occasionally eglandular); corollas blue(?), 
about 2 cm. long, sparingly puberulent, the lower portion of the tube 
1 mm. in diameter, the throat about 6 mm. broad, the lobes rounded, 



FLORA OF COSTA RICA 1251 

about 4 mm. wide; capsule 12 mm. long, 3 mm. broad, pointed at the 
tip, narrowed to a solid basal portion 6 mm. long and 1.5 mm. broad, 
glandular-puberulent, 4-seeded; seeds flat, ovate. 

Ruellia campestris (Oerst.) Hemsl. Biol. Centr. Amer. Bot. 
2: 504. 1882. Gymnacanthus campestris Oerst. Vid. Medd. Kjoeben- 
havn 1854: 126. 1855. Type collected at Hacienda Santa Rosa, 
Guanacaste, Oersted. Probably a small-leaved form of R. geminiflora 
HBK. 

Ruellia Cooperi Leonard, sp. nov. Herba vel suffrutex, caulibus 
quadrangularibus, glabris vel ad nodos parce puberulentis, angulis 
anguste alatis; lamina foliorum oblonga, breviter acuminata, basi 
angustata, supra glabra, subtus in costa et venis lateralibus pilis 
minutis brunnescentibus curvatis instructa; petioli pubescentes; 
paniculae laxae, axillares, dichotomo-ramosae, longe pedunculatae, 
pedunculis gracilibus, glabris, acute angulatis, floribus solitariis vel 
geminis, pedicellis glabris vel pilos paucos patentes glanduliferos 
gerentibus; calycis segmenta subulata, glabra vel sparse glanduloso- 
puberulenta; corolla infundibuliformis, minute et sparse pubescens, 
lobis rotundatis; stamina inclusa; capsulae clavatae, minute et parce 
retrorse pubescentes. 

Herbaceous or suffrutescent; stems quadrangular (the angles 
winged), glabrous, or the tips and nodes sparingly pubescent, the 
cystoliths numerous, minute, subpunctiform; leaf blades oblong, up 
to 16 cm. long and 5 cm. wide, short-acuminate (the tip blunt), 
narrowed at the base, glabrous above, the costa and lateral nerves 
minutely brownish-pubescent beneath, the hairs incurved; petioles 
up to 3.5 cm. long, minutely pubescent; flowers borne in peduncled, 
dichotomously branched, axillary panicles, the peduncles up to 30 
cm. long, slender, sharply angled, glabrous, the rachises (sometimes 
2 pairs at each node) slender, sharply angled, glabrous, often hori- 
zontally spreading, some of the internodes curved; flowers solitary 
or in pairs, borne at the nodes of the inflorescence or terminating its 
branches; pedicels up to 4 mm. long, glabrous or with a few minute, 
spreading, glandular hairs; calyx 6-7 mm. long, the segments 4-5 mm. 
long, subulate, 0.75 mm. wide at the base, glabrous or with a few 
minute, glandular hairs; corollas 2.5 cm. long, white, minutely and 
sparingly pubescent, the tube slender below, 1.5 mm. in diameter for 
about 12 mm. of its length, thence obliquely expanded and obconic, 
6-7 mm. broad at the mouth, the limb 8-12 mm. broad, subregular, 
the lobes rounded, about 5 mm. in diameter; stamens included; 
capsules about 15 mm. long, 4 mm. wide, clavate, the solid stipe 



1252 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

about 1 cm. long, the entire capsule minutely and rather sparingly 
retrorse-pubescent. Daytonia Farm, region of Almirante, Province 
of Bocas del Toro, Panama, January to March, 1928, Cooper 185 . 
(type in Herb. Field Mus., No. 579,478). Distinguishable from R. 
Pittieri by its pubescent capsules, more slender inflorescence, and 
nearly glabrous corollas. Although known only from Panama, its 
range probably extends into Costa Rica. 

Ruellia geminiflora HBK. Fields and waste places. Central 
America, West Indies, and northern South America. An herb, up 
to 50 cm. high, the stems erect or ascending, pilosulous; leaf blades 
ovate, oblong, or lanceolate, short-petioled, 1.5-5 cm. long, 0.5-2 cm. 
wide, obtuse or acutish at the apex, narrowed at the base, entire, 
hirtellous; flowers axillary; calyx 10 mm. long or less, pilosulous, the 
segments linear-lanceolate; corollas mauve, pubescent, usually 2.5-3 
cm. long, the lobes suborbicular, 5-10 mm. wide; capsules 8 mm. long, 

4 mm. wide, puberulent, 4-seeded. 

Ruellia inundata HBK. Dry forests and thickets. Mexico, 
Central America, and Colombia. Herbaceous or suffrutescent, up 
to 2 meters high; stems simple or branched, the tips glandular- 
pilose, the lower portions becoming whitish and glabrate; leaf blades 
lance-oblong to ovate, up to 15 cm. long and 7 cm. wide, acuminate, 
acute or rounded at the base, denticulate, repand-crenulate or sub- 
entire, hispidulous or occasionally grayish-pubescent beneath; 
petioles up to 9 cm. long, pilose; inflorescence axillary, the branches 
erect or ascending, dichotomous, glandular-pubescent, sometimes a 
dense, cylindric panicle; bractlets oblong, 3-8 mm. long, 1-3 mm. 
wide; calyx 8-15 mm. long, glandular-pilose, the segments linear; 
corollas purplish, pubescent, 2-2.5 cm. long, the lobes rounded, about 

5 mm. wide; capsule 8-9 mm. long, 3 mm. wide, flattened, pointed 
at the apex, narrowed to a slender, solid base, glabrous, 2-4-seeded; 
seeds flat, 4 mm. long, 3 mm. wide. 

Ruellia jussieuoides (Nees) Schlecht. Collected in a riverside 
thicket in the vicinity of El General and at Rodeo de Pacaca. Found 
also on the Rio Xucpec, Guatemala. Herbs or slender shrubs; 
stems erect or ascending (sometimes supported by other plants), 
up to 2.5 meters high, quadrangular, with curved hairs; leaf blades 
narrowly lanceolate, 6-15 cm., long, 1-3 cm. wide, slenderly acuminate 
(the tip obtuse), gradually narrowed from about the middle to the 
base, entire or undulate, both surfaces pilosulous, the hairs thickest 



FLORA OF COSTA RICA 1253 

on the costa and nerves (these conspicuous), the cystoliths promi- 
nent; petioles 1-2 cm. long, pubescent; flowers 1-3, subsessile, borne 
on peduncles up to 7 cm. long, these solitary or in pairs, sparingly 
pubescent; bracts subtending the flowers narrowly lanceolate, usually 
about 2.5 cm. long and 3-5 mm. wide; calyx 6-14 mm. long, sparingly 
pilosulous, the segments narrowly linear, 1 mm. wide at the base, 
with conspicuous cystoliths; corollas lavender or occasionally white, 
4-5 cm. long, minutely pubescent, the tube slender and usually 
curved, about 2 mm. in diameter at the base, 5 mm. at the mouth, 
the limb 2-5 cm. broad, the lobes elliptic, 1.5-2 cm. wide; capsules 
13 mm. long, 3-5 mm. broad, pointed, abruptly narrowed to a 
slender, solid base 4.5 mm. long and 1.5-2 mm. broad, obscurely 
puberulent, 12-seeded; seeds flat, suborbicular, 1.5 mm. in diameter. 

Ruellia malacosperma Greenm. Cultivated in the Meseta 
Central; also at La Hondura. Mexico to Nicaragua. Herbaceous; 
stem up to 60 cm. high, glabrous or sparingly pilose; leaf blades 
lanceolate to oblong-elliptic, up to 12 cm. long and 3.5 cm. wide, 
gradually narrowed from about the middle to an acute or obtuse tip, 
narrowed at the base, undulate, glabrous, or the costa and margins 
pilose; petioles 0.5-3 cm. long, glabrous or pilose; inflorescence 
axillary, once or twice dichotomous, the peduncles slender, 3-6 cm. 
long, glabrous; calyx 1.2-2 cm. long, more or less glandular-puberu- 
lent or sometimes glabrate, the segments linear-subulate; corollas 
mauve, finely pubescent, up to 6 cm. long, the slender portion of the 
tube 1-1.5 cm. long, the upper portion funnelform or campanulate, 
slightly oblique, the limb 3-4.5 cm. broad, the lobes rounded; 
capsules 2-3 cm. long, cylindric, glabrous (the apical calluses puberu- 
lent), usually 20-seeded. 

Ruellia metallica Leonard, sp. nov. Suffrutex, caulibus quad- 
rangularibus, glabris vel angulis pilosulis; lamina foliorum oblongo- 
elliptica, acuminata, basi angustata, in petiolum decurrens, paullo 
undulata, supra glabra, subtus in costa et venis depresso-pilosula, 
subtus metallico-purpurea; petioli pilosi, pilis curvatis; flores in 
fasciculis terminalibus et axillaribus sessiles, bracteis supremis 
linearibus, saepe longissimis, inferioribus foliosis; bracteolae tri- 
angulares, glabrae vel parce puberulentae; calycis segmenta subulata, 
glabra vel parce puberulenta; corolla purpurascens, infundibuliformis, 
lobis suborbicularibus; stamina inclusa; capsulae clavatae, minute 
puberulentae; semina rubescenti-brunnea, plana, orbicularia, glabra 
vel margine puberulenta. 



1254 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Suffrutescent, 50 cm. high or more; stems quadrangular, glabrous, 
or pilosulous on the angles; leaf blades oblong-elliptic, 5-15 cm. long, 
1-6 cm. wide, acuminate, often curved toward the tip, narrowed 
at the base and decurrent on the petiole, entire or shallowly undulate, 
glabrous except the costa and veins beneath, these pilosulous with 
curved, appressed hairs, the lower surface often a metallic purple; 
petioles 5-15 cm. long, pilosulous with curved, appressed hairs; 
flowers borne in terminal and axillary fascicles; uppermost bracts 
linear, 2-6 cm. long, 1-5 mm. wide, the lower intergrading into 
leaves; bractlets triangular, 2-2.5 mm. long, 0.75 mm. wide at the 
base; calyx 7-8 mm. long, glabrous or bearing a few minute hairs, 
the tube 1 mm. long, the segments subulate, 0.5 mm. wide at the 
base; corollas purplish, 2.5 cm. long, sparingly and minutely pubes- 
cent, the lower half of the tube 1.5 mm. in diameter, the upper portion 
obconic, 5 mm. broad at the mouth, the limb 1-2 cm. broad, the lobes 
suborbicular, 5-8 mm. in diameter; stamens included, one filament 
of each pair 3 mm. long, the other 5 mm., all glabrous; anthers 1.5 mm. 
long, 0.5 mm. wide; ovary minutely puberulent; style 15 mm. long, 
glabrous or bearing a few minute hairs near the base; stigma lobes 
very unequal, one about 1 mm. long, the other vestigial; capsules 
7-10 mm. long, 3 mm. broad, pointed at the apex, narrowed to a 
slender, solid base 5 mm. long and 1.25 mm. broad, minutely puberu- 
lent; retinacula 1.5 mm. long, bidentate or irregularly toothed at the 
apex; seeds reddish brown, orbicular, about 2 mm. in diameter, 
glabrous, or puberulent on the margins, mucilaginous when mois- 
tened. Zent Farm, Prov. Limon, at sea level, May 2, 1903, Cook & 
Doyle 456 (type in U. S. Nat. Herb., No. 474,307). Cuesta de La 
Vieja, Biolley 64 (in part). Banana plantation at Limon, Cook & 
Doyle 441- Forests of Hacienda Victoria, Plains of Zent, Pittier 
16029. Wet thicket, Finca Montecristo, Rio Reventazon below El 
Cairo, Prov. Limon, 25 meters, Standley & Valerio 48445. Moist 
forest, La Tejona, north of Tilaran, Guanacaste, 600-700 meters, 
Standley & Valerio 45880. Playa Blanca, Golfo Dulce, Manuel 
Valerio 474. NICARAGUA: Mosquito coast, Schramm. Sangsangta 
District, Schramm 28, 58. PANAMA: Changuinola Valley, Province 
of Bocas del Toro, Dunlap 380, 437, 461. Cacao plantation in 
western Panama, Stork 34^2- Readily distinguished from other 
Central American species by its sessile, fascicled flowers, the metallic 
luster of the lower surface of the leaf blades, and the long, narrow 
bracts of the inflorescence. In general appearance it must bear a 



FLORA OF COSTA RICA 1255 

close resemblance to R. barbillana, but that species has glabrous 
capsules and slightly larger flowers (3 cm. long). 

Ruellia nudiflora var. puberula Leonard. Collected in Guana- 
caste, at 25-70 meters. Oaxaca, Guatemala, and Salvador. Herbs, 
10-20 cm. high; stems ascending, branched, puberulent, the minute 
hairs mixed with longer, spreading ones, the younger stems rather 
densely pilose; leaf blades oblong-elliptic, 2-10 cm. long, 1-3 cm. 
wide, obtuse or rounded at the apex, gradually narrowed at the base 
and decurrent on the petiole, firm, undulate-crisped, gray-puberulent 
with minute, curved hairs, these interspersed with longer, spreading 
ones; petioles 1-3 cm. long; inflorescence of axillary, dichotomous 
cymes, with a flower in the forks; peduncles 1-8 cm. long, glandular- 
puberulent; bracts leaflike, 5-10 mm. long, 2-3 mm. wide, or occa- 
sionally much larger; pedicels 5-10 mm. long, or the one in the 
lowest fork becoming stout and 15-20 mm. long; calyx 15-30 mm. 
long, grayish-puberulent and ciliate, the segments linear, keeled, 
1 mm. wide at the base; corolla 3-4 cm. long, the lower portion of 
the tube 10-12 mm. long and 1.5 mm. in diameter, the upper part 
abruptly enlarged and broadly funnelform or narrowly campanulate, 
the throat 15 mm. broad, the limb 3-3.5 cm. broad, the lobes rounded, 
undulate, often emarginate; capsule 15-18 mm. long, 4 mm. broad, 
puberulent. 

Ruellia paniculata L. Waste places. Mexico to Colombia and 
Venezuela, and West Indies. Herbaceous or suffrutescent; stems up 
to 1 meter long, erect, ascending, or sometimes decumbent and root- 
ing at the nodes, the pubescence a mixture of minute, grayish hairs 
and longer, glandular ones; petioles slender, 1-4 cm. long; leaf blades 
ovate to oblong-ovate, 3-17 cm. long, 1.5-6.6 cm. wide, obtuse or 
acute, abruptly narrowed at the base, glandular-pubescent, grayish- 
puberulent or glabrate; inflorescence axillary, the branches divaricate 
or ascending, dichotomous; bracts leaflike, glandular-pubescent, 
those subtending the flowers 2-10 mm. long and 1-4 mm. wide; calyx 
1-1.5 cm. long, glandular-pubescent, the segments linear-subulate, 
unequal ; corollas blue, finely pubescent, 2-3 cm. long, the lobes oval, 
6-7 mm. long, 5 mm. wide; capsules cylindric, 10-13 mm. long, 2 mm. 
in diameter, erect, pointed, glabrous, 8-seeded; seeds flat, 2 mm. long 
and 1.5mm. wide. 

Ruellia Pittieri Lindau, Bull. Herb. Boiss. 6: 655. 1897. Cerro 
del Volcan, near Boruca, 1,200 meters, Pittier 6743. Forests of 
Cerro del Volcan and Santo Domingo de Golfo Dulce. Suffrutescent, 



1256 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the stems about 1 meter high, quadrangular, glabrous or puberulent; 
leaf blades ovate to oblong-elliptic, 6-21 cm. long, 1.5-6 cm. wide, 
acute or acuminate, narrowed at the base, entire or crenate-dentate, 
glabrous except the costa and veins of the lower surface, these 
puberulent, the cystoliths numerous; flowers borne in diffuse, axillary 
panicles, the peduncles up to 22 cm. long, quadrangular, narrowly 
winged, sparingly puberulent or glabrous, the cystoliths numerous; 
bracts up to 4 cm. long and 1.5 cm. wide, leaflike, rounded at the 
apex; bractlets triangular or subulate, about 1 mm. long; calyx about 
4 mm. long, the segments subulate, sparingly puberulent; corollas 
2 cm. long, puberulent, the tube slender, the lower portion 2 mm. in 
diameter, oblique above and 6 mm. in diameter at the throat, the 
lobes equal, obtuse, 8 mm. long, 5 mm. wide; style 15 mm. long, 
pubescent; capsules 13 mm. long, 4 mm. broad, acute at the apex, 
glabrous, 4-seeded, the sterile basal portion about 7 mm. long. 

Ruellia stemonacanthoides (Oerst.) Hemsl. Biol. Centr. Amer. 
Bot. 2: 507. 1882. Arrhostoxylum stemonacanthoides Oerst. Vid. 
Medd. Kjoebenhavn 1854: 130. 1855. Cerro de Aguacate, 480 
meters, Oersted. Forests, usually in wet situations. Campeche and 
British Honduras to Costa Rica. Suffrutescent, up to 3 meters tall; 
stems puberulent or glabrate; leaf blades oblong-ovate to oblong- 
lanceolate, 6-16 cm. long, 1.5-8 cm. wide, acuminate (the tip blunt, 
often minutely cuspidate), rounded or narrowed at the base, firm, 
entire, glabrous, the veins and costa of the lower surface puberulent; 
petioles 1-9 cm. long; inflorescence axillary, peduncled, dichotomous, 
sometimes a terminal panicle, the peduncles up to 17 cm. long, quad- 
rangular, horizontal or slightly ascending, curved, the flowers more or 
less secund; bracts leaflike; calyx up to 1 cm. long, puberulent or 
glabrate, the segments linear-subulate; corollas purple, salverform, 
3-4 cm. long, the lobes oval, about 9 mm. long and 8 mm. wide, 
shallowly emarginate; capsules 16 mm. long, glabrous, 12-seeded; 
seeds flat, 2.5 mm. in diameter. 

Ruellia tetrastichantha Lindau in Pittier, Prim. 2: 300. 1900. 
Corteza de venado. Banks of Rio Corozal, near Santo Domingo de Osa, 
Tonduz 9984. Wet forests of Costa Rica, Panama, and Colombia. 
Suffrutescent, 1-1.5 meters high; stems ascending, subquadrangular, 
minutely puberulent; leaf blades oblong-elliptic, 6-14 cm. long, 3-12 
cm. wide, shorfc-acuminate (the tip blunt), narrowed at the base and 
decurrent on the petiole, entire or undulate, glabrous except the costa 
and veins beneath, these puberulent, the lower surface with small, 



FLORA OF COSTA RICA 1257 

reddish, glandular dots, the cystoliths minute; petioles 1.5-2.5 cm. 
long, puberulent; flowers several, borne in terminal or lateral heads 
subtended by several leaflike bracts 2-5 cm. long and 1.5-2 cm. wide; 
peduncles 5-10 mm. long, the rachis conical, 5-12 mm. long, exposing 
(after the bracts, flowers, and fruits fall) vertical rows of conspicuous, 
corky scars; bractlets broadly triangular, obtuse, puberulent; pedicels 
2-3 mm. long; calyx 4 mm. long, puberulent, with glandular dots, 
the segments narrowly triangular; corolla white, yellow, or occasion- 
ally red, 7-8 cm. long, puberulent, the lower portion of the tube 
slender, 3-5 mm. in diameter, the upper half obconic or subcylindric, 
1-2 cm. broad at the mouth, the limb 2-3 cm. broad, the lobes oval 
or suborbicular, 10-15 mm. broad; stamens included; ovary puberu- 
lent and gland-dotted; style 6 cm. long, puberulent; capsules 15 mm. 
long, 5 mm. broad, pointed at the apex, narrowed to a solid base 2.5-3 
mm. long and 3 mm. broad, puberulent, 8-10-seeded; seeds brown, 
flat, orbicular, 5 mm. in diameter. 

Ruellia Tonduzii Lindau in Pittier, Prim. 2: 302. 1900. Punta 
Mala, Pacific coast, Tonduz 6778. Found on the Pacific coast at 
Punta Mala and Puerto Jimenez; also on low hills above Rio Paquita 
and flood plains of Rio Sandalo. Endemic. Stems obscurely quad- 
rangular, puberulent or glabrate, with cystoliths; leaf blades oblong, 
5-10 cm. long, 2-5 cm. wide, acuminate or obtuse, acute at the base 
and decurrent on the petiole, entire or shallowly crenate, both sur- 
faces glabrous except the costa and veins beneath, these puberulent 
with curved hairs, the cystoliths slender and prominent; petioles 
slender, 0.5-1.5 cm. long; flowers few, borne in peduncled, dichoto- 
mous, axillary cymes; peduncles up to 9 cm. long, quadrangular, 
glabrous; bracts oblong, 5-18 mm. long, 1-3 mm. wide; bractlets 
linear-subulate, 1-1.5 mm. long; calyx 4-5 mm. long, puberulent or 
glabrate, the segments subulate; corollas lilac, 2-2.5 cm. long, 
obscurely pubescent, the lower half of the tube slender, 1 mm. in 
diameter, the upper half narrowly campanulate, 7 mm. broad at the 
mouth, the lobes ovate, about 7 mm. long and 5 mm. wide; style 
about 17 mm. long, sparingly puberulent; capsules 11-12 mm. long, 
3 mm. broad, pointed at the apex, abruptly narrowed to a slender, 
solid base 5-6 mm. long and 1 mm. broad, glabrous; seeds flat, len- 
ticular, 1.5 mm. in diameter. 

SPATHACANTHUS Baill. 

Shrubs or small trees; leaves usually large, oblong or elliptic, 
petiolate; flowers large, few, borne on small, terminal or lateral 



1258 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

branches; bracts small; calyx spathaceous, valvate; corolla usually 
large, the tube salveriform, curved, the limb 2-lipped, the lobes sub- 
equal, rounded; stamens 4, didynamous; anther lobes parallel; ovules 

2 in each locule; capsules large, glabrous. 

Spathacanthus Hoffmann! Lindau, Bull. Herb. Boiss. 3: 370. 
1895. Near Aguacate, Hoffmann 842. Forests on the slopes of the 
central cordillera. Trees, up to 6 meters high; branches glabrous; 
leaf blades oblong, up to 28 cm. long and 11 cm. wide, acuminate, 
narrowed at the base and decurrent on the petiole, glabrous, both 
surfaces bearing numerous small cystoliths; petioles about 5 cm. long; 
flowers borne in short racemes, the rachis 1-4 cm. long; bracts and 
bractlets narrowly triangular, 1-3 mm. long, acute; spathe segments 

3 cm. long, connate below, the lobes ovate, short-acuminate, 13 mm. 
wide; corolla 6-7 cm. long, white, glabrous, the tube 5 mm. in diame- 
ter at the base, 2 cm. at the mouth, the limb 4 cm. broad, the lobes 
rounded; stamens included; style about equaling the corolla; cap- 
sules 6-6.5 cm. long, glabrate, woody, 4-seeded; seeds flat, ovate, 
about 10 mm. long, minutely roughened. 

STREBLACANTHUS Kuntze 



Herbs (sometimes shrubby); leaves petioled; flowers borne in 
lax, terminal and axillary spikes; bracts narrow; calyx segments 4, 
narrow, equal; corolla tube expanded at the base, gradually narrowed 
to a slender throat, the limb 2-lipped, spreading, the upper lip 2-lobed, 
the lower 3-lobed; stamens 2, attached to the throat of the corolla, 
the filaments short, the anther cells unequal, widely separated, one 
usually fertile and calcarate at the base, the other often abortive; 
staminodes none; style filiform; capsules slender, sterile and solid 
from middle to base, the upper portion flat, ovate, 2-4-seeded, 
the seeds lenticular. Two species not occurring in Costa Rica 
have been described, one from Panama the other from Bolivia. 

Limb of the corolla 15-20 mm. broad S. macrophyllus. 

Limb of the corolla 10 mm. broad or less. 

Superior anther lobe fertile S. longiflorus. 

Superior anther lobe sterile S. monospermus. 

Streblacanthus longiflorus Cufodontis, Archivio Bot. 10:48. 
1934. Description adapted from the original. Type collected 12 km. 
from the mouth of Rio Reventazon, Atlantic coast, Cufodontis 336. 
A shrub, up to 2 meters high; stems sub terete, subquadrangular 
toward the tip, pilose in 2 lines; leaf blades ovate-lanceolate, up to 



FLORA OF COSTA RICA 1259 

20 cm. long and 8 cm. wide, acuminate, unequally narrowed at the 
base, glabrous, or the younger ones sparingly setulose, the cystoliths 
inconspicuous; petioles 4-7 cm. long; flowers borne in a terminal, 
glandular- villous spike about 6 cm. long; bracts linear-lanceolate, 
25 mm. long, 3 mm. wide, obtuse; bractlets subulate, 1 cm. long; 
calyx segments subulate; corollas pale lilac, the tube 25 mm. long, 
3 mm. in diameter at the base, gradually narrowed to 1 mm. at the 
mouth, the lobes of the limb about equal, ovate, rounded, about 
5 mm. long and 3 mm. wide, the upper obsoletely bifid(?); filaments 
2 mm. long, the superior anther lobe subglobose, fertile, the inferior 
lobe distant, smaller and apparently sterile; ovary villous, 2-ovuled. 

Streblacanthus macrophyllus Lindau in Pittier, Prim. 2: 
306. 1900. Forests along Rio Zhorquin, Pittier 8547. Forests of 
southern Costa Rica. Endemic. Stems glabrous, or pubescent in 
2 lines toward the tip; leaf blades oblong-ovate or elliptic, up to 25 cm. 
long and 10 cm. wide, acuminate, narrowed at the base, glabrous, 
or especially in the upper leaves the costa and margins more or less 
hirsute; spikes 1 or 2, about as long as the leaves, softly pilose, the 
hairs brownish, glandular; bracts oblong, up to 15 mm. long, 2-3 mm. 
wide, rounded at the tip; bractlets 6 mm. long, 1 mm. wide, acute; 
calyx segments 4 mm. long, 1 mm. wide, linear, minutely glandular- 
pubescent; corollas purplish white, pubescent, the tube 20 mm. 
long and 4 mm. in diameter at the inflated base, 1.5 mm. at the 
mouth, the upper lip 8 mm. long, 2-lobed, the lobes 6 mm. long, 
3.5 mm. wide, acute, the lower lip 3-lobed, the middle lobe 9 mm. 
long and 6 mm. wide, the lateral ones 8 mm. long and 5 mm. wide, 
all rounded; anthers exserted, 1-celled, obtuse; styles 22 mm. long. 

Streblacanthus monospermus Kuntze, Rev. Gen. 2: 498. 
1891. South of San Jose", Kuntze. Wet forests of southern Costa 
Rica. An herb, up to 1 meter high; stem simple, erect, terete, fleshy, 
pubescent in 2 lines with small, curved hairs; leaf blades oblong- 
ovate to oblong-elliptic, up to 18 cm. long and 7.5 cm. wide, acumi- 
nate (the tip blunt), narrowed at the base, glabrous; petioles up to 
5 cm. long, glabrous; spikes terminal or borne in the upper axils, 
up to 25 cm. long, soft-pilose, the hairs glandular, about 1 mm. 
long; bracts lanceolate, 1.5-2.5 cm. long, 3-4 mm. wide, acute to 
obtuse at the tip, glandular-pilose; bractlets similar but smaller; 
calyx segments lanceolate, 5-6 mm. long, glandular-pilose; corolla 
white, rather sparingly pilose, the tube 2-3 cm. long, about 3 mm. 
in diameter at the base, 1 mm. at the mouth, the limb 7-8 mm. 



1260 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

broad; capsules 2-2.5 cm. long and 8 mm. wide, glandular-pilose, 
4-seeded, long-stipitate, the upper seed-bearing portion flattened and 
ovate; seeds brown, about 5 mm. in diameter, glabrous, the edges 
roughened. 

TETRAMERIUM Nees 

Fruticose or suffruticose; stems usually branched, the pubescence 
often disposed in 2 opposite, lateral lines; leaves sessile or petiolate, 
ovate to linear, entire, glabrous or pubescent; flowers borne in termi- 
nal or axillary, 4-angled spikes, the bracts conspicuous, usually 
closely imbricate, opposite, ciliate, mucronate; calyx 4-5-parted, 
the segments ciliate; corollas tubular-infundibuliform, the tube 
straight or slightly curved, the upper lip entire or emarginate, the 
lower 3-lobed, the lobes nearly equal; stamens 2, included; anther 
sacs slightly converging toward the apex, one a little longer than the 
other; capsules obovate, apiculate, contracted below into a solid 
base, usually hispid and 4-seeded, sometimes 2-seeded; seeds flattened, 
tuberculate or muriculate. A genus of about 23 species, extending 
from southern United States to northern South America. Many 
of the species frequent dry situations at rather low altitudes. 

Bracts rotund-ovate to ovate, 7-nerved T. nervosum. 

Bracts elliptic, 3-5-nerved T. hispidum. 

Tetramerium hispidum Nees. Dry, open places throughout 
tropical America. Herbs, up to 30 cm. high; stems terete, brittle, 
pilosulous or glabrate; leaf blades ovate, 1-2.5 cm. long, 0.5-1.2 cm. 
wide, obtuse, rounded at the base, pilosulous; petioles up to 8 mm. 
long, slender, pilose; flowers borne in terminal spikes up to 9 cm. 
long and about 8 mm. in diameter, the rachis glabrous or sparingly 
pilose; bracts elliptic, about 7 mm. long and 4 mm. wide, acute, 
ending in a mucro 0.5 mm. long, ciliate, 3-5-nerved, closely imbri- 
cate; bractlets narrowly lance-aristate, 2.5 mm. long, pilose at the 
tip; calyx segments similar to the bractlets but slightly longer; 
corolla 1 cm. long, white, glabrous, the tube slender, the lips 5 mm. 
long, the upper one entire, the lower 3-lobed, the lobes elliptic, 3 mm. 
long; capsules 4.5 mm. long, pubescent or glabrate; seeds papillose. 

Tetramerium nervosum Nees. Wet forests, Central America 
to northern South America. Found at 600 meters in Costa Rica, 
in the vicinity of Los Ayotes (Guanacaste) and Orotina. Herbs; 
stems sometimes glabrous, but usually evenly pubescent or the hairs 
arranged in 2 lines; leaf blades ovate or ovate-lanceolate, up to 4 cm. 
long and 3 cm. wide, acuminate or acute, subcordate to obtuse at 



FLORA OF COSTA RICA 1261 

the base, thin, sparingly pubescent on both surfaces; petioles up to 
2 cm. long, sparingly pubescent or glabrous; flowers borne in terminal 
spikes up to 4 cm. long; bracts rotund-ovate to ovate, 6-10 mm. 
long, 5-7 mm. wide, short-acuminate to cuspidate at the apex, 
rounded and abruptly contracted below the middle into a sub- 
petiolate base, 7-nerved, hispid-ciliate, the hairs up to 2 mm. long; 
bractlets linear-lanceolate, 2-3 mm. long; calyx lobes 4, linear- 
lanceolate, 2-3 mm. long, hispid-ciliate; corolla 10 mm. long, essenti- 
ally glabrous, purplish white, the lips about 5.5 mm. long, the upper 
one subemarginate; capsules 5 mm. long, 1.8 mm. broad, hirsute 
near the apex, 4-seeded. 

THUNBERGIA Retz. 

Herbaceous or woody vines (occasionally shrubs) with large, 
2-bracted flowers, these solitary in the axils of the leaves, or in termi- 
nal racemes; calyx short, annular, usually with many teeth; corolla 
tube oblique, the limb 5-lobed; stamens 4, didynamous; ovary 4- 
ovuled; capsules coriaceous, globose, abruptly beaked. An Old 
World genus, widely introduced into tropical America. Other spe- 
cies than the four here described are likely to occur in gardens of 
Costa Rica. 

Petioles winged; corollas orange or yellow, with a dark purple eye. 

Petioles not winged. T ' alata " 

Vines; flowers pure white T. fragrans. 

Erect or scandent shrubs; flowers usually blue or violet. 
Leaf blades thin, small, usually 2-4 cm. long; erect shrubs. 

T. erecta. 
Leaf blades firm, usually 7-13 cm. long; scandent shrubs. 

T. Vogeliana. 

Thunbergia alata Bojer. Ojo de poeta. Roadsides and waste 
places. Tropical America generally. Naturalized from eastern 
Africa. A trailing, pubescent vine up to 1 meter long; leaf blades 
ovate to triangular-ovate, 2-8 cm. long, remotely few-toothed to 
entire, acute, cordate or hastate at the base; petioles winged; flowers 
axillary, the slender peduncles longer than the petioles; bracts ovate- 
lanceolate, about 1.5 cm. long, acute to acuminate; calyx shorter 
than the bracts; corolla 2.5-4 cm. long, yellow or occasionally white, 
usually with a purple eye; capsules depressed-globose, pubescent, 
0.5-1 cm. in diameter, the stout beak about 1 cm. long. In culti- 
vated plants a number of color forms may appear, varying from white 
to yellow or bright orange, with or without the purple eye. 



1262 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Thunbergia erecta (Benth.) T. Anders. Cultivated or found 
as an escape in tropical America. Originally from Cape Coast, 
Africa. Erect shrubs, up to 3 meters high; stems quadrangular, 
narrowly winged, glabrous, or the nodes pubescent; petioles 5-10 mm. 
long; leaf blades oblong-ovate or oblong-lanceolate, up to 7 cm. long 
and 3.5 cm. wide, but usually much smaller (2-4 cm. long), thin, 
glabrous, entire or sinuately toothed; flowers solitary, axillary; 
peduncles up to 2 cm. long, glabrous; bracts oblong, oblique, 1-2 cm. 
long, 5-8 mm. wide, obtusish, glabrous or minutely pubescent at the 
tip, faintly striate; calyx teeth 8-12, small (1-4 mm. long); corollas 
4-8 cm. long, deep violet with a white or yellowish throat, or the 
entire corolla pure white. 

Thunbergia fragrans Roxb. Roadsides, thickets, and waste 
places; cultivated in tropical America generally. Native of India. A 
slender vine, up to 2 meters long; stem grooved, minutely strigose or 
glabrate; petioles slender, up to 4 cm. long; leaf blades ovate to ovate- 
lanceolate, 4-11 cm. long, 1.5-6 cm. wide, acuminate, acute, or 
occasionally obtuse, at the base subtruncate, cordate, or hastate, 
entire or remotely few-toothed toward the base, thin, minutely 
strigose or glabrate; flowers axillary; peduncles up to 7 cm. long; 
bracts ovate to ovate-lanceolate, 1.5-2 cm. long, 6-13 mm. wide, 
acute or acuminate, pubescent; flowers pure white, the lobes sub- 
truncate at the apex, about as long as the tube; capsules depressed- 
globose, about 8 mm. in diameter, puberulent, tipped by a stout, 
subulate beak 1-1.5 cm. long. 

Thunbergia Vogeliana Benth. Occasionally cultivated in 
tropical America. The type was collected on Fernando Po, Africa. 
A scandent shrub, up to 2 meters high; branches glabrous, or finely 
pubescent at the nodes; petioles about 5 mm. long; leaf blades oblong, 
up to 13 cm. long and 5 cm. wide, acuminate, narrowed at the base, 
firm, subscabrous or glabrous, serrulate or entire; flowers solitary; 
peduncles up to 2.5 cm. long, densely brownish-pubescent at the 
base; bracts white, ovate, 2-2.5 cm. long, about 1 cm. wide, obtuse, 
thick; calyx teeth (about 12) 8 mm. long; corollas about 4 cm. long, 
the limb dark blue, the throat yellow within and white without. 

TRIGHANTHERA HBK. 

Shrubs or trees; leaves petioled, ovate to oblong, entire or un- 
dulate, the cystoliths numerous; inflorescence a terminal panicle or 
corymb, usually dense and many-flowered; bracts small, triangular; 
calyx segments 5, obtuse; corollas regular, 5-lobed, tomentose, 



FLORA OF COSTA RICA 1263 

campanulate; stamens 4, exserted, didynamous, the anthers bearded, 
the basal lobes rounded; capsules 4-seeded. A single species occurs 
in Central America, an additional one in Colombia. 

Trichanthera gigantea (Humb. & Bonpl.) Nees. Palo de agua. 
A common tree in moist forests. Costa Rica to Venezuela, Colombia, 
and Peru. Shrubs or trees, up to 5 meters high; branchlets quad- 
rangular, the angles rounded, the tips brown-tomentose; leaf blades 
ovate to oblong, up to 26 cm. long and 14 cm. wide, acuminate, 
narrowed at the base, entire or undulate, glabrous except the costa 
and veins, these prominent, more or less pubescent; petioles up to 5 
cm. long; panicles 5-15 cm. long, 4-5 cm. broad, brown-tomentose, 
the flowers secund; bracts 3 mm. long; calyx 10-12 mm. long, brown- 
tomentose, the lobes erect, oblong, 7-10 mm. long, 5 mm. wide; 
corollas 3-4 cm. long, red and glabrous proximally, yellowish and 
silky-tomentose distally, red and glabrous within, the tube 1-1.5 
cm. long, 6 mm. broad below, campanulate above, the limb 2-3 cm. 
broad, the lobes oblong to oblong-ovate, 10 mm. long; styles 4-5 cm. 
long, glabrous; capsules oblong, 1.5-2 cm. long, 0.5 cm. broad, obtuse 
at the apex, silky with closely appressed hairs, 1-4-seeded; seeds 
lenticular, glabrous, 3-4 mm. in diameter. 

PLANTAGINACEAE. Plantain Family 
Reference: Pilger, Pflanzenreich IV. 269. 1937. 

In Central America this family is represented by a single genus 
which contains the vast majority of the plants of the family. 

PLANTAGO L. Plantain 

Low, annual or perennial herbs, acaulescent in the Central 
American species, the leaves often forming a basal rosette, entire or 
dentate, usually with longitudinal nerves; flowers small, greenish, 
in solitary heads or spikes at the ends of the naked scapes; calyx 4- 
parted, the segments equal or 2 of them larger; corolla salverform, 
the 4 lobes spreading or reflexed ; stamens 4 or 2, inserted in the tube 
or throat of the corolla, the anthers versatile, 2-celled, longitudinally 
dehiscent; style simple, filiform; ovary sessile, superior, usually 2- 
celled; fruit capsular, circumscissile at or below the middle, the seeds 
1 or more in each cell. Probably no other species occur in Central 
America. 

Plantago hirtella HBK. Llanten. P. Galeottiana Dene. 
Common in pastures and on open banks of the central region, mostly 



1264 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

at 1,000-2,000 meters. Mexico to Brazil and Bolivia. Perennial; 
leaves oblanceolate to narrowly elliptic or obovate-elliptic, acute or 
obtuse, attenuate to the base, 5-7-nerved, entire or nearly so, sparsely 
or densely villous with whitish hairs; scapes 12-30 cm. long, villous, 
the spikes dense, mostly 4-10 cm. long; corolla lobes erect in fruit. 
Pilger recognizes several varieties and forms, none of which appear 
to be of great importance. He refers some Costa Rican collections 
to f . minor Pilger. 

Plantago lanceolata L. Well established in pastures on the 
upper slopes of the volcanoes of Turrialba and Irazu. Native of 
the Old World, but generally naturalized in temperate regions of 
America. Perennial or biennial, pilose; leaves narrowly oblong- 
lanceolate, entire, acute or acuminate, gradually attenuate to the 
petiole, 3-5-nerved; scapes slender, striate, often 30 cm. long or even 
more; spikes very dense, at first ovoid, becoming cylindric, 1-10 cm. 
long; capsule with only 2 seeds. In the United States both this and 
the following species are often troublesome weeds. 

Plantago major L. Llanten. Naturalized in grassy places 
about San Jose". Native of the Old World; naturalized generally in 
temperate regions of America. Perennial, glabrous or inconspicuously 
pubescent; leaves petiolate, often lying flat upon the ground or 
nearly so, ovate or broadly ovate, entire or coarsely dentate, 3-11- 
nerved; scapes mostly less than 30 cm. high, the spikes linear-cylin- 
dric, dense, 5-20 cm. long; capsule with 5-16 seeds. 

RUBIACEAE. Coffee Family 
Reference: Standley, N. Amer. Fl. 32: 3-300. 1918-1934. 

Herbs, shrubs, or trees, sometimes scandent, often armed with 
spines; leaves opposite or verticillate, stipulate, simple and entire, 
pinnatifid in one genus; inflorescence various, the flowers perfect 
or rarely unisexual, usually regular and symmetric; calyx tube 
(hypanthium) adnate to the ovary; corolla gamopetalous, the lobes 
in bud valvate, imbricate, or contorted; stamens usually as many as 
the corolla lobes and alternate with them, the anthers 2-celled; ovary 
1-10-celled, the ovules 1-many in each cell; fruit capsular, baccate, 
drupaceous, or of dehiscent or indehiscent cocci. One of the largest 
families of tropical America, well represented in Costa Rica. A 
considerable number of additional genera are known from other 
parts of Central America, and a few of them may well be represented 
in Costa Rica. 



FLORA OF COSTA RICA 1265 

Key to the tribes and genera 
Ovules more than 1 in each cell of the ovary. 
Fruit fleshy. 

Corolla lobes valvate VI. Mussaendeae. 

Corolla lobes imbricate or contorted VII. Gardenieae. 

Fruit dry. 

Flowers arranged in compact, globose heads; shrubs or trees, 
armed with recurved spines V. Naucleeae. 

Flowers not in compact, globose heads. 

Seeds winged or appendaged, vertically imbricate. 

IV. Cinchoneae. 
Seeds exalate or, if winged, horizontal. 

Corolla lobes imbricate or contorted. . . .III. Rondeletieae. 
Corolla lobes valvate. 

Seeds horizontal, usually very numerous; stipules entire 
or biparted; large shrubs or trees with usually large 
leaves I. Condamineae. 

Seeds imbricate, vertical, usually few; stipules often 
setose-laciniate; herbs or small shrubs with small 
leaves II. Oldenlandieae. 

Ovules solitary. 

Seed pendulous, the radicle superior; shrubs or trees. 

Flowers in compact, globose heads V. Naucleeae. 

Flowers never in compact, globose heads. 
Stamens inserted in the throat of the corolla. 

VIII. Guettardeae. 
Stamens inserted at the base of the corolla tube. 

IX. Chiococceae. 
Seed ascending, the radicle inferior; plants often herbaceous. 

Corolla lobes contorted ; shrubs or trees X. Ixoreae. 

Corolla lobes valvate. 

Ovules basal, attached at the base of the cell; mostly shrubs 

or trees. 

Ovary 1-celled, or 2-celled but with a very thin septum, the 
fruit 1-seeded XIII. Coussareae. 

Ovary 2-celled, or the cells rarely more numerous, the 
septum thick. 



1266 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Stamens usually inserted in the base of the corolla tube; 
flowers often dioecious XII. Anthospermeae. 

Stamens usually inserted in the throat of the corolla; 
flowers perfect XL Psychotrieae. 

Ovules lateral, attached to the septum. 

Stipules neither foliaceous nor setiferous; trees or large 
shrubs; flowers confluent to form a dense head. 

XIV. Morindeae. 

Stipules either foliaceous (the leaves then appearing verti- 
cillate and estipulate) or setiferous; herbs or low shrubs; 
flowers never confluent into a true head. 

Stipules setiferous XV. Spermacoceae. 

Stipules foliaceous, resembling the leaves. .XVI. Galieae. 

I. CONDAMINEAE 

Calyx lobes dissimilar, one of them expanded into a large, colored, 

bractlike limb Pogonopus. 

Calyx lobes all similar, none of them expanded. 
Anthers dehiscent by longitudinal slits, dorsifixed; inflorescence 

axillary Chimarrhis. 

Anthers dehiscent by terminal pores, basifixed; inflorescence 
terminal Rustia. 

II. OLDENLANDIEAE 

Seeds angulate; plants herbaceous Oldenlandia. 

Seeds crateriform or concavo-convex; small shrubs. . . Arcytophyllum. 

III. RONDELETIEAE 

Corolla lobes contorted. 
Corolla tube very short, shorter than the lobes. 

Corolla glabrous within, 4-parted; filaments not appendaged. 

Deppea. 

Corolla villous within, 5-parted; filaments appendaged at the 
base Elaeagia. 

Corolla tube elongate, several times as long as the lobes. 

Lindenia. 
Corolla lobes imbricate. 

Calyx lobes dissimilar, one of them expanded into a large, petiolate, 
colored limb . . . Warszewiczia. 



FLORA OF COSTA RICA 1267 

Calyx lobes all similar, none of them expanded into a petiolate limb. 

Seeds very large, horizontal Sickingia. 

Seeds small or minute. 

Corolla salverform or funnelform Rondeletia. 

Corolla cylindric-campanulate. 

Corolla barbate in the throat Holtonia. 

Corolla glabrous within. Oregandra. 

IV. ClNCHONEAE 

Corolla lobes valvate. 

Plants scandent, herbaceous Manettia. 

Plants erect, woody. 

Flowers 4-parted; capsule subglobose Bouvardia. 

Flowers 5-parted ; capsule oblong to clavate. . 

Capsule splitting from below upward Cinchona. 

Capsule splitting from the apex downward. 

Capsule loculicidal Macrocnemum. 

Capsule septicidal Ladenbergia. 

Corolla lobes imbricate or contorted. 
Calyx lobes dissimilar, one of them expanded into a large, white, 

petiolate limb Calycophyllum. 

Calyx lobes all similar, none of them expanded into a petiolate 

limb. 
Corolla lobes contorted. 

Calyx tubular, deciduous; seeds not barbate Cosmibuena. 

Calyx parted to the base, or none; seeds barbate Hillia. 

Calyx lobes imbricate. 

Anthers sessile or nearly so; plants epiphytic; corolla red. 

Ravnia. 
Anthers borne on long filaments; plants terrestrial; flowers 

not red. 
Corolla symmetric, the tube very slender, the limb 4-5- 

lobate Exostema. 

Corolla asymmetric, the tube enlarged, the limb 6-8-lobate. 

Coutarea. 
V. NAUCLEEAE 
A single genus in Costa Rica Uncaria. 



1268 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

VI. MUSSAENDEAE 

Inflorescences terminal. 
Inflorescence a spikelike panicle; leaves membranaceous; ovary 

2-4-celled Gonzalagunia. 

Inflorescence a thyrsoid panicle; leaves coriaceous; ovary 4-6- 

celled . Isertia. 

Inflorescences axillary. 

Leaves finely lineolate between the veins; inflorescence not capitate. 
Calyx subtruncate, very shallowly and irregularly lobate. 

Nothophlebia. 

Calyx spathaceous or conspicuously and regularly lobate. 
Calyx spathaceous or lobate, the lobes not foliaceous; leaves 

often pinnatifid Pentagonia. 

Calyx deeply lobate, the lobes subfoliaceous; leaves entire. 

Sommera. 
Leaves not lineolate; inflorescence usually capitate or cymose- 

capitate. 
Plants herbaceous, usually prostrate or nearly so, not scandent; 

ovary 2-celled Coccocypselum. 

Plants woody or suffrutescent, sometimes scandent. 

Ovary 3-5-celled; plants scandent Sabicea. 

Ovary 2-celled; plants erect Xerococcus, 

VII. GARDENIEAE 
Corolla lobes imbricate. 

Ovary 4-5-celled Hamelia. 

Ovary 2-celled Hoffmannia. 

Corolla lobes contorted. 

Seeds minute, the testa foveolate Bertiera. 

Seeds large, the testa smooth or fibrous. 

Corolla somewhat irregular, curved in bud Posoqueria. 

Corolla regular, the buds not curved. 
Flowers perfect. 

Flowers terminal or terminal and axillary. 

Flowers in cymes Tocoyena. 

Flowers solitary or fasciculate Randia. 



FLORA OF COSTA RICA 1269 

Flowers axillary. 
Corolla tube villous in both throat and base; stigma 

attenuate Genipa. 

Corolla tube villous in the throat or the base, but not in 
both; stigma not attenuate. 

Ovary 2-celled; native plants Randia. 

Ovary 1-celled; cultivated plants Gardenia. 

Flowers dioecious. 
Stipules forming a conic cap, deciduous above a circular 

slit Duroia. 

Stipules free or united only near the base, not dehiscent 

by a circular slit. 
Staminate flowers lateral; plants usually armed with 

spines Randia. 

Staminate flowers terminal; plants unarmed . . Alibertia. 

VIII. GUETTARDEAE 

Fruit separating into 2 cocci at maturity; calyx lobes 4-5, persistent; 

corolla lobes not imbricate Machaonia. 

Fruit drupaceous, not separating into cocci at maturity. 
Corolla lobes valvate or but slightly imbricate. 
Corolla lobes corniculate-appendaged near the apex; basal lobes 

of the anthers acute or attenuate Chomelia. 

Corolla lobes not appendaged; basal lobes of the anthers obtuse. 

Anisomeris. 
Corolla lobes strongly imbricate, 1 or 2 of them exterior. 

Guettarda. 
IX. CHIOCOCCEAE 
A single genus in Costa Rica .Chiococca. 

X. IXOREAE 

Bractlets connate, calyx-like; calyx usually truncate Coffea. 

Bractlets distinct, not calyx-like; calyx lobate Ixora. 

XI. PSYCHOTRIEAE 

Inflorescence an involucrate, usually solitary head; heads sometimes 
branched in Cephaelis. 

Plants creeping herbs Geophila. 

Plants erect shrubs Cephaelis. 



1270 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Inflorescence not involucrate or rarely so, not a single involucrate 

head. 
Carpels of the fruit laterally compressed, the fruit didymous; 

plants low herbs Declieuxia. 

Carpels of the fruit not laterally compressed; plants mostly trees 
or shrubs. 

Seeds with an inrolled ventral surface; stipules pectinate-lobate 
or with setiform, dorsal appendages Rudgea. 

Seeds not with an inrolled ventral surface; stipules entire or 
bilobate. 

Corolla tube straight, not gibbous at the base; branches of 
the inflorescence usually green Psychotria. 

Corolla tube elongate, more or less curved, gibbous at the 
base; branches of the inflorescence usually pale or red or 
yellow Palicourea. 

XII. ANTHOSPERMEAE 
A single genus in Costa Rica Gomozia. 

XIII. COUSSAREAE 

Seeds vertical; ovules connate, borne upon a common basal column. 

Coussarea. 

Seeds horizontal; ovules separate in a 1-celled ovary, collateral, basal. 

Faramea. 

XIV. MORINDEAE 
A single genus in Central America Morinda. 

XV. SPERMACOCEAE 
Fruit not separating into cocci, the whole fruit circumscissile. 

Mitracarpus. 
Fruit separating into cocci, these not circumscissile. 

Cocci indehiscent. 

Cocci 3-4 Richardia. 

Cocci 2. 

Calyx lobate, the lobes elongate-subulate; cocci separating 

from a persistent axis Crusea. 

Calyx 2-10-dentate; fruit without a persistent axis. .Diodia. 
Cocci, at least one of them, dehiscent. 

Cocci opening only at the base; flowers axillary. . . . Hemidiodia. 



FLORA OF COSTA RICA 1271 

Cocci opening at the apex. 

Cells of the fruit alike, both opening Borreria. 

Cells of the fruit dissimilar, one opening, the other remaining 
closed Spermacoce. 

XVI. GALIEAE 

Calyx lobes developed, lanceolate; introduced plants. . . .Sherardia. 
Calyx lobes obsolete; native plants. 

Flowers surrounded by a calyx-like involucre Relbunium. 

Flowers not involucrate. 

Leaves opposite Didymaea. 

Leaves verticillate Galium, 

ALIBERTIAA. Rich. 

Shrubs or trees; leaves usually coriaceous; stipules acute, connate 
at the base; flowers small, terminal, sessile, dioecious, white, the 
staminate fasciculate, the pistillate usually solitary; hypanthium 
hemispheric or globose, the calyx short or tubular, truncate or den- 
tate; corolla coriaceous, salverform, the lobes contorted; ovary 2-8- 
celled, the ovules few or numerous in each cell; fruit baccate, usually 
rather large, the large seeds compressed. No other species are found 
in North America. 

Alibertia edulis (L. Rich.) A. Rich. Trompillo, Madrono. 
Forests and thickets of the Pacific tierra caliente; probably also on the 
Atlantic coast. Mexico to West Indies and the Amazon Basin. 
Usually a shrub or 4 meters or less, almost glabrous; stipules lance- 
triangular or deltoid, 8-15 mm. long, acute or acuminate; leaves 
short-petiolate, lance-oblong or ovate-oblong, 6-20 cm. long, usually 
short-acuminate, acute to rounded at the base, commonly barbate 
beneath in the axils of the nerves, otherwise glabrous; corolla 2.5 cm. 
long, minutely sericeous outside, the lobes acute or acuminate; 
calyx denticulate; fruit globose, yellowish, 2.5 cm. in diameter. 
Boys often use the fruits, with a stick thrust through them, as tops, 
hence the name trompillo. The leaves of seedling plants, often found 
abundantly in the forest, are narrower than those of adult plants, 
and handsomely colored with pink or purple. The fruits are reported 
edible, but they can not be very palatable. 

ANISOMERIS Presl 

Shrubs or small trees, often with spinose branchlets; stipules 
acuminate, deciduous; flowers small, white or yellowish, generally in 



1272 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

axillary, pedunculate cymes, rarely solitary; calyx 4-lobate, the lobes 
usually elongate, equal or unequal, persistent; corolla salverform or 
funnelform, with a slender tube, the limb 4-lobate, the lobes un- 
appendaged, valvate or slightly imbricate; stamens 4, the anthers 
dorsifixed, sessile; ovary 2-celled, the cells 1-ovulate; fruit small, 
drupaceous, with a single 2-celled stone. A few other species are 
known from Central America. 

Anisomeris microloba (Donn. Smith) Standl. Field Mus. Bot. 
4: 293. 1929. Chomelia microloba Donn. Smith, Bot. Gaz. 31: 114. 
1901. Forests of Santo Domingo de Golfo Dulce, Tonduz 7048 
(987 If). Known only from the original collection. An unarmed 
shrub; stipules 3-4 mm. long, deltoid, cuspidate; leaves short-petio- 
late, elliptic-oval to ovate-oblong, 5-10 cm. long, obtusely short- 
acuminate, rounded or very obtuse at the base, short-barbate beneath 
in the axils of the nerves, otherwise glabrous; cymes axillary, few- 
many-flowered, pedunculate, the flowers sessile or subsessile; calyx 
lobes minute, obtuse; corolla strigillose outside, the tube 8-10 mm. 
long, the lobes ovate, obtuse. 

Anisomeris sylvicola Standl. N. Amer. Fl. 32: 225. 1934. Cho- 
melia sylvicola Standl. Journ. Wash. Acad. 18: 182. 1928. Wet 
forest, Yerba Buena, northeast of San Isidro, Prov. Heredia, 2,000 
meters, Standley & Valerio 49196. Known only from the original 
collection. A glabrous shrub; stipules ovate-oval, 3 mm. long, 
obtuse; leaves slender-petiolate, the blades elliptic-oblong, 6.5-8 cm. 
long, 2-3 cm. wide, attenuate to an obtuse apex, acute and decurrent 
at the base, subcoriaceous, domatiate beneath in the axils of the 
nerves; inflorescence terminal, cymose-paniculate, rather few- 
flowered, on a peduncle 2 cm. long, the slender pedicels 10-12 mm. 
long; fruit obovoid, 18 mm. long, 7 mm. thick, acute at the base, 
2-celled; calyx persistent, cupular, 2 mm. long, undulate. It is 
fairly certain that this plant does not belong to the genus Anisomeris, 
although its relationship is apparently with Guettarda, Chomelia, etc. 
Until flowering material is available, the plant's true affinities must 
remain uncertain. 

ARCYTOPHYLLUM Schum. 

Small shrubs, erect or prostrate; leaves crowded and often 
densely imbricate, very small, thick-coriaceous; stipules entire, bifid, 
or setose; flowers small, solitary and axillary or in small cymes; 
hypanthium globose or turbinate; calyx 4-lobate, the lobes erect, 
remote; corolla funnelform or salverform, with a short or elongate 



FLORA OF COSTA RICA 1273 

tube, the lobes valvate; ovary 2-celled, the ovules few; fruit capsular, 
turbinate or globose, small; seeds concavo-convex, punctate. No 
other species are known in Central America. The genus probably 
should be united with Houstonia, since the only differences between 
the two groups seem to be habital, and even those are inconstant. 

Arcytophyllum lavarum Schum. Mallostoma lavarum Bonn. 
Smith. Dense, low thickets of the higher mountain slopes, especially 
on the central volcanoes, at 2,400-3,150 meters. Adjacent Panama. 
A very densely branched shrub, usually less than a meter high, 
glabrous except on the corolla; leaves broadly ovate to ovate-oblong, 
4-6 mm. long, 2-3 mm. wide, rounded or very obtuse at the apex, 
contracted into a short petiole, paler beneath; flowers in dense, small, 
terminal cymes, short-pedicellate; calyx lobes broadly ovate, obtuse 
or acute, with 1 or more setae between each pair; corolla 4-5 mm. long, 
the lobes ovate, obtuse, white- villous within; capsule subglobose, 
1.5-2 mm. long. One of the characteristic plants of the wet, moss- 
covered thickets about the lake on Pods, and in other similar localities. 

Arcytophyllum muticum (Wedd.) Standl. Paramos of Cerro 
de Las Vueltas, 3,000 meters; probably also on Cerro de La Muerte. 
Colombia. A small, prostrate shrub; leaves sessile, lance-oblong, 
obtuse, muticous, coriaceous, glabrous; flowers terminal, solitary; 
calyx lobes obovate-oblong, obtuse; corolla white. In North America 
this species (unknown from the region when the Rubiaceae of the 
North American Flora was published) is known only from sterile 
specimens collected by Juvenal Valeric and the writer. It is a 
characteristic species of the Colombian paramos. 

BERTIERA Aubl. 

Slender shrubs or small trees; leaves petiolate or sessile, mem- 
branaceous; stipules connate into a bifid sheath; flowers small, 
white, cymose, the cymes arranged in terminal, pedunculate panicles; 
calyx 5-dentate or truncate, persistent; corolla funnelform, usually 
sericeous, the limb 5-lobate, the lobes short, ovate, acute, contorted ; 
ovary 2-celled, the ovules numerous; fruit baccate, globose, fleshy; 
seeds numerous, very small, angulate, foveolate. One other Central 
American species has been described from Panama. 

Bertiera angustifolia Benth. Bot. Voy. Sulph. 103. 1845. 
Cocos Island, Barclay. Also in Panama. A shrub of 3 meters, the 
branches pilose-strigose; stipules 10-18 mm. long; leaves sessile or 
nearly so, oblong-lanceolate, 10-12 cm. long, acute or attenuate, 



1274 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

rounded or obtuse at the base, almost glabrous above, sericeous 
beneath, especially on the nerves; panicles 6-11 cm. long, the 
flowers secund, sessile, the linear bracts 6-16 mm. long; fruit globose, 
3-3.5 mm. in diameter. 

Bertiera guianensis Aubl. In forest or thickets of the Atlantic 
tierra caliente. Southern Mexico to Bolivia and Amazonian Brazil. 
A shrub of 2-4 meters; stipules 5-10 mm. long; leaves short-petiolate, 
8-17 cm. long, acute or acutish at the base; panicles narrow, elongate, 
the bracts linear, 6 mm. long or less; corolla 5-7 mm. long; fruit 
blue, 10-costate when dried, 3.5-4 mm. in diameter. 

BORRERIA Meyer 

Low, annual or perennial herbs, rarely suffrutescent; stipules 
united with the petioles into a setiferous sheath; flowers small, 
usually sessile in the leaf axils or in terminal heads; sepals 2 or 4, 
equal or unequal, often with interposed teeth; corolla very small, 
funnelform, the limb 4-lobate; stamens usually attached in the 
throat of the corolla; fruit a coriaceous or membranaceous capsule, 
2-celled, the cells septicidal; seeds 1 in each cell. A few other species 
are known from Central America. The plants usually are inconspicu- 
ous weeds of waste or cultivated ground. 

Borreria densiflora DC. B. spinosa Cham. & Schlecht. Waste 
ground of the tierra caliente, apparently not common. Widely 
distributed in tropical America. Plants stout, erect, less than a 
meter high, from a taproot, glabrous or nearly so; leaves linear or 
lance-linear, rough-margined, stiff; flower heads large, subtended 
by 4-8 large leaves; hypanthium densely pubescent above, the 2 
lobes linear; corolla white, slightly exceeding the calyx lobes. 

Borreria laevis (Lam.) Griseb. A common weed of waste and 
cultivated ground, often growing in pastures, usually at 1,300 meters 
or less; region of San Ramon; Guanacaste. Widely dispersed in 
tropical America. An annual or perennial, erect or decumbent, 
usually less than 60 cm. high, the branches glabrous, or puberulent 
on the angles; leaves ovate or oblong-ovate, 2.5-5 cm. long, acute 
or acuminate, abruptly decurrent into a slender petiole, more or 
less pilose, at least on the nerves, or glabrate; flower heads sessile, 
terminal and axillary, 8-15 mm. in diameter, the terminal ones 
subtended by large leaves; hypanthium puberulent at the apex; 
sepals minute, broadly triangular; corolla white, 2-2.5 mm. long; 
capsule 2 mm. long; seeds transverse-sulcate. One of the most com- 
mon rubiaceous plants of tropical America. 



FLORA OF COSTA RICA 1275 

Borreria latifolia (Aubl.) Schum. Mielcillo (Tonduz). Fre- 
quent in waste ground or pastures of the tierra caliente, ascending 
to about 1,000 meters; region of San Ramon. Widely distributed 
in tropical America. Annual or perennial, prostrate or ascending, 
conspicuously yellowish when dried, the stout branches 4-angulate, 
often narrowly winged above, glabrous, or pubescent on the angles; 
leaves thick, elliptic to oblong, almost sessile, 3.5-7 cm. long, acute 
or acuminate, acute at the base, more or less scaberulous or puberu- 
lent, at least on the nerves; hypanthium glabrous or pilose; flowers 
densely glomerate in the leaf axils; hypanthium glabrous or pilose; 
sepals 4, ovate-lanceolate, acute, ciliate, 2 mm. long. 

Borreria ocimoides (Burm.) DC. Chiquizacillo. Common in 
waste or cultivated ground, Meseta Central to the coasts, at 1,300 
meters or less; often growing in sand along stream beds; Cocos 
Island. Generally distributed in tropical America. A slender 
annual, usually much branched, 60 cm. high or less, erect or prostrate, 
the stems glabrous, or puberulent on the angles; leaves elliptic to 
linear-lanceolate, 1-3 cm. long, acute, attenuate at the base to a 
short petiole, or sessile, glabrous, or somewhat scabrous near the 
margins; flowers densely glomerate in the leaf axils, the heads 5 mm. 
in diameter; hypanthium glabrous or puberulent; sepals subulate, 
about equaling the hypanthium ; corolla white, shorter than the calyx ; 
capsule 1 mm. in diameter. The flower heads are much smaller than 
in the other species listed here. 

Borreria suaveolens Mey. B. tenella Cham. & Schlecht. Fre- 
quent in pastures and savannas of the Pacific slope, usually at 1,000 
meters or less; Nicoya; also in the region of Cartago. Widely dis- 
tributed in tropical America. Plants perennial, erect, stout, usually 
60 cm. high or less, commonly much branched, densely pubescent 
or glabrate; leaves linear or lance-linear, the margins often revolute, 
2-8 cm. long; flower heads 8-12 mm. in diameter, chiefly terminal 
and subtended by numerous leaves; sepals 4, linear-subulate, longer 
than the hypanthium, ciliate; corolla white, 2.5-5 mm. long; capsule 
oblong, pubescent at the apex; seeds not sulcate. This species, as 
interpreted by Schumann and the present writer, is a highly variable 
one, and it and its relatives are in need of careful revision. The 
Costa Rican material is uniform in its characters, and careful study 
may show it to represent a species distinct from the South American 
B. suaveolens. There probably are numerous names, now considered 
synonyms, available for it. 



1276 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Borreria verticillata (L.) Mey. I have seen only a single 
Costa Rican specimen apparently referable here, although its condi- 
tion makes the record uncertain: Between Guachipelin and Volcan 
de La Vieja, Guanacaste, Brenes 15539. Widely distributed in 
tropical America. Very much like B. suaveolens in general appear- 
ance, but the sepals only 2; plants usually glabrous or glabrate, the 
flower heads substantially smaller than in that species. 

BOUVARDIA Salisb. 

Herbs or shrubs, more or less pubescent; leaves opposite or 
verticillate; stipules connate with the petioles to form a sheath, 
this entire, laciniate, or aristate; flowers usually large and rather 
showy, white, yellow, or red, cymose or cymose-corymbose; calyx 
4-lobate, the lobes short or elongate, persistent; corolla tubular or 
salverform, the 4 lobes short or elongate, valvate; ovary 2-celled, 
many-ovulate; capsule didymous-globose, loculicidally bi valvate; 
seeds imbricate, peltate, compressed, winged. At least one other 
species has been collected in Central America. About 30 species 
are found in Mexico, some of them such handsome plants that they 
have been introduced into cultivation in Europe and elsewhere. 

Bouvardia glabra Polak. Linnaea 41 : 565. 1877. Jazmin, Jaz- 
min de la Virgen. B. glabra var. gracilis Polak. op. cit. 566 (thickets 
and hedges, San Jose", Polakowsky 172). In thickets, San Jose", Pola- 
kowsky (photos, of types of species and variety in Herb. Field Mus.). 
Common in cultivation in gardens of the Meseta Central; collected 
by the writer at Escazu. Mountains of Guatemala. A shrub of 
1.5-3 meters; leaves opposite, short-petiolate, ovate or lance-ovate, 
3-7 cm. long, acute to long-acuminate, obtuse or short-acuminate 
at the base, glabrous, paler beneath; cymes terminal or axillary, 
mostly 3-5-flowered, the pedicels 2-12 mm. long; hypanthium gla- 
brous; calyx lobes lanceolate or oblanceolate, foliaceous, 4-12 mm. 
long; corolla white, glabrous, the slender tube 1.5-3.5 cm. long, the 
lobes oblong or elliptic, 6-11 mm. long; capsule globose, 6 mm. in 
diameter. It is rather doubtful whether this plant is really native 
in Costa Rica, for there are no recent collections from plants that 
were certainly wild. It is quite possible that it did grow in forest 
areas that have been cleared for cultivation, and that it is now 
extinct in Costa Rica in a wild state. It is quite possible that the 
plant was brought to Costa Rica from Guatemala long ago and 
planted in gardens. The shrub is a rather handsome one, its flowers 
fragrant. 



FLORA OF COSTA RICA 1277 

CALYCOPHYLLUM DC. 

Large trees; stipules caducous; flowers small, 4-8-parted, in 
terminal cymes, the bracts and bractlets large, membranaceous, 
enclosing the buds; calyx almost obsolete or of short segments, one 
of the segments in some of the flowers expanded into a large, colored 
limb; corolla short-funnelform, lobed to the middle or more deeply, 
the lobes imbricate, the throat villous; stamens long-exserted ; capsule 
elongate, costate, 2-celled, septicidally bivalvate; seeds small, winged. 
A single species occurs in Central America. 

Calycophyllum candidissimum (Vahl) DC. SurrA, Salamo, 
Madrono. Forests of the Pacific coast, especially in Guanacaste, 
at low elevations. Southern Mexico to Cuba and Colombia. A 
tree 20 meters or less, the branchlets reddish brown; stipules tri- 
angular or lanceolate, 1 cm. long or less; leaves slender-petiolate, 
oval to ovate or elliptic, 5-13 cm. long, abruptly acuminate or 
cuspidate-acuminate, acute or obtuse and abruptly decurrent at the 
base, glabrous above, paler beneath and usually sparsely hirsute 
on the costa; hypanthium 2.5-3 mm. long, hirtellous or glabrate; 
expanded blade of the calyx creamy white, subreniform to ovate- 
rhombic, 1.5-3 cm. long, rounded at the apex; corolla white, the 
tube 3 mm. long, the lobes slightly longer; capsule 6-10 mm. long. 
When in flower, the tree is a conspicuous and handsome one, because 
of the great abundance of white calyx lobes, which give it the appear- 
ance of being covered with white flowers. The bracts retain their 
original coloring for a long time, but finally turn brown. The wood 
is pale brown, hard, heavy, strong, highly elastic, fine-textured, 
usually straight-grained, easy to work, finishes very smoothly. 
Under the name lancewood it is imported into the United States 
from Cuba, for archery bows. In Costa Rica and elsewhere in Central 
America it is utilized for tool handles and many other purposes. In 
some regions of the Pacific coast fine-toothed combs are made from it. 

CEPHAELIS Swartz 

Herbs, shrubs, or small trees; stipules free or connate, usually 
persistent; flowers capitate, the head subtended by an involucre 
of usually sessile, free or connate, often colored bracts, terminal 
or axillary; calyx short or elongate, 4-7-dentate, persistent; corolla 
funnelform or salverform, the tube straight, usually elongate, the 
throat villous or naked, the 4-5 lobes valvate in bud; ovary 2-celled; 
fruit drupaceous, containing 2 hard, smooth or costate nutlets, these 



1278 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

usually longitudinally sulcate on the inner face. Other species are 
known from Central America. 

Cephaelis axillaris Swartz. Frequent in forests of the central 
region, especially south of Cartago ; regions of Dota and San Ramon ; 
mostly at 1,000-1,500 meters. Honduras to Colombia and West 
Indies. A glabrous shrub, 1-1.5 meters high, sparsely branched; 
stipules bilobate, the lobes obtuse or acute; leaves petiolate, obovate- 
oblong, abruptly short-acuminate, acute or attenuate at the base; 
flower heads axillary, sessile, about 1 cm. in diameter or somewhat 
larger, the small bracts green or purplish; corolla white; fruits blue. 
Easily recognized among local species by the numerous small, 
sessile heads. 

Cephaelis chlorochlamys Standl., sp. nov. Frutex, ramulis 
gracilibus teretibus viridibus, internodiis brevibus vel elongatis; 
stipulae persistentes virides in vaginam subinflatam 6 mm. longam 
connatae, vagina in lobos erectos lineari-attenuatos usque 9 mm. 
longos desinente; folia inter minora graciliter petiolata membranacea, 
petiolo usque 2 cm. longo glabro; lamina elliptico-oblonga vel lanceo- 
lato-oblonga 5-11 cm. longa atque 2-3.5 cm. lata vel major longissime 
anguste attenuato-acuminata, basi acuta atque subabrupte decurrens, 
supra viridis glabra, costa gracillima prominente, subtus pallidior, 
glabra vel tantum ad costam obscure pilosula, costa elevata, nervis 
lateralibus utroque latere ca. 20 angulo recto abeuntibus subarcuatis, 
alio vix graciliore inter paria interposito, venulis prominulis laxe 
reticulatis; capitula terminalia longipedunculata dense multiflora 
ca. 1.8 cm. longa atque fere aequilata, pedunculo erecto 2.5 cm. longo 
glabro, bracteis viridibus liberis lanceolatis vel lanceolato-ovatis 
extus glabris intus sparse pilosulis, acute attenuatis adpressis, interi- 
oribus contracto-petiolatis; fructus 5 mm. longus glaber leviter 
bisulcatus atque costatus, calyce persistente 1 mm. longo acute 
profunde 5-dentato. La Hondura de San Jose", 1,300 meters, August, 
1933, Manuel Valeria 704 (type in Herb. Field Mus.). The single 
specimen is not in perfect condition for study, and the description 
may need revision in some details when better material is available. 

Cephaelis discolor Polak. Linnaea 41: 572. 1877. Evea guapi- 
lensis Standl. Journ. Wash. Acad. Sci. 15: 104. 1925 (wet forest near 
Guapiles, Prov. Limon, 500 meters, Standley 37025}. C. guapilensis 
Standl. Field Mus. Bot. 4: 295. 1929. In forest, Angostura, Polakow- 
sky 384 (photo, of type in Herb. Field Mus.). Known only from the 
localities mentioned. Endemic. Plants simple, suffrutescent, a 



FLORA OF COSTA RICA 1279 

meter high or less, the stems glabrous; stipules 1 cm. long or more, 
the lobes linear from a triangular base; leaves slender-petiolate, the 
blades elliptic-oblong, 14-21 cm. long, 6.5-9.5 cm. wide, abruptly 
short-acuminate at each end, glabrous above, paler beneath, sparsely 
hirtellous on the nerves; flower heads terminal, solitary, subsessile, 
2 3 cm. in diameter, the bracts purplish green, rounded and apiculate 
at the apex, covered on both surfaces with slender, multicellular 
hairs; fruit villous. In combining the two species listed I have had 
to depend upon photographs of the types and a fragment of C. dis- 
color, from the Berlin herbarium. While there is a faint possibility 
that two species are involved, I believe the reduction of C. guapilensis 
is correct, especially since both types were collected in the same region. 
Polakowsky describes the plant as "glaberrima," but erroneously, 
as shown by the type fragment. 

Cephaelis elata Swartz. Cephaelis costaricensis Schlecht. 
Linnaea 28: 546. 1856 (type from Costa Rica, the locality not indi- 
cated). Cephaelis Phoenicia Bonn. Smith, Enum. PI. Guat. 5: 39. 
1899, nomen. Uragoga phoenicia Schum. ex Bonn. Smith, loc. cit. 
in syn. Frequent in forests of the central region, at 1,550 meters or 
less, also in the tierra caliente; region of San Ramon; Atlantic slope 
of Guanacaste. Mexico to West Indies and Colombia. A glabrous 
shrub 1.5-3.5 meters high; leaves short-petiolate, oblong-lanceolate 
to elliptic-oblong, 10-25 cm. long, acuminate, acute to attenuate at 
the base; flower heads terminal, long-pedunculate, the 2 outer bracts 
2.5-4 cm. long, broadly rounded at the apex, dark red, rarely green; 
corolla white. The plant is a showy and handsome one. Cephaelis 
elata f. lutea Standl., f. nov. (a forma typica rubrobracteata non 
nisi bracteis luteis differt) is distinguished by having yellow rather 
than bright red bracts. It was collected at San Isidro de El General, 
750 meters, H. E. Stork 3084 (type in Herb. Field Mus.). 

Cephaelis glomerulata Bonn. Smith. Tortuguero, Prov. 
Limon, Lankester 1184- Extending to Guatemala. A glabrous shrub ; 
stipules small, truncate and with minute lobes; leaves short-petiolate, 
lance-oblong, long-acuminate, acute at the base, lustrous; flower 
heads terminal, sessile or nearly so, about 2 cm. in diameter, the 
numerous closely imbricate bracts very broadly rounded at the apex, 
pearly white, edged with violet-purple. 

Cephaelis latistipula Standl. Journ. Wash. Acad. Sci. 18: 281. 
1928. Moist forest, Orosi, Prov. Cartago, Standley 39695. Collected 
also at El Muneco, 1,500 meters. Endemic. Plants simple, suffrutes- 



1280 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

cent, 1 meter high or less, glabrous; stipules distinct, 14-22 mm. long 
and nearly or quite as wide, conspicuously nerved, short-bilobate at 
the apex; leaves slender-petiolate, oval-elliptic to broadly elliptic, 
14-21 cm. long, 7-9 cm. wide, rounded or very obtuse at the apex and 
abruptly short-acuminate, narrowed to the acute base; flower heads 
axillary, dark red, very dense, sessile or nearly so, about 1 cm. long 
and 2 cm. broad, the outer bracts broadly oblong, obtuse; corolla 
5 mm. long. 

Cephaelis Pittieri Krause, Bot. Jahrb. 54: Beibl. 119: 45. 1916. 
Canas Gordas, 1,100 meters, Pittier 11162 (photo, of type in Herb. 
Field Mus.). Collected also at San Isidro del General, 750 meters, 
in deep forest. A shrub or small tree, the branches glabrous; stipules 
united into a tube 2-2.5 mm. long, this bearing narrow setae 5-6 mm. 
long; leaves slender-petiolate, oblong-lanceolate, 14-20 cm. long, 
3-4.5 cm. wide, narrowly long-acuminate, attenuate at the base, 
thin, glabrous or with a few appressed hairs beneath on the nerves; 
flower heads terminal, short-pedunculate, somewhat 3-lobate, 
densely many-flowered, about 2 cm. long and 2.5-4 cm. wide, the 
bracts red, the outer ones ovate, acuminate, sparsely pilose. 

Cephaelis tomentosa (Aubl.) Vahl. Forests of the tierra 
caliente, at 900 meters or less, common in many localities. Mexico 
to Bolivia and Brazil. A slender shrub of 1-3 meters, sparsely 
branched, hirsute or at least pilose on almost all parts; leaves thin, 
lanceolate to ovate-elliptic, large, long-acuminate, acute to attenuate 
at the base; flower heads large, terminal, pedunculate, the bright red, 
acute or short-acuminate bracts much exceeding the flowers; corolla 
yellow; fruit blue. The shrub is an exceptionally showy and rather 
handsome one. It has a wide range in tropical America, and in 
most herbaria is represented by more specimens than perhaps any 
other tropical American member of the family. 

Cephaelis Tonduzii Krause, Bot. Jahrb. 54: Beibl. 119: 45. 
1916. Tuis, 650 meters, Tonduz 11461 (photo of type in Herb. Field 
Mus.). Stems simple, suflrutescent, 15-30 cm. high, glabrous; 
stipule tube short, with linear lobes; leaves long-petiolate, lanceolate 
or oblong-lanceolate, 14-20 cm. long, 6-7.5 cm. wide, long-acuminate, 
narrowed to the base, glabrous; flower heads terminal, subglobose, 
2.2-3 cm. broad, the peduncle 2-3 cm. long, the bracts red, broadly 
ovate, short-acuminate, sparsely short-pilose. This is probably 
synonymous with C. discolor. 



FLORA OF COSTA RICA 1281 

CHIMARRHIS Jacq. 

Trees; stipules caducous; leaves short-petiolate; flowers small, in 
axillary, long-pedunculate, corymbose cymes; hypanthium campanu- 
late, the 5 calyx lobes very short, persistent; corolla short-funnelform, 
the tube villous in the throat, the 5 lobes valvate; stamens exserted; 
ovary 2-celled, many-ovulate; fruit capsular, small, bisulcate, septi- 
cidally bivalvate from the apex; seeds angulate-globose. Two other 
Central American species are found in Panama. 

Chimarrhis parviflora Standl. Trop. Woods 11: 26. 1927. 
Manwood. Yema de huevo (Panama). Changuinola Valley, Atlantic 
coast, Cooper & Slater 120. Collected at various localities in the low- 
lands of Panama, and doubtless extending into Costa Rica. A tree 
of 18 meters, the trunk 25 cm. in diameter, with low buttresses; 
branches minutely puberulent or sericeous; stipules linear-lanceolate, 
1.5-2.5 cm. long; leaves oblong-obovate, 8-15 cm. long, 2.5-6 cm. 
wide, acute or abruptly acute, acute to abruptly attenuate at the 
base, glabrous above, densely sericeous beneath when young but in 
age glabrate; cymes 2-7 cm. broad, very dense and many-flowered, 
pedunculate, the branches densely puberulent; calyx and hypanthium 
together 1 mm. long; corolla 2 mm. long, glabrous outside. 

GHIOCOCCA P. Br. 

Shrubs or small trees, often scandent or sarmentose, usually 
glabrous; leaves petiolate, membranaceous to coriaceous; stipules 
broad, usually cuspidate, persistent; flowers small, whitish, pedi- 
cellate, in axillary, simple or paniculate, often secund racemes; 
hypanthium compressed, the calyx 5-lobate, persistent; corolla 
funnelform, glabrous in the throat, the 5 lobes spreading or reflexed, 
valvate; ovary 2-celled, the cells 1-ovulate; fruit small, drupaceous, 
white at maturity, usually orbicular and strongly compressed. 
No other species are known from Central America. 

Chiococca alba (L.) Hitchc. C. racemosa L. Frequent in 
thickets and forest, Meseta Central to the coasts, at 1,300 meters or 
less. Generally distributed in tropical America. A shrub, the 
branches usually recurved or trailing, often scandent; stipules 1-2 
mm. long; leaves short-petiolate, very variable in form, usually 
ovate or oval-ovate, sometimes lanceolate, mostly 3-6 cm. long, 
acute or short-acuminate, rounded and short-decurrent at the base, 
glabrous; racemes few-many-flowered, the flowers short-pedicellate; 
corolla white or yellowish, 6-8 mm. long, the short lobes triangular; 



1282 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

stamens included; fruit white, juicy, strongly compressed, 4-8 mm. 
in diameter. Called Lagrimas de Maria and Lagrimas de San Pedro 
in other parts of Central America. 

Chiococca phaenostemon Schlecht. Chiraquilla. Frequent in 
forests of the central region; El General; regions of San Ramon and 
Dota; at 750-2,000 meters. Mexico to Panama. A shrub or tree, 
sometimes 14 meters high, glabrous or nearly so, reported to be 
sometimes a large vine; leaves slender-petiolate, 5-12 cm. long, 
coriaceous; corolla white or yellowish, 5-8 mm. long; filaments and 
anthers conspicuously exserted. The trunk is sometimes 60 cm. in 
diameter at the base; bark thin, light to medium brown, nearly 
smooth; flowers fragrant. 

CHOMELIA Jacq. 

Shrubs or small trees, usually armed with stout, axillary spines, 
the leaves petiolate; stipules acuminate, deciduous; flowers small, 
white, in pedunculate, axillary cymes; calyx 4-lobate, the lobes 
narrow, equal or unequal, persistent; corolla salverform, with a 
slender, elongate tube, the 4 lobes elongate, valvate or with sub- 
imbricate margins; ovary 2-celled, the cells 1-ovulate; fruit small, 
drupaceous, the stone oblong, 2-celled, osseous. A single species is 
known from North America. 

Chomelia spinosa Jacq. Malacahuite. Guettarda costaricensis 
Schum. ex Tonduz, Bull. Herb. Boiss. 3: 7. 1895, nomen. Frequent 
in thickets of the Pacific tierra caliente. Southern Mexico to northern 
Brazil. A shrub or tree, sometimes 9 meters high, the branches 
reddish brown; stipules 4-8 mm. long; leaves often crowded on short, 
lateral branches, ovate-orbicular to oval or elliptic, 4-8 cm. long, 
abruptly short-acuminate, rounded to acute at the base, sparsely 
pilose above, pilose beneath, especially on the nerves, with appressed 
or spreading hairs; cymes few-flowered, slender-pedunculate, the 
flowers sessile or nearly so; calyx and hypanthium 2-3 mm. long, 
the calyx lobes lanceolate or triangular-subulate, half as long as the 
hypanthium or shorter; corolla yellowish white, sericeous outside, 
the tube 12-22 mm. long, the lobes 4-6 mm. long, each with a slender, 
hornlike appendage at or just below the apex; fruit black and juicy at 
maturity, 9-12 mm. long, sparsely strigillose. It is stated that the 
fruits are edible, and are employed for preparing refrescos. It is 
reported also that squirrels and pisotes eat the fruits and grow fat 
upon them at the principal fruiting season, in November. The fruits 
are said to be used as a remedy for fevers. The flowers are fragrant. 



FLORA OF COSTA RICA 1283 

CINCHONA L. 

Trees or shrubs, glabrous or variously pubescent; stipules large, 
distinct, caducous; leaves large or small, petiolate; inflorescence 
terminal, paniculate, with opposite branches, usually many-flowered, 
the flowers 5-parted; calyx campanulate, dentate or lobate; corolla 
salverform, pubescent outside, the spreading lobes valvate in bud, 
more or less villous within on the margins; stamens semi-exserted or 
included, the anthers linear; capsule ovoid or oblong to subcylindric, 
bisulcate, dehiscent from the base to the apex; seeds numerous, 
broadly winged. A single species is native in North America, the 
others inhabiting the South American Andes. The genus contains 
some of the most important drug plants of the earth, source of quinine, 
the universal remedy for malaria. 

Cinchona pubescens Vahl. Forests of the region of San Ramon, 
at about 1,100 meters; Tiquires, 1,700 meters. Extending to Bolivia. 
A medium-sized tree, the branchlets pubescent; stipules ovate, 
obtuse or acute, sericeous or almost glabrous; leaves slender-petiolate, 
large, the blades broadly ovate to orbicular, rounded to acute at the 
apex, cordate to acute at the base and often decurrent, usually gla- 
brate above, beneath densely short-pilose or tomentose to glabrate; 
panicles large, often leafy, many-flowered, the flowers subsessile; 
hypanthium densely sericeous; calyx teeth short and broad, acute; 
corolla red or pink, sericeous, 10-12 mm. long, the lobes half as long 
as the tube; capsule lanceolate or oblong, glabrate, usually 1.5-2.5 
cm. long. One of the forms of this species, cultivated in the East 
Indies, is probably the principal source of quinine. No information 
is available as to Costa Rican vernacular names, if any, or regarding 
any local knowledge of the tree's properties. In North America the 
species is known only from Costa Rica, and there, apparently, it has 
been collected only in recent years. 

COCCOCYPSELUM Swartz 

Prostrate, annual or perennial herbs, usually pubescent; leaves 
petiolate, small; stipules small, solitary; flowers small, purplish or 
blue, capitate, the heads axillary, solitary, sessile or pedunculate, 
few-many-flowered; calyx 4-lobate, the lobes narrow, persistent; 
corolla funnelform, the throat glabrous, the limb 4-lobate, the lobes 
oblong, valvate; ovary 2-celled, many-ovulate; fruit baccate, ovoid, 
juicy, blue; seeds small, subangulate, granulate. A few other species 
have been found in Central America. 



1284 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

i 

Coccocypselum guianense (Aubl.) Schum. Occasional in 
forests of the central region, chiefly at 1,400-1,800 meters. Southern 
Mexico to West Indies and Brazil. Stems densely pilose with long, 
spreading, soft hairs; leaves oval to ovate or rounded-ovate, 2.5-7 cm. 
long, rounded to subacute at the apex, truncate to obtuse at the 
base, densely pilose on the upper surface with long and short, mostly 
ap pressed hairs, often purplish beneath, densely soft-pilose; heads 
pedunculate, mostly 2-4-flowered; calyx lobes 3-4 mm. long; corolla 
blue, 7-10 mm. long; fruit pilose, prussian blue, 6-11 mm. in 
diameter. 

Coccocypselum hirsutum Bartl. Frequent in forests of the 
central region, chiefly at 750-1,800 meters; region of San Ramon; 
Buenos Aires. Mexico to Bolivia. Branches densely hirsute or 
hispid; leaves oval to oblong-ovate, rounded to subacute at the apex, 
rounded at the base, hirsute or hispid above with long, spreading 
hairs, hirsute beneath; heads 3-5-flowered, pedunculate; calyx lobes 
2.5-5 mm. long; corolla hirsute, 12-14 mm. long; fruit violet-blue, 
6-10 mm. in diameter. 

Coccocypselum hispidulum Standl. Tontanea hispidula 
Standl. In forest, Pejivalle, 900 meters; doubtless also in other 
localities. Panama. Branches slender, pilose with stiff, whitish, 
appressed or ascending hairs; leaves broadly ovate or oblong-ovate, 
2-3.5 cm. long, acutish to rounded at the apex, rounded and short- 
decurrent at the base, hispid above, sparsely or densely hispid 
beneath; flower heads mostly sessile, sometimes very shortly pedun- 
culate, few-flowered; calyx lobes 2-4 -mm. long; corolla bluish white, 
7-9 mm. long; fruit prussian blue, 8-15 mm. long. 

Coccocypselum lanceolatum (Ruiz & Pavon) Pers. C. canes- 
cens Willd.; Tontanea canescens Standl. Occasional in forests of the 
central region; region of San Ramon; mostly at 1,000-1,400 meters. 
Southern Mexico to Peru and Brazil. Branches densely pilose- 
sericeous when young; leaves oblong to ovate, 2-8 cm. long, acute 
or obtuse, obtuse to subcordate at the base, densely pilose above 
with short, mostly appressed, yellowish hairs, densely pilose beneath 
with longer, appressed or somewhat spreading hairs; heads densely 
many-flowered, the peduncles 1-6 cm. long; calyx lobes oblong or 
ovate, 2-3 mm. long; corolla purple-blue, 5 mm. long; fruit bright 
blue, densely pilose, 5-7 mm. in diameter. This species is frequent 
in the more open oak forests about Santa Maria de Dota. The 
brightly colored berries are handsome and rather showy. 



FLORA OF COSTA RICA 1285 

Coccocypselum pleuropodum (Bonn. Smith) Standl. El 
Muneco; region of San Ramon; 1,000-1,400 meters, in forest. Ex- 
tending to southern Mexico and Cuba. Stems slender, hirsute with 
long, spreading hairs; leaves reniform or orbicular, 1-3.5 cm. long, 
broadly rounded at the apex, cordate or truncate at the base, hirsute 
or hispid above with spreading hairs, often purplish beneath, sparsely 
or densely long-hirsute; heads pedunculate, mostly 2-4-flowered; 
calyx lobes 2-4 mm. long; corolla about 12 mm. long, bright blue; 
fruit densely hirsute. 

COFFEAL. Coffee 

Shrubs or small trees, usually glabrous throughout; stipules 
triangular, persistent; leaves short-petiolate; flowers sessile in the 
leaf axils, 4-7-parted; calyx very short; corolla salverform or funnel- 
form, the lobes dextrorsely contorted in bud; anthers subsessile, 
dorsifixed; fruit fleshy, indehiscent, 2-seeded; seeds semi-oval, flat 
and deeply furrowed on the inner face. No species of the genus 
are native in America. 

Coffea arabica L. Cafe. Native of Abyssinia, but now culti- 
vated generally in most tropical regions of suitable climate. It 
is said to have been introduced into Costa Rica by Governor Acosta 
about 1808, and soon became of great importance in local agriculture, 
being now the most important agricultural product of the country, 
at least for export. In Costa Rica coffee is grown principally in the 
central region, but also in the mountains of Guanacaste and in other 
remote areas, chiefly at elevations of 800-1,400 meters. It is planted 
upon a small scale also at lower elevations, and thrives if there is 
not too abundant moisture. The coffee of Costa Rica is of superior 
quality, and holds an enviable reputation in the markets of the 
world. Wild coffee plants are sometimes found in the forest in remote 
regions, to which the seeds have been carried probably by birds. 

Coffea liberica Hiern. Cafe de Liberia. Liberian coffee, native 
of West Africa, is planted occasionally for experimental purposes, 
or as a curiosity. It is grown extensively for market in some parts 
of the earth. In this species the corolla has 6 or 7 lobes; the some- 
what smaller corolla of C. arabica has only 5 lobes. The fruits of 
C. liberica are larger than those of C. arabica. 

COSMIBUENA Ruiz & Pavon 

Glabrous shrubs or small trees, usually epiphytic; leaves petiolate, 
rather thick and somewhat fleshy; stipules deciduous; flowers large, 



1286 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

terminal, solitary or cymose, pedicellate; hypanthium oblong or 
turbinate, the calyx tubular or campanulate, entire or 5-6-dentate, 
deciduous; corolla salverform or funnelform, the tube elongate, the 
limb 5-6-lobate, the lobes spreading, contorted; anthers included; 
ovary 2-celled, the ovules numerous; capsule narrowly oblong or 
cylindric, coriaceous, 2-celled, septicidally bivalvate from the apex; 
seeds numerous, oblong, produced at each end into a bifid wing. 
Two other species are known from Central America, in Nicaragua 
and Panama. 

Cosmibuena ovalis Standl. Region of El General. Panama. 
An epiphytic shrub or small tree, or sometimes terrestrial and 9 
meters high, glabrous throughout; stipules obovate-oval, 1.5-2 cm. 
long, rounded at the apex; leaves on stout petioles, the blades oval 
to broadly obovate, 6-14 cm. long, rounded or obtuse at the apex, 
acute or attenuate at the base; cymes 3-5-flowered, the stout pedicels 
1-2 cm. long; calyx 11 mm. long, shallowly 5-dentate, the teeth 
deltoid, acute; corolla white or tinged outside with pink, the tube 
6-7 cm. long, 3.5-5 mm. thick, the oblong lobes 2.5-3 cm. long, 
rounded at the apex; capsule oblong, 4.5 cm. long, 10-12 mm. thick. 
A specimen of Hoffmann 717 from Costa Rica, without specific 
locality, was indicated in the Berlin herbarium as a new species, 
still unpublished, by Schumann. 

COUSSAREA Aubl. 

Shrubs or trees, usually glabrous; stipules commonly ovate- 
triangular, muticous, apiculate, or truncate, never subulate-aristate 
nor connate into a long, intrapetiolar sheath; leaves mostly opposite, 
short-petiolate or subsessile, commonly more or less coriaceous; 
inflorescence terminal; calyx cupular and truncate, sometimes 4- 
dentate or rarely 4-lobate; corolla funnelform or salverform, the throat 
naked, the 4 lobes valvate in bud; anthers subsessile in the corolla 
tube; ovary 2-celled, the cells 1-ovulate; fruit baccate, by abortion 
commonly 1-seeded, longer than broad, the seed vertical. Two or 
three other species occur in Central America. 

Coussarea Austin- Smithii Standl., sp. nov. Frutex vel arbus- 
cula 4-5-metralis, trunco usque 10 cm. diam., ramulis sulcatis 
striatisque, novellis dense minute puberulis, internodiis brevibus vel 
elongatis; stipulae persistentes 6-7 mm. longae e basi rotundato- 
ovata abrupte breviter acuminatae, apice brevissime bilobae; folia 
mediocria breviter petiolata crasse membranacea, petiolo crassiusculo 
8-15 mm. longo puberulo; lamina lanceolato-oblonga usque late 



FLORA OF COSTA RICA 1287 

elliptica vel oblongo-obovata 6.5-14 cm. longa 3-6.5 cm. lata acuta 
vel subobtusa, saepe breviter abrupte acuminata, basi acuta usque 
subrotundata, supra in sicco fusca, ad costam dense minute puberula, 
subtus pallidior, ad nervos venasque dense minute pilosula, costa 
gracili elevata, nervis lateralibus utroque latere ca. 11 obliquis angulo 
lato adscendentibus arcuatis, venulis prominulis laxe reticulatis; 
inflorescentia corymbiformis vel thyrsoidea laxe vel dense pauci- vel 
multiflora, vulgo 4-6 cm, longa atque aequilata vel latior, 3-4.5 cm. 
longe pedunculata, ramis primariis subfastigiatis erectis vel adscen- 
dentibus minute puberulis, floribus sessilibus vel subsessilibus; hypan- 
thium ca. 1.5 mm. longum crassum minutissime puberulum, calyce 
aequilongo breviter inaequaliter dentato, dentibus late triangularibus 
obtusis vel acutiusculis; corolla alba extus sat dense minutissime 
pulverulenta, tubo 9 mm. longo superne paullo dilatato, lobis 
paten tibus linearibus attenuatis aequilongis vel paullo longioribus; 
fructus late obovoideus ca. 1 cm. longus. Zarcero, 1,590 meters, 
July, 1937, Austin Smith 4226 (type in Herb. Field Mus.); also 
A230, H226. Forests of La Palma de San Ramon, 1,275 meters, 
August, 1925, Brenes 4400; in 1925 and 1928, Brenes 4411, 5949. 
San Antonio de San Ramon, July, 1927, Brenes 5641- Tapesco de 
Zarcero, 2,000 meters, Austin Smith H427. Probably a good many 
other collections from the central mountains, determined in herbaria 
as C. paniculata (Vahl) Standl., are referable to this species. C. pani- 
culata is native in Trinidad, and apparently is not found in Central 
America. 

Coussarea impetiolaris Bonn. Smith, Bot. Gaz. 37: 418. 1904. 
La Colombiana, Llanuras de Santa Clara, 200 meters, Pittier 7582 
(13408). Panama to British Honduras. A small tree; stipules 4 mm. 
long, rounded, mucronulate; leaves subsessile, obovate-elliptic or 
obovate-oblong, 13-16 cm. long, 5-6.5 cm. wide, cuspidate-acuminate, 
cuneate to the base, the base itself narrowly rounded or obtuse, 
glabrous; inflorescence thyrsoid-paniculate, short-pedunculate, rather 
dense, minutely puberulent or pilosulous, the flowers congested; calyx 
and hypanthium puberulent or pilosulous, the calyx undulate, about 
2 mm. long; corolla white, pilosulous or tomentulose, about 2.5 cm. 
long or sometimes shorter, the tube almost twice as long as the lobes; 
fruit about 2 cm. long. 

Coussarea latifolia Standl. Journ. Wash. Acad. Sci. 18: 281. 
1928. Forests of Tsaki, Talamanca, 200 meters, Tonduz 9574. 
Known only from the original collection. A tree, glabrous through- 



1288 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

out; stipules semiorbicular, 5 mm. long, broadly rounded at the 
apex; petioles 2-2.5 cm. long; blades broadly elliptic or ovate- 
elliptic, 17-25 cm. long, 9-16 cm. wide, rounded and apiculate at 
the apex, acute or obtuse at the base, the lateral nerves about 9 on 
each side; inflorescence cymose-racemose, 6 cm. long, pedunculate, 
the cymules 2-3-flowered, the pedicels 1-4 mm. long; calyx tubular- 
campanulate, 5-7 mm. long, truncate; corolla yellowish white, the 
tube (in bud) 1 cm. long, the lobes 13-15 mm. long. Easily recog- 
nized among Costa Rican species by the very large leaves and calyx. 

Coussarea talamancana Standl., sp. nov. Ramuli crassiusculi 
virides sparse minute puberuli, internodiis brevibus; stipulae deci- 
duae, perfectae non visae; folia magna coriaceo-membranacea, 
breviter petiolata, petiolo 7-10 mm. longo crassiusculo glabro vel 
inconspicue puberulo; lamina late obovato-elliptica 15-20 cm. longa 
9-10 cm. lata, apice acuta et sensim lineari-caudata, acumine ipso 
fere lineari usque 2.5 cm. longo, basi acuta, glabra, supra fusco- 
viridis, costa nervisque prominulis, subtus pallidior, costa pallida 
elevata, nervis lateralibus utroque latere ca. 11 prominentibus angulo 
fere recto divergentibus arcuatis, venulis prominulis laxe reticulatis; 
inflorescentia parva sessilis thyrsoideo-paniculata (bene evoluta non 
visa) dense multiflora, ramis minutissime puberulis, floribus breviter 
pedicellatis; hypanthium pallidum sparse minute puberulum, calyce 
truncate fere 2 mm. longo; corolla in alabastro 7-8 mm. longa extus 
minute puberula apicem versus angustata, lobis tubo paullo longiori- 
bus. Talamanca Valley, June, 1927, G. P. Cooper T4 (type in Herb. 
Field Mus.). Easily recognizable among Central American species 
by the very long and narrow acuminations of the leaves. 

COUTAREA Aubl. 

Shrubs or small trees; leaves petiolate, membranaceous; stipules 
short, acute; flowers large, terminal or axillary, solitary or cymose, 
pedicellate; hypanthium obovoid-turbinate; calyx 5-8-lobate, decidu- 
ous; corolla funnelform-campanulate, more or less oblique, the tube 
often curved and gibbous-ventricose, the throat glabrous, the limb 
5-8-lobate, the lobes short, plicate-imbricate or contorted; anthers 
exserted; ovary 2-celled, many-ovulate; capsule oval or obovoid, 
more or less obcompressed, coriaceous or ligneous, loculicidally 
bivalvate; seeds imbricate, broadly winged. One other species is 
known from northern Central America. 

Coutarea hexandra (Jacq.) Schum. Thickets of the Pacific 
tierra caliente. Southern Mexico to Argentina. A shrub or small 



FLORA OF COSTA RICA 1289 

tree, 5 meters high or less, the branches conspicuously lenticellate, 
usually glabrous; stipules 2-4 mm. long; leaves short-petiolate, ovate 
to elliptic or oval, 5-12 cm. long, acute or cuspidate-acuminate, 
rounded to acute at the base, glabrous or nearly so, or sometimes 
rather densely short-pilose beneath; flowers mostly in 3-flowered, 
terminal and axillary cymes; calyx lobes 6, lance-subulate, 5-12 mm. 
long; corolla white or yellowish, often tinged with purple, 6-7.5 cm. 
long, the short lobes acute or obtuse; capsule 2.5-3.5 cm. long, 2 cm. 
wide, dark brown. Called Quina in Salvador, where the bitter bark 
is employed as a substitute for quinine in the treatment of fevers. 

CRUSEA Cham. & Schlecht. 

Annual or perennial herbs, usually low, the stems more or less 
tetragonous; leaves small, ovate or lanceolate, conspicuously nerved; 
stipules connate with the petioles to form a ciliate sheath; flowers 
small or medium-sized, usually pink, in dense heads surrounded by 
4 foliaceous bracts; hypanthium compressed; calyx lobes 4, elongate- 
subulate, alternating with minute teeth, persistent; corolla funnel- 
form, the slender tube glabrous in the throat, the lobes 4, spreading, 
valvate; anthers exserted; ovary 2-celled, the cells 1-ovulate; fruit 
capsular, didymous, 2-coccous, the cocci indehiscent, separating 
from the persistent axis. Most species of the genus are Mexican, 
but one or two others occur in northern Central America. 

Crusea brachyphylla Cham. & Schlecht. In potrero, Laguna 
de La Chonta, northeast of Santa Maria de Dota, 2,000 meters, 
Standley 42231. Ranging to Mexico. Plants annual, erect, 30 cm. 
high or less, the stems hispidulous, densely leafy; leaves oblong to 
oblong-ovate, 1-3 cm. long, obtuse or acute, usually abundantly 
hispidulous on both surfaces; corolla tube very slender, about 8 mm. 
long, minutely scaberulous or almost glabrous. The plant was 
plentiful in the Costa Rican locality, where it is perhaps introduced. 

Crusea coccinea DC. Brushy slope, above Los Lotes, north of 
El Copey, 2,100-2,400 meters, Standley 42572. Panama to Mexico. 
An ascending herb, the stems sometimes a meter long, glabrous; 
leaves petiolate, ovate or lance-ovate, 3-5 cm. long, long-acuminate, 
acuminate at the base, almost glabrous above, scabrous beneath on 
the nerves; corolla pale red, about 3 cm. long. 

Crusea parviflora Hook. & Arn. Dry forest or thickets, tierra 
caliente of Guanacaste. Extending to Mexico. Plants erect and 
often much branched, less than a meter high, the branches pale, 



1290 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

pilose with short, soft, white hairs; leaves lanceolate, mostly 2-4 cm. 
long, acute to long-acuminate, rather densely hispidulous on both 
surfaces; flower heads about 1 cm. in diameter, very dense and many- 
flowered; corolla white, scarcely over 3 mm. long. 

DECLIEUXIA HBK. 

Perennial herbs or low shrubs, usually fuscous when dried; 
stipules subulate-linear or obsolete, often decurrent; leaves opposite 
or verticillate, usually small and sessile, commonly coriaceous; 
flowers small, cymose, 4-parted; sepals free or nearly so and often 
remote; corolla funnelform, the lobes valvate; fruit didymous, 
indehiscent, 2-celled, the cells 1-seeded, laterally compressed. A 
single species is known from North America. 

Declieuxia fruticosa (Willd.) Kuntze, var. mexicana (DC.) 
Standl. D. mexicana DC. In grassy places, often in savannas, 
Pacific tierra caliente. Panama to Mexico, the typical form of the 
species occurring in northern South America. Plants herbaceous, 
usually with several simple or branched stems, 30 cm. high or less, 
glabrous throughout or nearly so; leaves subsessile, narrowly lance- 
oblong or linear-oblong, mostly 2-3 cm. long, acute or subobtuse, 
narrowed to the subacute base; cymes small and few-flowered, 
pedunculate, with small, persistent, linear bracts; corolla white, 
5-6 mm. long; fruit lustrous. The plant of Central America and 
Mexico differs so little, if at all, from that of Colombia and Venezuela 
that it scarcely deserves varietal designation. 

DEPPEA Schlecht. & Cham. 

Shrubs, more or less pubescent; leaves opposite, membranaceous; 
stipules small, deciduous; flowers small, bright yellow, in axillary 
or terminal cymes or umbels, pedicellate; hypanthium turbinate or 
hemispheric, the calyx 4-lobate, the lobes persistent; corolla rotate 
or short-funnelform, the throat glabrous, the 4 lobes spreading, 
contorted; anthers exserted; ovary 2-celled, the ovules numerous; 
capsule small, turbinate or obovoid, costate, loculicidal from the 
apex, the valves cleft; seeds numerous, subglobose, minute, granu- 
late. At least one other species is found in northern Central America. 

Deppea grandiflora Schlecht. D. costaricensis Polak. Linnaea 
41: 566. 1877 (Desengano, Polakowsky 148; photo, of type in Herb. 
Field Mus.). In forest, regions of Dota and Desengano, chiefly at 
1,500-1,800 meters. Panama to southern Mexico. A slender shrub, 
1.5-3 meters high, the branches densely puberulent when young; 



FLORA OF COSTA RICA 1291 

stipules minute, deltoid; leaves slender-petiolate, oblanceolate to 
oblong-elliptic, 5-15 cm. long, 1.5-6.5 cm. wide, acuminate or attenu- 
ate at each end, thin, glabrous or sparsely puberulent above, puberu- 
lent or villosulous beneath, especially on the nerves; inflorescence 
cymose-corymbose, usually many-flowered, long-pedunculate, the 
pedicels 1-6 mm. long; calyx lobes deltoid, acute, minute; corolla 
glabrous, 6-8 mm. long, the lobes obtuse; capsule 3-5 mm. long, 
glabrous, 8-costate, obtuse or acute at the base. 

DIDYMAEA Hook. f. 

Slender, fragile, flaccid herbs, the branches tetragonous; leaves 
small, opposite, petiolate; stipules geminate, subulate, persistent, 
finally recurved; flowers minute, axillary, pedicellate, the pedicels 
not articulate with the calyx, in fruit elongate, spreading or recurved; 
calyx entire; corolla glabrous, campanulate or rotate, 4-lobate, the 
lobes triangular, valvate; ovary 2-celled, the cells 1-ovulate; fruit 
didymous, lustrous, the lobes globose, fleshy, one of them often 
abortive. The genus consists of two species, the other Mexican. 

Didymaea alsinoides (Schlecht. & Cham.) Standl., comb nov. 
Nertera alsinoides Schlecht. & Cham. Linnaea 6: 413. 1831. D. 
mexicana Hook. f. Growing in forest, 1,500-2,400 meters; collected 
on the slopes of Barba and in the region of Dota. Ranging to Mexico. 
Plants procumbent or often scandent, the stems a meter long or less, 
glabrous or hispidulous; leaves mostly 1.5-3 cm. long, lanceolate or 
lance-oblong, sometimes ovate, acute to long-acuminate, glabrous or 
hispidulous above, usually glabrous beneath except on the nerves, 
there hispidulous; fruits very lustrous, dull, dark blue, the lobes about 
6 mm. long. From Costa Rica I have seen only three collections that 
seem referable to the presumably typical form of the species. There 
is some question even about these, and I have a suspicion that when a 
larger amount of material has been assembled from Costa Rica and 
Panama, it may be possible to separate varietally or perhaps even 
specifically all the southern collections of Didymaea. 

It is rather strange that the name Nertera alsinoides should have 
been overlooked so long, but it is not listed in the Biologia Centrali- 
Americana. In the Index Kewensis it is reduced to synonymy 
under Nertera depressa. The species was referred with doubt to Ner- 
tera by Schlechtendal and Chamisso. While I have seen no authentic 
material of Nertera alsinoides, the rather brief description seems to 
apply without any doubt to Didymaea, and I do not hesitate to make 
the transfer. 



1292 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Didymaea alsinoides var. australis Standl., var. nov. A 
forma typica specie! ut videtur non nisi foliis brevioribus atque 
latioribus differt; lamina late ovata usque rotundato-ovata, interdum 
suborbicularis, plerumque 1-1.8 cm. longa, 8-12 mm. lata, apice 
acuta vel abrupte acutata, basi late rotundata vel subtruncata atque 
subito breviter decurrens, supra breviter hispidula vel glabrata, sub- 
tus glabra vel ad nervos hispidula; fructus glaber. La Carpintera, 
1,700 meters, November, 1908, Alfred & A. C. Brade 2143 (type in 
Herb. Field Mus.). El Copey, 2,250 meters, Stork 1540. Oak forest 
near Quebradillas, north of Santa Maria de Dota, 1,800 meters, 
Standley 43027. Viento Fresco, Prov. Alajuela, 1,600-1,900 meters, 
Standley & Torres 47862. Hacienda Montecristo, 1,520 meters, Solis 
462. El Gallito de Heredia, Brenes 21706. PANAMA: Cerro Punta, 
Prov. Chiriqui, 1,500-2,000 meters, Seibert 255. Bajo Chorro, Prov. 
Chiriqui, 1,800 meters, M. E. Davidson 395. Inspection of the numer- 
ous collections of Didymaea from Costa Rica and Panama shows that 
most of them can be associated at a glance, and distinguished from 
the normal Mexican form by their relatively much broader and 
shorter leaf blades. It may be that this plant is a distinct species, 
but I find no satisfactory characters to justify specific rank, and the 
few specimens that approach the Mexican form seem to indicate 
that the form here described deserves no more than varietal status. 

Didymaea alsinoides var. mollis Standl., var. nov. Caules 
elongati dense pilosulo-tomentulosi; folia breviter petiolata, lamina 
oblongo-ovata vulgo 2-3 cm. longa, acuta vel subacuminata, basi 
late rotundata vel truncata, utrinque dense molliter pallido-pilosula; 
fructus dense pilis brevibus incurvis pilosulus. Cerro de Las Vueltas, 
wet forest, an ascending or subscandent herb, the fruit dark blue, 
common, Standley & Valerio 43949 (type in U. S. Nat. Herb.). This 
form is a very distinct one, because of the dense, short, soft pubes- 
cence that covers all parts, including the fruit (the fruit is glabrous 
in all other specimens of the genus examined). With more material 
this may well prove to be a distinct species. 

DIODIA L. 

Annual or perennial herbs, sometimes low shrubs; stipules united 
with the petioles to form a setiferous sheath; leaves opposite, usually 
small and narrow; flowers small or minute, axillary and glomerate or 
spicate or cymose; sepals 2 or 4, equal or unequal; corolla funnelform, 
4-lobate, the lobes valvate; fruit of 2 carpels, these indehiscent, mem- 
branaceous or ligneous, usually separating from the persistent column. 



FLORA OF COSTA RICA 1293 

A few other species are found in Central America, and some of 
them are to be expected in Costa Rica. 

Diodia brasiliensis Spreng. var. angulata (Benth.) Standl. 
Triodon angulatum Benth. Region of San Ramon, 1,000-1,200 
meters, in pastures or forest, usually along or near streams. Extend- 
ing to Mexico; typical form of the species in Brazil and southward. A 
slender shrub, 1 meter high or less, often densely branched, glabrous 
or sparsely puberulent; leaves oblong to elliptic, 5-20 mm. long, 
often appearing verticillate, obtuse or acute, short-petiolate; flowers 
very small, white, densely clustered in the axils of the upper leaves: 
calyx 4-dentate; corolla 2 mm. long. 

Diodia teres Walt. Nicoya. Widely distributed in tropical 
and temperate America; in Central America usually growing in 
grassland. An erect annual, simple or branched, usually 30 cm. 
high or less, hirsute or pilose; leaves linear or lance-linear, 1.5-4.5 cm. 
long, thick-margined; flowers axillary, solitary or geminate, sessile 
or nearly so; corolla 3-4 mm. long, white or pink; fruit 3 mm. long, 
usually hispidulous. 

DUROIA L. f. 

Shrubs or trees; leaves opposite or verticillate, sessile or petiolate; 
stipules oblong, deciduous; flowers often large, white or yellowish, 
dioecious, in terminal fascicles or cymes, the pistillate flowers rarely 
solitary; hypanthium oblong to hemispheric, the calyx cupular or 
tubular, persistent, truncate or 6-9-lobate; corolla salverform, 
sericeous outside, the throat pilose or naked, the limb 6-9-lobate, the 
lobes oblong, contorted; anthers included; ovary 2-4-celled, many- 
ovulate; fruit baccate, globose to oblong, 1-4-celled; seeds large, 
horizontal, compressed, embedded in pulp. No other species are 
known from North America. 

Duroia costaricensis Standl. Contr. U. S. Nat. Herb. 20: 208. 
1919. Sierpe, Pacific coast, Pittier 6803. Known only from the 
original collection. Branches hirsute; leaves opposite, short-petio- 
late, oblong-obovate, 10-17 cm. long, 3.5-6.5 cm. wide, obtuse and 
cuspidate-acuminate, cuneately narrowed to the base, abundantly 
hirsute; staminate flowers fasciculate-cymose, short-pedicellate; 
calyx and hypanthium densely hirsute, the calyx 4-4.5 mm. long, 
the 6-7 lobes linear-subulate, equaling the tube; corolla in bud 
14 mm. long, densely sericeous outside, the lobes longer than 
the tube. 



1294 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ELAEAGIA Wedd. 

Trees or shrubs, usually pubescent; stipules free, or connate at 
the base; leaves petiolate, usually large; flowers small, in terminal 
racemes or panicles; hypanthium small, hemispheric, sulcate; calyx 
5-lobate, persistent; corolla short-funnelform, the throat villous, the 
limb 5-lobate, the lobes oblong, recurved, contorted in bud; ovary 
2-celled, many-ovulate; capsule small, 2-celled, loculicidally bivalvate, 
the valves finally bifid; seeds minute, elongate. No other species 
are known from Central America. 

Elaeagia auriculata Hemsl. Diag. PI. Nov. 32. 1879. At 1,350 
meters, without definite locality, Endres 95. Mountains south of 
Cartago; region of San Ramon; at 1,100-1,500 meters. Honduras. 
A shrub or tree of 3-8 meters, with few branches, the branchlets 
tetragonous, densely tomentulose; stipules very large, at least 5 cm. 
long; leaves sessile or nearly so, ovate-elliptic to rounded-elliptic- 
obovate, as much as 40 cm. long and 23 cm. wide but often smaller, 
acute or abruptly short-acuminate, narrowed to the auricled base, 
densely and softly short-pilose, especially beneath; flowers white, 
in large, broad, sessile panicles; calyx and hypanthium puberulent, 
together scarcely more than 2 mm. long; corolla 3-4 mm. long, 
glabrous outside, lobed almost to the base; capsule subglobose, 3 mm. 
in diameter. 

Elaeagia Karstenii Standl. Tapanti, 1,300 meters, M. Valeria 
1665. Venezuela and Colombia. A tree, the branchlets hirtellous 
or glabrate; stipules caducous, about 3 cm. long, obtuse or rounded 
at the apex; leaves short-petiolate, the blades broadly elliptic or 
obovate-elliptic, 13-25 cm. long, short-acuminate, acute at the base, 
puberulent, pilosulous, or glabrate; inflorescence broadly paniculate; 
calyx glabrous, 1.2 mm. long, shallowly 5-lobate; corolla 3-3.5 mm. 
long; capsule globose, 2 mm. long. The single Costa Rican specimen 
is an incomplete one, almost surely referable to this genus. It repre- 
sents a species evidently different from E. auriculata, but probably 
it is not properly referable to E. Karstenii. 

EXOSTEMA Rich. 

Shrubs or trees, glabrous or pubescent, the branches usually 
terete; stipules entire or bifid, persistent or deciduous; leaves petiolate 
or subsessile, membranaceous or coriaceous; flowers small or large, 
axillary and solitary or in terminal panicles or corymbs; hypanthium 
cylindric or obovoid; calyx usually 5-lobate; corolla tube often 
greatly elongate, the lobes oblong to elongate-linear, imbricate; 



FLORA OF COSTA RICA 1295 

anthers commonly exserted; ovary 2-celled, many-ovulate; capsule 
usually oblong-cylindric, septicidally bivalvate, the valves entire or 
biparted; seeds imbricate, compressed, winged. One other species is 
found in northern Central America. 

Exostema caribaeum (Jacq.) Roem. & Schult. E. longicuspe 
Oerst. Vid. Medd. Kjoebenhavn 48. 1852 (Puntarenas, Oersted). 
Thickets of the Pacific coast. Extending to Mexico, West Indies, 
and southern Florida. A shrub or small tree, the branchlets glabrous; 
stipules 2.5-5 mm. long, the lobes cuspidate; leaves slender-petiolate, 
mostly ovate to elliptic-oblong, 5-11 cm. long, abruptly acuminate 
or long-acuminate, obtuse or acute at the base, membranaceous, 
barbate beneath in the axils of the nerves, otherwise glabrous; 
flowers axillary, solitary, white, pedicellate; calyx lobes 1 mm. long 
or shorter; corolla glabrous, the tube 3-5 cm. long, 2 mm. thick, the 
5 lobes linear, about equaling the tube; anthers linear, 2 cm. long; 
capsule oval or ellipsoid, 1-1.5 cm. long. 

FARAMEA Aubl. 

Shrubs or small trees, usually glabrous throughout; stipules 
persistent, short-triangular and long-aristate, or often united to form 
a narrow sheath; leaves petiolate or subsessile, most often coriaceous; 
inflorescence usually terminal, sometimes axillary, few- or many- 
flowered; calyx truncate or dentate; corolla salverform, the 4 lobes 
valvate, the throat naked; ovary 1-celled; fruit baccate or almost dry, 
by abortion 1-seeded, the seed horizontal, deeply excavate on the 
lower side. Several other species occur in Central America. 

Faramea eurycarpa Donn. Smith, Bot. Gaz. 44: 113. 1907. 
Forests near Las Vueltas, Tucurrique, 700 meters, Tonduz 12880. 
Collected also at La Hondura, Prov. San Jose", and at Pejivalle, 
700-1,700 meters. Endemic. A glabrous shrub of 2.5-3.5 meters; 
stipules united into a tube, but this caducous; leaves short-petiolate, 
oblong or oblanceolate-oblong, mostly 11-15 cm. long and 4-5 cm. 
wide, narrowly caudate-acuminate, acute or subobtuse at the base, 
yellowish when dried; panicles small, many-flowered, pedunculate, 
the flowers pedicellate; calyx 2 mm. long, lobate; corolla blue or 
violet, 10-11 mm. long, the lobes shorter than the tube; fruit 12 mm. 
broad, blue. 

Faramea hondurae Standl. Journ. Wash. Acad. Sci. 18: 169. 
1928. Wet forest, La Hondura, Prov. San Jose', 1,400 meters, 
Standley 37890. Endemic, and known only from the type locality. 



1296 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

A glabrous shrub of 3-4.5 meters; stipules sheathing, caducous, only 
3-4 mm. long; leaves short-petiolate, narrowly oblong or lance-oblong, 
12-19 cm. long, 3-5 cm. wide, caudate-acuminate, with a long, 
linear acumination, acute at the base; inflorescence cymose-panicu- 
late, branched and many-flowered, the pedicels 2-3 mm. long; calyx 
1-1.5 mm. long, shallowly lobate; corolla white, only 5 mm. long, 
the lobes shorter than the tube. 

Faramea occidentalis (L.) Rich. Forests of the Pacific tierra 
caliente; Guanacaste. Southern Mexico to West Indies and northern 
South America. A glabrous shrub or small tree with smooth, gray 
bark; stipules distinct, long-cuspidate, coriaceous, finally deciduous; 
leaves short-petiolate, coriaceous, oblong to oval or oblong-obovate, 
6-18 cm. long, usually abruptly short-acuminate, acute at the base; 
inflorescences terminal and axillary, laxly cymose-paniculate,- often 
few-flowered but also with numerous flowers; calyx truncate; corolla 
white, the tube about 15 mm. long, the lobes 12 mm. long; fruit 
black at maturity, broader than high, about 1 cm. in diameter. The 
flowers are fragrant. 

Faramea quercetorum Standl. Journ. Wash. Acad. Sci. 18: 
168. 1928. Oak forest near Quebradillas, north of Santa Maria de 
Dota, 1,800 meters, Standley 1*2999. Collected by Brenes between 
Guachipelin and Volcan de La Vieja, and by Endres, without locality. 
Endemic. A shrub or small tree of 2.5-4.5 meters, glabrous through- 
out; stipules persistent, short-connate, the subulate lobes 4-5 mm. 
long, the tube 2 mm. long; leaves short-petiolate, elliptic or elliptic- 
oblong, 7-9 cm. long, 2-4.5 cm. wide, acuminate, acute or obtuse 
at the base; flowers in terminal umbels, these sessile or pedunculate, 
mostly 5-flowered, the slender pedicels 8-18 mm. long; calyx truncate, 
1 mm. long; corolla violet, the tube 13 mm. long, the lobes 8-10 mm. 
long; fruit 8 mm. broad. Easily recognized by the simply umbellate, 
few-flowered inflorescence. 

Faramea suerrensis Donn. Smith, Bot. Gaz. 44: 112. 1907. 
F. trinervia var. suerrensis Donn. Smith, op. cit. 31: 115. 1901. 
Suerre, Llanuras de Santa Clara, 300 meters, J. D. Smith 6589. 
Collected also in El General and San Ramon. Endemic. A shrub 
of 3-4 meters, glabrous; stipules short-connate, the lobes rounded 
and mucronate, deciduous; leaves short-petiolate, narrowly oblanceo- 
late-oblong, up to 21 cm. long and 5.5 cm. wide, gradually attenuate 
into a long, narrow acumination, acute to attenuate at the base, 
very conspicuously 3-nerved; inflorescence cymose-corymbose, dense 



FLORA OF COSTA RICA 1297 

and many-flowered, long-pedunculate, the flowers short-pedicellate; 
calyx 1 mm. long, dentate; corolla deep blue, 6 mm. long. Pittier 
3912 in herb. Berlin, from El General, was indicated by Schumann 
as a new species, but is apparently referable here. 

Faramea talamancarum Standl. Field Mus. Bot. 4: 332. 1929. 
Talamanca Valley, Panama, M. A. Carleton 135. Common in wet 
forests of the tierra caliente; region of San Ramon; Guanacaste; 
at 1,200 meters or less. Panama. A slender, glabrous shrub, usually 
2 meters high or less; stipules united into a narrow sheath 8-9 mm. 
long, persistent; leaves short-petiolate, oblong-elliptic to narrowly 
oblong, mostly 11-16 cm. long, thin, cuspidate-acuminate, acute at 
the base; inflorescence cymose-corymbose, many-flowered, usually 
long-pedunculate, the slender pedicels mostly 8-10 mm. long; calyx 
truncate, 0.6 mm. long; corolla blue, the slender tube 10-12 mm. 
long, the lobes 5-7 mm. long. Like other blue-flowered Farameas, 
this must be a handsome shrub. Here probably belong F. salicifolia 
Presl, f. subumbellata and f. paniculata Kuntze (Rev. Gen. 1: 282. 
1891), described from Costa Rica. F. talamancarum is closely related 
to that South American species, and may, indeed, prove to be 
synonymous with it. 

Faramea trinervia Schum. & Donn. Smith, Bot. Gaz. 31: 115. 
1901. In forest, Boca Zhorquin, Talamanca, Tonduz 8571. Forests 
of Tsaki, Tonduz 9583. Endemic. Glabrous; stipules 6-8 mm. long, 
semiconnate, the lobes rounded and aristate; leaves subsessile, nar- 
rowly oblong, 20-25 cm. long, 6-9.5 cm. wide, cuspidate-acuminate, 
subcordate at the base, very conspicuously 3-nerved; inflorescence 
cymose-corymbose, rather dense and many-flowered, pedunculate, 
the pedicels 5-11 mm. long; calyx denticulate; corolla unknown. 
Easy of recognition on account of the sessile leaves, subcordate 
at the base. 

GALIUM L. 

Reference: Greenman, Proc. Amer. Acad. 33: 455. 1898. 

Annual or perennial herbs with slender, 4-angulate stems; leaves 
in verticels of 4 or more; flowers minute, 3-4-parted, in axillary and 
terminal, few-flowered cymes, ebracteate, the hypanthium articulate 
with the pedicel; calyx obsolete; corolla rotate, the lobes valvate; 
anthers exserted; ovary 2-celled, with 2 short styles, the cells 1-ovu- 
late; fruit didymous, dry or fleshy. A few other species probably 
occur in northern Central America. 



1298 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Galium Aschenbornii Schauer. Las Concavas, south of 
Cartago, Lankester 262. Extending to Mexico. Perennial, the stems 
reclining, 50 cm. long or less, glabrous or minutely hispidulous; 
leaves in 4's, linear-oblong or elliptic-oblong, 1.5 cm. long or less, 
acute or cuspidate, glabrous but hispidulous on the margins; corolla 
lobes cuspidate; fruit glabrous, orange-colored. 

Galium mexicanum HBK. Cartago, 1,650 meters, Cooper 
5797. Panama to Mexico. Perennial, the stems much branched, 
aculeolate-hispid on the angles; leaves 6-8 in a whorl, linear, 2 cm. 
long or less, the margins revolute, cuspidate, uncinate-hispidulous 
on the margins and on the nerves beneath; corolla white or purplish, 
pubescent outside; fruit covered with short, uncinate, whitish hairs. 

Galium obovatum HBK. Occasional in forests of the central 
region; region of Dota; at 1,450-3,300 meters. Guatemala to 
Bolivia. Plants slender and much branched, ascending or sometimes 
scandent, the stems pilose with weak, whitish, spreading hairs, some- 
times glabrate; leaves in 4's, ovate to elliptic or obovate, mostly 
5-15 mm. long, obtuse to acuminate, acute at the base, densely 
pilose on both surfaces or rarely glabrate, 3-nerved ; corolla yellowish 
or whitish, the lobes caudate-acuminate; fruit densely uncinate- 
hispid. The Costa Rican material is variable in shape and size of 
the leaves, and it is quite possible that it represents more than a 
single species. 

GARDENIA Ellis 

Shrubs or small trees, usually unarmed; stipules acute or acumi- 
nate, triangular; leaves commonly opposite; flowers large, axillary 
and solitary or rarely terminal and corymbose; calyx tubular, 
spathaceous, or parted; corolla salverform or funnelform, with an 
elongate tube, usually glabrous in the throat, the lobes 5-9; ovary 
commonly 1-celled; fruit usually baccate, the seeds numerous, 
horizontal. All the species are natives of the Old World. 

Gardenia augusta (L.) Merrill. Jazmindelcabo. G.jasminoides 
Ellis; G. florida L. Planted commonly for ornament. Native of 
southern China. A densely branched shrub, the branchlets scabrous- 
puberulent; stipules 1 cm. long; leaves short-petiolate, coriaceous, 
obovate or oblong-obovate, obtuse or acute, narrowed to the base, 
almost glabrous; flowers white, large and showy, usually double in 
cultivated plants, very fragrant; calyx lobes foliaceous, 2-2.5 cm. 
long. The gardenia, well known in the north as a hothouse plant, 
is one of the favorite garden shrubs of Central America. 



FLORA OF COSTA RICA 1299 

GENIPA L. 

Tall trees, glabrous or pubescent; stipules deciduous; leaves 
rather large, opposite, petiolate, subcoriaceous; flowers large, 5-6- 
parted, in terminal, few-flowered cymes; calyx tubular, truncate or 
shallowly lobate; corolla salverform or subrotate, coriaceous, the 
lobes contorted, the tube short, villous in the upper half; stamens 
exserted, the anthers linear; fruit baccate, large, ovoid or globose, 
2-celled; seeds large, horizontal or oblique, compressed. One or two 
other species are known from Central America. 

Genipa americana L. Guaitil. Frequent in forests of the tierra 
caliente. Southern Mexico to Peru and Brazil. A tree of 14 meters 
or less, the crown spreading, the branchlets glabrous, densely leafy; 
stipules triangular, acuminate, 8-12 mm. long; leaves short-petiolate, 
oblong to obovate, 15-30 cm. long, acute or short-acuminate, nar- 
rowed to the acute base, glabrous; inflorescence short-pedunculate, 
the pedicels 4-10 mm. long; calyx 5-8 mm. long, truncate or undulate, 
glabrous; corolla yellowish white, 2-4.5 cm. long, the lobes longer 
than the tube; fruit 6-7 cm. in diameter; seeds 6-12 mm. long. The 
wood is strong, resistant, and flexible, in its properties being some- 
what like the wood of hickory (Carya) of the United States. It is 
used for many purposes. The pulp of the fruit is edible, but dark and 
repulsive in appearance, and not particularly palatable. Its dark 
juice leaves an indelible stain upon every object that it touches. By 
some of the American aborigines it was used for painting their bodies, 
giving a dark blue or almost black color. 

Genipa americana var. Caruto (HBK.) Schum. The only 
Costa Rican specimen I have seen is from Guanacaste, but the tree is 
doubtless more widely distributed in the country. The variety has 
a narrower distribution than typical G. americana, and is by far the 
commoner form found in continental North America. Differs from 
typical G. americana in the abundant, dense pubescence of the 
branches and lower leaf surface. The TeYraba name is reported 
as Brir. 

Genipa codonocalyx Standl. Contr. U. S. Nat. Herb. 17: 446. 
1914. Jagua. Near Boca Matapalo, Pacific coast, Pittier 12085. 
Known only from the original collection. A tree, the branchlets 
glabrous or sparsely short-pilose; stipules triangular-ovate, acumi- 
nate, 10-12 mm. long; leaves short-petiolate, oblong-oblanceolate or 
narrowly oblong, 12-17 cm. long, 4-7 cm. wide, abruptly short- 
acuminate, attenuate to the acute base, glabrous and lustrous above, 



1300 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

paler beneath, short-pilose on the nerves; cymes subsessile, lax, the 
pedicels 3-7 mm. long; calyx and hypanthium glabrous, the calyx 
only 3-3.5 mm. long, truncate; corolla yellowish white, 2.5 cm. long, 
the lobes sericeous, longer than the tube. The wood is said to be 
very hard. 

Genipa venosa Standl. Journ. Wash. Acad. Sci. 18: 168. 1928. 
Dense, wet forest, El Arenal, Guanacaste, 485 meters, Standley & 
Valeria 45269. Known only from the original collection. A tree of 
9-15 meters; petioles 3.5-4 cm. long; leaf blades obovate-oblong, 
29-35 cm. long, 12-14 cm. wide, rounded or obtuse at the apex and 
linear-cuspidate, gradually attenuate to the obtuse base, glabrous or 
glabrate above, brown-tomentose beneath along the nerves or gla- 
brate, the veins prominent and closely reticulate; inflorescence few- 
flowered, the branches thick; fruits green, subglobose or oval, as 
much as 10 cm. long or even larger, smooth, rounded at the apex. I 
have not seen specimens of this tree recently, and have a suspicion 
that it may belong to some other genus than Genipa. 

GEOPHILA Don 

Creeping herbs; leaves long-petiolate, membranaceous, ovate- 
cordate or rounded-cordate; flowers small, in terminal, pedunculate, 
few-flowered heads, the heads subtended by 2 free bracts; calyx 
dentate or lobate; corolla tubular-funnelform, pilose in the throat; 
ovary 2-celled; fruit fleshy, drupaceous, the 2 nutlets plano-convex, 
dorsally compressed, usually costate. One other species has been 
found in Central America. 

Geophila herbacea (Jacq.) Schum. Frequent in shaded places 
of the tierra caliente, often a weed in banana plantations. Generally 
distributed in tropical America. Plants slender, almost glabrous, 
rooting at the nodes; leaves long-petiolate, the blades rounded-ovate 
or subreniform, deeply cordate at the base, 3-4 cm. long; heads long- 
pedunculate, usually 3-5-flowered; calyx lobes lanceolate, acuminate; 
corolla white or purplish, often 1 cm. long; fruit red or almost black, 
the nutlets somewhat spirally twisted. 

GOMOZIA Mutis 

Creeping, perennial herbs; stipules connate with the petioles to 
form a sheath, bidentate or entire and triangular; leaves small, 
opposite; flowers minute, axillary, sessile; calyx truncate or dentate, 
persistent; corolla tubular or funnelform, glabrous in the throat, the 



FLORA OF COSTA RICA 1301 

4 lobes valvate; anthers exserted; fruit drupaceous, containing two 
1-seeded nutlets. A single species grows in North America. 

Gomozia granadensis L. Nertera depressa Banks & Soland. 
Abundant in moist forest of the mountains, mostly at 1,600-3,000 
meters; region of San Ramon. Widely distributed in the higher 
mountains from Mexico to Chile. Plants small and slender, almost 
glabrous, forming dense, close mats on banks or logs, the stems much 
branched; leaves petiolate, the blades rather thick, mostly 1 cm. long 
or less, ovate to deltoid-ovate or ovate-orbicular, obtuse, puncticulate 
above; corolla white or greenish yellow; fruits bright red, juicy, 3-4 
mm. long. The plant is a pretty one when in fruit because of the 
abundance of small but brightly colored berries. In general appear- 
ance it suggests the partridge berry (Mitchella repens, Rubiaceae) of 
the United States. It is unfortunate that the long established generic 
name Nertera must be relegated to synonymy, but the genus is such 
a small and unimportant one that there is little reason for placing 
Nertera on the list of nomina conservanda. 

GONZALAGUNIA Ruiz & Pavon 

Shrubs or small trees, usually abundantly pubescent, the branches 
slender, terete; leaves opposite, petiolate or subsessile; flowers small, 
sessile or pedicellate, in slender, elongate, terminal spikes or thyrsi- 
form panicles; hypanthium globose or campanulate; calyx usually 
4-lobate, the lobes equal or unequal, persistent; corolla funnelform 
or salverform, the tube short or elongate, the lobes short, spreading, 
valvate or imbricate; anthers included; ovary 2- or 4-celled, the 
ovules numerous; fruit baccate, depressed-globose, the 2 or 4 lobes 
chartaceous or osseous, many-seeded; seeds minute, foveolate. One 
or two other species probably occur in Central America. 

Gonzalagunia bracteosa (Donn. Smith) Robinson, Proc. 
Amer. Acad. 45: 405. 1910. Gonzalea bracteosa Donn. Smith, Bot. 
Gaz. 33: 252. 1902. Duggena bracteosa Standl. Contr. U. S. Nat. 
Herb. 18: 125. 1916. Forests near Suerre, Llanuras de Santa Clara, 
300 meters, J. D. Smith 6583. Forests of the Atlantic tierra caliente. 
Endemic. A shrub of 3 meters, the branchlets pilose-sericeous; 
stipules 1.5-2 cm. long; leaves petiolate, lanceolate to narrowly 
elliptic-oblong, 9-21 cm. long, 2-7 cm. wide, acute to attenuate, 
acute at the base, sparsely appressed-pilose above or glabrate, 
sparsely appressed-pilose beneath; flower cymules sessile, subtended 
by foliaceous bracts 5-12 mm. long; flowers 4 mm. long; calyx lobes 



1302 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

short, triangular; corolla strigose; fruit black, 4 mm. in diameter, 
2-celled. 

Gonzalagunia Brenesii Standl., sp. nov. Ramuli teretes fusco- 
brunnei, breviter strigosi vel fere glabri, internodiis elongatis; stipulae 
ca. 8 mm. longae e basi triangulari subulato-attenuatae extus dorso 
marginibusque sericeae; folia magna breviter petiolata subcoriacea, 
petiolo crasso vix ad 5 mm. longo; lamina anguste oblongo-lanceolata 
12-19 cm. longa 3.5-5.5 cm. lata longissime sensim attenuata, basi 
anguste obtusa, supra glabra lucidissima, ad costam prominentem 
puberula, nervis impressis, subtus fere ubique breviter sericea, costa 
gracili elevata, nervis lateralibus utroque latere ca. 14 prominentibus 
angulo latiusculo adscendentibus, venulis prominentibus laxe reticu- 
latis; inflorescentia spiciformis pedunculata 21-25 cm. longa, cymulis 
sessilibus paucifloris remotis, bracteis minutis, floribus sessilibus vel 
breviter pedicellatis; ovarium vix 1 mm. longum dense minute 
sericeum 4-loculare; calyx vix 1 mm. longus brevissime dentatus 
glabratus, dentibus late obtusis; corolla extus dense adpresso- 
pilosula, tubo gracili 7 mm. longo. Coastal forests of Golfito de Osa, 
April, 1930, Brenes 12323 (type in Herb. Field Mus.). Conspicuous 
because of the very lustrous upper surface of the leaves, glabrous or 
nearly so, and the numerous pairs of lateral nerves. 

Gonzalagunia ovatifolia (Donn. Smith) Robinson, Proc. Amer. 
Acad. 45: 405. 1910. Gonzalea ovatifolia Donn. Smith, Bot. Gaz. 
27: 336. 1899. Duggena ovatifolia Standl. Contr. U. S. Nat. Herb. 
18: 126. 1916. Atirro, Prov. Cartago, 600 meters, J. D. Smith 6509. 
Forests of the Atlantic tierra caliente. Panama. A slender shrub 
of 2-3 meters, the branchlets densely pilose-sericeous; stipules 6-8 
mm. long; leaves subsessile, oval-ovate or broadly oblong-ovate, 
8-12 cm. long, 3.5-6.5 cm. wide, acuminate to long-attenuate, rounded 
at the base, almost glabrous above, sericeous-strigose beneath; 
cymules few-flowered, sessile, the bracts 5 mm. long or less; calyx 
lobes 1-2 mm. long, oblong or elliptic, obtuse; corolla white, strigose, 
3-3.5 mm. long; fruit white, 4-coccous, strigose. 

Gonzalagunia panamensis (Cav.) Schum. Frequent in forests 
and thickets of the central region, descending to the Pacific tierra 
caliente; region of San Ramon; at 1,500 meters or less. Southern 
Mexico to West Indies and northern South America. A slender 
shrub or small tree, the branches densely strigose at first; stipules 
3-8 mm. long; leaves short-petiolate, ovate or lanceolate, 7-14 cm. 
long, 2-6 cm. wide, acuminate or long-attenuate, obtuse or acute 



FLORA OF COSTA RICA 1303 

at the base, densely or sparsely strigillose above or glabrate, sparsely 
or densely pilose beneath or glabrate; inflorescence 10-24 cm. long, 
the cymules few-flowered, dense or remote, sessile or nearly so, the 
bracts 2-3 mm. long; calyx lobes deltoid, obtuse or acute; corolla 
white, 10-17 mm. long, the tube glabrous or sparsely pilose, the 
lobes 2-2.5 mm. long; fruit 4-coccous, 3-4 mm. in diameter, white. 

Gonzalagunia rosea Standl. In forest, regions of San Ramon 
and Zarcero, San Pedro Coronado, and doubtless elsewhere, 1,400- 
2,100 meters. Adjacent Panama. A slender shrub or small tree, 
1-4.5 meters high, the young branches densely pilose; stipules 6-7 
mm. long; leaves short-petiolate, lance-oblong, 9-14 cm. long, 2.5-5 
cm. wide, narrowly attenuate-acuminate, acute or subobtuse at the 
base, short-pilose above with mostly spreading hairs, densely soft- 
pilose beneath; inflorescence up to 30 cm. long, the cymules pedun- 
culate, the bracts minute; calyx lobes broadly ovate, acute or 
subobtuse; corolla pink, white-strigose, the tube 6-8 mm. long, lobes 
2 mm. long; fruit glabrate, 4-coccous, 2.5-3 mm. in diameter. Similar 
to G. panamensis, but distinguished by the mostly spreading pubes- 
cence of the leaves and by the pedunculate cymules. 

GUETTARDA L. 

Trees or shrubs; stipules deciduous; leaves opposite or ternate, 
petiolate or subsessile, membranaceous to rigid-coriaceous; flowers 
small or large, in axillary, bifurcate or congested cymes, usually 
secund; hypanthium ovoid or globose, the calyx tubular or cupular, 
truncate or very obscurely dentate, deciduous; corolla funnelform 
or salverform, the tube elongate, the throat naked, the 4-9 lobes 
obtuse, imbricate; anthers included; ovary 2-9-celled, the cells 
tubular, elongate, 1-ovulate; fruit drupaceous, globose to oblong, 
sometimes acutely angulate, the flesh very thin, the stone ligneous 
or osseous. A few other species have been found in Central America. 

Guettarda Brenesii Standl., sp. nov. Ramuli crassiusculi 
teretes fusci pallide lenticellati, novellis pilis brevibus patentibus 
dense hirtellis; stipulae deciduae ca. 8 mm. longae brunneae oblongo- 
ovatae acutae extus sericeae; folia breviter petiolata membranacea, 
petiolo usque 6 mm. longo dense adpresso-hirtello ; lamina late ovata 
vel late ovato-elliptica 4-6.5 cm. longa 3-4.5 cm. lata, apice obtusa 
vel subrotundata atque apiculata, basi subrotundata breviter cordata, 
supra viridis sparse hispidula, nervis vix elevatis, subtus pilis ple- 
rumque patentibus nitidis intertextis piloso-tomentosa; cymae axil- 
lares dense multiflorae 2.5-3 cm. longe pedunculatae, usque 2 cm. 



1304 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

latae, breviter ramosae, floribus sessilibus, pedunculo dense hispidulo, 
bracteolis calyce longioribus oblongo-ovatis brunneis longiciliatis; 
hypanthium subglobosum 1 mm. longum dense strigillosum; calyx 
truncatus ca. 1.2 mm. longus sparse strigillosus vel glabratus; corolla 
extus dense minute sericea, tubo gracili 9 mm. longo, lobis ca. 2 mm. 
longis. Isla de Caballo, Golfo de Nicoya, June, 1932, Brenes 15694. 
(type in Herb. Field Mus.). The available material is fragmentary, 
but adequate to show that it represents a species different from all 
others known from Central America. It is probably related to 
G. macrosperma, but differs in the abundant, spreading pubescence 
of the lower leaf surface, and in the much smaller flowers. 

Guettarda conferta Benth. Bot. Voy. Sulph. 106. 1845. Mathi- 
ola conferta Kuntze, Rev. Gen. 1: 288. 1891. Cocos Island, Barclay. 
Known only from the original collection. Plant with dense, ferru- 
ginous hairs on the branchlets, petioles, peduncles, and nerves of the 
leaves; stipules broadly obovate, 12 mm. long, hirsute outside at the 
base, otherwise glabrous, about equaling the petioles; blades ovate, 
10-15 cm. long, acuminate, acute at the base, hirsute on both sides; 
cymes subsessile, the branches recurved, 2.5 cm. long or less, the 
flowers 8 mm. long; calyx shallowly 3-4-dentate; corolla sericeous- 
hirtous, the 4 lobes short, obtuse, crispate; fruit ovoid-tetragonous, 
4 mm. long, hirsute, 4-celled. I know the species only from the 
description, and have seen no Cocos Island material of the genus. 

Guettarda crispiflora Vahl. Mosquito (mosqueta?) de montana 
(Tonduz). Frequent in forests of the central region; mountains of 
San Ramon and Zarcero; at 1,150-1,500 meters. Lesser Antilles and 
Trinidad. A shrub or tree, sometimes 8 meters high, the trunk to 
15 cm. in diameter, the bark gray; stipules ovate, 1-2 cm. long, acute 
or acuminate, setose-pilose at the base and along the costa; leaves 
petiolate, ovate to broadly oval, 9-12 cm. long, 4.5-7 cm. wide, 
rounded or obtuse at the apex and abruptly acuminate, broadly 
rounded at the base; cymes short-pedunculate, bifurcate, the branches 
1-3 cm. long, the bractlets minute; calyx and hypanthium densely 
fulvous-sericeous; corolla white or tinged with pink, the tube 12-15 
mm. long, densely retrorse-pilose, the lobes 3 mm. long, lacerate- 
undulate; fresh fruit as much as 2 cm. long, violet-blackish, the dried 
fruit acutely tetragonous, 5-7 mm. long. 

Guettarda macrosperma Bonn. Smith. In thickets or dry 
forest, Meseta Central to the Pacific coast; Guanacaste; Changuinola 
Valley. Panama to Guatemala. A shrub or small tree, sometimes 



FLORA OF COSTA RICA 1305 

7 meters high, with a trunk 10 cm. in diameter; stipules ovate-deltoid, 
about 4 mm. long, filiform-acuminate; leaves slender-petiolate, 
membranaceous, oval to oblong, 5-13 cm. long, 2.5-7 cm. wide, acute 
to acuminate, rounded to subacute at the base, hispidulous above 
when young but soon glabrate, minutely and usually sparsely 
appressed-pilose beneath; cymes few-flowered, at first dense, in fruit 
more open, pedunculate, the bractlets subulate, shorter than the 
calyx; calyx and hypanthium tomentulose, the calyx 2-2.5 mm. long; 
corolla white, sericeous outside, the tube 12 mm. long; fruit terete, 
subglobose, 1-1.5 cm. or more in diameter, often dark red, 3^4-celled. 

Guettarda poasana Standl. Journ. Wash. Acad. Sci. 18: 182. 
1928. Wet forest, Viento Fresco, slopes of Volcan de Pods, Prov. 
Alajuela, 1,800 meters, Standley & Torres 47807. Collected also at 
Rio Poas, and at Las Nubes, Prov. San Jose", at 1,800-2,100 meters. 
Endemic. Closely related to G. crispiflora, but differing in its gla- 
brous stipules and glabrous or nearly glabrous hypanthium and 
calyx; corolla pink, the tube 15-20 mm. long. 

HAMELIA Jacq. 

Shrubs or small trees, glabrous or pubescent; stipules deciduous; 
leaves opposite or verticillate, petiolate, usually membranaceous; 
flowers yellow or red, small or large, usually in terminal, scorpioid 
cymes, sessile or pedicellate; calyx 5-lobate, the lobes short or elon- 
gate, persistent; corolla tubular or funnelform, the tube 5-costate, the 
throat glabrous, the lobes short, imbricate; anthers included or semi- 
exserted; ovary 5-celled, many-ovulate; fruit small, baccate, ovoid 
to cylindric, 5-celled; seeds numerous, minute, angulate, foveolate. 
A few other species are native in Central America. 

Hamelia axillaris Swartz. Occasional in forests of the tierra 
caliente. British Honduras to West Indies, Brazil, and Peru. A 
slender shrub or small tree, the branches glabrous or minutely puberu- 
lent when young; leaves opposite, petiolate, elliptic or oblong-elliptic, 
5-12 cm. long, abruptly acuminate, acute at the base, glabrous above, 
glabrous beneath or minutely puberulent on the nerves; inflorescence 
few-many-flowered, lax, pedunculate, the flowers sessile or subsessile, 
secund; calyx and hypanthium 3 mm. long, minutely puberulent or 
glabrate, the calyx lobes oblong or linear-oblong, obtuse; corolla 
yellow, 1-1.5 cm. long, glabrous or nearly so, the limb dilated, 3-4 
mm. broad; fruit globose-ellipsoid, 5-6 mm. long. 

Hamelia costaricensis Standl. Contr. U. S. Nat. Herb. 20: 207. 
1919. Surubres, near San Mateo, Biolley 2656. Collected also at 



1306 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Taus and Carrillos de Poas; ascending to 1,600 meters. Endemic. 
Branchlets densely puberulent; leaves opposite, slender-petiolate, 
oval-ovate to oval-elliptic or obovate-oblong, 8-19 cm. long, 4-10 
cm. wide, acute or acuminate, rounded or obtuse and short-decurrent 
at the base, minutely puberulent beneath on the nerves or almost 
glabrous; inflorescence pedunculate, branched, many-flowered, the 
flowers secund, sessile; calyx and hypanthium densely puberulent, 
the calyx lobes subulate, 1-1.5 mm. long; corolla densely fulvous- 
puberulent, becoming glabrate, the tube 22 mm. long, ampliate 
above, 6-7 mm. wide in the throat, the lobes rounded, 5 mm. long, 
spreading. 

Hamelia magnifolia Wernham, Journ. Bot. 49: 210. 1911. 
Zorrillo Colorado. Rio Corozal, near Santo Domingo de Golfo Dulce, 
Tonduz 10091. Collected also at Tilaran, Guanacaste, 700 meters. 
Panama. A shrub or small tree, 6 meters high or less, the trunk 8 cm. 
or less in diameter, the branchlets glabrous or minutely puberulent 
when young; leaves opposite, slender-petiolate, oblanceolate or 
obovate, 10-28 cm. long, 3-11 cm. wide, abruptly acuminate or long- 
acuminate, acute to long-attenuate at the base, glabrous or minutely 
puberulent beneath on the nerves; cymes many-flowered, short- 
pedunculate, the branches short, the flowers sessile, subsecund; 
calyx and hypanthium 3 mm. long, puberulent or glabrate, the calyx 
lobes minute, broadly deltoid; corolla yellow, subcylindric, 15 mm. 
long, glabrous or obscurely puberulent, the lobes 1 mm. long; fruit 
oblong-ellipsoid, 5-8 mm. long. 

Hamelia nodosa Mart. & Gal. H. viridifolia Wernham, Journ. 
Bot. 49: 213. 1911 (without locality, Tonduz 13867}. Frequent in 
thickets and forest of the tierra caliente, at least on the Pacific slope; 
region of San Ramon; Guanacaste; at 1,100 meters or less. Panama 
to southern Mexico. A shrub or small tree, 6 meters high or less, the 
branches glabrous or when young sparsely puberulent: leaves mostly 
in whorls of 4, short-petiolate, the blades elliptic-oblong to elliptic 
or ovate, 4-9 cm. long, 2-4 cm. wide, acute or short-acuminate, 
rounded to attenuate at the base, glabrous above, beneath villosulous 
or puberulent on the nerves or glabrate; inflorescence many-flowered, 
the flowers sessile or nearly so, secund; calyx and hypanthium 3-4 
mm. long, sparsely puberulent, the calyx lobes minute, deltoid; 
corolla red, tubular, 18-22 mm. long, minutely puberulent, the lobes 
1 mm. long; fruit oblong-cylindric, 8-10 mm. long. This may be no 
more than a form or variety of H. patens. 



FLORA OF COSTA RICA 1307 

Hamelia patens Jacq. Zorrillo, Azulillo, Zorrillo real, Coralillo, 
Pissi, Palo camaron, Anileto. Frequent in forests and thickets of 
the central region, descending to the Pacific coast, at 1,700 meters 
or less; Guanacaste. Generally distributed in tropical America. A 
shrub or small tree, sometimes 6 meters high, the branchlets villous 
or puberulent; leaves mostly ternate, slender-petiolate, lance-oblong 
to elliptic or ovate, 6-20 cm. long, 2-9 cm. wide, short-acuminate, 
rounded to acuminate at the base, puberulent or villosulous above, 
sometimes glabrate, beneath usually abundantly villosulous or 
puberulent; inflorescence many-flowered, the branches often much 
elongate in fruit, the flowers secund, sessile or short-pedicellate; 
calyx and hypanthium 2.5-3 mm. long, puberulent or villous, the 
calyx lobes minute, deltoid; corolla orange-red, tubular, 15-20 mm. 
long, sparsely or densely puberulent or villosulous, the lobes minute; 
fruit red to almost black, juicy, 6-10 mm. long. As Indian names 
Pittier reports Tsus-kra (Brunka) and Pilii-tso (Guatuso). 

Hamelia Rovirosae Wernham. In thickets, Atlantic tierra 
caliente. Panama to southern Mexico. A shrub or small tree, 
sometimes 5 meters high, the branchlets sparsely or densely villous; 
leaves ternate, short-petiolate, elliptic-oblong to oval-elliptic, 5-15 
cm. long, 2-5 cm. wide, acute or short-acuminate, acute or acuminate 
at the base, glabrous or sparsely villous above, sparsely or densely 
villous beneath along the nerves; inflorescence few-many-flowered, 
short-pedunculate, lax, the branches short or elongate, the flowers 
sessile or subsessile, usually secund; calyx and hypanthium sparsely 
or densely villous, the calyx lobes narrowly oblong, obtuse, 2-5 mm. 
long, reflexed; corolla dull red or orange-red, villous, 18-24 mm. long, 
gradually ampliate above, 6 mm. wide in the throat, the lobes 1.5-2 
mm. long; fruit ovoid, 8 mm. long, red or purple, sparsely or densely 
villous. 

Hamelia Rowleei Standl. Journ. Wash. Acad. Sci. 15: 7. 1925. 
Livingston, on the Rio Reventazon, Rowlee & Stork 75. Changuinola 
Valley. Panama. A shrub, the branchlets pilose with long, spread- 
ing hairs; leaves opposite, slender-petiolate, elliptic-obovate or 
elliptic, 15-19 cm. long, 6-9 cm. wide, abruptly short-acuminate, 
cuneate-attenuate at the base or abruptly contracted and decurrent, 
villosulous above along the costa, sparsely pilose or glabrous else- 
where, copiously pilose beneath with long, stiff, spreading hairs; 
inflorescence with few or numerous branches, many-flowered, the 
flowers secund, sessile or nearly so; hypanthium oblong-turbinate, 
densely villous, the calyx lobes deltoid-subulate, 1.5 mm. long; 



1308 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

corolla puberulent or short-pilose, the tube 28 mm. long, slightly 
dilated above, the lobes acuminate, 4-5 mm. long, spreading. 

Hamelia Storkii Standl. Changuinola Valley. Panama. A 
shrub or small tree, as much as 4.5 meters high, with a trunk 5 cm. 
in diameter, the branches glabrous; leaves opposite, slender-petiolate, 
elliptic to oblanceolate-elliptic, 12-22 cm. long, 4-9 cm. wide, 
acuminate, acute at the base, sparsely barbellate beneath in the 
axils of the nerves, elsewhere glabrous, or sometimes sparsely short- 
pilose beneath; inflorescence with few or numerous branches, these 
often 10 cm. long, the flowers secund, sessile or nearly so; hypanthium 
glabrous; calyx lobes triangular-oblong, 1.5 mm. long; corolla yellow, 
2.5-3 cm. long, tubular, slightly dilated above, 4 mm. broad in the 
throat, the lobes ovate, 3-4 mm. long; fruit oblong, 10-12 mm. long. 

Hamelia xerocarpa Kuntze, Rev. Gen. 1: 284. 1891. "Baguar," 
600 meters, Kuntze. Nicaragua. A tall shrub, the branches fulvous- 
villous; leaves opposite, the petioles 2.5 cm. long, villous, the blades 
elliptic-obovate or elliptic-oblong, 13.5-15.5 cm. long, 6-6.5 cm. wide, 
acuminate, acute at the base, glabrous above or nearly so, fulvous- 
villous beneath; flowers sessile, secund; calyx and hypanthium densely 
fulvous- villous, the calyx lobes lance-subulate, 1-1.5 mm. long; 
corolla subfunnelform, 1.5 cm. long, 2.5 mm. wide in the throat, 
densely villous, the lobes broad, acuminate, 3-4 mm. long; fruit 
cylindric, 1.5 cm. long, 4 mm. thick, villous. 

HEMIDIODIA Schum. 

Perennial herbs, sometimes suffrutescent at the base; stipules 
united with the petioles into a setiferous sheath; leaves opposite, 
conspicuously nerved; flowers sessile and densely clustered in the 
leaf axils; sepals 4, equal, connate at the base; corolla small, white, 
funnelform, the 4 lobes valvate; stamens exserted; fruit of 2 carpels, 
these 1-seeded, separating from the central septum, opening near the 
base. The genus consists of a single species. 

Hemidiodia ocimifolia (Willd.) Schum. Frequent in thickets 
or waste ground of the tierra caliente, ascending to La Hondura, at 
1,200 meters; Guanacaste; region of San Ramon. Widely distributed 
in tropical America. Stems often much elongate, decumbent or 
ascending, sparsely puberulent or glabrate; leaves petiolate, lanceo- 
late or lance-oblong, 3-5 cm. long or larger, acuminate, puberulent 
or glabrate, scabrous on the margins; sepals ovate, acute, 0.5 mm. 
long; corolla glabrous, 3-4 mm. long; fruit 3-4 mm. long, puberulent 
or glabrous. 



FLORA OF COSTA RICA 1309 

HILLIA Jacq. 

Shrubs, usually epiphytic, glabrous; leaves opposite, petiolate, 
fleshy, coriaceous when dried; stipules membranaceous, caducous; 
flowers large, white, terminal, solitary, subsessile; hypanthium obo- 
void or cylindric; calyx none or of 2-4 foliaceous, caducous lobes; 
corolla salverform, the tube elongate, the throat ampliate, naked, 
the limb 3-7-lobate, the lobes spreading, contorted; anthers included; 
ovary 2-celled, many-ovulate; capsule elongate-oblong or cylindric, 
septicidally bivalvate; seeds imbricate, produced at the base into 
an appendage and at the apex into a tuft of hairs. At least one 
other species occurs in Central America. 

Hillia chiapensis Standl. Jazmin del vokan. Forests of the 
Pacific tierra caliente; region of Zarcero; Guanacaste; collected by 
Werckle" at some unspecified locality with an elevation of 1,600 
meters. Southern Mexico. A densely branched, epiphytic shrub; 
stipules oblong to obovate, 3-4 mm. long, rounded at the apex; 
petioles 2-5 mm. long; leaf blades elliptic or oval to oblong-elliptic, 
9-14 mm. long, 4-7 mm. wide, rounded at the apex, obtuse or acutish 
at the base, the lateral nerves inconspicuous; corolla tube 15 mm. 
long, the lobes 7 mm. long; capsule about 2 cm. long, the valves 
after dehiscence 3-4 mm. wide. 

Hillia loranthoides Standl. Journ. Wash. Acad. Sci. 18: 165. 
1928. Jazmin del monte, Montana. Moist forest, Quebrada Serena, 
southeast of Tilaran, Guanacaste, 700 meters, Standley & Valeria 
46152. Region of San Ramon, at about 1,100 meters. Endemic. 
A large, epiphytic shrub; stipules elliptic-oblong, 18 mm. long, 
6-8 mm. wide, obtuse; petioles stout, 8 mm. long or less; leaf blades 
elliptic to oblong-ovate, 5-7 cm. long, very thick, acutely narrowed 
to the obtuse apex, obtuse or acute at the base, the lateral nerves 
obscure, about 4 on each side, ascending at a narrow angle; corolla 
tube about 5 cm. long, the broad lobes 2 cm. long; capsule cylindric, 
3 cm. long, 7 mm. thick. Like other species of the genus, this is a 
very handsome plant when in flower. 

Hillia Maxonii Standl. Mountains of the central region and in 
Dota and San Ramon, at 1,250-2,400 meters. Nicaragua. An 
epiphytic shrub, the branches sometimes 5 meters long and pendent; 
stipules broadly obovate, 12 mm. long, rounded at the apex; petioles 
3-4 mm. long; leaf blades oval to oval-obovate, 2-3.5 cm. long, 1-2 
cm. wide, broadly rounded at the apex, somewhat narrowed to the 
obtuse or acute base, coriaceous, the costa and lateral nerves obscure; 



1310 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

calyx lobes oblong-linear, 8 mm. long, green, rounded at the apex; 
corolla tube 5 cm. long, the lobes almost 2 cm. long; capsule cylindric, 
3 cm. long. 

Hillia palmana Standl. Journ. Wash. Acad. Sci. 18: 164. 1928. 
Between La Palma and La Hondura, Prov. San Jose", 1,500-1,700 
meters, Maxon & Harvey 801+5. Known only from the original 
collection. Stipules oblong or spatulate-oblong, 13-20 mm. long, 
rounded at the apex; petioles 3 mm. long or less; leaf blades narrowly 
spatulate-oblong or oblong-cuneate, 1.5-3.5 cm. long, 5-11 mm. 
wide, broadly rounded at the apex, gradually narrowed to the long- 
attenuate base, the venation obsolete; corolla tube 3.5 cm. long, the 
lobes suborbicular, 1 cm. long. 

Hillia Valerii Standl. Journ. Wash. Acad. Sci. 18: 164. 1928. 
Azaharcillo. Between La Palma and La Hondura, Prov. San Jose*, 
1,500-1,700 meters, Maxon & Harvey 8092. Frequent in forests of 
the central region; region of San Ramon; at 1,200-1,700 meters. 
Endemic. An epiphytic shrub, sometimes 2 meters long; stipules 
spatulate-obovate, 3-4 cm. long, broadly rounded at the apex; 
petioles stout, 5-10 mm. long; leaf blades obovate-oblong or narrowly 
obovate, 5-8 cm. long, 2-3.5 cm. wide, broadly rounded at the apex, 
long-attenuate to the base, coriaceous, the lateral nerves about 6 on 
each side; calyx lobes ovate-oval, obtuse, 4-5 mm. long; corolla tube 
8 cm. long, the oval lobes 3 cm. long; capsule columnar, terete, 6-7.5 
cm. long, 8 mm. in diameter. Specimens of this species have been 
referred to H. tetrandra Swartz, a species of the West Indies and 
of other regions of Central America, but apparently unknown in 
Costa Rica. 

HOFFMANNIA Swartz 

Shrubs or herbs, glabrous or pubescent, the branches terete or 
tetragonous; leaves opposite or verticillate, usually membranaceous; 
stipules deciduous or persistent; flowers small, white, yellow, or red, 
cymose, the cymes axillary, sessile or pedunculate; hypanthium 
oblong or turbinate; calyx usually 4-lobate, the lobes short, persistent; 
corolla funnelform to almost rotate, the tube short or elongate, the 
throat glabrous, the limb commonly 4-lobate, the lobes obtuse or 
acute, imbricate; anthers exserted; ovary 2-celled, many-ovulate; 
fruit small, baccate, 2-celled; seeds numerous, minute, foveolate. 
The genus consists of a large number of species, most of them much 
alike in general appearance, and separated only by slight characters, 
which, however, appear to be usually constant. Because of the large 



FLORA OF COSTA RICA 1311 

number of species and the difficulty of separating them, there is 
reproduced here the key published in the North American Flora, so 
far as it applies to Costa Rican plants of the genus. In some of the 
species the form of the corolla is unknown or uncertain, and on this 
account they appear twice in the key. The genus ranges from 
Mexico to the southern Andes, but is represented in Costa Rica 
by a larger number of species than in any other region. 

Lobes of the corolla twice as long as the tube or longer. 

Cymes long-pedunculate, longer than the petioles, the peduncles 
equaling or longer than the cymes. 

Stems herbaceous, low; corolla bright red H. refulgens. 

Stems woody, elongate; corolla red, green, or yellow. 

Leaves sessile, the base dilated and clasping. H. subauriculata. 
Leaves petiolate. 

Fruit white; flowers long-pedicellate; corolla red. 

H. leucocarpa. 
Fruit red; flowers partly sessile; corolla pale green. 

H. pallidiflora. 
Cymes sessile or short-pedunculate, usually shorter than the 

petioles, the peduncles shorter than the cymes. 
Leaves conspicuously puberulent or villosulous beneath with 

ferruginous hairs; corolla more or less villous. 
Leaves essentially sessile, the blades decurrent to the base of 

the petiole. 
Cymes lax, many-flowered; base of the leaf blade deeply 

cordate and clasping H. amplexifolia. 

Cymes dense, few-flowered; base of the blade acute. 

H. ramonensis. 

Leaves conspicuously petiolate. 

Leaf blades chiefly obovate, broadest above the middle. 

H . hamelioides. 
Leaf blades mostly elliptic, broadest at the middle. 

Lateral nerves of the leaves 5-8 pairs H. Valerii. 

Lateral nerves of the leaves about 12-13 pairs. 

H. asclepiadea. 
Leaves glabrous beneath or obscurely grayish-puberulent; corolla 

glabrous or puberulent. 
Calyx lobes linear or narrowly triangular, very acute. 

H. Tonduzii. 



1312 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Calyx lobes broadly deltoid or ovate, obtuse. 
Leaves sessile or essentially so, the blade decurrent to the 

base of the petiole H. carpinterae. 

Leaves conspicuously petiolate. 

Leaf blades broadest above the middle . H. longepetiolata. 

Leaf blades mostly broadest at the middle . H. inamoena. 

Lobes of the corolla equaling or shorter than the tube, or but slightly 

exceeding it. 

Leaf blades dilated and clasping at the base . . . . H. subauriculata. 
Leaf blades not dilated and clasping at the base. 
Corolla pubescent outside. 
Leaves glabrous beneath. 

Calyx lobes elongate, acute H. nesiota. 

Calyx lobes broadly deltoid, obtuse H. arborescens. 

Leaves villosulous beneath, or rarely puberulent, at least along 

the nerves. 
Leaves merely puberulent beneath along the nerves. 

H. inamoena. 
Leaves villosulous beneath. 

Cymes long-pedunculate . . . H. affinis. 

Cymes sessile or short-pedunculate. 
Leaves sessile, the blade decurrent to the base of the 

petiole H. ramonensis. 

Leaves petiolate. 

Leaf blades broadest above the middle. 

H. hamelioides. 
Leaf blades broadest at the middle. 

Leaves short- villous beneath H. asclepiadea. 

Leaves minutely puberulent beneath on the nerves. 

H. inamoena. 
Corolla glabrous. 

Lobes of the corolla shorter than the tube. 
Leaf blades obovate or oblong-obovate, villosulous beneath 

along the nerves , H. josefina. 

Leaf blades elliptic to lance-oblong, glabrous beneath, at 

least at maturity H. piratarum. 

Lobes of the corolla about as long as the tube. 

Corolla 15 mm. long H. dotae. 



FLORA OF COSTA RICA 1313 

Corolla 6-12 mm. long. 

Leaves glabrous H. psychotriaefolia. 

Leaves villous beneath on the nerves H. trichocalyx. 

Hoffmannia aeruginosa Standl., sp. nov. Frutex metralis, 
caule suffrutescente subtereti crassiusculo densissime pilis intertextis 
brunneo-ferrugineis patentibus villoso, internodiis petiolis subaequa- 
libus; folia opposita longipetiolata crasse membranacea, petiolo 
2.5-5.5 cm. longo crassiusculo ut caule villoso; lamina elliptica vel 
oblongo-elliptica 11.5-19 cm. longa 4-8 cm. lata acuta vel acuminata, 
saepe abrupte longiacuminata, basi acuta vel interdum quoque 
decurrens, supra laete viridis glabra, nervis perspicuis, subtus paullo 
pallidior, ad costam nervosque prominentes dense breviter ferrugineo- 
villosa, aliter sparse puberula vel glabrata, nervis lateralibus utroque 
latere ca. 14 subarcuatis angulo lato divergentibus teneris; inflores- 
centiae axillares 3-10-florae congestiflorae cymosae vix 3 mm. longe 
pedunculatae vel subsessiles, ubique densissime ferrugineo- vel pur- 
pureo-villosae, floribus sessilibus vel breviter pedicellatis; hypanthium 
anguste obconicum 2.5-3 mm. longum, lobis lineari-triangularibus 
usque 2 mm. longis; corolla fere rotata, tubo brevissimo, lobis ellip- 
tico-oblongis 6 mm. longis; antherae exsertae. Zarcero, 1,500 meters, 
August, 1937, Austin Smith A86 (type in Herb. Field Mus.). Guada- 
lupe de Zarcero, 1,525 meters, Austin Smith H659. "Growing in 
deep forest shade; gregarious but very local. Base of the stem dull, 
dark brown. Buds deep chokeberry red. Petioles and some of the 
veins of the under surface of the leaves tinged with pinkish red." 
This species, material of which was received when these pages were 
in proof, is not included in the key to species. It is easily recognized 
by the very abundant, rusty pubescence of the stems and flowers. 

Hoffmannia affinis Hemsl. Diag. PI. Nov. 31. 1879. Type, 
Endres 150, without locality. Known only from the original collec- 
tion, and to the writer only from description. Branches terete, 
puberulent when young; petioles 6 mm. long, the blades ovate- 
oblong, 10-12.5 cm. long, obtusely acuminate, attenuate to the base, 
minutely puberulent beneath; flowers 6-8 mm. long, puberulent, 
umbellate-cymose, the cymes fasciculate, about 6-flowered, the 
slender peduncles 8-16 mm. long, the pedicels 2-4 mm. long; calyx 
lobes rounded ; corolla tube slightly shorter than the lobes. 

Hoffmannia amplexifolia Standl. Journ. Wash. Acad. Sci. 
15: 8. 1925. Moist forest near Orosi, Prov. Cartago, Standley 39869. 
Also at Santo Domingo de Vara Blanca, 2,200 meters. Endemic. 



1314 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Stems simple, herbaceous, 1-1.5 meters high, glabrous, acutely 
quadrangular; leaves ternate, sessile, oblanceolate or oblong-obovate, 
23-35 cm. long, 8-13 cm. wide, or larger, acuminate or long-acumi- 
nate, long-attenuate to the very narrow, deeply cordate, clasping 
base, glabrous above, minutely puberulent beneath on the nerves; 
cymes many-flowered, 3-8 cm. long, lax, sessile, the flowers slender- 
pedicellate; calyx short-villous, the lobes narrowly triangular, obtuse; 
corolla greenish yellow, puberulent or villosulous, the lobes longer 
than the tube; fruit red. 

Hoffmannia arborescens Donn. Smith, Bot. Gaz. 37: 417. 
1904. In forest, Santa Rosa del Copey, 1,800 meters, Tonduz 8121 
(12230}. Known only from the original material. A shrub or small 
tree, the branchlets quadrangular; petioles 3-5 mm. long; blades 
oblanceolate-elliptic, 15-19 cm. long, 3.5-5.5 cm. wide, acuminate, 
long-attenuate to the base, glabrous; cymes fasciculate, many- 
flowered, fuscous-pubescent, 2-3.5 cm. long, the bracts foliaceous, 
1 cm. long; hypanthium and calyx pubescent, the lobes minute, 
deltoid; corolla 12 mm. long, pale rose, pubescent, the lobes about 
equaling the tube. 

Hoffmannia asclepiadea Standl. Journ. Wash. Acad. Sci. 15: 
7. 1925. Wet forest at Las Nubes, Prov. San Jose*, 1,900 meters, 
Standley 38349. Forests of the slopes of Irazu, 1,900-2,300 meters. 
Endemic. A simple shrub or herb, 1.5 meters high, erect or ascending, 
the stems villosulous; leaves opposite, on stout petioles 2.5-4 
cm. long, the blades elliptic or ovate-elliptic, 15-20 cm. long, 7-9 cm. 
wide, abruptly acuminate or long-acuminate, cuneate at the base, 
glabrous above, copiously short-villous beneath, especially on the 
nerves; cymes sessile, shorter than the petioles, many-flowered, the 
flowers short-pedicellate; hypanthium villosulous, the calyx lobes 
ovate-deltoid, obtuse, 1 mm. long; corolla greenish yellow, densely 
short-villous; fruit subglobose, red, sparsely short-villous. 

Hoffmannia carpinterae Standl. N. Amer. Fl. 32: 199. 1934. 
H. macrophylla Standl. Journ. Wash. Acad. Sci. 15: 9. 1925, non 
Hemsl. Wet forest, Cerro de La Carpintera, Prov. Cartago, 1,700 
meters, Standley 35636. Known only from the type locality. A 
shrub 3 meters high, with few branches, glabrous throughout; leaves 
sessile or nearly so, oblong-obovate, 21-35 cm. long, 8-10 cm. wide, 
abruptly acute, long-attenuate to the base; cymes sessile or peduncu- 
late, 3-4 cm. long, lax, few-flowered, the pedicels 3-8 mm. long; 



FLORA OF COSTA RICA 1315 

calyx red, the lobes rounded-deltoid, 1.5 mm. long; corolla in bud 
6 mm. long, the lobes yellow within, twice as long as the tube. 

Hoffmannia decurrens Standl. Contr. U. S. Nat. Herb. 20: 
205. 1919. Forests of Santa Rosa del Copey, 1,800-2,000 meters, 
Tonduz 12230. Mountains of Dota, 1,650-2,000 meters. Endemic. 
A sparsely branched shrub of 1-2.5 meters, the branchlets rufous- 
villosulous or glabrate; petioles 5-30 mm. long; blades oblong- 
oblanceolate to narrowly elliptic, 7-20 cm. long, 2-6 cm. wide, acute 
or acuminate, long-attenuate to the base, glabrous above, paler 
beneath, villosulous along the nerves or finally glabrate; cymes 
axillary or at naked nodes below the leaves, usually many-flowered, 
sessile or short-pedunculate, the pedicels 5 mm. long or less; calyx 
and hypanthium rufous-villous or villosulous, the lobes triangular, 
1-1.5 mm. long, obtuse; corolla white, tinged with rose, 8-9 mm. 
long, sparsely villosulous, the lobes obtuse, slightly shorter than the 
tube; fruit oval, red, 6-8 mm. long. 

Hoffmannia dotae Standl. Journ. Wash. Acad. Sci. 18: 181. 
1928. Moist forest near Santa Maria de Dota, Prov. San Jose", 1,700 
meters, Standley & Valeria 43277. Region of the type locality. 
Endemic. An erect, branched shrub 2-4.5 meters high, the branches 
glabrous or sparsely villous; petioles 2 cm. long or less, the blades 
oblong-obovate to obovate-elliptic, 18-30 cm. long, 8-12 cm. wide, 
abruptly short-acuminate, narrowed toward the abruptly long- 
decurrent base, glabrous above, at first sparsely short-villous beneath 
on the nerves but soon glabrate; cymes lax, few-flowered, 5.5 cm. 
long or shorter, the peduncles to 3.5 cm. long, the pedicels 4-12 
mm. long; hypanthium sparsely short-villous, the calyx lobes 2.5- 
3.5 mm. long, triangular, obtuse or acutish; corolla red below, 
yellow above, 15 mm. long, glabrous, or sparsely villous on the lobes, 
the lobes equaling the tube; fruit red, oblong, 8-9 mm. long. 

Hoffmannia hamelioides Standl. Journ. Wash. Acad. Sci. 15: 
8. 1925. Moist forest between Aserri and Tarbaca, Prov. San Jose", 
1,800 meters, Standley 34149. Known certainly only from the 
original material. A sparsely branched shrub of 1.5 meters, the 
branches glabrous; leaves opposite, the stout petioles 1-3 cm. long, 
the blades obovate-elliptic or oblanceolate-elliptic, 12-21 cm. long, 
4.5-7 cm. wide, acuminate, cuneate-decurrent to the base, glabrous 
above, villosulous beneath on the nerves; cymes sessile, with few or 
many flowers, about equaling the petioles; calyx short-villous, the 
lobes narrowly triangular, 1-1.5 mm. long; corolla in bud 4 mm. 



1316 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

long, greenish white, short- villous; fruit dark red, subglobose, 7-8 
mm. long. 

Hoffmannia inamoena Standl. Journ. Wash. Acad. Sci. 18: 
179. 1928. Wet forest, Los Ayotes, near Tilaran, Guanacaste, 600 
meters, Standley & Valerio 454%1 - Mountain forests of Guanacaste. 
Endemic. A simple, erect shrub of 1-1.5 meters, the stems terete, 
at first minutely puberulent; leaves opposite, the petioles 1-4.5 cm. 
long, the blades mostly elliptic, 8-20 cm. long, 3.5-10 cm. wide, 
abruptly acute or acuminate, obtuse to rounded and abruptly long- 
decurrent at the base, glabrous above, densely and minutely puberu- 
lent beneath on the nerves, or even over the whole surface; flowers 
fasciculate in the leaf axils, or in sessile or short-pedunculate, few- 
flowered cymes, the pedicels 4 mm. long or less; calyx lobes triangular- 
oblong, 1-2 mm. long, obtuse, short- villous; fruit subglobose, 6-7 mm. 
long, white, villous. 

I lamella josefina Standl. Journ. Wash. Acad. Sci. 15: 8. 1925. 
Moist forest between Aserri and Tarbaca, Prov. San Jose", 1,800 
meters, Standley 34143. Also in the region of Dota. Endemic. A 
sparsely branched shrub 3 meters high, the branches terete, glabrous; 
petioles 1-1.5 cm. long, the blades obovate or oblong-obovate, 13-22 
cm. long, 5.5-9 cm. wide, acute or abruptly short-acuminate, cuneate- 
attenuate to the base, glabrous; cymes with few or many flowers, 
pedunculate, longer than the petioles, the branches glabrous, the 
pedicels 1-3 mm. long; calyx glabrous or with a few short, scattered 
hairs, the lobes 2 mm. long, narrowly triangular, acute; corolla 11 mm. 
long, glabrous, the lobes shorter than the tube; fruit subglobose, 
6 mm. long, dark red. 

Hamelia latifolia (Bartl.) Kuntze has been reported from Costa 
Rica, but doubtless in error, the species being a Peruvian one. 

Hoffmannia leucocarpa Standl. Journ. Wash. Acad. Sci. 15: 
9. 1925. Wet forest, Las Nubes, Prov. San Jose", 1,600 meters, 
Standley 38340. Forests of the mountains of the central region; 
Zarcero; at 1,300-2,400 meters. Endemic. A simple or branched 
shrub, 2.5 meters high or less, glabrous throughout; leaves opposite, 
the petioles 3-6.5 cm. long, the blades elliptic or lance-elliptic, 15-25 
cm. long, 6-12 cm. wide, rather abruptly acuminate, acute or cuneate 
at the base; cymes equaling or often exceeding the petioles, lax, 
with few or many flowers, the branches bright red, the pedicels 
5-12 mm. long; calyx red, 2-3 mm. long, the lobes deltoid, acute; 



FLORA OF COSTA RICA 1317 

corolla yellow and red, 7 mm. long, glabrous, the lobes longer than 
the tube; fruit subglobose, 1 cm. long, white. 

Hoffmannia longepetiolata Polak. Linnaea 41: 567. 1877. 
Forests of La Carpintera, Polakowsky 134 (photo, of type in Herb. 
Field Mus.). Mountain forests of the central region, at 1,600-2,200 
meters. Endemic. A slender, glabrous shrub, the branches sub- 
terete; leaves opposite, the petioles 2-4 cm. long, the blades obovate 
or obovate-oblong, 10-19 cm. long, 3-7 cm. wide, abruptly acuminate, 
long-cuneate or attenuate to the base; cymes 3-8-flowered, sessile, 
much shorter than the petioles, the flowers slender-pedicellate or 
subsessile; calyx lobes minute, obtuse; corolla yellow, almost 1 cm. 
long, the lobes obtuse, much longer than the tube; fruit subglobose. 
Var. minor Polak. op. cit. 568 may be a form of this, but it is more 
likely to be a distinct species. I have seen no material of the variety. 

Hoffmannia nesiota Bonn. Smith, Bot. Gaz. 61: 374. 1916. 
Wafer Bay, Cocos Island, Pittier 12387. Branchlets sub terete, 
glabrous; leaves opposite, the stout petioles 8 cm. long, the blades 
broadly elliptic, 22-26 cm. long, 11 cm. wide, acuminate at each 
end, glabrous; cymes with few or many flowers, lax, the peduncles 
mostly 3.5-5 cm. long, the pedicels 6-10 mm. long; calyx lobes 
narrowly triangular, 1.5-2 mm. long, acute; corolla 11 mm. long, 
puberulent, the linear lobes acute, about equaling the tube; fruit 
oval, 1 cm. long, 3-celled. 

Hoffmannia pallidiflora Standl. Journ. Wash. Acad. Sci. 15: 
9. 1925. Wet forest, La Hondura, Prov. San Jose', 1,500 meters, 
Standley 37877. Mountain forests of the central region, 1,400-1,500 
meters. Endemic. A simple shrub 1-2 meters high, the stems 
obtusely tetragonous, glabrous; leaves opposite, the slender petioles 
2-4 cm. long, the blades elliptic or elliptic-obovate, 15-25 cm. long, 
7-9 cm. wide, abruptly acuminate, abruptly decurrent at the base, 
glabrous; cymes few-flowered, long-pedunculate, often at naked 
nodes below the leaves, equaling the petioles, the branches glabrous, 
the flowers partly sessile and partly slender-pedicellate; calyx pale 
green, the lobes triangular, obtuse or acute, 1-1.5 mm. long; corolla 
pale green, 8 mm. long, the lobes almost twice as long as the tube; 
fruit oval, red, 1 cm. long. 

Hoffmannia piratarum Standl. Journ. Wash. Acad. Sci. 18: 
180. 1928. Wafer Bay, Cocos Island, Pittier 16259. Known only 
from the original collection. Branchlets obtusely tetragonous, 
glabrous; leaves opposite, the slender petioles 2.5-4.5 cm. long, the 



1318 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

blades lance-oblong, 10-20 cm. long, 5-6 cm. wide, long-acuminate, 
glabrous above, sparsely short-villous beneath when young but soon 
glabrate; cymes solitary or fasciculate, dense, few-flowered, 3 cm. 
long or shorter, the peduncles 2.5 cm. long or less, the pedicels 2-5 
mm. long; hypanthium glabrous or sparsely short-villous; calyx 
lobes triangular-oblong, 2-3 mm. long, acute, villous-ciliate; corolla 
1 cm. long, glabrous or with a few short hairs at the apex, the lobes 
slightly shorter than the tube; fruit subglobose, 2-celled, 6 mm. long, 
glabrous. 

Hoffmannia psychotriaefolia (Benth.) Griseb. Fl. Brit. W. 
Ind. 321. 1861. Higginsia psychotriaefolia Benth. in Oerst. Vid. 
Medd. Kjoebenhavn 50. 1852. Volcan de Barba, 1,800 meters, 
Oersted (photo, of type in Herb. Field Mus.). Frequent in wet 
forest of the central mountains; region of San Ramon; at 1,000-1,600 
meters. Panama to Guatemala. A shrub 1-2.5 meters high, gla- 
brous throughout, usually branched, the branches sub terete; leaves 
opposite, the slender petioles 1-4 cm. long, the blades elliptic-oblong 
or elliptic, 7-15 cm. long, 2.5-5 cm. wide, cuspidate-attenuate, acute 
or attenuate at the base, bright green above, pale yellowish green 
beneath; cymes sessile, few-flowered, equaling or shorter than the 
petioles, the flowers short-pedicellate; calyx 1.5-2 mm. long, 
the lobes minute, deltoid; corolla yellow, sometimes tinged with 
red, 10-11 mm. long, the lobes acute, about equaling the tube; 
fruit red. 

Hoffmannia ramonensis Standl. Journ. Wash. Acad. Sci. 18: 
180. 1928. Along the Rio Barranca at San Juan, near San Ramon, 
1,300-1,400 meters, Tonduz 17812. Region of Zarcero, 1,260-1,500 
meters. Endemic. A shrub or small tree, sometimes 6 meters high, 
with a trunk as much as 12 cm. in diameter, the branches terete, 
glabrous or nearly so; leaves opposite, sessile or almost so, obovate- 
oblong, 14-28 cm. long, 6-10 cm. wide, acute or abruptly short- 
acuminate, gradually narrowed below the middle then rather abruptly 
long-attenuate into a petioliform portion 3-6 cm. long, glabrous 
above, densely tomentose beneath when very young but in age 
glabrate except along the nerves; cymes solitary or fasciculate, 
umbelliform, mostly 2-5-flowered, the peduncles 6-15 mm. long, the 
pedicels 2-4 mm. long, densely villous- tomentose; hypanthium 
2.5-3 mm. long, brown-tomentose, the calyx lobes narrowly tri- 
angular, 1.5-2 mm. long, narrowed to an obtuse apex; corolla in 
bud 6-7 mm. long, villous-tomentose, yellow; fruit dark crimson. 



FLORA OF COSTA RICA 1319 

Hoffmannia refulgens (Hook.) Hemsl. Mountains of Guana- 
caste; region of San Ramon; at 600-900 meters, growing in deep, wet 
forest; Atlantic tierra caliente. Ranging to southern Mexico. 
Plants simple, herbaceous or suffrutescent, usually less than 40 cm. 
high, the stems ferruginous- villous or glabrate, densely leafy above; 
leaves opposite, the stout petioles 3.5 cm. long or less, the blades 
obovate to oblong-oblanceolate, 9-25 cm. long, 4-11 cm. wide, 
rounded or obtuse at the apex and abruptly apiculate, acute to long- 
attenuate at the base, glabrous or sparsely villous above, often 
purple or reddish beneath, ferruginous- villous, at least on the nerves, 
or finally glabrate; flowers subumbellate or subracemose, sometimes 
secund, short-pedicellate, the slender peduncles 2.5-12 cm. long; 
calyx and hypanthium 3-4 mm. long, the hypanthium glabrous, the 
calyx lobes 2 mm. long or less, acute, sparsely villosulous; corolla 
red or pale red, 1 cm. long, glabrous, the lobes 2-3 times as long as 
the tube; fruit bright red. The plant, although small, is rather 
handsome because of its red flowers and fruits. It has been intro- 
duced into cultivation in Europe. It grows usually in the deepest, 
darkest forest. 

Hoffmannia subauriculata Standl. Journ. Wash. Acad. Sci. 
18: 179. 1928. Moist forest, El Mufieco, Rio Navarro, Prov. Cartago, 
1,400 meters, Standley & Torres 50956. Known only from the type 
region. Endemic. A slender, decumbent shrub 1-1.5 meters long, 
the branchlets obtusely quadrangular, glabrous; leaves opposite, 
sessile, elliptic-obovate, 15-22 cm. long, 7-9.5 cm. wide, abruptly 
short-acuminate, abruptly narrowed near the base into a petioliform 
portion about 2 cm. long, the very base rounded to cordate and 
amplexicaul, glabrous; cymes at naked nodes below the leaves, 
many-flowered, on slender peduncles 6-9 cm. long, the slender 
pedicels 6-12 mm. long; fruit oval or globose, bright red, glabrous, 
8 mm. long; calyx lobes deltoid, subacute, 1 mm. long. 

Hoffmannia Tonduzii Standl. Contr. U. S. Nat. Herb. 20: 205. 
1919. Forests of Las Vueltas, Tucurrique, 700 meters, Tonduz 
13373. Frequent in mountains of the central region, at 1,800 meters 
or less, descending to the upper part of the Atlantic tierra caliente; 
regions of Dota and San Ramon; said to be sometimes epiphytic. 
Endemic. A slender, usually branched shrub, 1-2 meters high, the 
branchlets green, terete, glabrous; leaves short-petiolate, mostly 
obovate-elliptic or oblong-oblanceolate, 5-12 cm. long, 1.5-5.5 cm. 
wide, acute to cuspidate-acuminate, acute to long-decurrent at the 



1320 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

base, glabrous; cymes sessile or subsessile, few-flowered, usually 
shorter than the petioles, the pedicels 5 mm. long or less; calyx lobes 
narrowly triangular, 1-1.5 mm. long, acute; corolla 6-7 mm. long, 
glabrous, yellow, the lobes acute, twice as long as the tube; fruit 
dark cherry-red, subglobose, 1 cm. long. 

Hoffmannia trichocalyx Standl. Journ. Wash. Acad. Sci. 18: 
181. 1928. Wet forest, Fraijanes, Prov. Alajuela, 1,600 meters, 
Standley & Torres 47690. Known only from the type region. En- 
demic. A large, weak shrub, 1-2.5 meters long, often decumbent, 
the stout branches terete, the young branchlets sparsely villous or 
glabrous; leaves opposite, the slender petioles 1.5-7 cm. long, the 
blades elliptic to oblong-elliptic, 12-26 cm. long, 5-10 cm. wide, 
abruptly acuminate, cuneate to obtuse at the base, usually abruptly 
contracted and short-decurrent, glabrous above, usually villous 
beneath along the nerves; cymes pendent from naked nodes below the 
leaves, fasciculate, 3-14 cm. long, with few or many flowers, the long, 
slender peduncles usually villous, dark red, the pedicels 3-6 mm. long; 
calyx lobes broadly deltoid, subacute, 2 mm. long, villous; corolla 
1 cm. long, bright yellow or red and yellow, glabrous or sparsely 
villous, the lobes about equaling the tube; fruit oval, 8 mm. long, 
dark red, glabrous or sparsely villous. 

Hoffmannia Valerii Standl. Journ. Wash. Acad. Sci. 18: 178. 
1928. El Arenal, Guanacaste, 600 meters, Juvenal Valeria 57. 
Mountain forests of Guanacaste, 600-700 meters. Endemic. A 
branched shrub 1-1.5 meters high, the branches subterete, densely 
villous when young; leaves opposite, the slender petioles 1-2 cm. 
long, the blades elliptic or ovate-elliptic, 7-11 cm. long, 3.5-5.5 cm. 
wide, acute or acuminate, obtuse or rounded at the base and abruptly 
decurrent, sparsely villous above when young but soon glabrate, 
villous beneath along the nerves; cymes few-flowered, dense, fascicu- 
late, sessile or nearly so, the peduncles in fruit sometimes 1 cm. long, 
the pedicels 2-5 mm. long, glabrous or nearly so; hypanthium gla- 
brous or with a few short hairs, the calyx lobes narrowly triangular, 
1 mm. long, acute or obtuse; corolla in bud 5-6 mm. long, short- 
villous, the lobes obtuse, 3 times as long as the tube; fruit subglobose, 
6 mm. long, bright red, glabrous. 

HOLTONIA Standl. 

Trees, almost glabrous; leaves opposite, thick-membranaceous, 
petiolate; stipules resiniferous, persistent, connate to form a truncate 
or shallowly bilobate sheath; inflorescence terminal, many-flowered, 



FLORA OF COSTA RICA 1321 

paniculate, the flowers small, 5-parted; calyx cupular, shallowly and 
remotely denticulate; corolla white, tubular-campanulate, glabrous 
outside, barbate within at the insertion of the stamens, the short 
lobes broadly triangular, obtuse, valvate or subimbricate, one-third 
as long as the tube; anthers exserted; capsule small, subglobose, 
loculicidally bivalvate; seeds numerous, minute, angulate. The 
genus consists of a single species. 

Holtonia myriantha Standl. Sickingia myriantha Standl. 
El General, 975 meters, Skutch 2387. Colombia. A tree 23 meters 
high, the trunk 28 cm. in diameter; branchlets glabrous; petioles 1-3 
cm. long, the blades obovate to oblong-elliptic, 8-20 cm. long, 4-8 
cm. wide, abruptly short-acuminate, acute or attenuate at the base, 
densely short-barbate beneath in the axils of the nerves, otherwise 
glabrous; panicles 10-20 cm. long and broad, the branches sparsely 
and minutely puberulent, the flowers sessile; calyx 0.5 mm. long; 
corolla 4 mm. long, 3.5 mm. wide; capsule 2-2.5 mm. long. 

ISERTIA Schreb. 

Trees or shrubs, pubescent or glabrous, the branchlets terete; 
leaves opposite or ternate, petiolate; stipules 2 between each 2 leaves, 
narrow, erect, persistent; flowers usually large, short-pedicellate, 
mostly red and yellow, cymose-corymbose or cymose-paniculate; 
hypanthium subglobose or ovoid; calyx short, 4-6-dentate or trun- 
cate, persistent; corolla tubular-funnelform or salverform, coriaceous, 
barbate in the throat, the limb 4-6-lobate, the lobes short, valvate; 
ovary 4-6-celled, many-ovulate; fruit small, baccate, globose or ovoid, 
4-6-pyrenate; seeds minute, subglobose, foveolate. One other 
Central American species occurs in Panama. 

Isertia Haenkeana DC. I. Deamii Bartlett, var. stenophylla 
Bonn. Smith, Bot. Gaz. 61: 374. 1916 (plains near Boca Culebra, 
Prov. Puntarenas, 50 meters, Pittier 11989). Thickets of the Pacific 
coast; while I have seen no specimens from the Atlantic tierra caliente 
of Costa Rica, the species doubtless occurs there also, for it is common 
along the coast both north and south of Costa Rica. Guatemala to 
Colombia; Cuba. A shrub 2-3 meters high, the branches densely 
sericeous; stipules 6-12 mm. long; petioles stout, 2.5 cm. long or less, 
the blades obovate to oblong-obovate, 20-45 cm. long, 8-18 cm. 
wide, abruptly acuminate, acute or attenuate at the base, green and 
glabrate above, densely grayish-pilose and pale beneath; inflorescence 
usually thyrsiform, 7-20 cm. long, dense and many-flowered; corolla 



1322 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

17-35 mm. long, yellow or orange-red, sparsely or densely puberulent 
or tomentulose; fruit purple, depressed-globose, 5-6 mm. broad. 

IXORA L. 

Trees or shrubs, glabrous or pubescent, the branchlets terete or 
angulate; leaves opposite or rarely ternate, sessile or petiolate, 
usually coriaceous; stipules mostly acuminate from a broad base, 
persistent or deciduous; flowers in terminal or axillary corymbs, 
variously colored; calyx short, usually 5-lobate, persistent, the lobes 
short or elongate; corolla salverform, the throat glabrous or pilose, 
the lobes usually short, spreading, contorted; anthers wholly or par- 
tially exserted; ovary 2-celled, the cells 1-ovulate, the ovules peltately 
attached to the middle of the septum; fruit baccate, the 2 carpels 
concavo-convex. No other species are known from Central America. 

Ixora coccinea L. Jazmin. Planted commonly for ornament. 
Native of India. A small, dense shrub, glabrous or nearly so; 
leaves sessile, broadly oblong to suborbicular, obtuse or rounded 
at the apex, generally cordate at the base; inflorescence dense and 
headlike, the flowers sessile or nearly so; calyx lobes broadly ovate, 
obtuse or acutish; corolla deep red, minutely puberulent or glabrate, 
the filiform tube 2.5-3 cm. long, the spreading lobes acute. 

Ixora Finlaysoniana Wallich. Corona de la reina. Grown 
commonly as an ornamental shrub. Native of Siam. A dense shrub 
of 1-2 meters, glabrous or nearly so; leaves short-petiolate, oblong 
or oblanceolate-oblong, obtuse or acute, attenuate to the base; 
inflorescence dense and headlike, many-flowered; corolla white, the 
filiform tube about 3.5 cm. long, the obtuse lobes 6-7 mm. long. 

Ixora floribunda (A. Rich.) Griseb. Palo de Maria. Monte 
Aguacate, Friedrichsthal 1288. Extending to Salvador; Cuba; 
Colombia. A shrub or tree, sometimes 9 meters high; petioles 1-2.5 
cm. long, the blades elliptic to elliptic-oblong, 11-25 cm. long, 
4-11 cm. wide, acute or short-acuminate, attenuate to subacute at 
the base, glabrous; inflorescence cymose-paniculate, short-peduncu- 
late or sessile, 4-10 cm. broad, the branches densely pilose, the 
flowers sessile or short-pedicellate; calyx shallowly lobate; corolla 
white, glabrous, 6-10 mm. long, the lobes about equaling the tube; 
fruit globose, short-pilose, 4 mm. or more in diameter. 

Ixora nicaraguensis Standl. I. rauwolfioides Standl. Forests 
of the Atlantic tierra caliente. Panama to British Honduras. A 
shrub or tree, 2.5-7.5 meters high, the trunk sometimes 12 cm. in 



FLORA OF COSTA RICA 1323 

diameter; petioles 4-10 mm. long; blades elliptic-oblong or lance- 
oblong, 7-15 cm. long, 2-5.5 cm. wide, acute or short-acuminate, 
acute or acuminate at the base, glabrous; inflorescence cymose- 
paniculate, sessile or pedunculate, 5-9 cm. long, the branches mi- 
nutely puberulent, the pedicels 5 mm. long or less; calyx truncate or 
nearly so; corolla white, glabrous, the tube 5 mm. long, the lobes 
3-4 mm. long, rounded at the apex. 

LADENBERGIA Klotzsch 

Trees or shrubs; leaves mostly opposite, petiolate, coriaceous; 
stipules deciduous; flowers small or medium-sized, white or pink, in 
terminal panicles; hypanthium ovoid or turbinate, pubescent, the 
calyx cupular, 5-lobate, persistent or deciduous; corolla salverform, 
pubescent, the lobes valvate, papillose within and on the margins; 
ovary 2-celled, many-ovulate; capsule oblong or cylindric, septicidally 
bi valvate from the apex to the base; seeds numerous, imbricate, 
broadly winged. The other members of the genus are South 
American. 

Ladenbergia Brenesii Standl., sp. nov. Quina, Agujilla. Arbor 
5-10-metralis, ramulis crassis obtuse tetragonis ochraceis glabris; 
stipulae ovales 2-3 cm. longae obtusae coriaceae glabrae vel extus 
sparse minute strigillosae; folia magna coriacea, petiolo crasso glabro 
1-3.5 cm. longo; lamina oblonga usque oblongo-elliptica vel elliptico- 
obovata plerumque 10-25 cm. longa atque 5-10 cm. lata, obtusa vel 
subacuta, basi late rotundata usque subacuta, supra lucida glabra 
saltern in stato adulto glaberrima, costa elevata, nervis lateralibus 
utroque latere ca. 10 prominentibus obliquis; inflorescentia cymoso- 
paniculata dense multiflora, basi trichotoma, crasse pedunculata, 
ca. 11 cm. longa, ramis crassis dense sericeis, floribus sessilibus vel 
brevissime pedicellatis; hypanthium clavatum 4-6 mm. longum 
densissime pilis brunnescentibus subadpressis indutum; calyx 2 mm. 
longus, lobis late ovalibus apice rotundatis extus dense adpresso- 
pilosis; corolla extus dense fulvo-sericea, tubo 2 cm. longo sursum 
sensim dilatato atque 5-6 mm. lato, lobis lanceolato-oblongis 8 mm. 
longis acutis vel acutiusculis intus dense minute papillosis; capsula 
2-4.5 cm. longa, valvis ca. 1 cm. latis. San Ramon, March, 1932, 
Brenes 15146 (type in Herb. Field Mus.). Los Angeles y La Paz de 
San Ramon, March, 1928, Brenes 6094. Alto de La Palma de San 
Ramon, 1,150 meters, February, 1923, Brenes 3837. La Palma de 
San Ramon, 1,250 meters, July, 1927, Brenes 5591. Alto de La 
Estrella, Prov. Cartago, Standley 39298. La Palma, Prov. San Jose", 



1324 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

1,460 meters, Tonduz 12647 (J. D. Smith 7397). In the North 
American Flora (32: 95. 1921) I referred this tree incorrectly to 
L. undata Klotzsch, a species of Colombia and Venezuela, chiefly 
because at that time adequate South American material was not 
available for study. Comparison with authentic material of that 
species shows that the Costa Rican tree is altogether distinct, nor 
does it agree better with any other Colombian species of Ladenbergia. 

Ladenbergia sericophylla Standl., sp. nov. Arbor usque 36 m. 
alta, trunco 55 cm. diam., ramulis breviter pilosis vel glabratis; 
stipulae late oblongae 3.5 cm. longae obtusae extus sparse adpresso- 
pilosae; folia magna papyracea, petiolo usque 6 cm. longo glabrato; 
lamina elliptica vel late elliptica, 30 cm. longa atque 17 cm. lata vel 
minor, breviter acutata vel apice subrotundata, basi obtusa vel 
subrotundata atque subito breviter decurrens, supra lucida glabra, 
subtus in statu juvenili ubique minute subsparse strigillosa, ad 
nervos venasque densissime fulvo-sericea, in statu adulto glabrata, 
costa crassa, nervis lateralibus utroque latere ca. 10 angulo recto 
vel paullo latiore adscendentibus; inflorescentia perfecta non visa; 
hypanthium clavatum 7 mm. longum dense brunneo-sericeum; calyx 
6-7 mm. longus late campanulatus ferrugineus prope basin sericeus, 
superne fere glaber, ad tertiam partem lobatus, lobis semiorbiculari- 
bus subrecurvis; corolla extus subdense fulvo-strigosa, tubo crasso 
3 cm. longo subtereti, fauce 6 mm. lato. Two miles southeast of 
San Isidro del General, August, 1936, Ralph E. Danforth 32 (type 
in Herb. Field Mus.). El General, 880 meters, January, 1936, 
A. F. Skutch 2539 (detached flowers and tracing of a leaf, from U. S. 
Nat. Herb.). I once determined these two collections asL. calycina 
(Wedd.) Standl., a species of Colombia. While the Costa Rican 
and Colombian trees are related, because of the form of the calyx, 
more careful study shows that they are specifically distinct. The 
flowers are described by the collectors as white, pinkish, or yellowish 
white, and very fragrant. Mr. Danforth reports that the leaves are 
as much as 35 cm. long and 12 cm. wide, the lateral nerves being 
depressed so that the leaf, when fresh, appears to be slightly fluted. 

Ladenbergia Valerii Standl., sp. nov. Arbuscula 3-5-metralis, 
ramis elongatis, ramulis teretibus cinereo-brunnescentibus striatis, 
novellis glabris; stipulae subcoriaceae ca. 1.5 cm. longae ovali-ovatae 
obtusae ferrugineae glabrae; folia breviter petiolata coriacea modica, 
petiolo crasso 1-2 cm. longo glabro; lamina elliptica vel anguste 
elliptica 12-15 cm. longa 5-7.5 cm. lata breviter acuminata, basi 
acuta, supra in sicco fusca lucida glabra, costa nervisque subsulcatis, 



FLORA OF COSTA RICA 1325 

subtus brunnescens sparse secus costam nervosque hirtella, aliter 
glabra vel glabrata, costa crassiuscula elevata, nervis lateralibus 
utroque latere ca. 11 angulo latiusculo adscendentibus subarcuatis 
teneris prominentibus; paniculae parvae dense pauciflorae breviter 
pedunculatae, corollis exclusis vix 6 cm. longae, ramis crassis sparse 
strigosis, floribus sessilibus vel breviter pedicellatis; hypanthium 
anguste turbinatum 4-5 mm. longum dense fulvo-sericeum; calyx 
fere ad basin lobatus glaber vel glabratus, lobis 5-6 mm. longis late 
ovalibus apice rotundatis plus minusve recurvis; corolla extus dense 
adpresse fulvo-pilosa, tubo 2 cm. longo sursum sensim dilatato, 
fauce 5 mm. lato, lobis 1 cm. longis lineari-lanceolatis acutis, intus 
dense minute papillosis. In forest, La Paz de San Ramon, 1,225 
meters, June, 1925, Brenes 4296 (type in Herb. Field Mus.). The 
species is dedicated to Professor Juvenal Valerio Rodriguez, Director 
of the Museo National de Costa Rica. While related to L. seri- 
cophylla, it is, I believe, sufficiently distinct in characters of pubescence 
and inflorescence. 

LINDENIA Benth. 

Shrubs, the branches terete; petioles short, connate, cuspidate, 
more or less persistent; leaves short-petiolate, narrow; flowers large 
and showy, white, short-pedicellate, in few-flowered, terminal cymes; 
hypanthium elongate-turbinate, 5-costate, the calyx 5-lobate, the 
lobes elongate-subulate or lanceolate, persistent; corolla salverform, 
the tube greatly elongate, the throat naked, the 5 lobes oblong, 
spreading, contorted in bud; anthers exserted; ovary 2-celled, many- 
ovulate; capsule clavate or pyriform, septicidally bivalvate, the 
valves bifid, often coiled after dehiscence; seeds small, angulate, 
smooth. The genus consists of a single species. 

Lindenia rivalis Benth. Lirio. Along streams, usually on rocks 
at the edge of water, region of Tilaran, Guanacaste, and probably 
elsewhere. Panama to southern Mexico. A stout, simple or sparsely 
branched shrub, usually less than a meter high; leaves short-petiolate, 
oblanceolate to linear-elliptic, 5-15 cm. long, 1-2.5 cm. wide, acute 
or attenuate at each end, puberulent or pilosulous or almost glabrous; 
cymes mostly 3-flowered; calyx lobes 1-2 cm. long, green; corolla tube 
usually 10-16 cm. long, pilosulous, the lobes 2.5-3 cm. long; capsule 
1-2 cm. long. The plant is a conspicuous and showy one when in 
blossom. 

MACHAONIA Humb. & Bonpl. 

Shrubs or small trees, glabrous or pubescent, the branches terete, 
often spinose; leaves opposite or ternate, petiolate or sessile; stipules 



1326 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

small, deltoid, acute or acuminate; flowers small, white, the inflores- 
cence terminal; hypanthium turbinate, compressed; calyx 4-5-lobate, 
the lobes persistent; corolla short-funnelform, the tube short, villous 
in the throat, the 4-5 lobes rounded, imbricate; ovary 2-celled, the 
cells 1-ovulate; fruit small, dry, laterally compressed, dicoccous, the 
cocci indehiscent. Two other species are known from Central 
America, and one of them, M. acuminata Humb. & Bonpl., with 
pubescent leaves, is to be expected in Costa Rica. 

Machaonia rotundata Griseb. Thickets of the Atlantic coast. 
Panama and Jamaica. A shrub of 3 meters or less; stipules 3-4 mm. 
long; petioles 3-10 mm. long, the blades oval to broadly ovate, 5-9 
cm. long, 2-5.5 cm. wide, abruptly short-acuminate, rounded or 
obtuse at the base, glabrous, sometimes obscurely barbate beneath 
in the axils of the nerves; panicles pyramidal or rounded, 6-13 cm. 
broad, the flowers densely glomerate; calyx and hypanthium 3 mm. 
long, glabrous or sparsely puberulent, the calyx lobes oval-oblong to 
suborbicular, rounded at the apex, ciliate; corolla 4-5 mm. long, 
glabrous outside; fruit cuneate-oblong, 4.5-6 mm. long. 

Machaonia rotundata var. Dodgei Standl., var. nov. Petiolus 
gracilis 7-8 mm. longus; lamina glabra elliptica vel elliptico-oblonga 
4.5-6 cm. longa 2-2.8 cm. lata, apice obtusa vel rotundata atque 
emarginata, interdum acuta vel acuminata, basi acuminata. Beach 
between Rio Sandalo and Rio Tigre, May, 1930, C. W. Dodge 7752 
(type in Herb. Field Mus.; dupl. in herb. Mo. Bot. Card.). Note- 
worthy for the small leaves, acuminate at the base and often emargi- 
nate at the apex. At first I was inclined to regard it as a new species, 
but the flowers appear to be identical with those of M. rotundata, 
and the plant is probably better treated as a variety, for the present 
at least. 

MACROCNEMUM P. Br. 

Trees or shrubs, glabrous or pubescent, the branches terete; 
stipules oblong or obovate, deciduous; leaves opposite, petiolate; 
flowers small or medium-sized, white or pink, in terminal or axillary 
panicles; calyx cupular, 5-dentate, persistent; corolla funnelform or 
salverform, the tube elongate, the 5 lobes spreading, puberulent 
within, valvate or reduplicate; anthers included; ovary 2-celled, 
many-ovulate; capsule oblong or subcylindric, bisulcate, loculicidally 
bi valvate; seeds numerous, minute, compressed, winged. One other 
species grows in Central America, in Panama. 



FLORA OF COSTA RICA 1327 

Macrocnemum glabrescens (Benth.) Wedd. Ann. Sci. Nat. IV. 
1: 76. 1854. Palo cuadrado. Lasionema glabrescens Benth. Bot. Voy. 
Sulph. 105. 1845. Nicoya, Barclay. Thickets of the Pacific coast. 
Ranging to Colombia. A tree of 10-15 meters; stipules 1-2 cm. long; 
petioles 5-15 mm. long; blades obovate or obovate-elliptic, 7-20 cm. 
long, 3-8 cm. wide, acute or short-acuminate, rarely rounded, cuneate 
or attenuate at the base, glabrous above, sparsely strigose beneath 
or almost glabrous, usually sparsely barbate in the axils of the nerves; 
inflorescence 6-20 cm. broad, pedunculate, the flowers sessile or 
short-pedicellate; calyx teeth minute, acute; corolla pink, glabrous 
outside, the tube 4-10 mm. long, the rounded lobes 4 mm. long. 

MANETTIA Mutis 

Plants scandent, herbaceous or suffrutescent, glabrous or pubes- 
cent; stipules short, broad, persistent; leaves opposite, small, petio- 
late; flowers small or large, solitary and axillary or in cymes or 
panicles, white, red, or yellow; calyx usually 4-lobate, persistent, 
the lobes short or elongate, broad or narrow, often alternating with 
minute teeth or large lobes; corolla tubular or funnelform, the tube 
short or elongate, the limb 4-lobate, the lobes usually short, erect or 
recurved, valvate; anthers exserted or included; ovary 2-celled, many- 
ovulate; capsule obovoid or turbinate, septicidally bivalvate from 
the apex; seeds compressed, winged. One other species has been 
described from Central America. 

Manettia barbata Oerst. Vid. Medd. Kjoebenhavn 47. 1852. 
Lygistum barbatum Kuntze, Rev. Gen. 287. 1891. M. stenophylla 
Bonn. Smith, Bot. Gaz. 56: 58. 1913 (in thickets, Las Vueltas, 
Tucurrique, 635 meters, Tonduz 12969). Monte Aguacate, 600 
meters, Oersted (photo, of type in Herb. Field Mus.). Occasional in 
thickets, ascending to 2,450 meters in the region of Dota; region of 
San Ramon. Endemic. Stems glabrous; leaves short-petiolate, 
lance-linear to lanceolate, 3-5 cm. long, narrowly long-acuminate, 
acute at the base, glabrous, the lateral nerves obscure; calyx lobes 4, 
oblong-ovate, 2 mm. long; corolla white, 5-6 mm. long, glabrous 
outside, the lobes barbate within; capsule 4.5 mm. long. 

Manettia coccinea (Aubl.) Willd. M. costaricensis Wernham, 
Gen. Manettia 38. 1919 (Endres 2, without locality). Thickets of 
the Atlantic tierra caliente, at 900 meters or less; Guanacaste; 
region of San Ramon, 1,100 meters. Southern Mexico to West 
Indies and Guianas. Stems glabrous or short-pilose; leaves lanceo- 
late to ovate, 3-10 cm. long, conspicuously nerved, acute or acumi- 



1328 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

nate, acute or obtuse at the base, glabrous or scaberulous above, 
short-pilose or glabrous beneath; calyx lobes 8, linear or oblanceolate, 
5-14 mm. long, recurved; corolla 17-28 mm. long, pink or dull red, 
more or less pilose outside; capsule 6-10 mm. long. 

Manettia flexilis Brandegee. M. estrellae Standl. Journ. Wash. 
Acad. Sci. 15: 6. 1925 (wet forest, La Estrella, Prov. Cartago, 
Standley 3922 J^). Forests south of Cartago, 1,500 meters; region of 
San Ramon, about 1,100 meters. Extending to Mexico, and reported 
from Ecuador. Stems puberulent or villosulous; leaves ovate to 
lance-ovate, 3-7 cm. long, acute to long-attenuate, acute or obtuse 
at the base, glabrous to villosulous above, short-pilose or villosulous 
beneath; calyx lobes 4, linear to lance-ovate, 1.5-4 mm. long, acute, 
recurved; corolla rose-colored, glabrous or sparsely pilose, the tube 
6-7 mm. long, the lobes of equal or less length; capsule 5-7 mm. 
long. While I have not seen recently the type material of M. estrellae, 
examination of a photograph of the type and study of the descrip- 
tion lead to the conclusion that it is too close to M . flexilis to merit 
specific rank. 

MITRACARPUS Zucc. 

Annual or perennial herbs; stipules united with the petioles to 
form a setiferous sheath; leaves opposite, usually narrow; flowers 
small or minute, 4-parted, in terminal or axillary heads; sepals 
unequal, connate below; corolla funnelform, the lobes valvate; 
stamens included or subexserted; fruit 2-celled, the cells 1-seeded, 
transversely or obliquely circumscissile. At least one other species 
is known from northern Central America. 

Mitracarpus hirtus (L.) DC. In thickets or waste ground, 
Meseta Central to the coasts, 1,300 meters or less. Generally dis- 
tributed in tropical America. A low annual, usually less than 50 cm. 
high, simple or sparsely branched, the branches puberulent or villous; 
leaves subsessile, lanceolate to ovate-oblong, acute, 2-5 cm. long; 
flowers densely clustered in the leaf axils and at the ends of the 
branches; corolla white, 1-2 mm. long; capsule subglobose, bisulcate, 
short-pilose, scarcely 1 mm. long. 



The genus Morinda probably is represented in the tierra caliente, 
but I have seen no Costa Rican specimens. 

NOTHOPHLEBIA Standl. 

Trees, the branchlets thick, sub terete; leaves large, opposite, 
petiolate, subcoriaceous, finely striate-lineolate between the veins; 



FLORA OF COSTA RICA 1329 

stipules large, elongate; flowers large, sessile or pedicellate, the 
inflorescence axillary, cymose-corymbose, many-flowered, pedun- 
culate; hypanthium turbinate, the calyx campanulate, subtruncate, 
irregularly and very shallowly undulate-lobate, persistent; corolla 
funnelform, coriaceous, the throat glabrous, the 5 lobes short, valvate; 
anthers included; ovary 2-celled, many-ovulate; fruit not seen but 
doubtless baccate. The genus consists of a single species, which 
should perhaps be referred to Pentagonia. 

Nothophlebia costaricensis Standl. Contr. U. S. Nat. Herb. 
17: 438. 1914. Hills of Moin, Atlantic tierra caliente, Pittier 16024. 
Known only from the original collection. A tree with a depressed 
crown, the branchlets minutely sericeous or glabrate; stipules lance- 
oblong, 3.5^4 cm. long, attenuate, sparsely sericeous outside, gla- 
brous within; petioles 4-5.5 cm. long, the blades oval, 33 cm. long, 
17 cm. wide, very obtuse, rounded and short-decurrent at the base, 
glabrous above, minutely sericeous beneath along the veins or gla- 
brate, the lateral nerves about 12 on each side; inflorescence many- 
flowered, lax, short-pedunculate, the stout pedicels 7 mm. long or 
less; hypanthium 4-5 mm. long, sericeous; calyx 4-5 mm. long and 
broad, sparsely and minutely sericeous outside; corolla puberulent, 
the tube 2 cm. long, gradually dilated from the base to the apex, 
the lobes 4-5 mm. long. 

OLDENLANDIA L. 

Annual or perennial herbs, glabrous or pubescent; leaves opposite, 
sessile or petiolate, usually very small; stipules small, acute or acu- 
minate, sometimes setiferous; flowers in dichotomous, axillary and 
terminal panicles or cymes, or sometimes solitary; calyx 4-parted, 
the lobes usually erect and remote in fruit; corolla rotate or funnel- 
form, the tube short or elongate, the lobes obtuse, valvate; anthers 
commonly exserted; ovary 2-celled, many-ovulate; capsule small, 
globose or turbinate, loculicidally dehiscent at the apex or for the 
whole length; seeds few, angulate or subglobose, smooth or minutely 
granulate. One other species occurs in northern Central America. 

Oldenlandia corymbosa L. Reported from Costa Rica, prob- 
ably in the Atlantic tierra caliente. Widely distributed in tropical 
regions of both hemispheres. Annual, erect or decumbent, glabrous 
except on the leaves, usually much branched; stipules with 1 or more 
long setae; leaves sessile or subsessile, linear or linear-lanceolate, 
1-3.5 cm. long, acute, 1-nerved, usually scaberulous above; flowers 
in small, axillary cymes, the slender pedicels filiform; corolla white, 



1330 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

equaling or slightly exceeding the calyx lobes; capsule subglobose, 
1.5-2 mm. broad. 

Oldenlandia herbacea (L.) DC. Atlantic tierra caliente, in 
waste ground; pastures near San Jose'. Widely distributed in the 
tropics of both hemispheres. Annual, erect or procumbent, glabrous; 
leaves sessile, usually linear, mostly 2-5 cm. long; flowers axillary, 
usually solitary, the filiform pedicels 8-20 mm. long; corolla white, 
about equaling the calyx lobes; capsule 2.5-3 mm. long. 

OREGANDRA Standl. 

Glabrous trees; leaves opposite, short-petiolate; stipules decidu- 
ous; flowers pedicellate, in terminal, many-flowered panicles; hypan- 
thium obovoid; calyx short, 5-dentate; corolla cylindric-campanulate, 
the tube obconic, glabrous within, the 5 lobes short, rounded, im- 
bricate; stamens inserted near the base of the corolla tube, the 
glabrous filaments exserted; ovary 2-celled, many-ovulate. The 
genus consists of a single species. 

Oregandra panamensis Standl. Field Mus. Bot. 4: 265. 1929. 
Talamanca Valley, Panama, Cooper & Slater 144- Doubtless extend- 
ing into Costa Rica. Branchlets terete or subcompressed; petioles 
1-1.5 cm. long, the blades oblong, obovate-oblong, or rarely ovate, 
14-22 cm. long, 5.5-11 cm. wide, acute or abruptly short-acuminate, 
acute or almost rounded at the base, the lateral nerves 8-9 on each 
side; panicles 6.5-7.5 cm. long, 8-11 cm. broad, the stout pedicels 
2-4 mm. long; calyx 1.5 mm. high, 2.5-3 mm. broad, remotely 
5-denticulate; corolla tube 3.5 mm. long, the lobes 1.5-2 mm. long; 
anthers linear, 3 mm. long. The genus probably is related to Sickingia. 

PALICOUREA Aubl. 

Shrubs or small trees; leaves opposite or ternate, petiolate, 
usually somewhat coriaceous; stipules persistent, small or large, 
usually united below to form a short sheath, bilobate, the lobes 
broad or narrow; flowers small, cymose, corymbose, or paniculate, 
the panicles often thyrsiform, the inflorescence terminal, the branches 
usually colored or pale; calyx small, dentate; corolla tubular, with 
very short lobes, the tube commonly more or less curved and some- 
what dilated on one side at the base; ovary 2-celled, the cells 1- 
ovulate; fruit drupaceous, commonly 2-celled, containing 2 pyrenes. 
A few other species are known from Central America. The genus is 
an American one, and very large, but it is not separated from Psycho- 
tria by any constant character. Nevertheless it is a convenient 



FLORA OF COSTA RICA 1331 

group, and the species usually can be referred to the genus by their 
general appearance, especially by the more or less red, yellow, or 
blue coloring of the flowers, which extends to the bracts and the 
branches of the inflorescence. 

Leaves ternate P. triphylla. 

Leaves opposite. 

Inflorescence corymbiform, flat-topped, the branches erect or 
strongly ascending and fastigiate; corolla glabrous or minutely 
puberulent; leaves glabrous or nearly so P. fastigiata. 

Inflorescence paniculate, the branches usually spreading or ascend- 
ing at a rather wide angle, not fastigiate. 

Leaves glabrous beneath. 

Calyx 5 mm. long P. macrocalyx. 

Calyx 1-3 mm. long. 

Lobes of the stipules obtuse; corolla densely and minutely 
tomentulose P. guianensis. 

Lobes of the stipules acute to subulate; corolla not tomen- 
tulose; leaves small, most of them 2.5 cm. wide or 
narrower. 

Calyx lobes 1.5-2.5 mm. long, oblong to spatulate, 
rounded at the apex; lobes of the stipules 1.5-2.5 mm. 
long P. salicifolia. 

Calyx lobes less than 1 mm. long, obtuse or acute; lobes of 
the stipules about 1 mm. long P. montivaga. 

Leaves variously pubescent beneath, at least along the nerves 
and costa. 

Corolla glabrous outside. 

Calyx lobes conspicuous, 2.5-3 mm. long ... P. macrosepala. 
Calyx lobes inconspicuous, scarcely 1 mm. long. 

Leaves only 4-6 cm. long; corolla 8 mm. long, violet. 

P. adusta. 

Leaves mostly more than 10 cm. long; corolla more than 
1 cm. long, yellow P. Galeottiana. 

Corolla puberulent to villous, sometimes only sparsely hir- 
tellous at the apex. 

Lateral nerves of the leaves about 20 pairs P. vestita. 

Lateral nerves mostly 15 or fewer pairs. 



1332 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Leaves narrowly elongate-lanceolate; corolla merely 

puberulent P. angustifolia. 

Leaves elliptic-oblong to obovate-oblong; corolla hir- 

tellous or villous. 
Corolla very sparsely hirtellous, usually only near the 

apex P. costaricensis. 

Corolla densely short-villous with spreading or ap- 
pressed hairs P. lasiorrhachis. 

Palicourea adusta Standl. Journ. Wash. Acad. Sci. 18: 279. 
1928. Wet forest, Cerro de Las Vueltas, Prov. San Jose", 3,000 
meters, Standley & Valeria 43666. Occasional in forests of the central 
mountains; region of San Ramon and Zarcero; mostly at 1,800-3,000 
meters. Endemic. Sometimes a low, decumbent shrub a meter high 
or less, but reported also as a tree of 5.5 meters, with a trunk 20 cm. 
in diameter, the bark brown, slightly roughened, the branchlets 
sparsely pilose or glabrous; stipule lobes linear, acute, 1-2.5 mm. 
long; leaves slender-petiolate, elliptic to elliptic-oblong, mostly 4-6 
cm. long and 1.5-2.5 cm. wide, but sometimes as much as 3.5 cm. 
wide, gradually or abruptly acuminate or long-acuminate, obtuse or 
acute at the base, when young pubescent beneath with short, ap- 
pressed or spreading hairs, but in age almost completely glabrous; 
inflorescences cymose-paniculate, sparsely branched, almost as broad 
as long, the branches puberulent or glabrous; calyx lobes scarcely 
1 mm. long, triangular, acute to obtuse; corolla violet, 8 mm. long, 
glabrous. Noteworthy for the violet flowers. It is possible that two 
species are represented by the material now referred to this species. 

Palicourea angustifolia HBK. P. lanceolata Oerst. Vid. Medd. 
Kjoebenhavn 17. 1852 (Candelaria, 2,100 meters, Oersted 11556; 
photo, of type in Herb. Field Mus.). P. intermedia Oerst. ex Polak. 
Linnaea 41: 571. 1877 (forests of La Carpintera, Polakowsky 143; 
photo, in Herb. Field Mus.). Occasional in forests of the Central 
region, especially near Cartago; region of Dota; chiefly at 1,400-1,800 
meters. Extending to Peru. A slender shrub or small tree, the young 
branches usually densely puberulent; stipule sheath 4-6 mm. long, 
the lobes erect, subulate, equaling or longer than the sheath; leaves 
on very short petioles, mostly elongate-lanceolate, 6-20 cm. long, 
long-attenuate, obtuse or acute at the base, puberulent or pilose 
beneath, at least along the nerves; panicles pedunculate, narrow, 
thyrsiform, the branches pilose or puberulent; calyx very small, 
acutely dentate; corolla 10-15 mm. long, puberulent or short-pilose, 



FLORA OF COSTA RICA 1333 

purple or purplish; fruit 4-5 mm. long, purple-black. P. angustifolia 
is a widely ranging species that exhibits considerable variation, and 
there is no apparent reason why the two names based on Costa 
Rican material should not be referred to its synonymy. The species 
is well marked by its long and exceptionally narrow leaves. 

Palicourea Austin-Smithii Standl., sp. nov. Arbuscula 4- 
metralis, trunco 9 cm. diam., praeter inflorescentiam omnino gla- 
bra, ramis gracilibus, vetustioribus ochraceis vel brunneis, novellis 
viridibus, internodiis breviusculis; stipulae 3-4 mm. longae virides 
ad medium vel profundius lobatae, lobis approximatis erectis tri- 
angularibus acutis vel acuminatis; folia inter minora breviter 
petiolata firme membranacea, petiolo 8-12 mm. longo; lamina 
lineari-lanceolata vel anguste oblongo-lanceolata 7.5-12 cm. longa, 
1.3-2.3 cm. lata longissime anguste attenuata, basin angustatam 
versus sensim attenuata, supra viridis vix lucida, costa nervisque 
prominentibus, subtus fere concolor, costa gracili elevata, nervis 
lateralibus utroque latere ca. 17 arcuatis angulo fere recto diver- 
gentibus tenerrimis; inflorescentia terminalis sessilis late cymoso- 
paniculata, ca. 7 cm. longa atque 7 cm. lata, ramis oppositis late 
divergentibus basi bracteatis sparsissime minute pilosulis, bracteis 
linearibus viridibus usque 5 mm. longis, floribus in cymulas pau- 
cifloras laxas dispositis, pedicellis crassiusculis usque 7 mm. longis 
ut videtur viridibus; hypanthium basi articulatum latum ca. 1 mm. 
longum; calycis lobi virides erecti triangulari-ovati obtusi vel 
acutiusculi 2 mm. longi; corolla alba extus glabra usque 15 mm. 
longa, tubo crasso ima basi paullo uno latere dilatato fauce ca. 4 
mm. lato, lobis triangulari-ovatis obtusis ca. 5 mm. longis; antherae 
subexsertae; fructus didymo-globosus 5-6 mm. altus. Palmira, 
Prov. Alajuela, 2,300 meters, August, 1938, Austin Smith H1034- 
(type in Herb. Field Mus.). "Bark pale brown, shallowly striated; 
leaves highly polished and shining on the upper surface." Material 
of this species was received too recently to permit its inclusion in 
the key to species. It is well marked by the long and narrow leaves, 
and the relatively large, white corolla. 

Palicourea Brenesii Standl., sp. nov. Frutex metralis sparse 
ramosus, ramulis crassis obtuse tetragonis glabris, internodiis elon- 
gatis; vagina stipularis 5 mm. longa et aequilata truncata glabra, 
lobis remotis erectis subulato-linearibus aequilongis vel paullo longi- 
oribus; folia magna petiolata subcoriacea, petiolo crasso glabro 1.5-2 
cm. longo; lamina obovato-elliptica vel oblongo-elliptica 9.5-17 cm. 



1334 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

longa 4-8 cm. lata abrupte acuta vel breviter acuminata, basi acuta 
vel acuminata, glabra, costa nervisque supra prominentibus, costa 
subtus elevata gracili, nervis lateralibus utroque latere ca. 20 promi- 
nentibus gracillimis valde arcuatis angulo fere recto abeuntibus, 
venulis prominulis laxiuscule reticulatis; inflorescentia thyrsoideo- 
paniculata erecta dense multiflora 8-10 cm. longa basi ca. 6 cm. lata 
crasse 3-5 cm. longe pedunculata, ramis brevibus paten tibus crassis 
densiuscule breviter patenti-pilosis, bracteis inconspicuis lineari- 
lanceolatis usque 6 mm. longis, floribus plerumque breviter peticel- 
latis; hypanthium basi cum pedicello articulatum vix ultra 1 mm. 
longum crassum puberulum, calyce ca. 1.5 mm. longo profunde 
lobato, lobis latis obtusis; corolla caerulea extus praesertim ad lobos 
dense breviter subadpresso-pilosa, tubo crasso 8 mm. longo, lobis 
subpatentibus oblongis 3 mm. longis; stylus breviter exsertus. 
Palmira del Naranjo, in forest, 1,850 meters, April, l92l,Brenes 3512 
(type in Herb. Field Mus.). Palmira, on high, clay ridge, 1,800 
meters, September, 1937, Austin Smith A331 . 

Palicourea costaricensis Benth. ex Oerst. Vid. Medd. Kjoeben- 
havn 38. 1852. Volcan de Barba and Cerros de Candelaria, Oersted 
(photo, of type, Oersted 11540, in Herb. Field Mus.). Frequent 
in mountain forests of the central region, chiefly at 1,500-2,500 
meters; region of San Ramon. Panama. A stout shrub of 2-3 
meters, the branches usually villous, at least at first; stipule sheath 
short, the lobes linear-subulate, equaling or often longer than the 
tube, 3-5 mm. long; leaves short-petiolate, rigid, mostly elliptic- 
oblong or obovate-oblong and 3-4 cm. wide, acuminate, acute at 
the base, beneath pilose, at least along the nerves; inflorescence 
narrowly or broadly paniculate, many-flowered, villosulous or pilose, 
pale yellow; hypanthium pilosulous or glabrous; calyx lobes usually 
puberulent, scarcely more than 1 mm. long; corolla yellow, 12 mm. 
long or less, hirtellous or pilosulous, chiefly on the lobes, or often 
almost glabrous. The species is a rather vaguely marked one, and 
should perhaps be included with P. Galeottiana. 

Palicourea discolor Krause, Bot. Jahrb. 54: Beibl. 119: 40. 
1916. Cafias Gordas, 1,100 meters, Pittier 11161 (photo, of type 
in Herb. Field Mus.). Known only from the original material. 
Branchlets glabrous; stipule sheath 5-6 mm. long, the lobes linear, 
of equal length; leaves large, long-petiolate, glabrous, oblong-elliptic 
to oblong-obovate, 18-25 cm. long, 9-11 cm. wide, short-acuminate, 
cuneate-acute at the base, the lateral nerves 18-22 on each side; 



FLORA OF COSTA RICA 1335 

panicles long-pedunculate, about 18 cm. long and 7 cm. broad, rather 
lax and many-flowered, the flowers short-pedicellate; calyx lobes 1.5 
mm. long or less, ovate, acute; corolla tube 8-9 mm. long. 

Palicourea fastigiata Benth. Thickets or forest of the Atlantic 
tierra caliente. Extending to Amazonian Brazil. A slender shrub 
about 2 meters high, almost glabrous; stipules 5 mm. long or less, 
the broad lobes obtuse or acute; leaves short-petiolate, oblong- 
elliptic to oblong-ovate, small or medium-sized, thin, blackening 
when dried, acuminate, usually acute at the base, glabrous; inflores- 
cence long-pedunculate, small, very dense, fastigiately branched, 
flat- topped, the branches suberect, usually puberulent, the flowers 
slender-pedicellate; calyx lobes minute, ovate or rounded; corolla 
yellow, glabrous, 8-10 mm. long; fruit 4-5 mm. long, purple-black. 

Palicourea Galeottiana Martens. Frequent in mountain 
forests of the central region; regions of Dota and San Ramon; 1,000- 
2,000 meters. Extending to southern Mexico. A shrub or small 
tree of 2^4.5 meters, the young branches usually glabrous; stipules 
short, the lobes short or elongate, narrowly triangular to subulate, 
often much shorter than the tube; leaves short-petiolate, oblong or 
elliptic-oblong, 7-20 cm. long, acuminate, acute at the base, usually 
almost glabrous but more or less pilose at least along the costa; 
panicles usually almost as broad as long, the branches yellow or pale 
red; calyx lobes scarcely 1 mm. long; corolla pale yellow, commonly 
1-1.5 cm. long, glabrous; fruit purple-black, about 5 mm. long. 

Palicourea guianensis Aubl. Forests and thickets of the tierra 
caliente; Cerro de Aguacate. Widely distributed in tropical America. 
A shrub or small tree, rarely more than 4 meters high, almost 
glabrous; stipules 8-10 mm. long; leaves large, short-petiolate, 
elliptic-oblong to ovate or elliptic, mostly 15-25 cm. long and 7-11 
cm. wide, acuminate, acute to almost rounded at the base, glabrous 
or nearly so; panicles large, dense, many-flowered, thyrsoid or 
pyramidal, the numerous stout branches ascending or spreading, 
glabrous or puberulent, the flowers pedicellate; calyx lobes minute, 
deltoid, acute or obtuse; corolla yellow or orange-red, 10-18 mm. long, 
usually densely furfuraceous-tomentellous; fruit ovoid, 4-5 mm. long. 

Palicourea lasiorrhachis Oerst. Vid. Medd. Kjoebenhavn 39. 
1852. Zorrillo amarillo. Cerros de Candelaria, Oersted 11557 (photo, 
of type in Herb. Field Mus.). Frequent in mountain forests of the 
central region, 1,000-2,500 meters; region of San Ramon. Endemic. 



1336 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

A shrub of 2-3.5 meters, the young branches pilose or villosulous; 
stipular sheath truncate, the linear-subulate, erect lobes usually 
much longer; leaves slender-petiolate, elliptic-oblong or obovate- 
oblong, mostly 10-15 cm. long, acuminate, acute at the base, con- 
spicuously nerved, glabrous above or nearly so, beneath rather 
densely pilose or villosulous over almost the whole surface or some- 
times glabrate; panicles thyrsiform or narrowly pyramidal, many- 
flowered, pedunculate, villosulous, yellowish; calyx lobes less than 
1 mm. long; corolla yellow, about 1 cm. long, slender, villosulous 
or puberulent outside. 

Palicourea macrocalyx Standl. Journ. Wash. Acad. Sci. 18: 

278. 1928. Wet forest, Cerro de Las Lajas, north of San Isidro, 
Prov. Heredia, 2,200 meters, Standley & Valeria 51611. Also at 
La Palma de San Ramon, collected by Brenes. Endemic. A shrub 
of 2 meters, glabrous throughout; stipules 6-9 mm. long, the lobes 
triangular-oblong, longer than the sheath; leaves slender-petiolate, 
oblong-elliptic, 8-12 cm. long, 3.5-5 cm. wide, abruptly short-acu- 
minate, obtuse or acute at the base; panicle much branched, dense, 
pedunculate, about 6 cm. long and broad, the pedicels 4-6 mm. 
long; calyx pale yellow, 5 mm. long, the lobes ovate or lanceolate, 
conspicuously 3-nerved; corolla pale yellow, the tube 9 mm. long, 
the lobes 2-5 mm. long. 

Palicourea macrosepala Krause, Bot. Jahrb. 54: Beibl. 119: 
41. 1916. In forest, Alto del Pito, 1,000 meters, Tonduz 7704 (photo, 
of type in Herb. Field Mus.). Region of San Ramon, at 1,300 meters 
or less. Endemic. A shrub of 1.5-3 meters, the branches glabrous 
or nearly so; stipule sheath short, the lobes linear, erect, longer than 
the sheath; leaves slender-petiolate, elliptic-oblong or lance-oblong, 
10-25 cm. long, narrow-acuminate, acute or acuminate at the base, 
sparsely pilose beneath along the nerves; panicles many-flowered, 
pedunculate, 25 cm. long or less, usually thyrsiform, the branches 
glabrous or nearly so; calyx lobes ovate, subacute, 2.5-3 mm. long, 
usually recurved in age; corolla white or purplish, the tube 5-7 
mm. long. 

Palicourea montivaga Standl. Journ. Wash. Acad. Sci. 18: 

279. 1928. Wet forest, Yerba Buena, northeast of San Isidro, Prov. 
Heredia, 2,000 meters, Standley & Valeria 49850. Occasional in 
mountain forests of the central region, 1,800-2,900 meters. Endemic. 
A dense, slender shrub of 1.5-2.5 meters, the branches glabrous; 
stipule lobes less than 1 mm. long, the sheath longer; leaves small, 



FLORA OF COSTA RICA 1337 

slender-petiolate, lance-oblong, mostly 6-7 cm. long, long-acuminate, 
acute at the base, glabrous; inflorescence about as broad as long, 
rather lax, many-flowered, the pedicels 2.5 mm. long or less, the 
branches minutely puberulent or glabrous; calyx 0.6 mm. long; 
corolla yellow or greenish yellow, glabrous, the tube 7 mm. long. 

Uragoga pulchra var. irazuensis Kuntze (Rev. Gen. 1: 300. 
1891), described from Costa Rica, probably is a species of Palicourea, 
but its position is altogether doubtful at present. 

Palicourea salicifolia Standl. Journ. Wash. Acad. Sci. 18: 
280. 1928. Wet forest, Laguna de La Chonta, northeast of Santa 
Maria de Dota, Prov. San Jose", 2,000 meters, Standley 42174. Type 
region, at 2,000-2,400 meters; perhaps also in the region of San 
Ramon. Endemic. A slender shrub of 1.5-3.5 meters, glabrous 
throughout; stipule lobes linear, 1.5-2.5 mm. long; leaves short- 
petiolate, narrowly oblong to elliptic-oblong, 6-10 cm. long, long- 
acuminate, acute to attenuate at the base; inflorescence 4 cm. long 
and often much broader, many-flowered, pedunculate, the bracts 
rather conspicuous, green, 7 mm. long or less, usually obtuse, the 
pedicels 1 cm. long or less; calyx 1.5-2 mm. long; fruit 6 mm. broad. 
The plant may be better referable to Psychotria, the corolla being 
unknown. 

Palicourea subrubra Polak. Linnaea 41: 571. 1877. Cerro de 
La Carpintera, Polakowsky 200A (photo, of type in Herb. Field Mus.). 
Frequent in mountain forests of the central region; regions of Dota 
and San Ramon; 1,000-2,200 meters. Panama. A shrub or small 
tree, 4 meters high or less, glabrous; stipule sheath short, the lobes 
linear or subulate, equaling or longer than the sheath; leaves short- 
petiolate, oblong or elliptic-oblong, mostly 10-15 cm. long, narrow- 
acuminate, acute or subacute at the base; panicles rather large, 
usually thyrsoid-pyramidal, dense and many-flowered, the branches 
orange-yellow; calyx lobes minute; corolla yellow, 8 mm. long or less. 

Palicourea triphylla DC. Thickets of the Pacific tierra caliente, 
ascending to Villa Colon, at 1,300 meters. British Honduras to Peru. 
A shrub about a meter high, or sometimes larger, the branches terete, 
glabrous or puberulent; stipule lobes linear-attenuate, 6 mm. long; 
leaves ternate, subsessile, oblong or lance-oblong, large, long-acumi- 
nate, puberulent, especially beneath, or finally glabrate; inflorescence 
long-pedunculate, narrowly thyrsoid-paniculate, dense and many- 
flowered; calyx lobes obtuse; corolla yellow, 12-15 mm. long, fur- 
furaceous-puberulent; fruit black. 



1338 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Palicourea vestita Standl. Journ. Wash. Acad. Sci. 18: 277. 
1928. Wet oak forest near Quebradillas, about 7 km. north of Santa 
Maria de Dota, Prov. San Jose", 1,800 meters, Standley 42909. Col- 
lected also in the region of Navarro, 1,500 meters. Endemic. A 
shrub of 1.5-2.5 meters, the branches densely villous; stipular sheath 
5 mm. long, the lobes linear, 5-7 mm. long; leaves petiolate, lance- 
oblong or obovate-oblong, 10-18 cm. long, 2.5-6 cm. wide, acuminate 
or long-acuminate, narrowed to the acute or obtuse base, short- 
villous above along the costa, densely villous beneath on the nerves, 
the nerves conspicuously elevated; panicles narrowly pyramidal or 
thyrsiform, 7-9 cm. long or larger, many-flowered, the branches 
densely villous; calyx lobes 1 mm. long; corolla yellow, 1 cm. long, 
pilose; fruit blue, 5 mm. long, pilose. 

PENTAGONIA Benth. 

Shrubs or small trees, simple or branched, the branchlets terete 
or obtusely tetragonous; leaves very large, opposite, sessile or petio- 
late, usually coriaceous, entire or pinnatifid, finely lineolate-striate 
between the veins; stipules large, elongate; inflorescence axillary, 
sessile or pedunculate, cymose-corymbose, with few or many flowers, 
the large flowers red or yellow; hypanthium turbinate or campanu- 
late; calyx spathaceous or 5-6-lobate, persistent, coriaceous, the 
lobes equal or unequal, obtuse; corolla coriaceous, funnelform or 
tubular, the throat glabrous, the 5-6 lobes short, valvate; anthers 
included; ovary 2-celled, many-ovulate; fruit baccate, subglobose; 
seeds numerous, angulate. Several other species are native in 
Panama. The genus is one of the most interesting or outstanding of 
the family, noteworthy for the very large leaves, which are deeply 
pinnatifid in several species. They are notable also for the curious 
striation of the leaf tissue, which has been compared with that of 
moire or watered silk. The leaves, bracts, and stipules often are 
handsomely colored with red. 

Pentagonia Alfaroana Standl. Journ. Wash. Acad. Sci. 17: 171. 
1927. Watsonamra Alfaroana Standl. op. cit. 15: 287. 1925. Moist 
forest along the Rio Grande de Tarcoles, near Capulin, Pacific coast, 
80 meters, Standley 40194- Known only from the type region. A 
simple shrub 1-1.5 meters high; stipules 5 cm. long; leaves petiolate, 
the blades 40-50 cm. long, about 40 cm. wide, sparsely sericeous, 
especially when young, pinnatifid two-thirds the distance to the 
costa, the segments again pinnatifid; cymes few-flowered, sessile or 



FLORA OF COSTA RICA 1339 

nearly so; fruit globose, 12-15 mm. in diameter; calyx in fruit 2 cm. 
long, the lobes oblong. 

Pentagonia Donnell-Smithii Standl. Journ. Wash. Acad. Sci. 
17: 170. 1927. Watsonamra Donnell-Smithii Standl. Contr. U. S. 
Nat. Herb. 17: 442. 1914. Near La Emilia, Llanuras de Santa Clara, 
250 meters, J. D. Smith 6590. Frequent in forests of the Atlantic 
tierra caliente; Atlantic slope of the mountains of Guanacaste. 
Ranging to Honduras. A shrub or tree, sometimes 7.5 meters high, 
simple or branched; stipules ovate, 5 cm. long; petioles 9-11 cm. long; 
blades oval or obovate-elliptic, about 45 cm. long and 30 cm. wide, 
or often larger or smaller, entire, obtuse or acute, obtuse or acute at 
the base, almost glabrous; calyx 12-15 mm. long, the lobes oval or 
oblong; corolla yellow, sericeous outside, the tube 2.5 cm. long, the 
lobes 6-7 mm. long; fruit globose, 2.5 cm. in diameter. In all proba- 
bility, this should be reduced to synonymy under P. macrophylla 
Benth., a species of Panama. 

Pentagonia hirsuta Standl. Journ. Wash. Acad. Sci. 18: 165. 
1928. Forests above Tsaki, Talamanca, 500 meters, Tonduz 9415. 
Known only from the original collection. Young branches hirsute; 
leaves sessile, broadly obovate, about 60 cm. long and 27 cm. wide, 
narrowed to the short-acuminate apex, attenuate to a narrow, cor- 
date-clasping base, entire, hirsute on both surfaces; hypanthium 
densely hirsute; calyx 2.5 cm. long, hirsute. 

Pentagonia Wendlandi Hook. Bot. Mag. pi. 5230. 1861. 
Watsonamra Wendlandi Kuntze, Rev. Gen. 302. 1891. Described 
from Central American plants of uncertain origin, probably from 
Costa Rica or Panama. Occurring in the Changuinola Valley, 
on the border between the two countries; collected also at La Colom- 
biana. A shrub or small tree, simple or branched, the branchlets 
glabrate; leaves subsessile, oblong-obovate, very large, usually more 
than 60 cm. long, acutish, narrowed to the narrowly rounded or 
obtuse base, almost glabrous; calyx sericeous outside, the lobes oval 
or suborbicular, about as long as the tube; corolla yellow, the tube 
2.5 cm. long, the lobes obtuse, erect. 

POGONOPUS Klotzsch 

Trees or shrubs; leaves opposite, petiolate, the blades large, 
membranaceous; stipules small, cuspidate, tardily deciduous; flowers 
showy, pedicellate, in small cymes, these disposed in lax, terminal 
panicles; hypanthium turbinate; calyx short, 5-dentate, deciduous, 



1340 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

one of the lobes expanded into a large, foliaceous, bright-colored 
limb; corolla tubular, villous in the throat, the short limb 5-lobate, 
the lobes valvate; anthers exserted; ovary 2-celled, many-ovulate; 
capsule loculicidally bi valvate; seeds horizontal. A single species 
is found in Central America. 

Pogonopus speciosus (Jacq.) Schum. Macrocnemum exsertum 
Oerst. Vid. Medd. Kjoebenhavn 45. 1852 (Jaris, 900 meters, Oersted). 
P. exsertus Oerst. AmeY. Centr. 17. 1863. Guanacaste. Southern 
Mexico to Venezuela. A shrub or tree, 10 meters high or less; 
stipules 3-5 mm. long; leaves obovate to oval-obovate, mostly 10- 
20 cm. long, abruptly acuminate, acute to long-attenuate at the base, 
sparsely or densely puberulent beneath; pedicels sometimes 1 cm. 
long; calyx 1 mm. high, the dilated lobe petiolate, rounded-ovate, 
1-5 cm. long, rounded to acute at the apex, purple-red; capsule 5-7 
mm. long. A showy and handsome tree because of the brightly 
colored, enlarged calyx lobes. 

POSOQUERIA Aubl. 

Trees or shrubs, glabrous or pubescent, the branchlets terete; 
leaves opposite, petiolate, coriaceous; stipules large, deciduous; 
flowers large, white, in terminal corymbs; calyx short, 5-dentate, 
persistent; corolla tube slender and much elongate, the limb gibbous 
in bud, 5-lobate, the lobes obtuse, contorted; anthers exserted; 
ovary 1-2-celled, many-ovulate; fruit baccate, large, globose or ovoid, 
1-2-celled; seeds large, obtusely angulate. No other species are 
known from Central America. 

Posoqueria grandiflora Standl. Journ. Wash. Acad. Sci. 18: 
166. 1928. Wet thicket at Hamburg Finca, Rio Reventazon below 
El Cairo, Prov. Limon, 55 meters, Standley & Valeria 48706. Region 
of the type. Also in Panama. A shrub of 3-5 meters, the branchlets 
puberulent or scaberulous; stipules obtuse, almost 2 cm. long; leaves 
short-petiolate, rounded-ovate to broadly elliptic or oblong-elliptic, 
13-36 cm. long, 8-21 cm. wide, rounded or obtuse at the apex or 
abruptly short-acuminate, broadly rounded to acute at the base, 
glabrous above, densely and minutely pubescent beneath; calyx 
4 mm. long, shallowly lobate; corolla glabrous outside, the tube 19-22 
cm. long, 3-5 mm. thick, the oblong lobes rounded at the apex, 3.5- 
4 cm. long; fruit globose, 7 cm. in diameter. Called Wild Coffee by 
the West Indians of the banana plantations. 

Posoqueria latifolia (Rudge) Roem. & Schult. Fruta de mono, 
Guayabo de mico. Frequent in thickets of the Pacific tierra caliente; 



FLORA OF COSTA RICA 1341 

Guanacaste. Southern Mexico to Bolivia and Brazil. A shrub or 
tree of 3-6 meters, glabrous throughout; stipules 8-15 mm. long; 
leaf blades oval to oblong, 10-25 cm. long, 4-13 cm. wide, acute 
or short-acuminate, obtuse or rounded at the base; corolla tube 
12.5-16.5 cm. long, 2-4 mm. thick, the lobes 1.5-2.5 cm. long; 
fruit globose, yellow, 4-5 cm. in diameter. The flowers are fragrant. 
When in blossom, the tree is a handsome and conspicuous one. 

PSYCHOTRIA L. 

Shrubs or small trees, rarely low and merely suffrutescent plants, 
sometimes epiphytic; stipules persistent, then usually bilobate, or 
caducous and then commonly entire; leaves opposite; inflores- 
cence generally terminal, sometimes axillary, variable in form, very 
rarely headlike and then not or scarcely involucrate, never with a 
showy involucre of partially united bracts, the flowers mostly small 
and inconspicuous and white or yellowish; calyx lobes more or less 
united, short or elongate, sometimes obsolete; corolla short or elon- 
gate, the tube straight, often barbate in the throat, the lobes valvate; 
stamens included or exserted; ovary 2-celled, or very rarely 5-celled; 
fruit baccate, containing normally 2 nutlets. A rather large number 
of other species is found in Central America. The genus is the 
largest and most complicated of the American Rubiaceae, and is 
represented abundantly also in tropical regions of the other con- 
tinents. There are no really definite lines of separation between 
Psychotria, Palicourea, and Cephaelis, but all these genera are large 
ones, and in so complicated a group of plants it is convenient to 
recognize them. 

Inflorescence axillary; stems mostly simple and low, commonly 

herbaceous or only suffrutescent. 
Inflorescences capitate. 

Leaves very obtuse or rounded at the apex, hirsute beneath. 

P. polyphlebia. 

Leaves acute or acuminate, not at all hirsute P. emetica. 

Inflorescences cymose-paniculate, open. 

Leaves densely villous or villosulous beneath P. Siggersiana. 
Leaves glabrous or nearly so. 

Inflorescence sessile or subsessile P. Tonduzii. 

Inflorescences long-pedunculate. 

Leaves carnose, pale beneath; fruit red P. uliginosa. 

Leaves membranaceous, green beneath; fruit white. 

P. macrophylla. 



1342 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Inflorescences normally terminal, a few sometimes axillary; shrubs 

or small trees, usually much branched. 

A. Stipules caducous, entire or nearly so, brown or ferruginous; 
plants generally blackening when dried. Subgenus Mapouria. 

Leaf blades cordate at the base P. Wendlandiana. 

Leaf blades acute to rounded at the base. 

Young branches densely pilose or villosulous with long or 
short hairs. 

Leaves small, 3.5 cm. long or less P. parvifolia. 

Leaves large, 7-12 cm. long or more. 

Inflorescence sessile or nearly so; leaves densely puberu- 

lent beneath P. psychotriaefolia. 

Inflorescence long-pedunculate; leaves almost glabrous. 

P. Jimenezii. 
Young branches glabrous or nearly so, sometimes minutely 

puberulent. 

Flowers glomerate, the glomerules spicate, the spikes panic- 
ulate; leaf blades long-attenuate to the base . . P. viridis. 
Flowers glomerulate or scattered, but not in spicate glom- 
erules. 
Flowers in small, solitary, sessile heads subtended by 

large, brown, membranous bracts P. chagrensis. 

Flowers not capitate. 
Calyx deeply lobate, the lobes lance-linear. 

P. horizontalis. 
Calyx shallowly dentate, the teeth broad. 

Flowers all or chiefly on short or elongate pedicels. 
Leaves essentially sessile, the petioles broadly 

marginate to the base P. orosiana. 

Leaves conspicuously petiolate. 
Panicles long-pedunculate. 

Branches of the panicle puberulent; flowers 

short-pedicellate P. sylvivaga. 

Branches of the panicle glabrous; flowers long- 
pedicellate P. marginata. 

Panicles sessile or nearly so. 
Stipules mostly 3-4.5 cm. long. 

P. grandistipula. 
Stipules mostly less than 1 cm. long. 



FLORA OF COSTA RICA 1343 

Leaf blades 3-4 cm. wide. P. sarapiquensis. 
Leaf blades mostly 8-10 cm. wide. 

P. limonensis. 
Flowers all sessile or nearly so. 

Leaf blades all or mostly less than 3.5 cm. long. 

P. graciliflora. 
Leaf blades much larger. 

Base of the leaf blade sharply differentiated 
from the petiole, narrowed to an obtuse or 
subacute base, the petiole scarcely 5 mm. 

long P. quinqueradiata. 

Base of the leaf blade gradually narrowed into 
the elongate petiole. 

Panicles pedunculate P. carthaginensis. 

Panicles sessile or nearly so. 
Leaves 8-12 cm. wide; panicles large and 

open P. magna. 

Leaves less than 7 cm. wide; panicles small 

and dense P. granadensis. 

AA. Stipules persistent, usually green or greenish, variable in 
form, most often deeply bilobate or connate into a truncate 
sheath which is produced into short or elongate lobes. 
Fruit 5-celled ; leaves elliptic or oblong-elliptic, short-acuminate, 
decurrent at the base, glabrous or nearly so ... P. racemosa. 
Fruit 2-celled. 

Plants epiphytic; leaves thick and fleshy, glabrous; stipules 

very short, with obtuse lobes, these usually caducous. 

Leaves linear-lanceolate, 5 mm. wide or less. ... P. Maxonii. 

Leaves much broader and larger. 

Leaf blades 2-3.5 cm. wide, the nerves prominent 

beneath P. pithecobia. 

Leaf blades 6-15 mm. wide, the nerves obsolete. 

P. orchidearum. 

Plants terrestrial; leaves not fleshy; stipule lobes usually 
persistent and most often acute to subulate. 

Inflorescence composed of one or more dense heads sub- 
tended by large, often brightly colored bracts, the 
heads large or small; bracts of the small heads at least 
longer than the calyx. 



1344 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Flower heads few or numerous, more than one. 

Inflorescence cymose, the heads 1.5 cm. or more in 

diameter P. chiapensis.. 

Inflorescence pyramidal-paniculate, the heads less than 

1 cm. in diameter P. brachiata. 

Flower heads solitary at the ends of the branches. 

Peduncles glabrous P. ramonensis. 

Peduncles pilose or puberulent. 
Larger bracts 2-4, very unequal, long-attenuate. 

P. involucrata. 

Larger bracts numerous, subequal, obtuse or sub- 
acute P. suerrensis. 

Inflorescence usually open, never with large or colored or 
green, conspicuous bracts, never of heads subtended by 
broad or elongate bracts. 
Branches densely pilose or hirsute with long or short 

hairs; calyx dentate. 

Leaves abundantly hirsute on the upper surface. 
Inflorescence lax and openly branched . P. Brenesii. 

Inflorescence dense and small, the branches very 
short P. costaricensis. 

Leaves not hirsute on the upper surface. 

Corolla 12 mm. long or more; stipules entire. 

P. hondensis. 
Corolla 7 mm. long or less; stipules bilobate. 

Corolla glabrous; leaves mostly 8-10 cm. wide. 

P. Mortoniana. 
Corolla puberulent or pilose; leaves narrower. 

Branches conspicuously constricted at the nodes 
when dry; inflorescence dense and compact, 
the branches all very short. . .P. hebeclada. 

Branches not contracted at the nodes when 
dried; inflorescence open, the lower branches 
elongate. 

Pubescence of the lower leaf surface of long, 
closely appressed hairs P. Pittieri. 

Pubescence of the lower leaf surface of minute, 
spreading hairs P. dispersa. 



FLORA OF COSTA RICA 1345 

Branches glabrous or puberulent; calyx sometimes 
truncate. 

Stipules large, entire, often 2 cm. long P. grandis. 

Stipules less (usually much less) than 1 cm. long. 
Calyx truncate. 
Stipule lobes caducous, membranous; calyx 1 mm. 

long or more P. microdon. 

Stipule lobes persistent, stiff, subulate; calyx 

scarcely 0.5 mm. long. 
Branches of the panicle erect or strongly 

ascending P. Valeriana. 

Branches of the panicle spreading or somewhat 

reflexed P. cuspidata. 

Calyx evidently dentate. 

Inflorescence capitate P. haematocarpa. 

Inflorescence not capitate. 
Inflorescence cymose-umbellate, compact, sub- 
globose, 2-4.5 cm. broad, the primary 
branches only 4-7 mm. long . . P. Alfaroana. 

Inflorescence not cymose-umbellate, commonly 
paniculate or corymbiform. 

Lowest branches of the inflorescence not 
bracteate at the base, the inflorescence 
narrow, thyrsoid, the branches often 

reflexed P. patens. 

Lowest branches of the panicle bracteate at 

the base. 

Branches of the inflorescence densely short- 
pilose. 
Inflorescence corymbiform . P. pubescens. 

Inflorescence typically pyramidal-panicu- 
late P. Berteriana. 

Branches of the inflorescence glabrous or 
very minutely puberulent. 

Leaves lance-linear, 9-16 cm. long, 1.5- 
2.5 cm. wide P. Torresiana. 

Leaves relatively broader, oblong-lanceo- 
late to elliptic. 



1346 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Inflorescence open-paniculate; leaves 
mostly 1.5-2.5 cm. wide. 

P. Goldmanii. 

Inflorescence corymbiform; the leaves 
mostly 4-7 cm. wide . P. eurycarpa. 

Psychotria Alfaroana Standl. Journ. Wash. Acad. Sci. 18: 273. 
1928. Wet forest, El Arenal, Guanacaste, 500 meters, Standley & 
Valeria 4-5179. Mountains of Guanacaste, 500-700 meters; Pejivalle, 
at 900 meters. Endemic. An erect shrub of 30-60 cm., usually 
simple but sometimes sparsely branched; stipules 8-18 mm. long, 
oblong-ovate, cleft to below the middle, the lobes linear; petioles 1-2 
cm. long, the blades obovate-oblong to elliptic, 12-25 cm. long, 
4-9 cm. wide, acute or abruptly acute, usually long-attenuate to the 
base, glabrous or nearly so; inflorescence terminal, cymose-umbellate, 
the primary branches several, 4-7 mm. long, sordid-puberulent or 
glabrate, the pedicels 2-4 mm. long, the inflorescence pedunculate, 
subglobose, in fruit 2-4.5 cm. broad; fruit red, 8-10 mm. long. 

Psychotria Berteriana DC. P. angustiflora Krause, Bot. Jahrb. 
54: Beibl. 119: 43. 1916 (Las Vueltas, Tucurrique, 600-700 meters, 
Tonduz 12996; photo, of type in Herb. Field Mus.). Forests of the 
Atlantic tierra caliente; region of San Ramon; at 1,200 meters or less. 
West Indies. A slender shrub of 2-4.5 meters, the branches glabrous 
or minutely puberulent; stipules persistent, broad, shallowly bilobate; 
leaves petiolate, membranaceous, lance-oblong or oblong-ovate, 
15-18 cm. long and 5-6.5 cm. wide, or often smaller, long-acuminate, 
rounded to acute at the base and usually abruptly decurrent, sparsely 
villosulous beneath or often almost glabrous; panicles terminal, 
large and broad, usually long-pedunculate, lax, and open, generally 
pyramidal, as much as 16 cm. long and broad but often much smaller, 
the branches densely short-pilose, the flowers mostly short-pedicellate; 
calyx minute and remotely dentate; corolla greenish yellow, minutely 
pilose or almost glabrous, 10 mm. long or less. P. crebrinervia Standl., 
described from Honduras, to which Costa Rican material has been 
referred, probably is synonymous with P. Berteriana. 

Psychotria brachiata Swartz. Atlantic tierra caliente. Mexico 
to Peru. A shrub of 4 meters or less, the branches glabrous; stipules 
persistent, short, bilobate, the lobes obtuse or rounded; leaves 
petiolate, oblong-obovate to oblong-elliptic, short-acuminate, acute 
or attenuate at the base, glabrous or nearly so, often pilose beneath 
on the costa; inflorescence terminal, long-pedunculate, thyrsoid- 



FLORA OF COSTA RICA 1347 

paniculate or racemose, the opposite branches divaricate, subtended 
at the base by long, narrow bracts, the flowers in small, dense, many- 
bracteate heads; corolla white or ochroleucous; fruit blue. 

Psychotria Brenesii Standl., sp. nov. Frutex 2-3-metralis 
ramosus, ramis obtuse tetragonis crassis densissime pilis perlongis 
paten tibus sordidis pilosis, internodiis brevibus; stipulae persistentes 
1 cm. longae et ultra breviter bilobae, lobis acutis vel acuminatis, 
longipilosae vel glabratae; folia majuscula petiolata membranacea, 
petiolo crassiusculo 1.5-3.5 cm. longo dense longipiloso; lamina 
oblongo-obovata 11-19 cm. longa 5-8 cm. lata cuspidato-acuminata, 
basi acuta, supra dense pilis longissimis laxis pilosa, subtus paullo 
pallidior ubique longipilosa, costa gracili elevata, nervis lateralibus 
utroque latere ca. 18 teneris prominentibus arcuatis obliquis, venis 
laxe reticulatis; inflorescentia terminals sessilis atque basi tricho- 
toma laxa ca. 10 cm. longa atque aequilata, ramis primariis usque 
5 cm. longis dense pilosis, floribus cymosis sessilibus vel breviter 
pedicellatis, in cymas paucifloras laxe cymose dispositis, ramis basi 
bracteatis, bracteis angustis usque 7 mm. longis pilosis; calyx cum 
hypanthio usque 5 mm. longus ubique dense pilis longis laxis patenti- 
bus indutus, profunde lobatus; corolla lutea ca. 12 mm. longa, 
densissime pilis longis patentibus pilosa, lobis brevibus intus glabris. 
Los Angeles de San Ramon, Cerros de La Balsa, March, 1928, 
Brenes 6050 (type in Herb. Field Mus.). Wet forest, Cerros de San 
Antonio de San Ramon, 1,100-1,200 meters, Brenes 5648. Related 
to P. costaricensis, but conspicuously different in the form and details 
of the inflorescence. 

Psychotria carthaginensis Jacq. Forests and thickets of the 
tierra caliente, especially of the Pacific; region of San Ramon; Guana- 
caste; at 900 meters or less. Widely distributed in tropical America. 
An almost glabrous shrub of 1-3 meters; stipules large and thin, 
obovate, obtuse, brownish, caducous; leaves short-petiolate, lance- 
elliptic to oblong-obovate, usually broadest at or above the middle, 
acute or acuminate, acute to attenuate at the base, puberulent 
beneath along the nerves or glabrate; inflorescence pedunculate, 
terminal, paniculate, open, usually radiate-branched from the base, 
many-flowered, the minute bracts deciduous; flowers mostly sessile 
and glomerate; calyx obsoletely dentate; corolla white, glabrous or 
minutely puberulent, 4 mm. long; fruit at first yellow, turning red, 
4 mm. long. 

Psychotria chagrensis Standl. In forest, region of El Cairo, 
Atlantic coast. Panama to Nicaragua. A dense shrub of 1-2 meters, 



1348 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the slender branches glabrous; stipules 8-10 mm. long, triangular- 
ovate, long-cuspidate, thin, brown, caducous; petioles 4-12 mm. long, 
the blades elliptic-obovate, 5-8 cm. long, 2-3 cm. wide, abruptly 
acute-acuminate, cuneately decurrent to the base, glabrous; flowers 
in few-flowered, terminal heads, the head surrounded by bracts 
similar to the stipules, the flowers sessile; calyx lobes linear; corolla 
white, glabrous outside, villous in the throat, the tube 4 mm. long; 
fruit 5 mm. long. 

Psychotria chiapensis Standl. Cephaelis tetragona Donn. 
Smith, Bot. Gaz. 61 : 376. 1916, non Psychotria tetragona Seem. 1865- 
67 (Tuis, Tonduz 11352). Thickets and forest of the tierra caliente. 
Panama to southern Mexico. A shrub or sometimes a small tree 
with a dense, rounded crown, the branches glabrous or obscurely 
villosulous; stipules persistent, small, bilobate, the lobes broad, 
obtuse; leaves short-petiolate, elliptic-oblong, mostly 12-20 cm. long 
and 5-10 cm. wide, short-acuminate, acute or acuminate at the base, 
glabrous; inflorescence terminal, the flowers subcapitate, the large 
heaols corymbose or cymose, the large, broad bracts rounded at the 
apex; calyx 5-6 mm. long, puberulent, denticulate; corolla white, 
almost glabrous, the slender tube as much as 3 cm. long, the narrow 
lobes 8 mm. long. This plant is extraordinary because of the 
extremely elongate corolla, fantastically large when compared with 
that of other species. It is not placed very satisfactorily in Psy- 
chotria, and might be referred equally well to Cephaelis. Quite 
possibly it should be placed in a distinct genus. Cufodontis has 
reported the plant as scandent, but such a habit in this group would 
be most unusual; other collectors report it as an erect shrub or tree. 

Psychotria costaricensis Polak. Linnaea 41: 571. 1877. In 
forest, Angostura, Polakowsky 384A (photo, of type in Herb. Field 
Mus.). Forests of the Atlantic tierra caliente; region of San Ramon; 
at 1,150 meters or less. Ranging to Guatemala. A slender shrub of 
1.5-3 meters, the stems densely long-pilose; stipules persistent, 
about 1 cm. long, deeply bilobate; leaves slender-petiolate, mem- 
branaceous, oblong-elliptic, sometimes 30 cm. long and 11.5 cm. wide 
but usually considerably smaller, abruptly long-acuminate, acute at 
the base, densely long-pilose or hirsute on both surfaces; inflorescence 
terminal, pedunculate, usually thyrsoid and very dense but some- 
times rather open, the flowers crowded into headlike glomerules, the 
bracts ovate, green, like all parts of the inflorescence densely long- 
pilose; fruit blue. A rather isolated species, noteworthy for the 



FLORA OF COSTA RICA 1349 

abundant, long pubescence on all parts. It is questionable whether 
the Central American plant is different from the Peruvian P. pilosa 
Ruiz & Pavon, with which Polakowsky originally compared it. 

Psychotria cuspidata Bredem. Forests of the Atlantic tierra 
caliente, ascending to the region of Navarro; region of San Ramon; 
Guanacaste; at 1,400 meters or less. Widely distributed in tropical 
America. A slender, glabrous shrub of 1.5-3 meters; stipules green, 
persistent, short, bicuspidate; leaves thin, short-petiolate, bright 
green when dried, the nerves often reddish, ovate to elliptic or oblong- 
elliptic, very narrowly long-acuminate, the acumination usually 
somewhat curved, acute to obtuse at the base; inflorescence terminal, 
pedunculate, thyrsoid-paniculate, much shorter than the leaves, 
rather few-flowered, the few pale branches often reflexed, not brac- 
teate at the base; calyx minute, truncate; corolla glabrous, 5 mm. 
long, white or pale yellow; stamens exserted; fruit didymous-globose, 
smooth, black at maturity. 

Psychotria dispersa Standl. Journ. Wash. Acad. Sci. 18: 184. 
1928. Wet forest, Los Ayotes, near Tilaran, Guanacaste, 600 meters, 
Standley & Valeria 45548. Common in forests of the tierra caliente; 
region of San Ramon; Guanacaste; at 1,000 meters or less. Panama 
to Guatemala. A slender shrub of 1-2 meters, the branches short- 
pilose with spreading hairs; stipule sheath 2.5-3 mm. long, the lobes 
linear, 3-6 mm. long; leaves short-petiolate, oblong-elliptic to lance- 
oblong, 6-10 cm. long, 2^4 cm. wide, abruptly acuminate, acute and 
often decurrent at the base, beneath pilose on the nerves with short, 
spreading hairs; panicles terminal, small, 2-4 cm. long and equally 
broad, the branches hirtellous, the flowers sessile or subsessile; calyx 
less than 1 mm. long, the teeth triangular; corolla white, puberulent, 
3-3.5 mm. long; fruit blue, 3 mm. long. The panicles are usually 
recurved or reflexed. 

Psychotria emetica L. f. Raicilla. Cephaelis emetica Pers.; 
Uragoga emetica Baill. Occasional in deep forests of the Atlantic 
tierra caliente. Guatemala to Bolivia. A slender, usually simple 
shrub, 60 cm. high or less, with rather thick, somewhat nodose roots, 
the stems puberulent; stipules 3-4 mm. long, triangular-lanceolate, 
acuminate; leaves short-petiolate, dark when dried, elliptic-oblong 
to oblanceolate-oblong, about 11 cm. long and 3.5 cm. wide, acute or 
acuminate, acute to long-attenuate at the base, glabrous above, 
puberulent or glabrate beneath; flowers capitate, the heads small 
and few-flowered, axillary, pedunculate, about equaling the petioles, 



1350 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

or shorter, solitary or geminate, the bracts minute; calyx obscurely 
dentate; corolla small, white, 5 mm. long, glabrous; fruit 6 mm. 
long, bright blue. The roots of this species yield a kind of ipecac, 
and are sometimes exported from tropical America for medicinal use. 

Psychotria eurycarpa Standl. Journ. Wash. Acad. Sci. 18: 275. 
1928. Cafecillo. Moist forest, Quebrada Serena, southeast of 
Tilaran, Guanacaste, 700 meters, Standley & Valerio 1+6237. Moun- 
tains of Guanacaste; Atlantic tierra caliente; region of San Ramon; 
at 1,100 meters or less. Endemic. A shrub or small tree, 2.5-5 
meters high, the branches glabrous; stipules persistent, the sheath 
1-2 mm. long, the lobes subulate, 1-2.5 mm. long; leaves short- 
petiolate, elliptic or broadly elliptic, 8-15 cm. long, 3-9 cm. wide, 
abruptly acuminate, acute to narrowly rounded at the base and often 
short-decurrent, glabrous, the lateral nerves 6-7 pairs; inflorescence 
terminal, cymose-paniculate, usually corymbiform, pedunculate, 
usually broader than long, the branches glabrous or nearly so, the 
flowers sessile; calyx scarcely 1 mm. long, shallowly dentate or 
sub truncate; corolla in bud 6 mm. long, pulverulent; fruit blue-black, 
9-12 mm. long. Noteworthy for the very large fruits. 

Psychotria Goldmanii Standl. In forest, region of San Ramon, 
about 1,200 meters; mountain forests of the central region, at 1,600 
meters or less. Panama. A slender shrub of 2 meters, the branches 
glabrous; stipules persistent, green, small, bilobate, the short lobes 
subulate; leaves on short, slender petioles, lance-oblong, 5-8 cm. 
long, 1-2 cm. wide, narrowly long-attenuate, attenuate at the base, 
glabrous or sometimes puberulent beneath along the costa, paler 
beneath; panicles terminal, slender-pedunculate, 3-7 cm. long and 
of equal breadth, the branches sparsely puberulent, the flowers 
sessile, the bracts ovate, acute; corolla 5-6 mm. long, white, glabrous; 
fruit 4 mm. long, blue. 

Psychotria graciliflora Benth. ex Oerst. Vid. Medd. Kjoeben- 
havn 35. 1852. Naranjo, 1,350 meters, Oersted (photo, of type in 
Herb. Field Mus.). Frequent in mountain forests of the central 
region; region of San Ramon; Dota; Atlantic tierra caliente; chiefly 
at 1,500 meters or less. Panama to Honduras. A rather dense, 
very leafy shrub, usually 1-2 meters high, glabrous almost through- 
out; stipules small, caducous, acuminate, entire or bidentate; leaves 
blackening when dried, elliptic-oblong or lance-oblong, 8 cm. long 
and 3 cm. wide or smaller, acuminate, attenuate to the base; inflores- 
cence terminal, cymose-paniculate, pedunculate, small and few- 



FLORA OF COSTA RICA 1351 

flowered, trichotomous, the flowers mostly sessile, white; calyx less 
than 1 mm. long, subtruncate; corolla 4 mm. long; fruit blue. 

Psychotria granadensis Benth. Thickets of the Atlantic 
tierra caliente; probably also along the Pacific coast. Panama to 
Guatemala. A shrub of 1-2 meters, glabrous almost throughout; 
stipules large, ovate, obtuse, ferruginous, caducous; leaves petiolate, 
oval-oblong, elliptic, or ovate, 11-15 cm. long, acuminate or long- 
acuminate, attenuate to the base or abruptly decurrent; panicles 
terminal, sessile, small, dense, at least in flower, the flowers sessile 
or subsessile; calyx very short, subtruncate; corolla white, 4 mm. 
long or less, densely villous in the throat; fruit red. Perhaps only 
a glabrous form of P. undata Jacq. 

Psychotria grandis Swartz. P. subsessilis Benth. ex Oerst. 
Vid. Medd. Kjoebenhavn 32. 1852 (Turrialba, 900 meters, Oersted). 
Forests of the Atlantic tierra caliente. Guatemala to Colombia and 
West Indies. A large shrub or small tree, seldom more than 4 meters 
high, glabrous almost throughout; stipules broadly ovate, large, 
cuspidate-acuminate, subpersistent; leaves almost sessile, very large, 
cuneate-obovate or oblong-obovate, abruptly short-acuminate, long- 
attenuate to the base, blackening when dried; inflorescence terminal, 
large, long-pedunculate, radiately branched at the base, the branches 
minutely puberulent, the flowers sessile; calyx obscurely denticulate; 
corolla white, 3 mm. long, glabrous outside, barbate in the throat. 
The flowers are said to be fragrant. This species really belongs to 
the subgenus Mapouria (treated as a genus by many authors), 
but the stipules are long-persistent, rather than caducous as in most 
Mapourias. They also are much larger and broader than is usual 
in that group. 

Psychotria grandistipula Standl. Journ. Wash. Acad. Sci. 18: 
276. 1928. Moist forest near Santa Maria de Dota, Prov. San Jose", 
1,600 meters, Standley & Valerio 43268. Regions of Dota and San 
Ramon; Zarcero; Atlantic tierra caliente. Endemic. A shrub of 
3 meters, or sometimes a tree of 7 meters, with a trunk 10 cm. or 
more in diameter, the bark smooth, the branchlets very minutely 
puberulent; stipules 1 at each node, forming a sheath about the young 
leaves, cleft along one side, 3-4.5 cm. long, attenuate to a subulate 
apex, membranous, brown; leaves slender-petiolate, lance-oblong 
to oblong-ovate or elliptic, 6-14 cm. long, 2.5-6 cm. wide, abruptly 
long-acuminate, usually very obtuse to truncate at the base but 
sometimes acute, minutely puberulent beneath on the nerves, other- 



1352 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

wise glabrous; inflorescence cymose-paniculate, usually sessile, 4-13 
cm. long, the branches minutely puberulent, the pedicels 3-12 mm. 
long; calyx less than 1 mm. long, shallowly dentate; corolla white, 
5 mm. long, glabrous, not barbate in the throat; fruit bright red, 7 
mm. long. 

Psychotria haematocarpa Standl. Journ. Wash. Acad. Sci. 18: 
274. 1928. Moist forest, Naranjos Agrios, near Tilaran, Guanacaste, 
600-700 meters, Standley & Valerio 46407. Collected also at Pejivalle, 
Prov. Cartago, 900 meters. Endemic. A shrub of 1-2.5 meters, the 
branches glabrous; stipules persistent, green, the short sheath bicuspi- 
date, the lobes linear-subulate, 3-4 mm. long; leaves on very short 
petioles, elliptic-oblong, 9-14 cm. long, 2.5-5 cm. wide, long-acumi- 
nate, acute or attenuate at the base and decurrent, glabrous; inflores- 
cence terminal, capitate, dense and few-flowered, the peduncle 3-4 
mm. long, puberulent, the outer bracts lance-linear, 2 mm. long, the 
flowers sessile or subsessile; fruit bright red, 5 mm. long. 

Psychotria hebeclada DC. Forests of the Atlantic tierra 
caliente; Aguacate. Mexico to Ecuador. A shrub, usually 2 meters 
high or less, the branches terete, densely puberulent or short-pilose; 
stipules short, green, persistent, bidentate; leaves membranaceous, 
short-petiolate, ovate to oblong-elliptic, acuminate, acute or attenu- 
ate at the base, finely pubescent, at least beneath; inflorescence 
terminal, thyrsoid-paniculate, small, dense, many-flowered, densely 
short-pilose; calyx lobes short, ovate; corolla whitish, 5 mm. 
long, short-pilose; fruit glabrate, subglobose, 4 mm. long. 

Psychotria hondensis Standl. Journ. Wash. Acad. Sci. 18: 183. 
1928. Forests of Rio Hondo, Atlantic slope, 100 meters, Pittier 
16161. Atlantic tierra caliente. Endemic. A shrub of 3 meters, 
the young branches densely puberulent or short-pilose; stipules 
persistent, broadly triangular, narrowed to an obtuse apex; petioles 
short and stout, the blades broadly elliptic to elliptic-oblong, 18-29 
cm. long, 6-15 cm. wide, acute or acuminate, acute or acuminate at 
the base, glabrous above, densely velutinous-pubescent beneath; 
inflorescence terminal, cymose-paniculate, pedunculate, open, rather 
few-flowered, 5-11 cm. broad, the branches short-pilose, the flowers 
sessile or subsessile; calyx 2.5 mm. long, subtruncate; corolla in bud 
11 mm. long, puberulent-tomentose; fruit 13 mm. in diameter. In 
preparing the present account of Costa Rican Rubiaceae, I have 
had at hand only a photograph of the type of this species. I judge 



FLORA OF COSTA RICA 1353 

that the plant is really a Coussarea, but delay its transfer until the 
type specimen may be re-examined. 

Psychotria horizontalis Swartz. P. longicollis Benth. ex Oerst. 
Vid. Medd. Kjoebenhavn 1852: 33. 1853 (based in part on specimens 
collected on Isla de San Lucas, Golfo de Nicoya, by Oersted). 
Thickets and forest of the tierra caliente; region of San Ramon; 
Guanacaste; at 1,000 meters or less. Widely distributed in tropical 
America, as far south as Ecuador, and north to Mexico. A shrub 
of 1-3 meters, glabrous or nearly so; stipules ovate- triangular, acute 
or obtuse, caducous, 3-8 mm. long; leaves short-petiolate, elliptic 
to oblong-lanceolate, acuminate, acute or obtuse at the base, bright 
grayish green when dry, glabrous or nearly so; inflorescence terminal, 
pedunculate, small and rather few-flowered, usually 5-rayed at the 
base, the branches puberulent, the flowers mostly sessile; calyx 
deeply 5-lobate, the lobes linear-lanceolate; corolla white, 3^4 mm. 
long, glabrous; fruit red, glabrous, 4 mm. long. 

Psychotria involucrata Swartz. Forests of the tierra caliente; 
Guanacaste. Widely distributed in tropical America. An almost 
glabrous shrub, usually 3 meters high or less; stipules short, green, 
persistent, the lobes very short; leaves on very short petioles, lanceo- 
late to elliptic-ovate, long-acuminate, abruptly contracted and 
decurrent at the base; inflorescence pedunculate, subcapitate, com- 
monly 1-2 cm. long, short-pedunculate, few-flowered, the short 
branches purplish, pubescent, the flowers sessile or nearly so; outer 
bracts narrow, much exceeding the flowers, purple; calyx minute, 
denticulate; corolla white or ochroleucous; fruit globose, 4 mm. long, 
black. 

Psychotria Jimenezii Standl. Journ. Wash. Acad. Sci. 15: 288. 
1925. Wet forest, La Hondura, Prov. San Jose", 1,400 meters, 
Standky 37892. Regions of Zarcero and San Ramon; 1,000-2,000 
meters. Endemic. A shrub or small tree of 2.5-4.5 meters, the 
branches ferruginous- villosulous; stipules triangular, acute, 1 cm. 
long, brown, caducous; leaves almost sessile, oblong-oblanceolate, 
9-14 cm. long, 2.5-4 cm. wide, long-acuminate, attenuate to the 
base, puberulent beneath along the nerves, otherwise glabrous; 
inflorescence terminal, long-pedunculate, cymose-paniculate, open 
and many-flowered, the branches fulvous- villosulous, the flowers 
sessile in small, headlike cymes; calyx 1 mm. long, very shallowly 
lobate; corolla creamy white, 2.5 mm. long, glabrous. 



1354 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Psychotria limonensis Krause, Bot. Jahrb. 54: Beibl. 119: 43. 
1916. Uvita Island, Limon, Pittier 12681 (photo, of type in Herb. 
Field Mus.). Region of Limon, Atlantic tierra caliente. Southern 
Mexico to Colombia. An almost glabrous shrub or small tree; 
stipules ovate, acuminate, large, ferruginous, caducous; leaves petio- 
late, elliptic-ovate or oblong-elliptic, 6-12 cm. wide, abruptly short- 
acuminate, abruptly contracted at the base; inflorescence terminal, 
sessile, much branched, rather large, usually as broad as long, the 
flowers mostly short-pedicellate; calyx minutely denticulate; corolla 
white, 5-6 mm. long, glabrous outside, the throat white-barbate. 

Psychotria macrophylla Ruiz .& Pavon. P. anomothyrsa 
Schum. & Donn. Smith. Atlantic tierra caliente; Atlantic slope of 
the mountains of Guanacaste; regions of Dota and Cartago; at 
1,700 meters or less. Southern Mexico to Bolivia. Stems usually 
simple but sometimes branched, 2.5 meters high or less, glabrous 
or nearly so; stipules short, broad, apiculate, deciduous; leaves large, 
slender-petiolate, elliptic-oblong to lance-oblong, thin, acuminate, 
acute to attenuate at the base, glabrous or pruinose-puberulent 
beneath on the nerves; inflorescences axillary, long-pedunculate, 
paniculate, rather few-flowered, the branches often reflexed, glabrous 
or minutely puberulent, the flowers sessile, the bracts minute; calyx 
denticulate; corolla whitish or yellowish, glabrous, 4-5 mm. long; 
fruit 4-5 mm. long, glabrous, white. 

Psychotria magna Standl. P. compressicaulis Schum. Bot. 
Jahrb. 54: Beibl. 119: 44. 1916 (forests of Tuis, 600 meters, Pittier 
12412). Known in Costa Rica only from the collection cited. 
Panama. A shrub of 3 meters, the young branches minutely puberu- 
lent or glabrous; stipules caducous; petioles 2-4 cm. long, the blades 
oblong-elliptic, 18-25 cm. long, 8-11 cm. wide, caudate-acuminate, 
obtuse or acute at the base, glabrous or nearly so; inflorescence 
terminal, sessile, cymose-paniculate, large and open, about 10 cm. 
long and broad, the branches very minutely puberulent, the flowers 
sessile; calyx undulate, ciliate; corolla yellowish white, 3-4 mm. 
long, glabrous outside, white-barbate in the throat. 

Psychotria marginata Swartz. P. nicaraguensis Benth. ex 
Oerst. Vid. Medd. Kjoebenhavn 1852: 34. 1853 (based in part upon 
material collected at Jaris by Oersted) . Forests of the tierra caliente. 
Widely distributed in tropical America. An almost glabrous shrub, 
about 2 meters high; stipules large, ferruginous, oblong to ovate, 
entire, caducous; leaves short-petiolate, obovate-oblong to oblong- 



FLORA OF COSTA RICA 1355 

oblanceolate, blackening when dried, abruptly acute or acuminate, 
long-attenuate to the base, minutely puberulent beneath or glabrate; 
inflorescence terminal, cymose-paniculate, long-pedunculate, lax, the 
slender pedicels 2-5 mm. long; calyx teeth short, obtuse; corolla 
ochroleucous, 3 mm. long, glabrous outside; fruit globose, 3-4 
mm. long. 

Psychotria Maxonii Standl. Proc. Biol. Soc. Wash. 37: 53. 
1924. On mossy tree trunk, Santa Clara de Cartago, 1,950 meters, 
Maxon & Harvey 8250. Occasional in mountain forest of the central 
region; region of San Ramon; at 1,000-2,000 meters. Endemic. A 
small, epiphytic shrub, a meter long or less, the branches often 
reddish, glabrous throughout; stipules united to form a sheath 
1-1.5 mm. long; leaves almost sessile, linear-lanceolate, 2.5 cm. long 
or less, 2.5-5 mm. wide, acute or acuminate, acute at the base, pale 
beneath, the nerves obsolete; inflorescence cymose, 6-8-flowered, 
terminal, the pedicels 2-3 mm. long; calyx less than 1 mm. long, 
dentate; corolla white, the tube 2.5-4 mm. long, glabrous; fruit 
black. This has smaller leaves than any other Central American 
species of the genus. 

Psychotria microdon (DC.) Urban. P. pinularis Sesse" & Moc. 
Thickets of the Pacific tierra caliente; Guanacaste. Mexico to 
Ecuador. A shrub of 3.5 meters or less, glabrous almost throughout; 
stipules very short, the lobes caducous, the sheath indurate and pale 
in age; leaves mostly oblong-obovate, 4-7 cm. long, obtuse to short- 
acuminate, thin, attenuate to the base, barbate beneath in the axils 
of the nerves, otherwise glabrous or nearly so; inflorescences terminal, 
pedunculate, cymose-corymbose, few-flowered, the flowers pedicel- 
late; calyx pale, truncate; corolla white, as much as 10 mm. long. 

Psychotria Mortoniana Standl., sp. nov. Arbor 6-metralis, 
ramulis subteretibus crassiusculis ubique dense villosulis, internodiis 
elongatis; stipulae persistentes ca. 1 cm. longae sparse villosulae 
liberae, fere ad basin bilobae, lobis erectis e basi late triangulari sub- 
abrupte longiacuminatis; folia magna breviter petiolata firme mem- 
branacea, petiolo crassiusculo 1-2.5 cm. longo dense villosulo; lamina 
late elliptico-obovata vel obovato-oblonga ad 30 cm. longa et 11 cm. 
lata, subabrupte longe angusteque attenuato-acuminata, basi acuta, 
supra in sicco olivacea ad costam villosula, aliter subscabrido-asper- 
ata, nervis pallidis, subtus fere concolor, praesertim ad venas molliter 
villosula, costa gracili elevata, nervis lateralibus utroque latere ca. 25 
gracilibus prominulis leviter arcuatis juxta marginem obscure junctis, 



1356 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

venulis obscuris laxe reticulatis; inflorescentia terminalis cymoso- 
paniculata erecta crasse 3 cm. longe pedunculata, ca. 8 cm. longa 
atque 11 cm. lata, dense multiflora, basi trichotoma, ramis primariis 
ca. 7 angulo fere recto divergentibus crassiusculis dense fulvo-villosu- 
lis, ramis omnibus basi bracteatis, bracteis ad 1 cm. longis oblongo- 
lanceolatis acuminatis fere glabris, bracteis ultimis multo latioribus 
interdum fere orbicularibus apice obtusis vel rotundatis, floribus 
arete sessilibus aggregatis; hypanthium hirtellum vel glabratum post 
anthesin valde costatum, calyce vix ultra 1 mm. longo, lobis ovalibus 
vel ovatis obtusis apice ciliatis; corolla alba extus glabra apice in 
alabastro late rotundata ca. 7 mm. longa, tubo late obconico, lobis 
oblongo-ovatis tubo fere duplo brevioribus acutiusculis. Edge of 
forest, vicinity of El General, Prov. San Jose", 950 meters, Alexander 
F. Skutch 2697 (type in U. S. Nat. Herb.; photo, in herb. Field Mus.). 

Psychotria orchidearum Standl. Journ. Wash. Acad. Sci. 18: 
276. 1928. On tree in wet forest, Cerros de Zurqui, northeast of 
San Isidro, Prov. Heredia, 2,200 meters, Standley & Valeria 50863. 
Also at Palmira and Cerro Gallito, 1,800-2,000 meters. Endemic. 
A small, epiphytic shrub, 15-30 cm. high, glabrous throughout; 
stipules forming a truncate, indurate sheath 1-2 mm. long; petioles 
2-4 mm. long, the blades elliptic-oblong, 1.5-3 cm. long, mostly 6-12 
mm. wide, obtuse or acute, apiculate, cuneate-acute or attenuate at 
the base, thick and fleshy, paler beneath, the nerves obsolete; inflores- 
cence terminal, cymose-paniculate, lax, few-flowered, pedunculate, 
1.5 cm. broad or smaller, the pedicels mostly 3-5 mm. long; calyx 
1 mm. long, dentate to the middle or more deeply; fruit red, sub- 
globose, 3 mm. long. 

Psychotria orosiana Standl. Journ. Wash. Acad. Sci. 15: 288. 
1925. Moist forest near Orosi, Prov. Cartago, Standley 39803. Also 
in the region of Navarro, 1,400 meters. Endemic. A shrub of 1-3 
meters, the branchlets glabrous; stipules 5 mm. long, bilobate, 
caducous; leaves essentially sessile, oblong-obovate or elliptic-obo- 
vate, 6-12 cm. long, 2-4.5 cm. wide, abruptly acute or acuminate, 
gradually or abruptly long-attenuate to the base, glabrous; inflores- 
cence terminal, long-pedunculate, cymose-paniculate, 4-6 cm. broad, 
open, the branches glabrous, the flowers sessile or short-pedicellate; 
calyx 1 mm. long, shallowly dentate; corolla greenish white, glabrous, 
2.5-3 mm. long. 

Psychotria parvifolia Benth. ex Oerst. Vid. Medd. Kjoebenhavn 
1852: 36. 1853. Volcan de Barba, 1,800 meters, and Naranjo, at 



FLORA OF COSTA RICA 1357 

1,350 meters, Oersted (photo, of Oersted 11637, from Naranjo, in Herb. 
Field Mus.). Frequent in mountain forests of the central region; 
regions of Dota and San Ramon; Atlantic tierra caliente; at 1,800 
meters or less. Panama. A rather densely branched, very leafy 
shrub of 1-2.5 meters, the slender branches puberulent or short- 
pilose; stipules small, ovate, ferruginous, deciduous, about 4 mm. 
long; leaves small, slender-petiolate, ovate to oblong-oblanceolate, 
mostly 3.5 cm. long or less, acute or subobtuse, usually long-attenuate 
to the base and decurrent, glabrous, paler beneath; flowers 4-parted, 
cymose, the cymes sessile, few-flowered, the flowers mostly sessile; 
calyx obscurely dentate; corolla white, 4 mm. long or less, glabrous 
outside, barbate in the throat; fruit small, globose, red. 

Psychotria patens Swartz. P. flexuosa Willd. Apparently 
common in the region of San Ramon, at about 1,000 meters, and 
doubtless also in the Atlantic tierra caliente. Widely distributed in 
the lowlands of tropical America. A slender, glabrous shrub, com- 
monly 1-1.5 meters high; stipules short, green, persistent, biaristate; 
leaves medium-sized or rather small, usually yellowish green when 
dried, short-petiolate, lanceolate to lance-oblong, long-acuminate, 
obtuse or acute at the base; inflorescence terminal, slender-peduncu- 
late, thyrsoid-paniculate, often elongate, the short, slender branches 
mostly reflexed, not bracteate at the base; calyx minute, the teeth 
triangular; corolla whitish, glabrous, 7 mm. long; fruit white or pale 
blue, 3 mm. long. 

Psychotria pithecobia Standl. Field Mus. Bot. 8: 187. 1930. 
La Palma de San Jose", 1,550 meters, Tonduz 12432. Collected also 
at Zarcero, Matina, Santo Domingo de Vara Blanca, and in the 
region of San Ramon, 1,000-2,200 meters. Panama. An epiphytic, 
glabrous shrub about 50 cm. high, the branches brown; stipules short, 
connate to form a truncate, persistent, indurate sheath; leaves 
petiolate, elliptic-oblong, 5-8 cm. long, 2-3.5 cm. wide, rather long- 
acuminate, acute at the base, coriaceous when dry, the 10 pairs of 
lateral nerves prominent on both surfaces; inflorescence terminal, 
long-pedunculate, cymose-corymbose, 5-7 cm. broad, trichotomous 
at the base, the pedicels 2-4 mm. long; calyx 1 mm. long, the lobes 
narrowly triangular, acute; corolla 8 mm. long, white or tinged with 
pink; fruit dark red or black, 4-5 mm. long, globose. To this species 
probably is referable all the Costa Rican material that has been 
referred previously to P. pendula (Jacq.) Urban, although it is pos- 
sible that that West Indian species does occur in the country. 



1358 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Psychotria Pittieri Standl. Forests of the Atlantic tierra cali- 
ente, apparently rare; collected at La Colombiana. Panama. A 
slender shrub, 1-2 meters high, the branches pilose with rather long, 
pale, appressed or spreading hairs; stipules persistent, 4-6 mm. long, 
bilobate, the lobes subulate; leaves small, short-petiolate, elliptic- 
oblong, 4-5.5 cm. long, 1.5-2.5 cm. wide, hirsutulous above along the 
costa, appressed-pilose beneath; inflorescence terminal, reflexed in 
age, cymose-paniculate, about 2 cm. long, few-flowered, the branches 
pilose, the flowers subsessile; calyx 1 mm. long, the lobes obtuse; 
corolla white, 4-5 mm. long, minutely appressed-pilose; fruit blue. 

Psychotria polyphlebia Donn. Smith, Bot. Gaz. 33: 253. 1902. 
Forests of Suerre, Llanuras de Santa Clara, 300 meters, J. D. Smith 
6854- Frequent in forests of the Atlantic tierra caliente, ascending 
to 900 meters; region of San Ramon, at 1,200 meters or less. Panama 
and Colombia. Plants suffrutescent, simple, usually 50 cm. high or 
less, the stems villous with long, ferruginous hairs or glabrate; stipules 
bifid, soon deciduous; leaves long-petiolate, oblong-obovate, 11-16 
cm. long and 5-7.5 cm. wide or smaller, rounded or very obtuse at the 
apex, attenuate to the base, glabrous above and conspicuously 
white-punctate, hirsute beneath, the lateral nerves 25-34 pairs, 
elevated and very conspicuous on both surfaces; flowers capitate, the 
peduncles axillary, shorter or longer than the petioles, the heads 
dense and many-flowered, 1.5-2.5 cm. broad or in fruit larger, the 
outer bracts oblong-ovate, green; calyx denticulate; corolla white, 
pilose, naked in the throat; fruit red, 7-9 mm. long. An isolated 
species, in habit similar to some species of Cephaelis, but in fact 
probably related rather to Psychotria macrophylla. 

Psychotria psychotriaefolia (Seem.) Standl. Forests of the 
tierra caliente. Extending to Colombia. A shrub of 4 meters or less, 
the young branches ferruginous- tomentose; stipules caducous, thin, 
ferruginous, 1.5-2 cm. long, attenuate; leaves short-petiolate, obovate 
to oblong-oblanceolate, acute or short-acuminate, attenuate to the 
base, puberulent beneath, at least on the nerves; inflorescences 
chiefly terminal, sessile, dense, the branches 2 cm. long or less, 
ferruginous- tomentose, the flowers sessile, glomerate; calyx dentate; 
corolla white, 3 mm. long; fruit red, 4-5 mm. long, glabrous. 

Psychotria pubescens Swartz. P. glauca Polak. Linnaea 41: 
569. 1877 (in hedges, San Jose", Polakowsky 377; fragment of the type 
seen, ex herb. Berlin) . Frequent in hedges and thickets of the Meseta 
Central; region of San Ramon; Atlantic slope of Guanacaste; chiefly 



FLORA OF COSTA RICA 1359 

at 600-1,200 meters; probably in the Atlantic tierra caliente. Mex- 
ico, Central America, and West Indies. A shrub, commonly 1-2 
meters high, the branches minutely pilose or puberulent, terete; 
stipules small, persistent, the subulate lobes longer than the short 
sheath; leaves slender-petiolate, thin, oblanceolate to lance-oblong, 
6-14 cm. long, acuminate, usually attenuate to the base, finely 
puberulent beneath; inflorescence corymbiform, pedunculate, small, 
many-flowered, densely short-pilose; calyx very small, lobate, the 
lobes ovate, acute; corolla white or pale yellow, 4 mm. long, puberu- 
lent; fruit red or black, 4 mm. long, pilose. 

Uragoga pulvinigera Kuntze (Rev. Gen. 1: 300. 1891) is listed 
in the card index of the Gray Herbarium as doubtfully Costa Rican, 
but for no evident reason. Kuntze gives no locality for his new 
species, the type of which has not been found in the. herbarium of the 
New York Botanical Garden, where it should exist, and there is no 
certainty that the plant is even American. 

Psychotria quinqueradiata Polak. Linnaea 41: 570. 1877. P. 
Morae Polak. loc. cit. (San Jose*, Polakowsky 171; the species was 
named for J. Rafael Mora, President of Costa Rica). In hedges near 
San Jose* and in forests of Cerro de La Carpintera, Polakowsky 94, 
135, 199 (photo, of No. 135, from La Carpintera, in Herb. Field Mus.). 
Frequent in hedges and thickets about San Jose" and Cartago, extend- 
ing to the Pacific coast; region of San Ramon; Guanacaste; at 1,400 
meters or less. Endemic. A shrub of 4 meters or usually less, gla- 
brous; stipules caducous, very obtuse or rounded, 7 mm. long, 
ferruginous; leaves almost sessile, the petiole less than 5 mm. long, 
elliptic-oblong or oblanceolate, mostly 7-10 cm. long, acute or 
acuminate, attenuate to the base, the base itself narrowly truncate 
to subacute; inflorescence terminal, cymose-paniculate, sessile, 
umbelliform, radiately branched from the base, the flowers glomerate, 
sessile, the rays of the umbel mostly simple; calyx obscurely dentate; 
corolla white, the tube 2 mm. long. Noteworthy for the form of the 
leaves, the narrow base often truncate. 

Psychotria racemosa (Aubl.) Willd. Forests of the Atlantic 
tierra caliente, ascending to Pejivalle, 900 meters; region of San 
Ramon. Widely distributed in tropical America. A shrub of 1-1.5 
meters, the branches puberulent; stipules persistent, biparted, the 
stiff, subulate lobes 8-10 mm. long; leaves green when dried, short- 
petiolate, elliptic-oblong, medium-sized, short-acuminate, acute or 
abruptly contracted at the base, glabrous or nearly so; inflorescence 



1360 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

terminal, short-pedunculate, thyrsiform, small and dense, minutely 
hirtellous, the flowers sessile; calyx lobes linear-lanceolate; corolla 
whitish, hirtellous; fruit 5-celled, 4 mm. long, at first orange-red, 
becoming black. Differing from all other Central American species 
in its 5-celled fruit. The species has been referred to a separate genus 
as Nonatelia racemosa Aubl. 

Psychotria ramonensis Standl., sp. nov. Frutex gracilis 
metralis, ramis teretibus viridibus glabris; stipularum vagina 6 mm. 
longa viridis glabra, lobis approximatis oblongo-linearibus attenuatis 
5 mm. longis erectis; folia modica vel subparva graciliter petiolata 
firme membranacea, petiolo 7-20 mm. longo glabro; lamina elliptico- 
oblonga vel lanceolato-oblonga 6.5-13 cm. longa 2.5-4.5 cm. lata 
subabrupte longe anguste acuminata, apice ipso obtuso, basi sub- 
inaequali acuta vel subrotundata atque abrupte decurrens, supra in 
sicco viridis glabra, costa prominente, subtus paullo pallidior secus 
costam minute pilosula, aliter glabra, costa gracillimo prominente, 
nervis lateralibus utroque latere ca. 15 tenerrimis prominulis sub- 
arcuatis angulo fere recto divergentibus, nervo altero paullo teneriore 
inter paria subparallelo, venulis inconspicuis laxe reticulatis; inflores- 
centia terminalis sessilis vel breviter pedunculata, laxe subcapitata, 
ca. 2 cm. longa, pauciflora, pedunculo glabro, bracteis foliaceis ut 
videtur viridibus oblongo-lanceolatis usque 12 mm. longis, interiori- 
bus brevioribus abrupte petiolatis extus glabris, intus adpresso- 
pilosulis, floribus sessilibus; calyx ad apicem fructus persistens ca. 
1.5 mm. longus glaber, lobis anguste triangularibus acutis; fructus 
in statu vivo caeruleus 12 mm. longus, pyrenis vix ultra 4 mm. lon- 
gis. Forest of La Palma de San Ramon, by a small stream, 1,250 
meters, November, 1925, Brenes 4599 (type in Herb. Field Mus.). 

Psychotria sarapiquensis Standl., sp. nov. Subgenus Ma- 
pouria. Arbor 6-metralis praeter inflorescentiam omnino glabra, 
ramis crassiusculis subteretibus viridibus, internodiis superioribus 
3-4 cm. longis; stipulae caducae, non visae; folia mediocria longius- 
cule petiolata subpapyracea, petiolo gracili 2.5-3 cm. longo; lamina 
oblonga vel anguste elliptico-oblonga 9-11 cm. longa 3-4 cm. lata 
subabrupte acuminata, basi acuta vel basin versus angustata atque 
interdum breviter decurrens, supra in sicco cinereo-viridis, costa 
subimpressa, nervis planis, venis obsoletis, subtus paullo pallidior, 
costa pallida prominente, nervis lateralibus utroque latere ca. 8 
prominentibus angulo lato adscendentibus subarcuatis, venis obso- 
letis; inflorescentia terminalis sessilis cymoso-paniculata basi tri- 
chotoma, 7 cm. longa atque aequilata, ramis primariis angulo lato 



FLORA OF COSTA RICA 1361 

adscendentibus minutissime pulverulento-puberulis, bracteis deciduis, 
floribus ad apices ramulorum umbellato-aggregatis pedicellatis, pedi- 
cellis usque 3 mm. longis; hypanthium minutissime puberulum 
turbinatum ca. 1 mm. longum basi attenuatum, calyce albido trun- 
cate fere 1 mm. alto subpatente, fere 2 mm. lato; corolla viridescens 
extus minutissime sparse puberula in alabastro apice obtusa, tubo 
crasso 3 mm. longo superne vix dilatato, lobis triangulari-oblongis 
2 mm. longis obtusis subpatentibus intus non barbatis; stylus breviter 
exsertus. In forest, Vara Blanca de Sarapiqui, 1,500 meters, Alex- 
ander F. Skutch 3330 (type in U. S. Nat. Herb. ; photo, in Herb. 
Field Mus.). 

Psychotria Siggersiana Standl. Journ. Wash. Acad. Sci. 15: 
289. 1925. Wet forest near Guapiles, Prov. Limon, 500 meters, 
Standley 37210. Forests of the Atlantic tierra caliente; collected 
also at Tuis and in the Changuinola Valley. Panama. A simple- 
stemmed shrub, 2 meters high or less, the stems green, sparsely 
villous-hirsute or glabrous; stipules green, rounded-deltoid, 7 mm. 
long, finally deciduous; petioles 4-9 cm. long, the blades elliptic 
or broadly oval, 20-35 cm. long, 11-17 cm. wide, abruptly short- 
acuminate at the broad, often rounded apex, acute at the base, 
villous-hirsute on both surfaces or sometimes glabrate above; in- 
florescences axillary, cymose-paniculate, many-flowered, sometimes 
20 cm. broad, pedunculate, the branches villous-hirsute, the flowers 
sessile, glomerate; calyx 1 mm. long, the lobes deltoid-ovate; corolla 
greenish white, 3 mm. long, sparsely short- villous; fruit subglobose, 
red, 5-6 mm. long. 

Psychotria suerrensis Bonn. Smith, Bot. Gaz. 27: 337. 1899. 
Forests of Suerre, Llanuras de Santa Clara, 300 meters, J. D. Smith 
6580. Forests of the Atlantic tierra caliente. Nicaragua; probably 
also in Panama. A shrub of 4.5 meters or less, the branches glabrous; 
stipules 4-5 mm. long, the lobes subulate; leaves very shortly petio- 
late, oblong-elliptic to lance-oblong, mostly 12-20 cm. long and 
4-9 cm. wide, long-acuminate, acute or acuminate at the base, gla- 
brous, usually tinged with red when dried; inflorescence terminal, 
short-pedunculate, subcapitate, dense and many-flowered, 2.5-4 cm, 
wide, the peduncle short-pilose, the bracts red, oblong-elliptic, 
pubescent, mostly 1-1.5 cm. long; calyx minute, subtruncate; corolla 
white, 12 mm. long; fruit violaceous, puberulent, 4 mm. long. 

Psychotria sylvivaga Standl. Journ. Wash. Acad. Sci. 18: 274. 
1928. Wet forest, Yerba Buena, northeast of San Isidro, Prov. 



1362 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Heredia, 2,000 meters, Standley & Valeria 1+9989. Also in the region 
of El Copey, at 2,200 meters. Panama. A shrub of 1-3 meters, 
the young branches minutely puberulent or glabrous; stipules cadu- 
cous, broadly ovate, 8-10 mm. long, ferruginous; leaves slender- 
petiolate, oblong-oblanceolate or elliptic-oblong, usually broadest 
above the middle, 9-17 cm. long, 2.5-5.5 cm. wide, abruptly acumi- 
nate, long-attenuate to the base, sparsely barbate beneath in the 
axils of the nerves, otherwise glabrous; inflorescence terminal, cymose- 
paniculate, long-pedunculate, open, many-flowered, 4.5-9.5 cm. long, 
the branches minutely puberulent, the pedicels 1-3 mm. long, in 
fruit sometimes 8 mm. long; calyx 1 mm. long, shallowly dentate or 
subtruncate; corolla greenish white, glabrous, the tube 5 mm. long; 
fruit 5-6 mm. long. 

Psychotria Tonduzii Standl. Journ. Wash. Acad. Sci. 15: 287. 
1925. Wet forest, La Estrella, Prov. Cartago, Standley 39461. 
Frequent in mountain forests of the central region, also in the 
Atlantic tierra caliente, at 1,500 meters or less. Endemic. A simple 
shrub about a meter high, the stems green, glabrous; stipules broadly 
triangular, green, 4 mm. long, at least the base persistent; leaves 
long-petiolate, elliptic-oblong to obovate-oblong, 19-35 cm. long and 
7-14 cm. wide, obtuse or rounded at the apex and cuspidate-acumi- 
nate, acute to long-decurrent at the base, glabrous; panicles axillary, 
usually sessile and branched from the base, the stout branches 
puberulent, the flowers sessile, densely glomerate; calyx 1.5 mm. 
long, the lobes broadly ovate, subacute; corolla ochroleucous, 3 mm. 
long, minutely puberulent. 

Psychotria Torresiana Standl. Journ. Wash. Acad. Sci. 15: 288. 
1925. Wet forest near Orosi, Prov. Cartago, Standley 39769. En- 
demic, and known only from the original locality. A shrub of 3 
meters, the slender branches glabrous; stipules persistent, green, 
4 mm. long, the lobes very short, obtuse; leaves slender-petiolate, 
lance-linear, 9-16 cm. long, 1.5-2.5 cm. wide, narrowly long-attenu- 
ate, acute or attenuate at the base, glabrous, pale beneath; inflores- 
cence terminal, slender-pedunculate, open-paniculate, many-flowered, 
about 11 cm. long and almost as broad, the primary branches divari- 
cate or refracted, glabrous, the flowers sessile or short-pedicellate; 
calyx 1 mm. long, shallowly dentate; fruit subglobose, blue, 5 mm. long. 

Psychotria uliginosa Swartz. Wet forest of the Atlantic tierra 
caliente; region of San Ramon; Atlantic slope of Guanacaste; at 
1,000 meters or less. Southern Mexico to West Indies and Ecuador. 



FLORA OF COSTA RICA 1363 

Plants usually simple, herbaceous or suffrutescent, 1.5 meters high 
or less, glabrous or nearly so; leaves petiolate, obovate to oblong or 
elliptic-oblong, 25-30 cm. long, 10-12 cm. wide or smaller, fleshy 
when fresh, acuminate, usually acute or attenuate at the base, very 
pale beneath; panicles axillary, long-pedunculate, small, trichotom- 
ous, the flowers sessile, densely aggregate; calyx 5-dentate; corolla 
white, 6 mm. long, barbate in the throat; fruit bright red, 6 mm. 
long, the nutlets strongly compressed. 

Psychotria Valeriana Standl., sp. nov. Frutex gracilis 1-1.5 m. 
altus praeter inflorescentiam omnino glaber, ramulis teretibus vel 
subcompressis, internodiis elongatis; stipularum vagina truncata vix 
2 mm. longa, dentibus 2 approximatis erectis subulatis vulgo vagina 
brevioribus; folia inter minora breviter petiolata papyracea, petiolo 
gracili 8-12 mm. longo; lamina lanceolato-oblonga 7-12.5 cm. longa 
2-4 cm. lata longe anguste acuminata, acumine saepe subfalcato, 
basi acuta vel attenuata, interdum obtusa atque subito decurrens, 
supra in sicco olivacea lucida, costa nervisque prominentibus, subtus 
paullo pallidior in sicco vulgo rubrotincta, costa tenui elevata, nervis 
lateralibus utroque latere ca. 10 arcuatis angulo lato adscendentibus, 
venulis prominulis laxe reticulatis; inflorescentia terminalis cymoso- 
paniculata, saepe corymbiformis, vulgo longe graciliter pedunculata 
sed saepe sessilis 1.5-3.5 cm. longa atque aequilata pauciflora vel 
multiflora, saepius saltern in statu florifero densa, ramis suberectis 
vel valde adscendentibus, rare subdivaricatis minutissimepuberulis vel 
glabratis crassiusculis, basi minute bracteatis, floribus sessilibus 
vel breviter pedicellatis; hypanthium minus quam 1 mm. longum gla- 
brum, calyce vix ultra 0.3 mm. longo truncate vel obsolete minute den- 
ticulate; corolla alba extus minutissime pulverulenta 3-4 mm. longa, 
fauce non barbata, tubo crassiusculo, lobis 5 brevissimis; antherae 
breviter exsertae. In forest, Cerros de San Pedro de San Ramon, 
1,075 meters, June, 1926, Brews 4886 (type in Herb. Field Mus.); 
also Nos. 18981, 14309, 4265, from the same locality. Alto de La 
Calera de San Ramon, Brenes 5871. La Palma de San Ramon, 
1,050-1,100 meters, Brenes 20641, 6200, 5605. Without locality, 
Endres 188. The species is named for Juvenal Valeric Rodriguez, 
Director of the Museo Nacional de Costa Rica. 

Psychotria viridis Ruiz & Pavon. P. microdesmia Oerst. Vid. 
Medd. Kjoebenhavn 1852: 36. 1853. Jaris, 900 meters, Oersted 
(photo, of type in Herb. Field Mus.). Collected also in the Sixaola 
Valley. British Honduras to Cuba and Bolivia. A shrub or small 



1364 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

tree, glabrous or nearly so; stipules caducous, large, thin, ferruginous; 
leaves blackening when dried, short-petiolate, obovate or obovate- 
oblong, acute or short-acuminate, cuneate-attenuate to the base; 
inflorescence pedunculate, spicate-paniculate, open, many-flowered, 
the minute flowers sessile in distant glomerules; corolla greenish 
white; fruit red, 4-5 mm. long. Easily recognized by the spicate 
branches of the open panicles. 

Psychotria Wendlandiana Standl. Journ. Wash. Acad. Sci. 
18: 9. 1928. San Miguel (de Naranjo?), Wendland 781 (photo, of 
type in Herb. Field Mus.). Collected also at Guapiles and La 
Hondura de San Jose", 300-1,300 meters. Endemic. A shrub of 
2.5-3 meters, the young branches densely short- villous; stipules 
caducous, 5-6 mm. long, rounded and bimucronate at the apex; 
leaves petiolate, ovate-oblong to oblong or oblanceolate-oblong, 
9-15 cm. long, 3-5.5 cm. wide, acuminate, narrowed to a truncate or 
deeply cordate base, more or less short-villous and puberulent be- 
neath; inflorescence terminal, cymose-paniculate, pedunculate, the 
flowers sessile; calyx dentate; corolla yellow, 2.5-3 mm. long, yellow. 
Easily distinguished by the cordate leaf bases. 

RANDIA L. 

Erect trees or shrubs, often armed with axillary or supra-axillary 
spines; leaves opposite, sessile or petiolate; stipules small, often sub- 
connate; flowers small or large, perfect or unisexual, axillary or 
terminal, solitary or fasciculate, usually white; hypanthium terete 
or costate; calyx commonly tubular, lobate or truncate, the lobes 
often elongate or foliaceous, persistent or deciduous; corolla funnel- 
form to campanulate or salverform, with a short or elongate tube, 
the throat glabrous or villous, the usually 5 lobes short or elongate, 
acute or obtuse, contorted; ovary normally 2-celled, the cells many- 
ovulate; fruit baccate, globose or oval, the pericarp usually hard and 
thick; seeds numerous or few, immersed in the pulp, commonly 
horizontal, compressed. A few other species occur in Central 
America. 

Randia aculeata L. Occasional in thickets of the tierra caliente; 
known from both coasts. Florida and Mexico to West Indies and 
Venezuela. A shrub or small tree, rarely more than 3 meters high, 
armed with stout spines 1.5 cm. long or less; leaves mostly clustered 
at the ends of the branches or on short, lateral spurs; leaves sessile or 
on very short petioles, very variable in shape and size, mostly oblong- 
obovate to obovate-orbicular, and about 3 cm. long, acute to broadly 



FLORA OF COSTA RICA 1365 

rounded at the apex, rounded to acute at the base, glabrous and 
lustrous above, glabrous beneath or sparsely pilose on the costa; 
flowers perfect, terminal, sessile, usually aggregate; calyx and hy- 
panthium 2-3 mm. long, commonly glabrous, the lobes linear to 
ovate; corolla white, 6-8 mm. long, glabrous outside, the 5 lobes 
acute or acuminate, equaling or longer than the tube, the throat 
densely white-barbate; fruit globose, 6-13 mm. in diameter. 

Randia armata (Swartz) DC. Mostrenco, Crucillo. Thickets 
of the Pacific tierra caliente; Guanacaste; to be expected also in the 
Atlantic lowlands. Widely distributed in tropical America. A shrub 
or small tree, rarely as much as 6 meters high, the branches usually 
bearing at the apex 4 stout spines 2.5 cm. long or less; leaves slender- 
petiolate, mostly ovate to oval or obovate, thin, usually 6-15 cm. 
long, acute or acuminate at each end, glabrous or puberulent above, 
puberulent or appressed-pilose beneath or glabrate; flowers dioecious, 
aggregate at the ends of the branches, slender-pedicellate; calyx lobes 
linear to obovate-oblong, 4-9 mm. long; corolla white or ochroleu- 
cous, glabrous or pilose outside, the tube 2.5 cm. long, the throat 
naked, the lobes rhombic-obovate, 1 cm. long; fruit oval or globose, 
1.5-3.5 cm. long, filled with a black, sweetish pulp. The pulp of this 
and other species is sometimes eaten, but it is unpleasant in appear- 
ance, and the flavor is not agreeable. Birds are very fond of the 
fruits, and it usually is difficult to find a ripe fruit that has not been 
punctured and robbed of its pulp and seeds. 

Randia Brenesii Standl., sp. nov. Frutex 3-4-metralis, ramis 
gracilibus teretibus ferrugineis, novellis dense pilis brevibus ple- 
rumque paten tibus hirtellis, internodiis brevibus; spinae secus ramos 
sparsae solitariae rigidae, graciles vel crassae, usque 1.5 cm. longae; 
stipulae minutae; folia parva brevissime petiolata membranacea, 
petiolo vix ad 2 mm. longo; lamina lanceolato-oblonga 2-4 cm. longa 
0.8-1.5 cm. lata acuta vel sensim acuminata, basi acuta vel obtusa, 
supra viridis glabra, costa prominente, nervis obsoletis, subtus paullo 
pallidior sparsissime praesertim ad costam hirtella vel fere glabra, 
costa gracili prominente, nervis lateralibus utroque latere ca. 4 
obscuris; flores dioeci ut videtur terminales atque solitarii, sessiles; 
hypanthium oblongum 5 mm. longum glabrum; calyx 5-partitus, 
lobis foliaceis viridibus rotundato-ovatis ca. 5 mm. longis atque 
aequilatis vel latioribus acutiusculis glabratis ciliatis; corolla alba 
extus glabra, tubo gracili 2-3.5 cm. longo superne paullo dilatato 
fauce glabro, lobis patentibus ovali-oblongis vel ovato-oblongis 



1366 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

obtusis vel subrotundatis intus glabris 13-20 mm. longis; fructus 
globosus glaber laevis 2 cm. diam. La Palma de San Ramon, March, 
1932, Brenes 15048 (type in Herb. Field Mus.); also Nos. 3748 and 
683 1 from the same locality. Entre San Ramon y La Palma de San 
Ramon, Brenes 6180. Tonduz 12481 from La Palma de San Jos in 
the Berlin herbarium also seems to be referable to this species, which 
is related to R. calycosa Standl., of Panama. 

Randia grandifolia (Donn. Smith) Standl. Journ. Wash. Acad. 
Sci. 18: 166. 1928. Basanacantha grandifolia Donn. Smith, Bot. 
Gaz. 55: 436. 1913. Forests near Santo Domingo, Golfo Dulce, 
Prov. Puntarenas, Tonduz 9878. Forests of the tierra caliente, 
ascending to the region of Cartago, and on the slopes of La Carpin- 
tera. Endemic. A shrub or small tree, 4.5 meters high or less, 
unarmed, glabrous throughout or nearly so; leaves large, mostly 
crowded at the tips of the branches, the petioles 3.5 cm. long or less, 
the blades elliptic to lance-oblong or elliptic-ovate, 12-40 cm. long, 
6-15 cm. wide, acute or acuminate at each end, lustrous; flowers 
dioecious, the staminate ones aggregate, subsessile; calyx 3 mm. long, 
sparsely short-pilose, the lobes subulate or lanceolate, 1-2 mm. long; 
corolla white, glabrous outside, the stout tube 12 mm. long, the 
throat naked, the lobes ovate, 7 mm. long; fruit yellow, globose, 2.5 
cm. in diameter. 

Randia Karstenii Polak. Linnaea 41: 568. 1877. Espino 
bianco, Horquetilla. In hedges near San Jose", Polakowsky 74 and 
302. Frequent from the Meseta Central to the coasts, usually in 
rather dry thickets; often in roadside hedges; region of San Ramon; 
Zarcero, at 2,250 meters. Endemic. A shrub or tree, sometimes 
7.5 meters high, with a trunk 25 cm. in diameter, the bark thick, 
gray-brown, slightly roughened, the branches appressed-pilose when 
young; spines terminal or lateral, stout, 1.5 cm. long or less; leaves on 
petioles 6 mm. long or less, obovate-oblong to oblong-elliptic, obovate, 
or ovate, 1.5-7 cm. long, 1-3.5 cm. wide, rounded to acute at the 
apex, acute at the base, glabrous above, minutely pilose beneath 
along the nerves or glabrous; flowers perfect, terminal, solitary, 
sessile; calyx and hypanthium 2-3 mm. long, glabrous, the lobes 
linear or oblanceolate; corolla white, 6 mm. long, glabrous outside, 
the throat naked, the lobes broadly ovate, short-acuminate; fruit 
globose, usually 1 cm. in diameter, but sometimes as much as 2.5 cm. 
Austin Smith states that the cambium layer stains the hands and 
clothing yellow. This shrub is much used about the Meseta Central 



FLORA OF COSTA RICA 1367 

as a hedge plant, a purpose for which it is admirably adapted because 
of its dense growth and abundant spines. 

Randia monantha Benth. has been reported from Costa Rica, 
probably in error, since at present it is known definitely only from 
Guatemala and southern Mexico. 

Randia panamensis Standl. In forest, region of San Ramon, 
about 1,100 meters. Panama. A shrub of 1.5-2.5 meters, with 
slender branches; spines lateral, slender, 5-7 mm. long; petioles 
3-6 mm. long, the blades obovate-oblong or oblanceolate-oblong, 
mostly 8-15 cm. long and 3-6.5 cm. wide, acute or short-acuminate, 
acute or acuminate at the base, glabrous above, glabrate beneath or 
strigose on the nerves, short-barbate in the axils of the nerves; 
flowers few, terminal, fasciculate, the pedicels 5-6 mm. long; calyx 
lobes 5-7 mm. long, linear-subulate; corolla ochroleucous, the tube 
2 cm. long, strigillose, the lobes oval or rounded, 5-6 mm. long, 
rounded at the apex. 

Randia Pittieri Standl. Contr. U. S. Nat. Herb. 20: 201. 1919. 
Horquetilla, Basanacantha Pittieri Standl. op. cit. 18: 134. 1916. 
Zent Junction, near Matina, Prov. Limon, Pittier 16036. Collected 
also at Escazu, and in Guanacaste. Endemic. Spines terminal in 
4's, 3-5 mm. long; petioles 9-14 mm. long, the blades thin, oval 
or obovate-oval, 12-16 cm. long, 7-10 cm. wide, abruptly acute, 
rounded or subcordate at the base, shortly setose-pilose above, 
paler beneath and abundantly setose-pilose; flowers dioecious, 
solitary or fasciculate, terminal; pedicels of the staminate flowers 
4-5 mm. long; calyx lobes subulate, 4 mm. long; corolla glabrous 
outside, the tube 6 cm. long, glabrous in the throat, the lobes lanceo- 
late or oblong-lanceolate, 2.5-3 cm. long, attenuate. 

Randia subcordata Standl. Contr. U. S. Nat. Herb. 20: 201. 
1919. Crucilla. Basanacantha subcordata Standl. op. cit. 18: 133. 
1916. Thickets and roadsides, Nicoya, Tonduz 13934- Known in 
Costa Rica only from the region of Nicoya. Extending to Honduras. 
A shrub or small tree, the spines borne at or near the apices of the 
branches, stout, 5-7 mm. long; leaves slender-petiolate, oval to 
broadly ovate, 4-10 cm. long, 2.5-6 cm. wide, acute or abruptly 
acute, subcordate or rounded at the base, membranaceous, minutely 
appressed-setose-pilose on the upper surface, abundantly appressed- 
pilose beneath; flowers dioecious, white, fragrant; staminate flowers 
in terminal fascicles of 2 or more, sessile; calyx and hypanthium 
strigose-sericeous, the lobes linear-subulate, 5 mm. long; corolla 



1368 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

densely hirtellous, the tube 5-6 cm. long, the lobes ovate or lan- 
ceolate, attenuate, 2.5-4 cm. long; pistillate flowers solitary, the 
calyx lobes oblanceolate-linear, 15-18 mm. long, the corolla tube 
4.5 cm. long. 

RAVNIA Oerst. 

Glabrous, epiphytic shrubs, the branchlets terete; leaves opposite, 
short-petiolate, somewhat carnose when fresh, coriaceous when dried ; 
stipules large, oblong, obtuse; flowers large, red, ternate at the ends 
of the branches, subsessile; hypanthium narrowly turbinate; calyx 
5-6-lobate, the lobes unequal, linear; corolla tubular, slightly curved, 
the tube slender, somewhat ampliate in the throat, constricted below 
the limb, the 5-6 lobes short, ovate, obtuse, spreading, imbricate; 
anthers subexserted; ovary 2-celled, many-ovulate; capsule slender- 
cylindric; seeds very numerous, with a tuft of long, slender hairs 
at the apex. The genus consists of only the two species listed here. 

Ravnia Pittieri Standl. N. Amer. Fl. 32: 114. 1921. Santa 
Clara, Pittier 13461. Collected also at Pejivalle, 900 meters; near 
El Cairo; and in the region of San Ramon, 1,000 meters. Endemic. 
A small shrub; stipules oblong-oblanceolate, 3.5 cm. long, acute, 
green; petioles scarcely 4 mm. long, the blades linear-lanceolate, 5- 
20 cm. long, 1.2-4 cm. wide, narrowly long-attenuate, obtuse or 
acute at the base, the lateral nerves obsolete or obscure; calyx 
lobes linear, 5-8 mm. long, one or more linear or subulate teeth 
between each 2 lobes; corolla 5.5 cm. long, the tube 12 mm. wide 
at the middle, the lobes 5 mm. long. This is probably only a narrow- 
leaved variety of R. triflora. 

Ravnia triflora Oerst. Vid. Medd. Kjoebenhavn 49. 1852. 
Near Cartago and Candelaria, 1,800-2,400 meters, Oersted 11696 
(photo, of type in Herb. Field Mus.). Frequent in forests of the 
central region; region of San Ramon; at 1,100-2,400 meters. Chiri- 
qui, Panama. A small, epiphytic shrub, usually a meter long or less; 
stipules 1.5 cm. long; leaf blades elliptic to elliptic-oblong or lance- 
oblong, mostly 5-10 cm. long, long-acuminate, obtuse or acute at 
the base; calyx lobes unequal, without interposed teeth; corolla 
bright red, 5-5.5 cm. long, 7-10 mm. wide above. The plant is a 
handsome one, although the flowers always are rather few. In 
general appearance it is more suggestive of the Gesneriaceae than of 
the Rubiaceae. 

RELBUNIUM Hook. f. 

Annual or perennial herbs, in habit and general appearance 
closely resembling Galium; leaves verticillate, narrow, small; flowers 






FLORA OF COSTA RICA 1369 



minute, the pedicels articulate with the involucre, this consisting of 
2-4 leaflike bracts; ovary didymous, the cells 1-ovulate; sepals none; 
corolla rotate, 4-lobate, the lobes valvate; stamens exserted; fruit 
more or less fleshy and juicy, smooth or tuberculate, normally 
2-seeded. One other species occurs in northern Central America. 
The genus is closely related to Galium, differing in having foliaceous 
bracts at the base of the flower. 

Relbunium hypocarpium (L.) Hemsl. Frequent in forest or 
thickets of the mountain regions, chiefly at 1,300-3,000 meters. 
Widely distributed in the mountains of America, from Mexico far 
southward. A weak-stemmed perennial, the stems sometimes a 
meter long or more, often subscandent, retrorse-hispidulous; leaves 
in 4's, oblong to elliptic or obovate, 5-15 mm. long, mucronate, 
narrowed to the base, more or less pilose and ciliate, the margins 
revolute; flowers pedicellate, equaling or shorter than the leaves; 
corolla white, 1-1.5 mm. broad; fruit orange-red, 2-3 mm. long. 

RICHARDIA L. 

Erect or prostrate annuals, usually with rough pubescence; 
stipules connate with the petioles into a setiferous sheath; leaves 
opposite; flowers small or large, densely crowded in terminal, involu- 
crate heads; ovary 3-4-parted, the cells 1-ovulate; sepals 4-6, more 
or less connate at the base; corolla funnelform, the lobes valvate; 
fruit consisting of 3-4 indehiscent, 1-seeded cocci. The genus is 
easy of recognition by its fruit, consisting of normally 3 or 4, rather 
than 2, nutlike cocci that separate at maturity. No other species 
are known from Central America. 

Richardia scabra L. Common as a weed in waste or cultivated 
ground, Meseta Central to the coasts; region of San Ramon. Gen- 
erally distributed in tropical America. Plants prostrate or ascending, 
often forming mats, pilose or hispid throughout; leaves petiolate, 
oblong or lanceolate, 2-9 cm. long, acute and mucronate; heads 
dense, few-many-flowered; sepals lanceolate; corolla white, 4-6 mm. 
long; carpels 3, muriculate, sulcate on the inner face, 2-3 mm. long. 

RONDELETIA L. 

Shrubs or trees, glabrous or pubescent; leaves opposite or rarely 
verticillate, sessile or petiolate; stipules usually broad, sometimes 
foliaceous, commonly persistent; inflorescence terminal or axillary, 
cymose, corymbose, or paniculate; calyx 4-5-lobate, the lobes often 
unequal, persistent; corolla funnelform or salverform, white, yellow- 



1370 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ish, or red, the tube usually slender and elongate, the throat often 
annular-thickened, sometimes barbate, the lobes spreading, broad, 
obtuse, imbricate; ovary 2-celled, many-ovulate; capsule small or 
large, generally globose, bisulcate, loculicidally or septicidally bi- 
valvate; seeds commonly minute, compressed or angulate, often 
winged or appendaged. Other species are known from Central 
America. 

Rondeletia amoena (Planch.) Hemsl. Teresa, Quina. R. 
rugosa Benth. ex Oerst. Vid. Medd. Kjoebenhavn 43. 1852 (Cartago, 
1,680 meters, Oersted). Frequent in forests of the central region, 
1,000-2,000 meters; regions of Dota and San Ramon. Panama to 
southern Mexico. A shrub or small tree, sometimes 5.5 meters high, 
with a trunk 15 cm. in diameter, but usually smaller, the bark dark 
cinnamon-brown, smooth; stipules triangular-oblong, obtuse, 1-1.5 
cm. long, reflexed; leaves on very stout, short petioles, ovate-oval to 
oblong-ovate, 6-15 cm. long, abruptly short-acuminate, rounded or 
subcordate at the base, glabrous or sparsely pilose above, densely 
short-pilose beneath; inflorescence terminal and axillary, cymose- 
corymbose, rather dense and many-flowered, about as broad as long; 
calyx lobes oblong or obovate-oblong, 1-2 mm. long; corolla pink, 
appressed-pilose, the tube 8-12 mm. long, the throat densely yellow- 
pilose, the 5 lobes 2-2.5 mm. long; capsule globose, 5-6 mm. in 
diameter. The shrub is a handsome one because of its abundance of 
bright pink flowers. It is cultivated at La Sabana in the garden of 
Dona Amparo de Zeledon, and long ago was introduced into cultiva- 
tion in Europe. 

Rondeletia aspera Standl. N. Amer. Fl. 32: 54. 1918. Puente 
del Rodeo, below Pacaca (Villa Colon), Prov. San Jose", Pittier 3243. 
In forest, region of San Ramon, 600-1,000 meters. Endemic. A 
shrub with slender branches; stipules linear to oblong-lanceolate, 6-10 
mm. long, erect; leaves short-petiolate, oblong-elliptic to lance-oblong, 
6-10 cm. long, acute or short-acuminate, acute or acuminate at the 
base, green above, very scabrous with short, slender hairs, densely 
and closely white-tomentose beneath when young, the tomentum 
more or less deciduous in age, the surface then short-pilose; inflores- 
cence terminal, cymose-paniculate but often much condensed and 
headlike, as broad as long; calyx lobes oval to ovate-deltoid, obtuse 
or rounded, 1-2 mm. long; corolla arachnoid-tomentose, the tube 8-9 
mm. long, the throat naked, the 4 lobes rounded, 3 mm. long. 

Rondeletia Brenesii Standl., sp. nov. Ramuli teretes brunnes- 
centes primo dense strigoso-sericei, internodiis brevibus vel elongatis; 



FLORA OF COSTA RICA 1371 

stipulae 8 mm. longae brunneae erectae extus sericeo-strigosae e basi 
triangular! subulato-acuminatae; folia majuscula petiolata mem- 
branacea, petiolo usque 1.5 cm. longo sed vulgo fere ad basin anguste 
alato; lamina oblanceolata vel oblongo-oblanceolata 14-22 cm. longa 
4-6.5 cm. lata longissime anguste falcato-attenuato-acuminata, basin 
versus longe sensim attenuata, supra viridis sparse pilis longis laxis 
albidis pilosa, subtus paullo pallidior, primo pilis longis laxis sub- 
adpressis albidis pilosa, cito glabrata, costa gracili prominente, nervis 
lateralibus utroque latere ca. 16 obliquis teneris leviter arcuatis; 
inflorescentia terminalis paniculata, sessilis vel breviter pedunculata, 
panicula spiciformi ad 16 cm. longa ca. 2.5 cm. lata dense multiflora, 
floribus sessilibus vel subsessilibus in cymulas parvas paucifloras 
densas breviter pedunculatas dispositis, ramis dense pilosis, bracteis 
parvis linearibus viridescentibus inconspicuis; calycis lobi vix 1.5 mm. 
longi viridescentes late ovales vel semiorbiculares glabrati, hypanthio 
adpresso-pilosulo; corolla ochroleuca, tubo gracili 5-5.5 mm. longo 
extus glabro vel infra lobos sparse hirtello, lobis 4 brevibus sub- 
orbicularibus basi tantum sparse hirtellis. Camino de San Ramon 
a La Calera, March, 1929, Brenes 6757 (type in Herb. Field Mus.). 
Rio Jesus de San Ramon, February, 1937, Brenes 22036. Cuesta de 
La Vieja, road to San Carlos, 800 meters, April, 1903, Pittier 16693; 
Cook & Doyle 39 (both specimens in U. S. Nat. Herb.). Closely 
related to R. stachyoidea Bonn. Smith, of northern Central America, 
and confused with that species by the writer when he prepared the 
account of the genus for the North American Flora. R. stachyoidea 
differs in its narrower, more abundantly and persistently pubescent 
leaves, its greatly elongate and conspicuous bracts, long and narrow 
calyx lobes, and longer corolla tube. 

Rondeletia buddleoides Benth. Frequent in mountain forests 
of the central region; Dota and San Ramon; 1,000-2,000 meters. 
Panama to southern Mexico. Usually a shrub of 2-3 meters, but 
sometimes a tree of 12 meters, with a trunk 20 cm. in diameter, the 
bark light gray; stipules lanceolate or oblong, 3-8 mm. long, obtuse 
to attenuate, erect; leaves on very short petioles, oval-elliptic to 
elliptic-oblong or lanceolate, 5-12 cm. long, rather thick, acute to 
long-acuminate, obtuse to long-acuminate at the base, green above, 
pilose or glabrous, covered beneath with a very dense and close, 
white tomentum; inflorescence a terminal, spikelike panicle 10-15 
cm. long, very dense and many-flowered; calyx lobes oblong or oval, 
obtuse, 1 mm. long; flowers fragrant; corolla dull white, tomentose 



1372 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

outside, the slender tube 4-8 mm. long, naked in the throat, the 

4 lobes 1-1.5 mm. long; capsule 3-4 mm. long. 

Rondeletia calycosa Bonn. Smith, Bot. Gaz. 56: 59. 1913. In 
forest, La Palma de San Jose", 1,460 meters, Tonduz 11633. Known 
also from forests on the slopes of nearby Barba, at about the same 
elevation. Endemic. A shrub of 2-3 meters; stipules triangular, 
cuspidate, 3-4 mm. long; leaves very shortly petiolate, lance-elliptic, 
6-9 cm. long, 2-3 cm. wide, abruptly acuminate or very long-acumi- 
nate, attenuate to the base, glabrous above, pilose beneath along the 
nerves; inflorescence terminal, cymose-corymbose, 5-6 cm. long and 
about as wide; calyx lobes linear-lanceolate, 4-7 mm. long; corolla 
red, strigillose-pilose, naked in the throat, the tube 15-17 mm. long, 
the 4 lobes 3 mm. long. 

Rondeletia costaricensis Standl. N. Amer. Fl. 32: 61. 1918. 
Near Buena Vista, San Carlos, 900 meters, Pittier 16696. Known 
only from the original collection. Stipules foliaceous, suborbicular, 

5 mm. long; leaves short-petiolate, elliptic or oblong-elliptic, 14-18 
cm. long, 6.5-8 cm. wide, acuminate, acute at the base, thin, glabrous; 
inflorescence terminal, cymose-paniculate, rather few-flowered, 
slender-pedunculate, calyx lobes narrowly triangular, acute, erect, 
shorter than the hypanthium; corolla minutely appressed-pilose, the 
slender tube 9 mm. long, the 4 lobes 3 mm. long. 

Rondeletia Pittieri Schum. & Krause ex Schum. Bot. Jahrb. 
40: 316. 1908. Teresa. Rio Poros and Rio Segundo, 2,000 meters, 
Pittier 1729. Occasional in mountain forests of the central region. 
Endemic. A shrub or tree of 3-6 meters, similar to R. amoena; 
differing in having the leaves strigose beneath on the nerves, rather 
than softly and densely pilose over the whole surface. 

Rondeletia stenostachya Standl., sp. nov. Rami graciles 
teretes ferruginei, novellis densissime pilis subadpressis ferrugineis 
pilosis, internodiis brevibus vel elongatis; stipulae ca. 7 mm. longae 
extus dense strigosae, e basi rotundato-ovata subulato-mucronatae 
persistentes erectae; folia modica sessilia vel brevissime crasse petio- 
lata firme membranacea lanceolata vel anguste oblongo-oblanceolata 
8-16 cm. longa 2-4 cm. lata longe anguste attenuato-acuminata, 
basin versus longe sensim attenuata, supra viridia sparse strigosa, 
subtus paullo pallidiora ubique dense strigoso-sericea vel in statu 
adulto subglabrata, nervis lateralibus valde obliquis; inflorescentia 
terminalis paniculata breviter pedunculata, panicula spiciformi ca. 



FLORA OF COSTA RICA 1373 

20 cm. longa vix 2 cm. lata solemniter interrupta, floribus in cymulas 
parvas paucifloras sessiles dispositis sessilibus vel usque ad 3 mm. 
longe pedicellatis, bracteis parvis angustis inconspicuis, rhachi dense 
subadpresso-pilosa; hypanthium dense adpresso-hispidulum; calycis 
lobi 4 subulati usque triangulares vix ultra 1 mm. longi; corolla extus 
dense hispidulo-pilosa, tubo crassiusculo 4 mm. longo fauce nudo, 
lobis 4 late ovalibus ca. 3.5 mm. longis. La Hondura de San Jose", 
1,300 meters, August, 1933, Manuel Valeria 769 (type in Herb. Field 
Mus.), 780. Similar in appearance to the Guatemalan R. rufescens 
Robinson, which, however, differs in almost all details of pubescence 
and flower. 

Rondeletia Torresii Standl. Journ. Wash. Acad. Sci. 18: 163. 
1928. Wet forest, Viento Fresco, Prov. Alajuela, 1,800 meters, 
Standley & Torres 47839. Frequent in mountain forests of the central 
region; region of San Ramon; 1,000-2,000 meters. Endemic. A 
slender shrub of 2-3.5 meters; stipules triangular, cuspidate-attenu- 
ate, 3 mm. long; leaves on slender petioles 1-3 cm. long, lance-oblong 
to ovate-oblong or elliptic, 8-12 cm. long, abruptly acuminate or 
long-acuminate, subobtuse to acute at the base, above green, sparsely 
puberulent on the costa and often sparsely short-pilose elsewhere, 
puberulent beneath on the nerves, short-barbate in the axils of the 
nerves; inflorescence terminal, cymose-corymbose, long-pedunculate, 
lax; calyx lobes unequal, 3 of them linear or subulate and 1.5-2.5 mm. 
long, the fourth elliptic to ovate, obtuse, 4-5 mm. long; corolla white, 
densely short-strigillose, naked in the throat, the tube 13-14 mm. 
long, the 4 lobes 3 mm. long; capsule 6-7 mm. long. 

RUDGEA Salisb. 

Shrubs or small trees, most often glabrous or nearly so; stipules 
bearing on the margins or at the apex or on the dorsal surface subu- 
late or aculeoliform teeth, these commonly pale and somewhat carti- 
laginous, the stipules sometimes laciniate; leaves opposite, short- 
petiolate or sessile; inflorescence terminal; calyx segments almost 
free or variously united; corolla salverform or funnelform, the lobes 
valvate; fruit baccate, containing 2 one-seeded nutlets. Probably 
no other species are known from Central America, but there is some 
uncertainty regarding division of the material into species. 

Rudgea cornifolia (Humb. & Bonpl.) Standl. R. fimbriata 
Standl. Apparently frequent in forests of the region of San Ram6n; 
mountains of Guanacaste; Rio Naranjo; chiefly at 600-900 meters. 
Extending to Bolivia and Brazil, and probably to southern Mexico. 



1374 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

A glabrous shrub of 2-4 meters; stipules deltoid-ovate, incised - 
laciniate; leaves sessile or nearly so, elliptic to elliptic-oblong, 8-15 
cm. long, acuminate, commonly narrowed to an obtuse or narrowly 
rounded base; inflorescence usually small, cymose-paniculate, broad, 
many-flowered, rather open, the flowers sessile or nearly so; calyx 
short, minutely 5-dentate; corolla white, 5-7 mm. long, the lobes 
longer than the tube; fruit ellipsoid, 5-8 mm. long (when fresh 
probably much larger), white. It is probable that R. ceratopetala 
Donn. Smith, described from Guatemala, is synonymous with 
R. cornifolia. 

Rudgea Skutchii Standl., sp. nov. Frutex 3-metralis omnino 
glaber, ramis gracilibus, novellis olivaceis plus minusve compressis, 
internodiis elongatis; stipulae diutius persistentes 7-8 mm. longae 
ovales coriaceae fere liberae apice obtusissimae et fasciculo setularum 
numerosarum brevium incrassatarum onustae; folia mediocria 
brevissime petiolata papyracea, petiolo 2-5 mm. longo; lamina 
elliptico-oblonga vel lanceolato-oblonga 8-10 cm. longa 3-4 cm. 
lata, subabrupte in acumen anguste longiattenuatum ad 2.5 cm. 
longum contracta, basin versus angustata, basi ipsa anguste obtusata, 
supra in sicco viridis, nervis pallidis, subtus fere concolor lucida, 
costa gracili elevata, nervis lateralibus utroque latere ca. 7 remotis 
prominentibus arcuatis prope marginem arcuato-conjunctis, venulis 
prominulis laxe reticulatis; inflorescentia terminalis graciliter 3.5-5 
cm. longe pedunculata cymoso-paniculata, basi trichotoma, laxe 
pauciflora, ca. 3 cm. longa atque 5 cm. lata, ramis primariis angulo 
fere recto divergentibus gracilibus rigidis viridibus, cymulis paucis 
trifloris, flore centrali sessili, lateralibus ad 1 cm. longe pedicellatis, 
bracteis obsoletis; hypanthium obconicum 2 mm. longum, calyce 
viridi 3-3.5 mm. longo primo subtruncato serius irregulariter breviter 
lobato; corolla alba glabra, tubo crasso cylindraceo 5 mm. longo, 
lobis oblongis patentibus obtusis tubo fere aequilongis. In forest, 
vicinity of El General, Prov. San Jose", 915 meters, Alexander F. 
Skutch 2836 (type in U. S. Nat. Herb.; photo in herb. Field Mus.). 

Rudgea thyrsiflora Donn. Smith, Bot. Gaz. 61: 375. 1916. 
Forests of Tsaki, Talamanca, 200 meters, Tonduz 9579. Collected 
also by Hoffmann (No. 767} at San Miguel. Endemic. Branches 
ferruginous-pubescent; stipules connate into a sheath, this bisetose 
on each side, aculeoliferous between the setae; leaves short-petiolate, 
thin, lance-elliptic, 19-23 cm. long, 6-8 cm. wide, narrowly long- 
acuminate, acute or acuminate at the base, glabrous; inflorescence 
thyrsiform, dense, ferruginous-pubescent, pedunculate, 5.5 cm. long, 



FLORA OF COSTA RICA 1375 

1.5 cm. wide; calyx lobes ovate; corolla sparsely ferruginous-strigil- 
lose, the tube 2 mm. long, the lobes corniculate at the apex. I have 
not seen material of this species recently, and have available only 
a photograph of the type (which does not show the inflorescence) and 
a leaf from another collection. I am rather doubtful that the plant 
belongs to this genus. 

RUSTIA Klotzsch 

Glabrous shrubs or trees, the branchlets terete or obtusely tetra- 
gonous; leaves large, opposite, petiolate; stipules large, caducous; 
flowers in terminal panicles; hypanthium campanulate or turbinate; 
calyx short, 5-dentate or 5-lobate, persistent; corolla funnelform or 
campanulate, the throat glabrous or pilose, the 5 lobes valvate, 
short; ovary 2-celled, many-ovulate; capsule thick-coriaceous, oblong- 
ovoid or clavate, loculicidally bivalvate; seeds minute, horizontal. 
Another species is known from Panama. 

Rustia occidentalis (Benth.) Hemsl. Cocos Island. Nicaragua 
to Colombia; reported from Guatemala, but the report probably is 
based upon a Nicaraguan specimen. A glabrous shrub or small tree; 
stipules lance- triangular, 5-8 mm. long; petioles 1-2.5 cm. long, 
the blades oblanceolate, 10-19 cm. long, 2.5-5 cm. wide, acuminate 
to attenuate, attenuate to the base, subcoriaceous; panicles peduncu- 
late, narrowly pyramidal, few-flowered, 6-9 cm. long, the pedicels 13 
mm. long or less; calyx 1 mm. long, obscurely dentate; corolla salver- 
form, pale violet, the tube 6-7 mm. long, the lobes ovate-oblong, 
obtuse, 5-6 mm. long; capsule rounded-obovoid, 1 cm. long, 5-8 mm. 
wide, dark brown. On Cocos Island the tree is reported to grow 
along streams, and to reach a height of 15 meters. 

SABICEA Aubl. 

Reference: Wernham, A monograph of the genus Sabicea, 1914. 

Plants ligneous or suffrutescent, usually scandent, pubescent; 
leaves opposite; stipules persistent, erect or often reflexed; flowers 
usually small, capitate or cymose, axillary, bracteate; calyx 3-6- 
lobate, the lobes mostly narrow and elongate, persistent; corolla 
funnelform or salverform, the tube short or elongate, the throat 
villous, the 4-5 lobes short, valvate; stamens included; ovary com- 
monly 4-5-celled, many-ovulate; fruit baccate, 2-5-celled, the numer- 
ous seeds minute, ovoid or angulate. Perhaps two additional species 
occur in Central America. 

Sabicea costaricensis Wernham, Monogr. Sabicea 31. 1914. 
Buenos Aires, in hedges, Pittier 6712. Occasional in thickets of 



1376 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the tierra caliente. Endemic (?). Suffrutescent, the young branches 
densely yellowish-sericeous; leaves petiolate, elliptic to oblong, 12 
cm. long and 4.5 cm. wide or usually smaller, acuminate, obtuse to 
subcordate at the base, strigose beneath; inflorescence rather lax, 
2 cm. broad or larger, the flowers conspicuously pedicellate; corolla 
sparsely strigose, the tube 8 mm. long, the lobes 2 mm. long; fruit 
greenish red. 

Sabicea panamensis Wernham, described from Panama and 
reported also from Guatemala, is probably identical with S. costari- 
censis. Wernham described the former as an erect shrub, but all 
the Central American Sabiceas, as I have seen them, are elongate 
vines when well developed, and there is no reason to suppose that 
so closely related a plant is different in habit from S. costaricensis. 

Sabicea villosa Roem. & Schult. S. hirsuta HBK. Thickets of 
the Atlantic tierra caliente. Widely distributed in tropical America. 
Similar in habit and general appearance to the preceding species; 
stems densely hirsute with spreading hairs; leaves 5-12 cm. long; 
flowers capitate, the heads sessile, few-flowered; corolla white, hirsute; 
fruit deep purple, about 1 cm. long, juicy. 

Sabicea villosa var. adpressa (Wernham) Standl. Thickets of 
the tierra caliente; known from both coasts. Distributed like the 
typical form of the species. Distinguished by having the pubescence 
of the stems and corollas of closely appressed rather than spread- 
ing hairs. 

SHERARDIA L. 

Low annuals; leaves verticillate, narrow; flowers small, subsessile, 
in terminal and axillary, involucrate heads; calyx segments 4-6, 
persistent; corolla funnelform, the tube equaling or longer than the 
lobes; stamens exserted; fruit 2-celled, didymous, the carpels 1-seeded, 
indehiscent. No species of the genus is native in America. 

Sherardia arvensis L. Common in potreros on the upper slopes 
of Turrialba and Irazu, doubtless introduced with grass seed. Native 
of Europe. Stems hispidulous, short or elongate and branched; 
leaves in whorls of 4-6, obovate to lanceolate or linear, acute and 
mucronate, 6-15 mm. long, ciliate; flowers heads pedunculate; corolla 
pink or bluish. An inconspicuous plant, much like Galium in habit 
and general appearance. 

SICKINGIA Willd. 

Trees or shrubs; stipules often large; leaves opposite; flowers 
small or medium-sized, paniculate, 4-5-parted; calyx cupular or 



FLORA OF COSTA RICA 1377 

campanulate, truncate or dentate; corolla tubular or funnelform, 
usually pilose at the insertion of the stamens, the lobes short, imbri- 
cate or open in bud ; stamens exserted ; capsule usually globose, often 
very large, 2-celled, bivalvate; seeds large, horizontal, broadly winged. 
-Two other species, at least, occur in Central America. The trees 
have fine-grained wood that is pink or red when freshly cut, but the 
handsome color, unfortunately, disappears in age. 

Sickingia Maxonii Standl. Inkwood. Collected only at Monte 
Verde, Atlantic tierra caliente, by Stork. Panama. A tree of 9-12 
meters, with a spreading crown, the trunk 12-25 cm. in diameter; 
leaves very large, sometimes 60 cm. long and 40 cm. wide, but 
mostly smaller, sessile or short-petiolate, somewhat reddish when 
dried, obovate or rhombic-obovate, short-acuminate, narrowed to 
a rounded or sub truncate base, somewhat puberulent when young; 
inflorescence rather small and dense; corolla coriaceous, narrowly 
campanulate, 7-8 mm. long; fruit globose, the walls very thick and 
hard, about 6 cm. in diameter; seeds 1.5-2.5 cm. long. Called Ink- 
wood by the English-speaking people of the Atlantic coast. Known 
in Panama as Guaiatil, Guaiatil Colorado, and Jagua de montana. 

SOMMERA Schlecht. 

Shrubs or small trees, more or less pubescent, the branchlets 
terete; leaves opposite, petiolate, membranaceous, conspicuously 
striolate-lineolate between the veins; stipules large, thin, caducous; 
flowers small, white, in axillary, pedunculate cymes, corymbs, or 
racemes; calyx 4-5-lobate, the lobes subfoliaceous, persistent; corolla 
funnelform or subcampanulate, sericeous, the throat villous, the 
4-5 lobes short or elongate, valvate; ovary 2-celled, many-ovulate; 
fruit baccate, globose or ovoid; seeds numerous, minute, obtusely 
angulate. One other species is found in Guatemala. 

Sommera grand is (Bartl.) Standl. S. Donnell-Smithii Standl. 
Contr. U. S. Nat. Herb. 17: 436. 1914 (near Alajuelita, Prov. San 
Jose", 990 meters, J. D. Smith 4771). S. mesochora Standl. Frequent 
in mountain forests of the central region; Dota and San Ramon; 
chiefly at 1,000-1,600 meters, but collected also in the Changuinola 
Valley, at a much lower elevation. Panama to Mexico. A shrub or 
tree, sometimes 8 meters high, with a dense crown; stipules 1.5-3.5 
cm. long; petioles 1-2.5 cm. long, the blades obovate to rhombic- 
obovate or oblanceolate-oblong, 10-20 cm. long, abruptly acuminate, 
rounded to attenuate at the base, minutely and sparsely appressed- 
pilose above, sparsely or densely pilose beneath with chiefly appressed 



1378 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

hairs; inflorescences usually many-flowered, short-pedunculate, the 
pedicels 8 mm. long or less; calyx lobes suborbicular to ovate-oblong, 
rounded or obtuse, 2.5-4 mm. long; corolla white, sericeous, 5-8 mm. 
long; fruit subglobose, almost 1 cm. in diameter. Noteworthy for 
the fine striation of the leaf tissue. 

SPERMACOCE L. 

Erect or spreading annuals; stipules connate with the petioles 
to form a setiferous sheath; leaves opposite, herbaceous; flowers 
minute, densely fasciculate in the leaf axils, 4-parted; sepals short- 
connate; corolla funnelform, the lobes valvate; stamens attached 
at the base of the corolla tube, the anthers versatile; fruit dry, of 
2 one-seeded carpels, these coherent at the base, one carpel opening, 
the other remaining closed. At least two other species are found 
in Central America. 

Spermacoce glabra Michx. Waste or brushy places about the 
Meseta Central, and in the Atlantic tierra caliente. Generally 
distributed in tropical America. Plants glabrous throughout or 
nearly so, erect or procumbent, rarely 60 cm. high; leaves short- 
petiolate, lanceolate or linear-lanceolate, 3-6 cm. long, acuminate; 
inflorescences many-flowered, one-sided; sepals triangular, 1 mm. 
long; corolla white, 2 mm. long; capsule 3 mm. long. One other 
species, S. confusa Rendle (S. tenuior of authors, not L.) is almost 
certainly to be found in Costa Rica, but I find no specimens or 
records. It is distinguished by having pubescent capsules, and the 
leaves are scabrous on the upper surface. Both species are incon- 
spicuous weeds. 

TOCOYENA Aubl. 

Unarmed shrubs or small trees; stipules mostly caducous, glan- 
dular within; leaves opposite, petiolate; flowers large and showy, 
terminal, cymose, perfect, 4-6-parted; calyx cupular, dentate, glan- 
dular within; corolla funnelform or salverform, the tube slender 
and greatly elongate, glabrous or pubescent in the throat, the lobes 
contorted, obtuse or acute; fruit baccate, 2-celled, globose to oblong; 
seeds numerous, large, compressed, horizontal. One other Central 
American species is native in Panama. 

Tocoyena obliquinervia Standl. Contr. Arnold Arb. 5: 152. 
1933. Posoqueria obliquinervia Standl. Journ. Wash. Acad. Sci. 
18: 167. 1928. Forests of Rio Naranjo, 200-250 meters, Tonduz 
9528. Known only from the original material. Branchlets glabrous; 
petioles 1-1.5 cm. long, the blades cuneate-obovate to oblanceolate- 



FLORA OF COSTA RICA 1379 

oblong, 20-28 cm. long, 8-13 cm. wide, rounded and abruptly short- 
acute at the apex, cuneately long-attenuate to the base, short-barbate 
beneath in the axils of the nerves, otherwise glabrous; inflorescence 
terminal, corymbiform, dense, many-flowered, glabrous; calyx 2 mm. 
long, shallowly lobate; corolla tube (in bud) 12-18 mm. long, the 
limb in bud globose-ovoid, 7 mm. long, obtuse. 

UNCARIA Schreb. 

Scandent shrubs, glabrous or pubescent, climbing by means of 
stout, uncinate spines, these formed from abortive peduncles; leaves 
opposite, coriaceous, short-petiolate; stipules entire or bifid; flowers 
small, yellowish white, usually pubescent, in dense, globose heads, 
these axillary and solitary or paniculate; corolla tubular-funnelform, 
the 5 lobes valvate; ovary 2-celled, the ovules numerous; fruit 
capsular, elongate, septicidally bi valvate; seeds produced at each 
end into a wing. A single species is native in Central America. The 
flower heads are similar to those of the related genus Cephalanthus 
found farther north, in Mexico and the United States. 

Uncaria tomentosa (Willd.) DC. Rangayo. Thickets and 
swamps of the Atlantic tierra caliente. Guatemala to the Guianas. 
A large, woody vine; stipules 6-11 mm. long, obtuse or rounded; 
leaves oval or ovate-oval, 10-15 cm. long, obtuse-acuminate, rounded 
or cordate at the base, glabrous above, whitish-tomentose or glabrate 
beneath; heads 6 mm. in diameter (excluding the corollas); corolla 
6 mm. long; capsule trigonous, 2-3 mm. long. The plant is a trouble- 
some weed in banana plantations of the Atlantic coast. 

WARSZEWICZIA Klotzsch 

Trees or shrubs, more or less pubescent; stipules large, persistent; 
leaves opposite, petiolate, large, membranaceous or coriaceous; 
inflorescence terminal, the small flowers 5-parted, cymose, the cymes 
arranged in racemiform panicles; calyx lobes persistent, one of them 
dilated into a large, foliaceous, petiolate, bright-colored limb; corolla 
short-funnelform, villous in the throat, the lobes obtuse, imbricate 
in bud; anthers exserted; ovary 2-celled, many-ovulate; capsule 
small, oblong or globose, septicidally bi valvate; seeds minute, 
horizontal, marginate, reticulate. A single species is found in North 
America. 

Warszewiczia coccinea (Vahl) Klotzsch. Forests of the 
Atlantic tierra caliente. Extending to Peru and Brazil. A shrub 
or tree, sometimes 12 meters high; stipules 1.5-2 cm. long, narrowly 



1380 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

triangular-oblong, acuminate or attenuate; petioles 2.5 cm. long or 
less, the blades oblanceolate-oblong to oval-obovate, mostly 20-50 
cm. long, acute or short-acuminate, sparsely pubescent or almost 
glabrous; inflorescence 30-40 cm. long or larger, narrow; expanded 
calyx lobe with a bright red, oblong limb 3.5-7 cm. long; corolla 
yellow or orange, 5-8 mm. long; capsule 4-5 mm. long. This is one 
of the most showy and handsome plants of Costa Rica, but it is 
of apparently infrequent occurrence here. 

XEROCOCCUS Oerst. 

Low, simple shrubs; leaves opposite, large, petiolate; flowers 
numerous, forming large, dense, sessile heads in the leaf axils, the 
bracts and calyces dark red; calyx much exceeding the hypanthium, 
4-lobate, the lobes linear-elongate, ciliate, persistent; corolla short, 
tubular, pubescent, the throat glabrous, the 4 lobes short, valvate; 
anthers included; ovary 2-celled, many-ovulate; fruit small, baccate, 
2-celled, when dry separating into 2 cocci; seeds numerous, angulate, 
reticulate. The genus consists of a single species. 

Xerococcus congestus Oerst. Vid. Medd. Kjoebenhavn 52. 
1852. Turrialba, 900 meters, Oersted 11776 (photo, of type in Herb. 
Field Mus.). Frequent in mountain forests of the central region, 
mostly at 900-2,400 meters; region of San Ramon. Endemic. A 
simple shrub, sometimes as much as 3.5 meters high, but usually 
much lower, and often less than a meter in height; leaves long-petio- 
late, broadly ovate or elliptic, as much as 30 cm. long, sparsely 
villous or glabrate; flower clusters as much as 2.5 cm. in diameter, 
dark red; calyx lobes about 8 mm. long; corolla 8-10 mm. long; 
fruit white, 4 mm. long. When a part of the Rubiaceae of the 
North American Flora was published, in 1921, this plant was still 
known only from Oersted's type, and it is only in recent years that 
a considerable quantity of herbarium material has accumulated. In 
fact, only the writer seems to have had much success in finding the 
plant, although it grows abundantly in many forested areas through- 
out the central mountain region. It is such a showy and handsome 
plant that it is hard to understand how it could escape the attention 
of other collectors. 

CAPRIFOLIACEAE. Honeysuckle Family 

Shrubs or trees, sometimes woody vines; leaves opposite, simple 

or compound, entire or dentate, without stipules; flowers perfect, 

usually cymose; calyx tube adnate to the ovary, the limb 3-5-dentate 

or 3-5-lobate; corolla gamopetalous, usually white, the limb 5-lobate, 



FLORA OF COSTA RICA 1381 

sometimes bilabiate; stamens normally 5, inserted on the corolla 
tube alternate with the lobes, the anthers versatile; ovary inferior, 
1-6-celled, the style slender, the stigma capitate or 2-5-lobate; fruit 
baccate or drupaceous in the Central American genera. No other 
genera are known from Central America. 

LONICERA L. Honeysuckle 

Shrubs, erect or scandent; leaves entire, those of young shoots 
sometimes lobate; flowers variously arranged, often accompanied 
by large, leaflike bracts; corolla irregular, the limb more or less 
bilabiate; fruit baccate. No species are native in Central America. 

Lonicera japonica Thunb. Madreselva. A common ornamental 
plant of gardens. Native of eastern Asia. A slender vine with very 
fragrant flowers; corolla at first white or pink, soon turning yellow; 
berries black. The Japanese honeysuckle is a popular ornamental 
plant in most parts of Central America, especially above the tierra 
caliente. 

SAMBUCUSL. Elder 

Shrubs or small trees, the branches with abundant pith; leaves 
pinnate or bipinnate, the leaflets serrate or laciniate; flowers small, 
white, in usually depressed and broad cymes; calyx tube ovoid or 
turbinate, the limb dentate; corolla rotate or nearly so, regular, 
3-5-lobate; ovary 3-5-celled; fruit small, drupaceous, berry-like, 
containing 3-5 one-seeded nutlets. No other species are known 
from Central America. 

Sambucus mexicana Presl. Sauco. Occasional in forest or 
hedges of the central region, chiefly at 900-1,600 meters; region 
of Dota; often seen in cultivation. Ranging northward to south- 
western United States. Usually a shrub but often a tree as much 
as 6 meters high ; leaflets mostly 5-7, oblong or lanceolate, acuminate, 
rather coarsely appressed-serrate, setulose-pilose beneath along the 
costa, otherwise generally glabrous or nearly so; flowers fragrant, 
in small and dense or broad and lax cymes; ripe fruit almost black. 
Most of the Central American specimens of this species are from 
cultivated plants, but this species is apparently native in some parts 
of Costa Rica. It is also planted, partly for ornament and partly 
for its flowers, whose infusion is much used in domestic medicine. 
The juicy, agreeably flavored fruits are edible, like those of other 
species of the genus. The typical form of S. mexicana, rare in Costa 
Rica, has pinnate leaves. The common form is var. bipinnata 



1382 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

(Schlecht. & Cham.) Schwerin, in which the lowest leaflets of each 
leaf are replaced each by 2 or 3 sessile leaflets. 

Sambucus oreopola Donn. Smith, Bot. Gaz. 25: 146. 1898. 
Sauco. S. canadensis L. var. oreopola Rehder, Trees & Shrubs 2: 
188. 1911. Forests of Rancho Flores, Volcan de Barba, Tonduz 
2107. Frequent in forest or clearings on upper slopes of the central 
volcanoes, 1,800-2,400 meters; Zarcero. Endemic. A shrub, or 
sometimes a tree of 9 meters; bark light brown with green streaks, 
the wood pale yellow; leaflets 9-11, averaging larger than those 
of S. mexicana and much more finely and closely serrate, pubescent 
or almost completely glabrous; cymes very broad and rather lax; 
fruit red, becoming almost black. Both Rehder and Schwerin con- 
sider this merely a variety of S. canadensis, a common species of the 
United States. In a genus in which the species, for the most part, 
are so notoriously "feeble," it would seem that this Costa Rican 
plant, far removed from the nearest region in which true S. canadensis 
is known to occur, might well be given specific rank. Personally 
I believe that it has quite as good characters upon which to base 
such a separation as have most of the species maintained or described 
by the two authors mentioned. 

VIBURNUM L. 

Reference: C. V. Morton, The Mexican and Central American 
species of Viburnum, Contr. U. S. Nat. Herb. 26: 339-366. 1933. 

Trees or shrubs; leaves entire or dentate; flowers small, white, 
cymose; calyx lobes 5; corolla campanulate or rotate, 5-lobate; 
ovary 1-celled, 1-ovulate; fruit a red or black drupe. Other species 
occur in Central America. 

Viburnum costaricanum (Oerst.) Hemsl. Biol. Centr. Amer. 
Bot. 2: 2. 1881. Conchudo. Oreinotinus costaricanus Oerst. Vid. 
Medd. Kjoebenhavn 1860: 290. 1861. Volcan de Irazu, 2,700 
meters, Oersted 7808. Frequent in forests of the mountains, mostly 
at 1,100-3,000 meters. Adjacent Panama. A large shrub or a tree, 
sometimes 22 meters high, with a trunk 75 cm. in diameter, the bark 
scaly, rusty brown; leaves opposite or ternate, petiolate, elliptic to 
oblanceolate or obovate, 12 cm. long and 4.5 cm. wide or smaller, 
acute or acuminate, cuneate at the base, entire, glabrate above, 
sparsely stellate-pubescent beneath or glabrate; flowers sweet- 
scented; calyx tube glabrous, sparsely red-glandular; fruit black at 
maturity, 4-8 mm. long. 



FLORA OF COSTA RICA 1383 

Viburnum stellate- tomentosum (Oerst.) Hemsl. Biol. Centr. 
Amer. Bot. 2: 3. 1881. Tirra, Curd, Sura. Oreinotinus stellato- 
tomentosus Oerst. Vid. Medd. Kjoebenhavn 1860: 283. 1861. 0. 
Wendlandii Oerst. loc. cit. (Volcan de Irazu, 2,600 meters, Wend- 
land 657). V. Wendlandii Hemsl. loc. cit. V. stellato-pilosum Polak. 
Linnaea 41: 564. 1877 (forests near Barba, Polakowsky 31+9}. Volcan 
de Irazu, 3,000 meters, Oersted 7818. Frequent in forest and thickets 
of the central region, chiefly at 1,000-2,700 meters. Adjacent 
Panama. A shrub or tree of 2-8 meters; leaves opposite, short- 
petiolate, ovate to elliptic or obovate, acute or short-acuminate, 
obtuse to rounded at the base, usually conspicuously dentate, densely 
stellate- tomentose; fruit black, 5-6 mm. long. The fruit, like that 
of other species, is sweet and edible. 

Viburnum venustum Morton, Contr. U. S. Nat. Herb. 26: 
350. 1933. Cerros de Zurqui, northeast of San Isidro, Prov. Heredia, 
2,000-2,400 meters, Standley & Valeria 50545. Frequent in forest 
on the slopes of Barba and Poas; Palmira; at 1,500-2,600 meters. 
Endemic. A large shrub or a tree of 9 meters, the trunk to 15 cm. 
in diameter, the bark cinnamon-brown; leaves short-petiolate, ovate 
or broadly ovate, 15 cm. long or less, abruptly long-acuminate, 
rounded or truncate at the base, entire or rarely serrate, barbate 
beneath in the axils of the nerves, otherwise glabrous or nearly so; 
fruit ovoid, 6 mm. long, black. The wood is salmon-red when first cut. 

VALERIANACEAE. Valerian Family 
VALERIANAL. 

Perennial herbs, sometimes scandent, the roots with a strong, 
distinctive odor; leaves opposite, simple or compound, without 
stipules; flowers small, paniculate or cymose, perfect or polygamo- 
dioecious; calyx limb represented by 5-15 setiform, plumose teeth; 
corolla gamopetalous, 5-lobulate; stamens usually 3; ovary inferior; 
fruit indehiscent, dry, compressed, 1-celled, 1-seeded. Other species 
are native in northern Central America. 

Valeriana Candolleana Gardner. V. Mikaniae Lindl. Common 
in forests and thickets of the temperate region, and in Guanacaste, 
at 600-1,800 meters; region of San Ramon. Mexico to Brazil. 
Stems elongate and scandent, glabrous; petioles long, the blades 
cordate, undulate-dentate, narrowly acuminate; flowers white or 
greenish. Some authors treat this plant as a mere variety of V. 
scandens, but it appears to be a perfectly good species. 



1384 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Valeriana longifolia HBK. Thickets in the upper region of 
Volcan de Turrialba. Colombia. Plants erect, almost glabrous, 
as much as a meter high; leaves chiefly basal, linear or lanceolate, 
entire; flowers tinged with pink, arranged in a large, terminal panicle. 

Valeriana prionophylla Standl., sp. nov. Herba perennis 
erecta ut videtur simplex atque 1 m. alta vel ultra, caule subfistuloso 
1 cm. crasso et ultra striato pallide viridi, internodiis valde elongatis 
bisulcatis, tan turn in sulcis sparse setuloso-pilosis; folia basalia ca. 
26 cm. longa lineari-oblonga, lamina 13-20 cm. longa 1.5-2.5 cm. 
lata obtusa vel acuta basin versus longiattenuata, grosse dentata, 
petiolis latis vaginantibus, foliis supra glabris, subtus sparse setuloso- 
pilosis; folia caulina 2-3 paria sessilia lineari-lanceolata 9-25 cm. 
longa plerumque 1.5-2 cm. lata, superiora basi paullo dilatata et 
subamplexicaulia, omnia versus apicem subobtusum angustata, 
grosse serrata vel saepe inciso-serrata; inflorescentia terminalis atque 
corymbosa longipedunculata ca. 15 cm. longa basi trichotoma, 
cymis longissime pedunculatis compactis saepe ex axillis foliorum 
superiorum nascentibus, floribus dense aggregatis sessilibus, bracteis 
lineari-lanceolatis usque 6 mm. longis, ramis saltern ad nodos dense 
albido-barbatis; corolla pallide violacea 2.5 mm. longa glabra, lobis 
tubo fere aequilongis subovalibus apice rotundatis; antherae breviter 
exsertae. Cerro de La Muerte, 3,000 meters, in swampy places 
beside streams, June, 1932, H. E. Stork 3040 (type in Herb. Field 
Mus.). Volcan de Irazu, 3,150 meters, northeastern slope, only a 
few plants observed among shrubbery, Stork 2897. The species is 
well marked, among Central American ones at least, by the very 
long and narrow, coarsely serrate or dentate leaves. 

Valeriana scandens L. Reported from Atirro, and probably 
to be found in various parts of the tierra caliente. Widely distributed 
in tropical America. Plants scandent, glabrous; leaves compound, 
the 3 leaflets broadly ovate. 

Valeriana scorpioides DC. Common in pastures and forests 
of the temperate region, 1,000-1,800 meters; region of San Ramon. 
Ranging to Mexico. Plants erect, perennial or annual, simple or 
sparsely branched, pubescent; leaves small, cordate-ovate, acute or 
obtuse, undulate or crenate; flowers very small, white. 

Valeriana sorbifolia HBK. Occasional in forests of the central 
region; region of San Ramon; chiefly at 1,100-2,400 meters. Extend- 
ing to Mexico. Plants erect, almost glabrous; leaves pinnate, the 






FLORA OF COSTA RICA 1385 



few leaflets serrate or dentate. It is possible that there may be 
referable to this species a collection from Cerro de Las Vueltas, at 
3,000 meters, reported under the name V. affinis Mart. & Gal. 

DIPSACACEAE. Teasel Family 
All plants of the family are natives of the Old World. 

SCABIOSA L. 

Annual or perennial herbs; leaves opposite; flowers in peduncu- 
late, involucrate heads; bracts of the involucre distinct or slightly 
united, herbaceous; flowers crowded upon a receptacle, this bearing 
small scales, or naked; calyx limb 5-dentate; limb of the corolla 
4-5-cleft, oblique or bilabiate; stamens 4; fruit an achene, crowned 
by the persistent calyx. 

Scabiosa atropurpurea L. Bambali. S. maritime, L. Often 
grown in gardens for ornament. Native of southern Europe. An 
erect annual, about 60 cm. high; basal leaves lance-ovate, lyrate- 
lobate and coarsely dentate; cauline leaves pinnate-parted, with 
narrow lobes; flowers dark purple, pink, or white. There are many 
horticultural forms of the plant, varying in color and size of the 
flowers. 

CUCURBITACEAE. Gourd Family 

References: Cogniaux in DC. Monogr. Phan. 3: 325-978. 1881; 
Pflanzenreich IV. 275, I, II. 1916-1924. 

Herbaceous or very rarely woody vines, provided with coiled 
tendrils; leaves alternate, usually petiolate, often palmately lobate 
or dissected, without stipules; flowers monoecious or dioecious, small 
or large; calyx tube adnate to the ovary, the limb commonly 5-lobate, 
the petals normally 5, inserted on the limb of the calyx, distinct, 
or united to form a gamopetalous corolla; stamens mostly 3, some- 
times 1, 2 of them with 2-celled anthers, the other with a 1-celled 
anther, the filaments short, often united; ovary 1-3-celled, the style 
terminal, simple or lobate; fruit a pepo, but very variable as to form, 
indehiscent or rarely dehiscent at the apex, sometimes dry; seeds 
usually compressed, without endosperm. A few other genera are 
known from Central America. The family is still poorly understood 
in tropical America, chiefly because of the absence of adequate 
herbarium material. Large collections of well prepared specimens 
of the family are greatly to be desired. 



1386 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ANGURIA L. 

Reference: Cogniaux, Pflanzenreich IV. 275, I: 178. 1916. 

Scandent herbs, glabrous or nearly so; leaves entire, lobate or 
composed of 3-5 leaflets; flowers usually dioecious, medium-sized, 
racemose or spicate, red or orange; receptacle elongate and narrow, 
the sepals short and dentiform; stamens 2, the anthers linear or 
oblong, straight or replicate, the connective narrow, usually produced 
as an appendage. A few other species are known from Central 
America. 

Anguria Dunlapii Standl. Field Mus. Bot. 4: 298. 1929. Type 
collected in the Changuinola Valley, Panama, but doubtless occurring 
also in Costa Rica. Plants glabrous; leaves ovate-oblong, shortly 
cuspidate-apiculate, shallowly cordate at the base; flowers 8 mm. 
long, forming a long-pedunculate head. 

Anguria limonensis Pittier, Contr. U. S. Nat. Herb. 13: 119. 
/. 25. 1910. Between Limon and Moin, Pittier 16112. Forests and 
thickets of the Atlantic coast. Endemic. Leaflets 3, glabrous, ovate 
to lanceolate, acuminate, entire or somewhat sinuate, acuminate at 
the base; receptacle 8-10 mm. long; anthers straight, the appendage 
rounded and glabrous. 

Anguria longipedunculata Cogn. Forests of the Atlantic 
coast. Mexico. Leaves broadly oblong, acuminate, rounded at the 
base, glabrous, entire or somewhat denticulate; flowers spicate, the 
receptacle 8-10 mm. long; anthers straight, the appendage narrow, 
papillose. 

Anguria ovata Bonn. Smith, Bot. Gaz. 31: 112. 1901. Las 
Vueltas, Tucurrique, 635 meters, Tonduz 13006. Also Rio Yurquin. 
Endemic. Leaves broadly ovate, cuspidate-acuminate, rounded and 
shallowly retuse at the base, glabrous; flowers racemose, the pedicels 
1-2 mm. long; receptacle 12-13 mm. long; anthers straight, the 
appendage glabrous. 

Anguria pachyphylla Bonn. Smith, Bot. Gaz. 54: 236. 1912. 
Tuis, Prov. Cartago, 650 meters, Tonduz 11535. Also at Las Vueltas. 
Endemic. Leaves almost orbicular, glabrous, deeply emarginate at 
the base, shallowly or deeply trilobate, the lobes acuminate, entire 
or undulate; flowers spicate, the receptacle 10-12 mm. long; anthers 
straight, the appendage narrow, obtuse, glabrous. The leaves are 
coriaceous; in the other species they are membranaceous. 



FLORA OF COSTA RICA 1387 

Anguria plurilobata Cogn. Pflanzenreich IV. 275, I: 191. 1916. 
Rio Tuis, 600 meters, Tonduz 8175. Stems somewhat pilose; leaves 
glabrous, pinnate-lobate, the 7-9 lobes triangular; flowers spicate, 
the receptacle 5-6 mm. long; anthers straight, obtuse and muticous. 
Easy of recognition because of the pinnate-lobate leaves. 

Anguria Tonduzii Cogn. Pflanzenreich IV. 275, I: 191. 1916. 
Rio Corozal, Santo Domingo, Tonduz 1999. Leaflets 3, glabrous, 
long-petiolulate, acuminate, narrowed to the base; flowers spicate, 
the receptacle 10 mm. long; anthers straight, obtuse and muticous. 

Anguria Warscewiczii Hook. Frequent in forests of the tierra 
caliente; region of San Ramon, at 1,200 meters or less. Mexico to 
Venezuela. The species was based upon cultivated plants, probably 
of Costa Rican or Panaman origin. Leaves glabrous, the 3 leaflets 
petiolulate, acute or short-acuminate, entire or somewhat undulate 
or denticulate; flowers spicate, the receptacle 8-10 mm. long; anthers 
straight, the appendage narrow, glabrous. In some specimens there 
are found on the same branch trifoliolate leaves and others with 
simple, trilobate blades. 

CAYAPONIA Manso 

Scandent, more or less pubescent herbs; leaves commonly 3-7- 
lobate and rough; flowers monoecious, small or large, whitish, 
greenish, or yellowish, solitary, fasciculate, racemose, or paniculate; 
calyx of the staminate flower campanulate, with 5 teeth or lobes, 
the corolla campanulate or rotate; stamens 3, the anthers coherent; 
fruit globose or oval, indehiscent, 1-12-seeded. A few other species 
are known from Central America. 

Cayaponia americana (Lam.) Cogn. C. americana var. 
Oerstedii Cogn. in DC. Monogr. Phan. 3: 787. 1881 (Ujarras, Oersted 
3312; photo, of type in Herb. Field Mus.). Thickets and forests of 
the Atlantic tierra caliente. Nicaragua and West Indies. Leaves 
scabrous, 3-5-lobate, not decurrent at the base; flowers small, race- 
mose or paniculate; calyx tube 12-18 mm. long, glabrous; fruit oval, 
2.5-3.5 cm. long. 

Cayaponia attenuata (Hook & Arn.) Cogn. Frequent in 
thickets of the tierra caliente. Mexico to Panama. Leaves scabrous, 
decurrent at the base, trilobate, the upper ones sometimes entire; 
flowers paniculate; calyx tube 8-10 mm. long, 1.5-2 cm. in diameter. 

Cayaponia Austin-Smithii Standl., sp. nov. Scandens, cauli- 
bus crassiusculis sulcatis dense breviter villosulis, internodiis elonga- 



1388 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

tis; folia 2-5 cm. longe petiolata trifoliolata crassa; foliola breviter 
petiolulata, terminalia obovato-oblonga acuminata, basi acuta vel 
attenuata, lateralia lanceolato-oblonga basi valde obliqua, omnia 
5-12.5 cm. longa 1.5-6.5 cm. lata subdense serrato-denticulata, 
lateralia interdum prope basin subhastatolobata, supra viridia 
aspero-scabra, subtus vix pallidiora ubique dense submolliter breviter 
pilosa; cirrhi elongati breviter pilosi; flores mediocres in racemos 
breves paucifloros dispositi, racemis saepe paniculas angustas sub- 
nudas elongatas efformantibus, pedicellis plerumque 3-4 mm. longis 
dense breviter pilosulis; calyx anguste campanulatus 5 mm. longus 
pilosulus vel scaber basi rotundatus, dentibus ovato-triangularibus 
acutis 2.5-3 mm. longis; corolla 8 mm. longa extus puberula, intus 
tomentosa; fructus juvenilis globosus glaber fere 1 cm. diam. 
Zarcero, 1,350 meters, in forest, September, 1937, Austin Smith 
A446 (type in Herb. Field Mus.). The collector's notes furnish the 
following information: A vigorous vine, climbing over trees to a 
height of 8 meters; leaves rather stiff and rough; flowers open- 
campanulate, 2.5 cm. broad, the corolla inside pale yellow, externally 
yellow-green, with deep green striations. 

Cayaponia leucosticta Standl., sp. nov. Herba elongata 
scandens, caulibus gracilibus sulcatis glabris, internodiis valde 
elongatis; folia valde variabilia papyracea petiolata, inferiora fere 
ad basin triloba, lobo terminali elliptico vel oblongo-elliptico, later- 
alibus valde obliquis, omnibus abrupte subcuspidato-acuminatis; 
folia superiora elobata vel saepe breviter subhastato-lobata, ovato- 
deltoidea, plerumque 6.5-12 cm. longa atque 4-8 cm. lata, sub- 
abrupte longe anguste caudato-acuminata, basi vulgo truncata, 
interdum breviter late cordata, remote denticulata vel integra, supra 
tactu laevis dense maculis magnis albis conspersa glabra, subtus 
epunctata glabra; flores solitarii vel breviter racemosi, pedicellis 
crassiusculis glabris usque 1.5 cm. longis, in statu fructifero incras- 
satis; calyx magnus globosus ca. 1 cm. longus et fere aequilatus glaber 
truncatus, margine dentibus 5 subulatis vix ultra 1 mm. longis 
patentibus onusto; corolla (in alabastro tantum visa) extus densis- 
sime brunneo-sordido-tomentosa; fructus ovalis glaber ca. 2.5 cm. 
longus atque 1.5 cm. diam. La Palma de San Ramon, September, 
1928, Brenes 6336 (type in Herb. Field Mus.); also, from the same 
locality, Nos. 6251, 6332, 6321, 5948. Los Ayotes, near Tilaran, 
Guanacaste, 600 meters, wet forest, Standley & Valeria 4.5439. 

Cayaponia macrantha Pittier, Contr. U. S. Nat. Herb. 13: 118. 
1910. Las Vueltas de Tucurrique, Reventazon Valley, 635 meters, 



FLORA OF COSTA RICA 1389 

Tonduz 12840. Known only from the original collection. Stems 
slightly puberulent at the nodes, otherwise glabrous; leaves long- 
petiolate, the blades ovate in outline, deeply trilobate, 7-12 cm. long, 
attenuate and subemarginate at the base, scabrous above, pale 
beneath and minutely white-punctate, remotely dentate, the lobes 
ovate-lanceolate, the lateral ones oblique, often bilobate; staminate 
flowers long-pedunculate, the peduncles 4-6 cm. long; calyx cam- 
panulate, 18 mm. long, villous, the teeth 3-5 mm. long; corolla 
yellowish white, 4 cm. long, lanate within; fruit globose, 5 cm. in 
diameter, yellow with green stripes. 

Cayaponia microdonta Blake. Thickets at Pejivalle, Atlantic 
tierra caliente, 900 meters, and doubtless elsewhere. Ranging to 
British Honduras. A large or small, herbaceous vine, the slender 
branches rough; leaves long-petiolate, tuberculate and hispidulous or 
glabrate above, tuberculate, glanduliferous, and sparsely hispidulous 
beneath, trilobate to the middle or almost to the base, the lateral 
lobes again deeply lobate, the blade contracted and decurrent at the 
base, the lobes acute or acuminate, finely or coarsely dentate; calyx 
campanulate, 14 mm. long, the triangular teeth 1.5 mm. long; corolla 
cream-colored or greenish, 1.5 cm. long, viscid-puberulous; fruit 
globose, at maturity as much as 5 cm. in diameter. 

Cayaponia racemosa (Swartz) Cogn. Frequent in thickets of 
the Meseta Central and on the Pacific slope; region of San Ramon; 
at 1,300 meters or less. Mexico to northern South America and 
West Indies. A large or small vine, the stems glabrous or nearly so; 
leaves petiolate, almost glabrous, or often very scabrous, mostly 
3-5-lobate, abruptly long-decurrent at the base, the lobes obtuse or 
acute, dentate; calyx broadly campanulate, about 3 mm. long, 
rounded at the base, glabrous or nearly so, the teeth minute; corolla 
papillose outside, tomentose within, 4-5 mm. long; fruit oval, 16-18 
mm. long, 8-10 mm. wide, glabrous. Var. scaberrima Cogn. in DC. 
Monogr. Phan. 3: 769. 1881, was based in part upon Oersted 1 from 
San Jos and Oersted 49 from Ujarras. 

CITRULLUS Forsk. Watermelon 
The few species of the genus are natives of the Old World. 

Citrullus vulgaris Schrad. Sandia. The watermelon, native of 
Africa, is grown commonly in Costa Rica, as in most warmer parts 
of America. The watermelons of the tierra caliente are often large 
and of superior quality, but many of those produced, especially on 



1390 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the Pacific slope during the dry season, are small and inferior, largely 
because inferior seeds are planted. There are numerous local vari- 
eties, differing in shape and in color of the flesh. Plants sometimes 
are found in waste places, where the seeds have fallen by accident, 
but such plants rarely if ever produce fruits. 

CUCUMIS L. 

Annuals or perennials, scandent or prostrate; tendrils simple; 
flowers yellow, mostly small, monoecious, the staminate fasciculate 
or rarely solitary, the pistillate usually solitary; staminate calyx 
campanulate or turbinate, the lobes subulate, remote, the corolla 
rotate or subcampanulate; stamens 3, free; fruit variable, usually 
fleshy and indehiscent, many-seeded; seeds ovate or oblong, com- 
pressed, smooth. No other species are known in Central America. 

Cucumis Anguria L. Thickets in the lowlands of Guanacaste, 
and perhaps elsewhere on the Pacific coast. Widely distributed in 
tropical America. Plants annual, prostrate, hispid; leaves long- 
petiolate, broadly ovate to rounded in outline, 5-10 cm. long, deeply 
3-5-lobate, scabrous and hispidulous, very rough, the lobes mostly 
obovate; calyx 6 mm. long; corolla 1 cm. broad, the lobes ovate; 
fruit ellipsoid, yellow, 4-7 cm. long, densely covered with long, 
flexible prickles. 

Cucumis Melo L. Melon. Cantaloupe, Muskmelon. Native of 
the Old World tropics, this well known plant, so generally cultivated 
in the United States, is seldom planted in Central America. I do 
not know whether its scarcity there results from dislike of the fruit 
or from difficulty in propagation. 

Cucumis sativus L. Pepino. Cucumber. Grown commonly in 
gardens, as in most regions of the earth. Probably of East Indian 
origin. 

CUCURBITA L. 

Annual or perennial herbs, usually with rough pubescence; 
leaves cordate at the base; tendrils branched; flowers monoecious, 
large, yellow, the staminate solitary or fasciculate, the pistillate 
solitary; calyx campanulate, 5-lobate; corolla campanulate, 5-lobate 
to the middle; stamens 3, distinct, the anther sacs flexuous; fruit 
large, fleshy, indehiscent. No species are native in Central America, 
but one or more are found in Mexico. 

Cucurbita ficifolia Bouche". Chiverre, Chilacayote (Pittier). 
Planted commonly in the temperate regions, especially at rather 



FLORA OF COSTA RICA 1391 

high elevations; naturalized in thickets and forest in the region of 
Dota. Probably native of tropical Asia. A large, scandent vine, 
perennial; leaves subreniform, usually deeply 5-lobate but often 
only shallowly lobate, the lobes broadly rounded; calyx villous- 
hirsute, the lobes linear, slightly shorter than the tube; corolla 6-8 cm. 
long; fruit globose-ovoid, 20-30 cm. in diameter or larger, pale green, 
with white, fibrous flesh. The chiverre is one of the well known culti- 
vated plants of Costa Rica, and often figures in literature, or especi- 
ally in jokes. Often when one has made a long and hard trip on 
horseback to some finca high in the mountains, the owner welcomes 
his guest with a promise, in appreciation of his visit, to give him a 
chiverre when he returns home. The fruit looks much like a water- 
melon. Its flesh is used for preparing a variety of desserts. 

Cucurbita Pepo L. Ayote, Sapayo. A large number of varieties 
of squashes are grown in Costa Rica, as in other parts of Central 
America, and they constitute one of the favorite vegetables, being 
prepared for the table in various ways. These plants are of American 
origin, but they are no longer found in a wild state, an indication 
that their cultivation must be very ancient. Pumpkins like those 
grown in the United States I have never seen in Central America, 
most of the forms being rather of the squash type. The name ayote 
is of Mexican origin; sapayo is said by Pittier to be of Quechua 
derivation. Indigenous names reported by Pittier are: Pis (Cabe"- 
cara); Api (Bribri); Bat (Brunka). 

GYCLANTHERA Schrad. 

Reference: Pittier, Contr. U. S. Nat. Herb. 13: 120. 1910. 

Scandent herbs, usually glabrous or nearly so; tendrils simple 
or branched; leaves angulate to pedately compound; flowers usually 
very small, yellowish or white, monoecious, the staminate racemose 
or paniculate; calyx tube cupular or subrotate, the 5 teeth subulate, 
sometimes obsolete; corolla rotate, deeply 5-parted, the lobes gen- 
erally acute; stamens coalescent to form a column, the anthers con- 
nate into a head, usually 1-celled and annular; pistillate flowers 
solitary, inserted in the same axil with the staminate inflorescence; 
fruit obliquely ovoid, gibbous, fleshy, usually echinate, containing 
1 or more seeds. A few other species probably occur in Central 
America. The young fruits of some species, known in Costa Rica 
as Caifas, often are cooked and eaten. The young shoots or quelites 
also are used as a vegetable. 



1392 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Cyclanthera explodens Naud. Chanchitos (fruits). C. costari- 
censis Cogn. Diagn. Cucurb. 2: 73. 1877 (San Jose", Oersted 23; photo, 
of type in Herb. Field Mus.). C. costaricensis var. angustiloba Cogn. 
loc. cit. (Ujarras, 1,000 meters, Oersted 21+). Thickets of the Meseta 
Central, chiefly at 1,100-1,300 meters; region of Dota. Salvador 
to Ecuador. Stems more or less villous, at least at the nodes; leaves 
short-petiolate, the upper ones often almost sessile, 6-9 cm. long, 
shallowly or deeply trilobate, acuminate, shallowly or deeply cordate 
at the base, remotely denticulate, scaberulous-pubescent on both 
sides; staminate racemes usually short, the calyx tube glabrous, 
scarcely more than 1 mm. broad; corolla 1 mm. long; fruits on very 
short peduncles, usually 8-seeded, 2-2.5 cm. long, somewhat glau- 
cous, bearing a few short, thick, fleshy spines. The ripe fruits dehisce 
elastically when squeezed in the hand. Cogniaux and Pittier both 
maintained C. costaricensis as a distinct species, but the key characters 
are extremely feeble, and a photograph of the type specimen shows 
that it is exactly like the Costa Rican collections cited by Pittier 
under C. explodens. 

Cyclanthera Langaei Cogn. Frequent in thickets and forest 
of the tierra caliente, ascending the slopes of the mountains to 3,000 
meters. Ranging to Mexico. A slender, much branched vine, 
the stems villous at the nodes, elsewhere glabrous; leaves long- 
petiolate, 5-7-foliolate, the leaflets lanceolate or linear-lanceolate, 
5-10 cm. long or even larger, acutely dentate, acute, sparsely villous 
or glabrate; staminate racemes long-pedunculate, the pedicels fili- 
form; calyx tube 3 mm. long; corolla white, 2-2.5 mm. long; fruits 
ovoid or oblong, 2.5-3 cm. long, short-rostrate, densely aculeate, 
the peduncles 3-6 cm. long. C. Langaei subsp. gracillima Pittier 
(Contr. U. S. Nat. Herb. 13: 124. 1910; hedges at Ochomogo, 1,500 
meters, Tonduz 10904) is a form with large leaves and very slender 
stems. 

Cyclanthera Naudiniana Cogn. Slopes of Volcan de Barba, 
and region of Zarcero, 2,000-2,300 meters, in forest. Panama to 
Texas. Stems slender, glabrous; leaflets 3-5, usually petiolulate, 
lanceolate or oblong, mostly 3-5 cm. long, acuminate, dentate or 
lobulate, almost glabrous; staminate racemes or panicles sometimes 
longer than the leaves, the calyx tube 2 mm. broad; corolla green, 
1.5-2 mm. long; fruiting peduncles 1-2 cm. long; fruit ovoid-oblong, 
2-2.5 cm. long, densely short-setose. 

Cyclanthera pedata Schrad. Caifa, Pepino. Region of Cartago, 
but perhaps only in cultivation for its edible fruits. Mexico to 



FLORA OF COSTA RICA 1393 

Bolivia. Stems glabrous; leaflets 5-7, sessile or nearly so, lanceolate 
or oblong-lanceolate, 7-15 cm. long, acute, attenuate to the base, 
denticulate or crenulate, somewhat scaberulous; staminate inflores- 
cences paniculate, 10-20 cm. long, the pedicels filiform; calyx 
glabrous, the tube 3-4 mm. broad; corolla 1.5-2 mm. long; fruits 
oblong-lanceolate, 5-10 cm. long, long-attenuate, bearing a few 
short, fleshy spines toward the base, the peduncle very short and 
thick. Illustrated by Pittier, Contr. U. S. Nat. Herb. 13: pi. 18. 

Cyclanthera Tonduzii Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 
274. 1892. Hedges near San Jose", Pittier 1^9. In thickets or forest, 
Meseta Central and the Pacific slope; Guanacaste; region of San 
Ramon; Dota; at 2,400 meters or less. Reported also from Guate- 
mala. Plants slender and much branched, the stems puberulent at 
the nodes, otherwise glabrous; leaflets 5, scaberulous above, glabrous 
beneath, 4-12 cm. long, obtuse or acute, dentate and often lobate; 
staminate inflorescences slender, usually longer than the leaves, 
the calyx glabrous, 3 mm. broad; corolla white, 2 mm. long; fruiting 
peduncles 2-3 cm. long; fruits 2-2.5 cm. long, 1-1.5 cm. thick, 
densely setose. Illustrated by Pittier, Contr. U. S. Nat. Herb 13: 
f. 27, 28. 

ECHINOCYSTIS Torr. & Gray 

Scandent herbs, annual or perennial; leaves long-petiolate, usually 
5-7-angulate or lobate; tendrils 2-5-fid; flowers usually small and 
white, the staminate racemose or paniculate; calyx tube campanu- 
late, the 5 teeth subulate; corolla commonly rotate, deeply 5-parted, 
the segments oblong or linear, papillose; stamens 3, the filaments 
united to form a column, the anthers connate or free, the cells 
flexuous; pistillate flowers solitary or arising from the same axil as 
the staminate; fruit dry, 1-3-celled, densely covered with long and 
rather stout spines, opening at the apex by 1-2 pores, sometimes 
irregularly rupturing. Probably no other species occur in Central 
America. 

Echinocystis Coulteri (Gray) Cogn. Tacaquillo, Chayotillo. 
Echinopepon horridus Naud. Thickets or forest of the Meseta 
Central, chiefly at 1,000-1,200 meters. Extending to Mexico. A 
slender, much branched vine, the stems sparsely villous-hirsute; 
leaves angulate or shallowly 5-lobate, mostly 8-10 cm. long, acute 
or short-acuminate, deeply cordate at the base, sparsely villous- 
hirsute, remotely and minutely denticulate; staminate racemes longer 
than the leaves, 5-15-flowered, the pedicels 5-9 mm. long; calyx 



1394 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

sparsely villous, 7-8 mm. broad, the narrowly triangular teeth 2 mm. 
long; corolla sparsely villous, 6-7 mm. long; fruit oblong, 4 cm. long, 
2 cm. broad, long-rostrate, the spines 4-18 mm. long. 

ELATERIOPSIS Ernst 

Similar to Cyclanthera, but the stamens 5, the anthers connate 
into a head, the anther cells flexuous; in Cyclanthera there is a 
single anther; fruit smooth, unarmed. No other species are known 
from Central America. 

Elateriopsis Oerstedii (Cogn.) Pittier, Contr. U. S. Nat. 
Herb. 13: 125. pi 19; f. 31-3 b. 1910. Caifa, Chayotillo. Cyclanthera 
Oerstedii Cogn. in DC. Monogr. Phan. 3: 856. 1881. San Jose", Oer- 
sted 20 (photo, of Oersted 18, from Cerro de Catalina, one of the 
other specimens cited by Cogniaux, in Herb. Field Mus.). In thickets 
and forest, Meseta Central to the coasts, at 1,300 meters or less; 
Guanacaste. Endemic. Stems rather stout, 5-sulcate, glabrous or 
slightly pubescent; leaves long-petiolate, the petioles glabrous; 
blades 6-15 cm. long, angulate or shallowly trilobate, almost gla- 
brous, punctate above, cordate at the base, remotely denticulate; 
staminate racemes 5-20 cm. long, the pedicels 1-2 cm. long; calyx 
sparsely pubescent, 6-7 mm. broad, the teeth subulate; corolla green- 
ish white, pubescent outside, 5-6 mm. long; fruiting peduncles 2-3 
cm. long; fruit obliquely ovoid, 6-8 cm. long, 3-4 cm. thick, smooth, 
light green with darker stripes, 3-celled, the seeds several in each 
cell, flattened, brownish, 17 mm. long. E. Oerstedii var. Biolleyi 
Pittier (Contr. U. S. Nat. Herb. 13: 126. /. 81. 1910; Cyclanthera 
Oerstedii var. Biolleyi Cogn., ined.) is a form with slightly larger 
flowers, the corolla lobes being longer and narrower than in the type. 
The fruits are a rather common vegetable of central Costa Rica, and 
the plants are often cultivated. 

ELATERIUM Jacq. 

Slender, scandent herbs, glabrous or rather sparsely pubescent; 
leaves cordate, entire, lobate, or parted; tendrils simple or branched; 
flowers small, white or yellow, monoecious, the pedicels capillary; 
staminate flowers racemose, the calyx tube elongate-cylindric, narrow, 
the usually 5 lobes subulate or filiform, the corolla rotate, parted 
almost to the base; filaments connate into an elongate column, the 
anther cells linear, sigmoid-flexuous; pistillate flowers solitary; fruit 
obliquely ovoid, rostrate, gibbous, fleshy, 1-many-celled, at maturity 
rupturing elastically; seeds marginate, the margin usually crenulate. 
One or two additional species are found in Central America. 



FLORA OF COSTA RICA 1395 

Elaterium ciliatum Cogn. Chanchitos (fruits). Frequent in 
thickets and forest, Meseta Central to the Pacific tierra caliente; 
region of San Ramon; at 1,300 meters or less. Panama to Guate- 
mala. Leaves thin, the upper ones sessile or nearly so, mostly ovate- 
cordate or rounded and 6-8 cm. long, somewhat angulate and 
remotely dentate, sparsely and rather softly pilose, the basal sinus 
densely ciliate with long, white hairs; tendrils branched; flowers 
green, the calyx nearly or quite 2 cm. long; petals linear, 12-15 mm. 
long; fruit at first setose, 2-2.5 cm. long. The young and tender 
fruits sometimes are cooked and eaten. The ripe fruits, if taken in the 
hand and pressed gently, explode and wriggle in one's hand like a 
caterpillar, in much the same fashion as the seed pods of Impatiens. 
These properties are well known in Central America, where the 
children delight in picking the fruits. 

Elaterium gracile (Hook. & Arn.) Cogn. Chanchitos. Common 
in thickets and forest, Meseta Central to the Pacific coast; region of 
San Ramon; at 1,300 meters or less. Mexico to Venezuela and 
Ecuador. Similar to the preceding species, but the leaves not ciliate 
in the basal sinus. 

Elaterium pauciflorum Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 
276. 1891. Piedra del Convento, Pittier 3907. Known only from the 
original material. Tendrils simple, rather than branched as in the 
preceding species; staminate racemes only 2-3-flowered. 

FEVILLEA L. 

Scandent vines, the leaves thin, angulate or lobate, the flowers 
small, paniculate, dioecious; calyx and corolla similar in both kinds 
of flowers, the calyx 5-lobate, the corolla divided into separate petals; 
stamens 5, accompanied by 5 staminodia, the anthers 1-celled; ovary 
completely 3-celled; fruit very large, imperfectly 3-celled, the seeds 
large, compressed. A single species is found in Central America. 

Fevillea cordifolia L. Cabalonga, Chichimora, Contraveneno. 
I have seen no Costa Rican material, but the plant is reported from 
the tierra caliente. Widely distributed in tropical America. Stems 
stout, glabrous; leaves long-petiolate, broadly oval to ovate-oblong, 
8-15 cm. long, entire or lobate, short-acuminate, subcordate at the 
base, 5-nerved; staminate panicles large, many-flowered, the calyx 
2-3 mm. long, the lobes ovate, rounded at the apex; petals yellow or 
brown, 3-4 mm. long, orbicular-ovate; fruit globose, about 10 cm. 
in diameter, green and lustrous; seeds 12, brown, very large. The 



1396 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

handsome and distinctive seeds often are displayed for sale by the 
curanderos who frequent the markets. The seeds have a high reputa- 
tion throughout Central America as a cure for various diseases. 
Pittier reports the Brunka name as Sran-ua. 

FRANTZIA Pittier 

Scandent herbs, almost glabrous; leaves entire or deeply lobate; 
flowers small, whitish or yellowish; staminate inflorescence racemose; 
calyx subrotate, the 5 teeth thick, rounded; corolla lobes 5, spreading, 
ovate-lanceolate; stamens 5, the filaments connate to form a column, 
the anthers forming an irregular head; pistillate flowers solitary, 
in the same axils with the staminate inflorescence; ovary 1-celled, 
1-ovulate; fruit fibrous or ligneous, ovate-rounded, somewhat acule- 
ate, indehiscent; seed ovate, smooth. The genus consists of only the 
following species. It was named for Alexander von Frantzius, 
investigator of the fauna and physiography of Costa Rica. 

Frantzia montana Pittier, Contr. U. S. Nat. Herb. 13: 128. /. 
35, 36. 1910. Climbing on old trunks in pastures on the slopes of 
Volcan de Turrialba, 2,500 meters, Pittier 13192. Also in the region 
of El Copey, 1,800 meters. Endemic. Stems glabrous; leaves long- 
petiolate, 8-10 cm. long, thick, white-punctate above, almost gla- 
brous, paler beneath and rather densely glandular and hispidulous, 
3-lobate almost to the base, the lobes narrow, long-acuminate, the 
outer ones hastate at the base; staminate inflorescence 15-25 cm. 
long, the filiform pedicels 3-5 mm. long; calyx 4 mm. broad, sparsely 
short-pilose; corolla pale yellow, sparsely brownish-pilose outside, 
the lobes 3.5 mm. long; fruit ovoid, 4.5 cm. long, 3 cm. broad, sparsely 
beset with short, hard spines; seed compressed, smooth, 2 cm. long, 
1 cm. broad. 

Frantzia Pittieri (Cogn.) Pittier, Contr. U. S. Nat. Herb. 13: 
129. /. 37-39. 1910. Chayotillo, Taca, Tacaco. Cyclanthera Pittieri 
Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 275. 1891. Elateriopsis Pittieri 
Cufodontis, Archivio Bot. 10: 50. 1934. Carrillo, Rio Sucio Valley, 
300 meters, Pittier 1212. Apparently frequent in forests of the Atlantic 
tierra caliente; region of Zarcero; ascending to 1,800 meters. En- 
demic. Plants glabrous throughout; leaves about 10 cm. long, entire 
or shallowly trilobate, deep green above, pale beneath, deeply cordate 
at the base, acuminate, the lobes remotely denticulate or subentire; 
staminate inflorescence 10-25 cm. long, the filiform pedicels 3-7 mm. 
long; calyx 5 mm. broad; corolla 1 cm. in diameter, green; fruit 4-6 
cm. long, 3-4 cm. broad, with short, hard spines throughout or only 



FLORA OF COSTA RICA 1397 

at the base; peduncle 3-5 cm. long. F. Pittieri var. quinqueloba 
(Cogn.) Pittier (Contr. U. S. Nat. Herb. 13: 130. 1910; Cyclanthera 
Pittieri var. quinqueloba Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 276. 
1891; Boruca, Pittier 3506} is a form with the leaves deeply 3-5- 
lobate, the lobes being narrower and more narrowly acuminate than 
in the typical form. The fruits are much like those of Polakowskia. 
They have an odor resembling that of cucumber. 

GURANIA Cogn. 

Reference: Cogniaux, Pflanzenreich IV. 275, I: 192. 1916. 

Scandent herbs, sometimes with woody stems, pilose; leaves 
entire or lobate; flowers small or medium-sized, the calyx red or 
orange, the petals small, yellow; flowers dioecious, the staminate 
spicate, capitate, or racemose, the sepals elongate and very narrow; 
stamens 2, the anthers narrow or broad, usually appendiculate; 
fruit oblong, containing many compressed seeds. Other species are 
known from Central America. 

Gurania costaricensis Cogn. Diagn. Cucurb. 1: 37. 1876; 
Pflanzenreich 275, I : /. 50. Type collected by Warscewicz, without 
indication of the locality (photo, in Herb. Field Mus.). Forests of 
the Atlantic tierra caliente, ascending to 1,350 meters. Endemic. 
A large, scandent herb, the stems hirsute; leaves short-petiolate, 
ovate-cordate or oblong-cordate, acuminate, dentate, pilose; stami- 
nate flowers pedicellate and umbellate, the sepals 7-9 mm. long, 
the receptacle sericeous-pilose. Var. subtrilobata Cogn. op. cit. 38, 
the type collected by Oersted (No. 7), differs in having somewhat 
trilobate leaves. In this as in other species the flowers are handsome 
and very conspicuous because of their brilliant colors, especially 
when the plants grow in places where there are few other plants 
that produce brilliant flowers. 

Gurania Levy ana Cogn. Common in the tierra caliente, ascend- 
ing to the Meseta Central; Guanacaste; region of San Ramon; at 
1,300 meters or less. Nicaragua to Panama. A large vine, the stems 
short-pilose and rough; leaves long-petiolate, broad, usually trilobate, 
the lobes obtuse or acuminate, denticulate; staminate flowers disposed 
in a long-pedunculate head; sepals linear, pilose, 3-4 cm. long, the 
receptacle tomentose; connective of the anther broad, prolonged into 
a papillose appendage, the anther oblong, replicate at the base. 

Gurania Makoyana (Lam.) Cogn. Frequent in forests of the 
Atlantic tierra caliente, ascending to Juan Vifias. Guatemala to 



1398 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Panama. Stems short- villous; leaves deeply trilobate, sometimes 
5-lobate, soft-pilose, the lobes narrow, remotely denticulate; stami- 
nate flowers in dense, long-pedunculate heads; sepals narrowly 
linear, about 3 cm. long, the receptacle densely tomentose; anthers 
oblong, replicate at the base, the connective narrow, prolonged into 
a glabrous appendage. 

Gurania megistantha Donn. Smith, Bot. Gaz. 33: 251. 1902. 
Suerre, Llanuras de Santa Clara, 300 meters, J. D. Smith 6649. 
Also at Talamanca and Pejivalle; ascending to 900 meters. Panama. 
A woody vine, the stems as much as 7 cm. thick, covered with a 
thick, corky bark, the wood small, with extremely large vessels, 
the stems climbing to the tops of tall trees; leaves unknown; stami- 
nate flowers racemose; receptacles glabrous, the sepals 8 mm. long, 
the whole flower as much as 4 cm. long. Easy to recognize by the 
form of the flower, with a long, narrow, glabrous receptacle. This 
plant is perhaps the only Costa Rican member of the Cucurbitaceae 
with truly woody stems. 

Gurania racemifera Standl., sp. nov. Caulis scandens her- 
baceus dense longihirsutus; folia profunde triloba basi breviter 
cordata utrinque sparse hirsuta, petiolo dense hirsute, lobis oblongis 
breviter abrupte acuminatis fere integris, lateralibus paullo latioribus; 
flores masculi racemosi, racemis dense multifloris ad 5 cm. longis 
longipedunculatis, pedicellis incrassatis 4-7 mm. longis glabris vel 
sparse hirsutis; receptaculum ovoideum 5 mm. longum pilis paucis 
longis laxis albis onustum, sepalis lineari-attenuatis erectis 4 mm. 
longis sparse hirsutis; corollae segmenta linearia sepalis paullo 
breviora attenuata; antherae ovatae rectae, connective angusto, 
appendice anguste triangulari papillose. Moist forest, Naranjos 
Agrios, Prov. Guanacaste, 600 meters, Standley & Valeria 43508 
(type in U. S. Nat. Herb.). Flowers orange. Easy of recognition 
among Costa Rican species by its racemose, very small flowers. 

Gurania Pittieri Cogn. Pflanzenreich IV. 275, I: 198. 1916. 
Forests of Tsaki, Talamanca, 200 meters, Tonduz 9589. Stems 
densely short-pubescent; leaves deeply trilobate, sparsely pilose, 
the lobes acuminate, remotely denticulate; staminate flowers forming 
a dense head; receptacle ovoid, short- villous, the sepals narrowly 
linear, 14 mm. long; anthers broad, replicate at the base, the connec- 
tive broad, not produced at the apex. 

Gurania Seemanniana Cogn. Frequent in forests of the 
Atlantic tierra caliente, ascending to 1,500 meters. Honduras to 



FLORA OF COSTA RICA 1399 

Panama. Stems villous with long, soft hairs; leaves 5-7-lobate, 
pilose, the lobes broad or narrow, acuminate, denticulate; staminate 
flowers forming dense heads; receptacle densely white- tomentose, 
the sepals pilose, very narrow, 2 cm. long; anthers replicate at the 
base, the connective narrow, prolonged as a pilose appendage. 

Gurania Tonduziana Donn. Smith, Bot. Gaz. 31: 112. 1901. 
Shirores, Talamanca, 100 meters, Tonduz 9332. Stems pubescent; 
leaves deeply 7-parted, pubescent, the lobes very narrow, acuminate, 
remotely denticulate; staminate flowers densely capitate; receptacle 
densely short-pubescent, the sepals narrowly linear, 2.5 cm. long; 
anthers oblong, replicate at the base, the connective narrow, produced 
into a glabrous appendage. 

LAGENARIA Seringe 

Large, scandent herbs; flowers large, white, monoecious, solitary; 
staminate calyx almost funnelform, the corolla of 5 distinct, spread- 
ing petals; stamens 3, distinct, the anther sacs flexuous; fruit large, 
indehiscent, dry or nearly so, with a hard, brittle shell; seeds white, 
obovate, marginate. The genus consists of a single species. 

Lagenaria siceraria (Molina) Standl. Calabaza. Cultivated 
commonly in temperate and lowland regions. Probably native in 
the tropics of the Old World. Plants softly and finely pubescent; 
leaves large, long-petiolate, reniform-cordate, denticulate, rarely 
lobate; petals crispate, 3-4 cm. long; staminate calyx 2-3 cm. long. 
The hard-shelled gourds or calabashes borne by this plant are of 
importance in the household economy of rural regions. They are 
used as containers for all sorts of substances, and for fashioning 
ladles, spoons, and other articles for kitchen use. The fruits, accord- 
ing to Pittier, fall into three classes, as regards form: large and 
globose, called nambiro in Nicoya; elongate and sausage-shaped, 
known as calabaza dulce; and the most common, bottle-shaped. The 
second form is used particularly for fashioning the sounding-boxes 
of marimbas. The young and tender fruits of this vine are cooked 
and eaten as a vegetable in some regions. Indian names listed by 
Pittier are: Deka (Cabcara), Dshiun, lun-kra, lo-kra (Brunka), 
Igug (Te"rraba). 

LUFFA Adans. 

Annual vines, usually scandent, with broad leaves and branched 
tendrils; flowers bright yellow, the staminate racemose, the pistillate 
solitary; staminate calyx 5-lobate, the 5 petals distinct; stamens 3, 



1400 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

distinct, the anther cells linear, flexuous; fruit dry, 3-celled, fibrous 
within, cylindric or oblong, many-seeded, opening at the apex by 
an operculum. One other species is known to occur in Central 
America. 

Luff a cylindrica (L.) Roem. Estopa, Paste. Often planted 
about the Meseta Central and in the tierra caliente, and frequently 
wild in thickets. Native of the Old World tropics, but widely natur- 
alized in America. A large vine with glabrous stems; leaves long- 
petiolate, rounded-ovate, as much as 25 cm. wide, deeply cordate at 
the base, scabrous, lobate and sinuate-dentate, the lobes triangular, 
acute; staminate flowers racemose at the end of a long peduncle, 10- 
20 in a raceme, the pedicels 2 cm. long; petals 2-3 cm. long; fruit 
cylindric or fusiform, 10-30 cm. long, smooth, ecostate. The mass 
of fiber that fills the interior of the fruit looks much like a sponge, 
and is used locally for the same purposes. The tender young fruits 
sometimes are cooked and eaten. 

Luff a operculata (L.) Cogn. Reported by Cogniaux from Costa 
Rica, on the basis of an Oersted collection, without locality; the 
record is doubtful, but the plant is to be expected in Costa Rica. 
Mexico to Brazil. Similar to the preceding species, but the fruits 
much smaller, about as large as a hen's egg, 10-costate, spinose- 
tuberculate along the ribs; petals 1 cm. long. 

MELOTHRIA L. 

Reference: Cogniaux, Pflanzenreich IV. 275, I: 75. 1916. 

Slender, usually scandent vines; tendrils simple or rarely bifid; 
leaves small, thin, angulate or lobate; flowers very small, white or 
yellow, monoecious, the staminate fasciculate, the pistillate often 
solitary; calyx campanulate, 5-dentate; corolla campanulate, deeply 
5-lobate; stamens 3, the anthers distinct or slightly united, their cells 
not flexuous; fruit small, resembling a small melon or cucumber, 
fleshy, smooth, indehiscent. One or two additional species are native 
in Central America. 

Melothria guadalupensis (Spreng.) Cogn. Chiverrillo. M. 
fluminensis Gardn. Frequent in the tierra caliente, ascending to the 
Meseta Central, and even to higher elevations in the mountains; 
region of San Ramon; at 1,500 meters or less, growing in forest or 
thickets. Mexico to West Indies and Venezuela. A small, slender 
vine, scandent or often prostrate, much branched, the branches 
glabrous; leaves slender-petiolate, ovate-cordate or triangular-sub- 



FLORA OF COSTA RICA 1401 

sagittate, 6-10 cm. long, acute or short-acuminate, entire or shallowly 
3-5-lobate, the margin undulate or remotely dentate, scabrous; 
stamina te peduncles 2-6-flowered, the pedicels 2-3 mm. long, the 
calyx tube 2.5 mm. long; corolla 3-4 mm. broad; fruit oval, about 1.5 
cm. long and 1 cm. thick. 

Melothria scabra Naud. Thickets and forest of the Atlantic 
tierra caliente, and in Guanacaste. Panama to Mexico. Leaves 
rather stiff, ovate-oblong to triangular-subsagittate, 5-10 cm. long, 
acute or short-acuminate, subentire or 5-lobate, remotely denticulate, 
very scabrous, short-hirsute beneath on the nerves; staminate 
peduncles 5-7-flowered, the pedicels 1-3 mm. long; calyx tube 3 mm. 
long; corolla 5 mm. broad; fruit 2.5 cm. long, 1.5 cm. thick. In this 
species the anthers are about as broad as long; in M. guadalupensis 
they are much longer than broad. 

Melothria trilobata Cogn. M. trilobata var. costaricensis Cogn. 
Pflanzenreich IV. 275, I: 81. 1916 (Rio Zhorquin, 50 meters, Tonduz 
8504}. Changuinola Valley. Ranging to Surinam. Plants coarser 
than in the other species; leaves 3-5-lobate almost to the base, 
scabrous; calyx tube 3-4 mm. long; corolla villous; fruit 4-5 cm. long, 
2.5-3 cm. thick, pale green, striped with dark green. 

MICROSECHIUM Naud. 

Scandent herbs, somewhat puberulent or villous-hirsute, peren- 
nial; tendrils 3-6-fid; leaves petiolate, large and broad, thin, cordate 
at the base, usually 3-5-lobate; flowers small, monoecious, the stami- 
nate racemose; calyx tube broadly campanulate, the 4 teeth subulate; 
petals triangular-ovate; stamens 4, the filaments connate into a col- 
umn, the anthers free, 2-celled, the cells flexuous; pistillate flowers 
capitate-umbellulate, 3-parted; fruit fleshy, indehiscent, very small, 
sparsely spinulose, 1 -seeded. One other species has been described 
from Guatemala, but its generic status is uncertain. 

Microsechium Helleri (Peyr.) Cogn. In forest, region of Zar- 
cero, 1,650 meters; material from Dota and Pejivalle also may be 
referable here; Oersted specimens, without locality, cited by Co- 
gniaux, probably were collected in Costa Rica. Ranging to Mexico. 
Plants slender, much branched, the stems glabrous or puberulent; 
leaves long-petiolate, bright green, commonly shallowly 3-5-lobate, 
sparsely short-hirsute above, punctate-scaberulous in age, sparsely 
short-villous-hirsute beneath, denticulate, mostly 8-15 cm. long, the 
lobes subacute to acuminate; staminate inflorescence 10-30-flowered, 



1402 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

the pedicels filiform; calyx sparsely pilose, 4 mm. broad or less; 
corolla 8-11 mm. broad, white; pistillate peduncles 1-2 cm. long, 2-5- 
flowered; fruit ovoid, obscurely angulate, 12-16 mm. long, short- 
spinulose. 

MOMORDICA L. Balsam-apple 

Slender, scandent herbs; tendrils simple or branched; flowers 
dioecious or monoecious, yellow, the staminate solitary or fasciculate, 
the pistillate solitary; calyx 5-lobate; corolla subrotate, 5-parted; 
stamens 3, with short, distinct filaments, the anther cells flexuous; 
fruit ovoid to cylindric, 3-valvate, fleshy. No other species are 
known from Central America. 

Momordica Charantia L. Sorosi, Pepino, Pepinillo. Common 
in thickets of the tierra caliente, often forming dense tangles over 
old stumps, especially along open river banks; Guanacaste. Widely 
distributed in the tropics of both hemispheres. Stems very slender, 
much branched, somewhat pubescent; leaves long-petiolate, thin, 
4-12 cm. wide, deeply 5-7-lobate, pubescent or glabrate, the lobes 
narrow, dentate, acute or obtuse; staminate peduncle bearing a 
broad, cordate bract at or below the middle; corolla 1.5-2 cm. long; 
fruit orange-yellow, obtusely tuberculate, 2-12 cm. long, the pulp 
bright red. The ripe fruits are sometimes eaten by children, but the 
flavor is not agreeable. 

PITTIERA Cogn. 

Scandent herbs; tendrils 4-5-fid; leaves rather large, thin, long- 
petiolate, ovate-cordate; flowers large, pale yellow, monoecious, all 
axillary and solitary; calyx of the staminate flower oblong-subcylin- 
dric, the 5 lobes triangular-subulate; corolla campanulate, lobate to 
the middle or more deeply, the lobes ovate, acute; stamens 3, the 
elongate filaments free; anthers linear, connate to form a head, one 
1-celled, the others 2-celled; fruit fleshy, indehiscent, many-seeded; 
seeds broadly obovate, complanate, smooth, narrowly marginate. 
The genus consists of about four species, in Central America and 
Mexico. It was named for Henry Pittier, whose name will ever be 
associated with Costa Rica and its flora. 

Pittiera longipedunculata Cogn. Bull. Soc. Bot. Belg. 30, pt. 
1: 272. 1891. Chiverrillo, Guillotilla, Sandillita. Roadside hedge, 
San Jose", Pittier 3200. Frequent in thickets about San Jose", 1,000- 
1,200 meters. Probably endemic. A large vine, the stems pilose; 
leaves mostly 5-10 cm. long, not lobate, remotely and minutely 
dentate, densely and minutely rough-pubescent on both surfaces, 



FLORA OF COSTA RICA 1403 

usually rounded and apiculate at the apex, deeply cordate at the 
base; staminate peduncles as much as 20 cm. long, the calyx densely 
puberulent, 14-17 mm. long, the lobes 12-18 mm. long; corolla punc- 
tate-furfuraceous, 2.5 cm. long; pistillate peduncles 1 cm. long or less; 
fruit elliptic-ovoid, about 7 cm. long and 4 cm. broad, green with pale 
yellow stripes; seeds 1 cm. long, 6-7 mm. wide, rounded at the apex, 
attenuate at the base. The Bribri name is recorded as Urmeme-uo. 

POLAKOWSKIA Pittier 

Plants herbaceous or suffrutescent, scandent, glabrous or nearly 
so, the roots fleshy; leaves large, membranaceous, more or less 
trilobate, cordate at the base; tendrils 5-fid; flowers small, white, 
monoecious; staminate inflorescence racemose; calyx broadly cam- 
panulate, the teeth small, subulate; corolla lobes 5, spreading, ovate- 
triangular; stamens 5, the filaments connate to form a slender, 
elongate column, the anthers free, 4 of them connate in 2 pairs, 
the cells forming a depressed head; pistillate flowers solitary in the 
same axils with the staminate racemes; ovary 1-celled, 1-ovulate; 
fruit rather fleshy, obovoid-ellipsoid, somewhat compressed, 5-sulcate, 
with few rather hard, short spines near the base; seeds elliptic, 
smooth. The genus consists of a single species. It was named for 
H. Polakowsky, an early collector in the mountains of Costa Rica. 

Polakowskia Tacaco Pittier, Contr. U. S. Nat. Herb. 13: 131. 
pi. 20; /. 40, 41. 1910. Tacaco, Tlacaco. San Jose", 1,200 meters, 
Werckle 16674- Frequent in thickets and forest of the central region; 
region of San Ramon; Guanacaste; at 1,900 meters or less. Endemic. 
Leaves long-petiolate, 5-10 cm. long or even larger, shallowly or 
deeply lobate, the lobes acuminate, dentate, often angulate-lobate 
at the base, white-punctate above; staminate racemes 6-20 cm. long, 
the filiform pedicels 1-2.5 cm. long; calyx tube 1.5 mm. long, the 
subulate teeth 1 mm. long; corolla 4 mm. long; fruiting peduncles 
3-4 cm. long; fruit green, 4-6 cm. long, 1.5-2.5 cm. broad, narrowed 
at each end; seed about 2.5 cm. long and 1.5 cm. broad. The fruits 
are one of the common vegetables of central Costa Rica. In flavor 
and appearance they are much like small chayotes, but they vary 
greatly in quality, according to the amount of fiber present. The 
vines are often planted, but little attention is given them, other 
than gathering the fruits. 

SECHIUM P. Br. 

A scandent, perennial vine with very large, tuberous roots; 
tendrils branched; leaves large, lobate or angulate; flowers small, 
white, monoecious, the staminate racemose, the pistillate solitary 



1404 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

or geminate; calyx hemispheric, 5-lobate; corolla rotate, 5-parted, 
the lobes ovate-lanceolate; stamens 3, the short filaments connate, 
the anther cells flexuous; fruit large, obovoid, sulcate, echinate or 
smooth, indehiscent, containing a single large seed. The genus 
consists of one species. 

Sechium edule (Jacq.) Swartz. Chayote. Cultivated abun- 
dantly in the temperate regions; also at lower and higher elevations; 
sometimes found more or less naturalized. Probably native in 
Mexico and Central America, but perhaps no longer known in a truly 
wild state. Plants often 10 meters long or more, much branched; 
leaves long-petiolate, the blades broadly ovate to rounded, mostly 
10-20 cm. long, angulate or shallowly lobate, deeply cordate at the 
base, scabrous or almost smooth; staminate racemes long-peduncu- 
late, the flowers short-pedicellate; calyx lobes 5-7 mm. long; corolla 
1.5 cm. broad; fruit green, usually 8-12 cm. long. In some regions 
of the uplands there are large plantations of this vine, the plants 
growing over trellises or poles. The fruits are one of the favorite 
vegetables of Costa Rica, as also of other parts of Central America. 
There are many varieties, differing in size, shape, and color of the 
fruit. In some forms the fruits are covered with long, flexible spines, 
while in others the spines are lacking. Practically every part of the 
plant is useful for food. The young shoots are one of the quelites 
cooked and eaten like spinach. The large roots, somewhat resem- 
bling sweet potatoes in gross appearance, may be taken from the 
plant without apparent injury to it. Known in Costa Rica simply 
as raiz ("root"), they are one of the usual ingredients of the plato 
national, the national dish that appears upon the table for dinner 
almost every evening, and consists of a great variety of vegetables, 
boiled in plain water. The sliced roots are employed also for making 
a common dulce or dessert, boiled in sirup. When eaten they are 
found to be very crisp, and crunch between the teeth like shavings, 
or some similar substance. The chayote has been grown in the 
southern United States, without finding much favor there, perhaps 
because the American housewife tends to avoid new fruits or vege- 
tables. Indian names reported by Pittier are: Pis (Cabe"cara), Se-uak 
(Bribri), Tsua-ua (Brunka), Suru (T&raba), Pok-pogi-ku (Guatuso). 

SICYDIUM Schlecht. 

Reference: Cogniaux, Pflanzenreich IV. 275, I: 256. 1916. 
Climbing herbs, tomentose or glabrate; leaves ovate, entire, 
cordate at the base; tendrils bifid; flowers minute, green or whitish, 



FLORA OF COSTA RICA 1405 

racemose or fasciculate along the branches of a usually lax and large 
panicle, dioecious; staminate calyx tube rotate, the corolla rotate, 
deeply 5-parted; stamens 3, free, 2 of the anthers 2-celled, the other 
1-celled; fruit very small, globose, indehiscent, 1-seeded. One other 
species is found in northern Central America. 

Sicydium coriaceum Cogn. La Hondura de San Jose* and 
region of San Ramon; at 1,100-1, 400 meters. Colombia. Plants gla- 
brous throughout; leaves subcoriaceous, short-acuminate, 5-nerved, 
8-15 cm. long; panicles usually much exceeding the leaves, many- 
flowered, the pedicels capillary; corolla lobes linear, 2.5 mm. long. 

Sicydium tamnifolium (HBK.) Cogn. Thickets of the tierra 
caliente; Guanacaste; Zarcero, at 1,600 meters. Mexico to West 
Indies and Ecuador. Plants abundantly pubescent, the leaves, 
especially, usually densely velutinous-pilose, 5-7-nerved; corolla 
lobes 1.5 mm. long; fruit black at maturity, 5-6 mm. long. 

SICYOS L. 

Slender, climbing vines, usually pubescent; leaves petiolate or 
the upper sessile, thin, angulate or lobate; tendrils branched; flowers 
monoecious, the staminate racemose or subcorymbose, very small; 
calyx tube broadly campanulate or cupular, the 5 small teeth remote, 
subulate; corolla rotate or subcampanulate, 5-parted; stamens 3, 
the filaments connate into a column, the anthers connate or free, 
the cells flexuous; pistillate flowers capitate, the peduncle in the same 
axil with the staminate flowers; fruit small, compressed or angulate, 
hard, indehiscent, usually aculeate. A few other species have been 
collected in Central America. 

Sicyos sertuliferus Cogn. Bull. Soc. Bot. Belg. 30, pt. 1: 277. 
1891. Climbing on trees, bridge over the Rio Tiliri, near San Jose", 
Pittier 3199. Occasional about San Jose" and elsewhere on the Pacific 
slope. Endemic. Branches densely villous at the nodes, elsewhere 
pilosulous or glabrate; leaves short-petiolate, cordate-orbicular, sub- 
lobate or 5-lobate almost to the middle, punctate-scabrous above, 
densely puberulent beneath, the lobes triangular, the terminal one 
narrowly long-acuminate, the lateral ones acute; tendrils 5-fid; 
umbels 20-30-flowered, the flowers minute; fruit ovoid, compressed, 
short-rostrate, sparsely furnished with very long and short intermixed 
setae, the margins angulate-dentate. 

LOBELIACEAE. Lobelia Family 

Herbs or shrubs, sometimes more or less scandent, often with 
milky sap; leaves alternate, entire or dentate, without stipules; 



1406 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

flowers small or large, solitary and axillary or in racemes or umbels; 
calyx tube united with the ovary, the limb 5-lobulate or of 5 sepals; 
corolla gamopetalous, irregular, generally bilabiate, the tube often 
cleft on one side to the base; stamens 5, inserted with the corolla, 
the filaments sometimes united to form a tube, the anthers united; 
ovary 2-5-celled; fruit capsular or baccate. No other genera are 
known in Central America. 

BURMEISTERA Karst. & Triana 

Reference: Wimmer, Repert. Sp. Nov. 30: 1-52. pi. 123-126. 1932. 
Terrestrial or sometimes epiphytic herbs; leaves mostly petiolate 
and denticulate; flowers large, axillary; corolla tube broad at the 
base, attenuate upward and narrow below the lobes; anthers glabrous, 
or the 2 inferior ones barbate at the apex; fruit baccate, sometimes 
greatly inflated or swollen and bladder-like. A few other species are 
known from Central America. 

Burmeistera aurobarbata Wimmer, Repert. Sp. Nov. 30: 35. 
1932. Centropogon aurobarbatus Wimmer, op. cit. 19: 251. 1924. 
Volcan de Barba, Brade 2315. Common in forests of Barba and 
Irazu, 1200-2,400 meters, frequent in the region of San Ramon. 
Endemic. A terrestrial or epiphytic herb, 1 meter long or less, gla- 
brous; leaves somewhat coriaceous when dry, ovate-lanceolate, 
usually 3-9 cm. long, pale beneath; sepals green or reddish, 4-5 mm. 
long; corolla reddish green, 2 cm. long; lower anthers barbate at 
the apex. 

Burmeistera coleoides (Vatke) Wimmer, Repert. Sp. Nov. 30: 
35. 1932. Siphocampylus coleoides Vatke, Linnaea 38: 732. 1874. 
Centropogon coleoides Zahlbr. Ann. Hofmus. Wien 6: 439. 1891. Type 
collected by Warscewicz, probably in Costa Rica or Panama. Similar 
to B. aurobarbata, but the leaves broader, with more conspicuous 
teeth, the corolla only 15 mm. long. I have seen no material referable 
to this species, but I suspect it is the same plant as B. aurobarbata. 

Burmeistera cyclostigmata Bonn. Smith, Bot. Gaz. 20: 291. 
1895. Near La Estrella, Prov. Cartago, 2,200 meters, J. D. Smith 
5845. Frequent in the region south of Cartago, 1,200-2,200 meters; 
La Carpintera and La Palma de San Jose 1 ; region of San Ramon. 
Endemic. An almost glabrous herb of 1-1.5 meters, usually some- 
what scandent; leaves large, lance-elliptic, inconspicuously denticu- 
late, membranaceous; calyx green or dark red, the sepals linear, 6 mm. 
long or sometimes as much as 15 mm., usually entire; corolla green 



FLORA OF COSTA RICA 1407 

and red or dark purple, 2.5 cm. long; anthers glabrous; berry not 
inflated, 1.5 cm. long, white and purple or pink. 

Burmeistera estrellana Wimmer, Repert. Sp. Nov. 30: 24. 
pi. 123, f. 8. 1932. La Estrella, Prov. Cartago, 2,300 meters, Stork 
1537. Frequent on the slopes of Barba, in moist forest, 2,000-2,400 
meters. Plants a meter high, erect, somewhat pubescent or almost 
glabrous; leaves elliptic, rather large, narrow-acuminate, denticulate, 
membranaceous; sepals small, triangular, 3-4 mm. long; corolla 
purplish or yellow, 18 mm. long. 

Burmeistera macrocalyx Wimmer, Repert. Sp. Nov. 30: 19. 
pi. 124, /. 22. 1932. Desengano, Wendland 839. Common in forests 
of Barba and Irazu, 1,400-2,400 meters. Reported also from Ecua- 
dor. A glabrous herb, as much as 2 meters high, erect or supported 
by other plants; leaves oblong or ovate-oblong, rather large, sinuate- 
dentate, narrow-acuminate; calyx purplish, the sepals oblong or 
narrowly lanceolate, 18-25 mm. long, denticulate; corolla green and 
very dark purple, 2 cm. long or larger; anthers glabrous; berries green, 
conspicuously inflated, 3-6 cm. long. This species is little different 
fromB. Pittieri. 

Burmeistera marginata Karst. & Triana. One collection 
obtained between La Estrella and Copey is referred here doubtfully 
by Wimmer, and material from Poas and the region of San Ramon is 
perhaps conspecific; at 1,850-2,500 meters. Colombia. Plants 
glabrous, subscandent; leaves short-petiolate, elliptic, about 11 cm. 
long and 4 cm. wide, cuspidate-acuminate, densely obtuse-dentate, 
with a conspicuous, submarginal nerve; flowers solitary, long-pedi- 
cellate; sepals triangular-oblong, 2-4 mm. long, subobtuse, entire; 
corolla greenish or purple, 2 cm. long. 

Burmeistera microphylla Bonn. Smith, Bot. Gaz. 25: 146. 
1898. La Palma, 1,520 meters, Werckle 11600. Known only from 
the original collection. (t)Centropogon scandens Planch, & Oerst. 
Vid. Medd. Kjoebenhavn 157. 1857. A small plant, probably 
epiphytic, pubescent; leaves ovate-elliptic, small, only 18-20 mm. 
long; sepals linear, 2 mm. long; corolla green, 2 cm. long; anthers 
barbate. 

Burmeistera obtusifolia Wimmer, Repert. Sp. Nov. 29: 58. 
1931. Near El Copey, 2,400 meters, Stork 1633. Known only from 
moist forests of the type region, 2,000-2,400 meters. Plants terres- 
trial, conspicuously pubescent; leaves small, short-petiolate, oblong- 



1408 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ovate, obtuse; sepals reflexed, oblong, 6-7 mm. long; corolla very 
dark purple-red, as much as 3.5 cm. long. The leaves are distichous. 

Burmeistera parviflora Wimmer in herb., sp. nov. Suffrutex 
epiphyticus, ramulis gracilibus flexuosis sparse pilis laxis villosulis; 
folia inter minora membranacea graciliter petiolata, petiolo usque 
1.5 cm. longo glabro; lamina ovato-oblonga vel elliptico-oblonga 
7-12.5 cm. longa 2-4.5 cm. lata longe anguste acuminata, basi plus 
minusve obliqua acuta vel obtusa, supra viridis glabra, subtus palli- 
dior tantum ad nervos venasque villosula, nervis lateralibus utroque 
latere ca. 8 prope marginem in nervum collectivum regularem con- 
junctis; flores ex axillis superioribus nascentes, pedicellis gracillimis 
usque 6 cm. longis glabris folia fere aequantibus; hypanthium 
glabrum late turbinatum 3.5 mm. longum; sepala ovato-triangularia 
obtusa erecta fere 3 mm. longa; corolla lutea extus glabra, tubo usque 
labium inferiorem 1 cm. longo basin versus sensim paullo dilatato, 
corolla tota 17 mm. longa, lobis anguste acuminatis; antherarum 
tubus intus apice sparse barbatus. In forest, Cerros de San Antonio 
de San Ramon, 1,100 meters, July, 1926, Brenes 5636 (type in Herb. 
Field Mus.). San Antonio, west of La Palma de San Ramon, 
1,400 meters, Brenes 3802. Cerros de La Balsa de San Ramon, 
1,200 meters, Brenes 4968. 

Burmeistera Pittieri Wimmer, Repert. Sp. Nov. 30: 19. pi. 
124, /. 20a. 1932. B. Pittieri var. decorans Wimmer, op. cit. 20 (La 
Palma, Tonduz 12522). Volcan de Turrialba, 2,200 meters, Pittier 
13073. Frequent in forests of the central region, 2,000-2,400 meters; 
region of San Ramon, 1,200 meters. Panama. A glabrous herb, 
erect or somewhat scandent, 1-1.5 meters long; leaves oblong, 
acuminate, denticulate; calyx purplish green, the sepals linear, 14-20 
mm. long; corolla green and dark red, 2-2.5 cm. long; anthers gla- 
brous; berry little inflated. 

Burmeistera suerrensis (Bonn. Smith) Wimmer, Repert. Sp. 
Nov. 30: 14. 1932. B. cyclostigmata var. suerrensis Donn. Smith, Bot. 
Gaz. 24: 394. 1897. Suerre, Llanos de Santa Clara, 300 meters, J. D. 
Smith 6623. Frequent in forests of the central region, especially 
on the slopes of Barba; 300-2,400 meters; region of San Ramon. 
Panama. An erect or somewhat scandent herb, about 1 meter high, 
almost glabrous; leaves membranaceous, rather large, ovate-oblong 
or elliptic, narrow-acuminate, denticulate; sepals oval or triangular, 
only 2-3 mm. long; corolla purplish green, 2.5 cm. long; anthers gla- 
brous; berry little inflated. 



FLORA OF COSTA RICA 1409 

Burmeistera tenuiflora Donn. Smith, Bot. Gaz. 25: 147. 1898. 
Alto del Roble, Volcan de Barba, 2,000 meters, Pittier 19. Collected 
also at La Palma and Yerba Buena, 1,400-2,000 meters. Endemic. 
A pubescent herb; leaves oblong or elliptic, narrow-acuminate; sepals 
narrowly linear, 10 mm. long; corolla yellowish, 18 mm. long; anthers 
glabrous. 

Burmeistera vulgaris Wimmer, Repert. Sp. Nov. 30: 27. pi. 123, 
f. 14. 1932. Finca La Cima, above Los Lotes, near El Copey, 2,100- 
2,400 meters, Standley 42686. Region of Dota and slopes of Barba, 
2,000-2,500 meters. Panama. An herb 1.5 meters or less in height; 
leaves oblong, narrowly acuminate, denticulate, membranaceous; 
sepals 4-5 mm. long; corolla green, 2.5 cm. long; berries much inflated, 
purplish, 3 cm. long. 

CENTROPOGON Presl 

Plants herbaceous or suffrutescent, sometimes more or less 
scandent; leaves membranaceous, small or large, usually dentate; 
flowers pedicellate, axillary or grouped in terminal, usually umbelli- 
form racemes, large and showy, red or purple; anthers not barbate; 
fruit baccate, not inflated. A few other species are known from 
Central America. The genus contains some of the most showy and 
brilliantly flowered plants of Costa Rica, notable for their abundant 
flowers, which often are of an intense, bright red. The genus is closely 
related to Burmeistera but differs in the form of the corolla, its tube 
being narrow above the ovary and gradually dilated upward. 

Centropogon augostanus Wimmer, Repert. Sp. Nov. 38: 9. 
1935. Prov. Cartago, forests near "Augosta," Polakowsky 1$1. 
Plants scandent, glabrous or nearly so; leaves broadly ovate, acute, 
somewhat pubescent beneath; flowers forming a short raceme; sepals 
8 mm. long; corolla 3.5 cm. long, papillose. It is unfortunate that 
the species should have received such a specific name, based upon a 
word that certainly is not a Costa Rican locality name. I am not 
certain, but it seems probable that the word interpreted by Wimmer 
as the place name is really "Agosto," i.e., the month August. In 
addition, he gave the country as Mexico. 

Centropogon Austin-Smithii Standl., sp. nov. Suffrutex 
metralis, caulibus crassiusculis brunnescentibus fere omnino glabris, 
internodiis brevibus; folia modica petiolata herbacea, petiolo gracili 
1-2.5 cm. longo glabro; lamina lanceolata vel oblongo-lanceolata 
11-16 cm. longa 3.5-5.5 cm. lata longe anguste attenuato-acuminata, 
basi acuta vel subobtusa, interdum abrupte breviter decurrens, 



1410 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ubique obscure adpresse remote crenata, supra viridis glabra, subtus 
paullo pallidior ad nervos venasque minutissime dense puberula, 
nervis lateralibus utroque latere ca. 7 obliquis arcuatis marginem 
fere attingentibus; flores ex axillis superioribus nascentes solitarii, 
pedicellis gracilibus ca. 3.5 cm. longis puberulis; hypanthium semi- 
globosum usque 5 mm. longum atque paullo latius; sepala viridia 
anguste triangularia 6 mm. longa acuminata Integra; corolla intense 
coccinea 5 cm. longa extus glabra, tubo supra hypanthium paullo 
contracto, deinde faucem versus sensim contracto, fauce fere 1 cm. 
lato, lobis suberectis ovato-triangularibus caudato-acuminatis; an- 
therae longiexsertae 7 mm. longae apice sparse breviter hispidulae. 
Palmira, 1,800 meters, on open, recently cleared land, September, 
1937, Austin Smith A327 (type in Herb. Field Mus.). The branches 
are hard and ligneous. 

Centropogon brumalis Standl., sp. nov. Herba elongata 
debilis subscandens omnino glabra, ramis gracilibus, internodiis 
brevibus; folia inter minora oblongo-ovata 5-8.5 cm. longa 2-4 cm. 
lata longe sensim anguste acuminata, basi breviter cordata, in toto 
margine arete pectinato-laciniatula, dentibus angustis patentibus 
fere glanduliformibus, subtus paullo pallidiora, venis fuscis valde 
conspicuis; flores axillares, pedicellis gracillimis 3-6 cm. longis; 
hypanthium oblongo-campanulatum 6 mm. longum 3.5 mm. latum 
basi rotundatum glabrum; sepala anguste lineari-attenuata 5-6 mm. 
longa erecta; corolla purpurea extus glabra 3.5 cm. longa, tubo 
supra basin angustato, dein versus faucem paullo ampliato, ad 
faucem 5 mm. lato. Wet forest, Cerro de Las Vueltas, Prov. San 
Jose*, 2,700-3,000 meters, Standley & Valeria 43984 (type in U. S. 
Nat. Herb.). A species well marked by its complete lack of pubes- 
cence. In general appearance it is similar to C. palmanus. 

Centropogon coccineus (Hook.) Regel. Forests of the Atlantic 
tierra caliente. Panama. Plants herbaceous, ascending or sub- 
scandent, the branches minutely puberulent; leaves slender-petiolate, 
ovate, 5-8 cm. long, acute or acuminate, obtuse or rounded at the 
base, densely, acutely, and unequally serrate, glabrous above, pilo- 
sulous beneath on the veins; flowers axillary, the pedicels 2.5-6.5 cm. 
long; sepals linear, 8-10 mm. long, attenuate, remotely serrate; 
corolla purple-red, 4.5 cm. long, sparsely and minutely villosulous. 

Centropogon cordifolius Benth. var. dentatus Wimmer in 
herb., var. nov. Herba debilis subscandens, caulibus gracilibus 
glabris; folia graciliter petiolata herbacea, petiolo ca. 2 cm. longo; 



FLORA OF COSTA RICA 1411 

lamina ovata 3.5-10 cm. longa 3-6 cm. lata acuminata, basi truncata 
vel breviter cordata, ubique grosse inaequaliter salienter serrata, 
supra viridis glabra, subtus pallida praesertim ad venas villosula; 
flores ex axillis superioribus nascentes solitarii longipedicellati; 
hypanthium glabrum, sepalis lanceolato-oblongis 5-6 mm. longis 
subrecurvis glabris; corolla rubra extus glabra fere 2.5 cm. longa. 
Socorro de San Ramon, 1,300-1,400 meters, September, 1926, 
Brenes 4991 (type in Herb. Field Mus.). Cerros de San Antonio de 
San Ramon, 1,100 meters, Brenes 5635. Piedades de San Ramon, 
1,400 meters, Brenes 3716. The typical form of the species occurs in 
Guatemala and southern Mexico. 

Centropogon costaricanus Planch. & Oerst. Vid. Medd. Kjoe- 
benhavn 156. 1857. Siphocampylus Regelii Vatke, Linnaea 38: 732. 
1874 (Irazu, Wendland 679}. C. affinis var. costaricanus Zahlbr. Ann. 
Hofmus. Wien 6: 437. 1891. C. costaricanus var. Cufodontidis and 
var. tomentellus Wimmer, Ann. Naturhist. Mus. Wien 46: 240. 
1933. Type collected in Costa Rica by Oersted (photo, of type, 
Oersted 9240, collected near Cartago, in Herb. Field Mus.). Frequent 
in forests of the slopes of the central mountains, 2,100-2,800 meters. 
Endemic. An erect herb of 1-1.5 meters, with abundant pubescence 
of branched hairs; leaves oval or elliptic, sometimes oblong, obtuse 
to short-acuminate, with minute teeth, more or less tomentose be- 
neath, the tomentum usually grayish; sepals very short; corolla red, 
somewhat tomentose, the lobes yellow. 

Centropogon erastus Wimmer, Repert. Sp. Nov. 38: 8. 1935. 
La Palma, A. C. Erode 2395 (photo, of type in Herb. Field Mus.). 
Plants scandent, the stems glabrous; leaves broadly ovate, 10-15 cm. 
long, short-acuminate, obtuse to cordate at the base, somewhat 
pubescent beneath; flowers arranged in a short raceme; sepals 
narrow- triangular, 1.5 cm. long; corolla red or purple, puberulent, 
3.5 cm. long. 

Centropogon Gutierrezii (Planch. & Oerst.) Wimmer, Repert. 
Sp. Nov. 22: 202. 1926. Siphocampylus Gutierrezii Planch. & Oerst. 
Vid. Medd. Kjoebenhavn 155. 1875. S. thysanopetalus Vatke, 
Linnaea 38: 731. 1874 (type collected in Costa Rica or Veraguas 
by Warscewicz). Volcan de Irazu, 2,700 meters, Oersted (photo, of 
type, Oersted 9239, in Herb. Field Mus.). Forests of Pods, Barba, 
and Irazu, 2,500-3,150 meters. Endemic. An herb about a meter 
high, somewhat pubescent; leaves short-petiolate, oblong, acute or 
acuminate, conspicuously serrulate, the veins very prominent 



1412 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

beneath; flowers axillary, long-pedicellate; sepals linear, as much as 
1.5 cm. long; corolla pale red or purple, 5 cm. long, somewhat 
tomentose. 

Centropogon nutans Planch. &Oerst.Vid. Medd. Kjoebenhavn 
156. 1875. Type collected by Oersted. Very common in mountain 
forests of the central region, 1,200-2,000 meters; region of San Ramon; 
Guanacaste, 500-700 meters. Panama. An herb of 1-2 meters, 
almost glabrous, erect or subscandent, somewhat succulent; leaves 
large, oval or broadly ovate, acute, rounded or obtuse at the base, 
almost entire; flowers arranged in a short, umbelliform raceme which 
is usually pendent; sepals linear, 1 cm. long or larger; corolla bright 
red, 5 cm. long, papillose. A characteristic and very handsome plant 
of the central mountains, often growing in great abundance. 

Centropogon palmanus (Bonn. Smith) Wimmer, Repert. Sp. 
Nov. 38: 7. 1935. C. nematosepalus var. palmanus Bonn. Smith, 
Bot. Gaz. 44 : 115. 1907. La Palma, 1,500 meters, Pittier 271 . Forests 
of Barba and Irazu, 1,500-2,400 meters. Endemic. A rather slender, 
procumbent herb; leaves small, ovate, acuminate, somewhat cordate 
at the base, finely serrate; flowers axillary, the pedicels elongate; 
sepals about 4 mm. long; corolla bright red, 3.5 cm. long, sparsely 
pubescent. 

Centropogon poasensis Gleason, Torreya 25: 92. 1925. Volcan 
de Pods, Prov. Alajuela, 2,500 meters, J. D. Smith 6626. Known 
only from the upper slopes of Poas, at 2,100-2,600 meters. An herb a 
meter high, the leaves almost coriaceous, ovate-oblong, narrow- 
acuminate, rounded or obtuse at the base, 4-11 cm. long, glabrous 
above, densely tomentose beneath, the tomentum composed of 
branched, brownish hairs; flowers axillary or forming a short raceme; 
ovary densely tomentose, the sepals 3 mm. long; corolla red, 3-4 cm. 
long, tomentose. 

Centropogon porphyrodontus Bonn. Smith, Bot. Gaz. 44: 114. 
1907. Valle de Los Arcangeles, Prov. San Jose", 1,900 meters, Pittier 
12307. Region of Copey and Cerro de Las Vueltas, at 3,000 meters 
or less. Endemic. Leaves somewhat coriaceous, lance-ovate, 7-13 
cm. long, narrow-acuminate, rounded or subcordate at the base, 
somewhat puberulent beneath; flowers axillary; sepals linear; corolla 
pink, glabrous, 4.5 cm. long. 

Centropogon Smithii Wimmer, Ann. Naturhist. Mus. Wien 46: 
240. 1933. Siphocampylus discolor Bonn. Smith, Bot. Gaz. 23: 248. 
1897, non Kunth & Bouche". Forests of Rancho Flores, Volcan 



FLORA OF COSTA RICA 1413 

Barba, 2,000 meters, Tonduz 2149. Forests of Barba and Irazu, 
1,800-2,500 meters; region of Zarcero. Endemic. A usually simple 
herb of 1-3 meters, sometimes more or less suffrutescent; leaves 
large, short-petiolate, obovate-oblong, acuminate, narrowed toward 
the base, finely dentate, somewhat furfuraceous-pubescent; stems 
narrowly winged; flowers axillary, the pedicels greatly elongate; 
sepals linear, 1.5-2 cm. long; corolla pale green, at times somewhat 
yellowish, about 5 cm. long, more or less furfuraceous. The species of 
this genus are sometimes described by writers as shrubs, but they are 
really herbs, at most merely somewhat suffrutescent near the base. 

Centropogon solanifolius Benth. Pavoncillo. C. solanifolius 
var. hirtellus Wimmer, Ann. Naturhist. Mus. Wien 46: 241. 1933 
(Volcan de Irazu, 2,400 meters, Cufodontis 285). Frequent in moun- 
tain forests of the whole central region, 1,500-2,800 meters; region 
of San Ramon. Extending to Ecuador. An herb of 1-1.5 meters, 
erect or somewhat scandent, glabrous or nearly so; leaves oblong- 
elliptic, narrow-acuminate, crenate, acute at the base; flowers in 
terminal racemes, these elongate or umbelliform; sepals linear, short; 
corolla orange-red, 4.5-5 cm. long, somewhat pubescent. A highly 
ornamental plant, abundant in many regions. It has been reported 
from Costa Rica as C. grandis (L.) Presl. 

Centropogon tortilis Wimmer, Repert. Sp. Nov. 22: 217. 1926. 
Rio Chirripo, Pittier 16033. A scandent herb of 2.5 meters, the stems 
glabrous; leaves very large, obovate-elliptic, rounded and short- 
acuminate at the apex, narrowed toward the base, glabrous; flowers 
forming a short, terminal raceme; sepals 5 mm. long; corolla red, 
3 cm. long, pilose. 

Centropogon Valerii Standl., sp. nov. Herba erecta metralis, 
caule tenuiter villosulo vel puberulo glabrato, internodiis brevibus; 
folia membranacea, petiolo 3-8 mm. longo; lamina lanceolato- 
linearis vel lineari-oblonga 5.5-13 cm. longa 1-2.5 cm. lata anguste 
acuminata, basi acuta, in toto margine arete minute dentata, supra 
glabra, subtus pallidior minutissime puberula vel fere glabra; flores 
axillares, pedicellis gracilibus 3-3.5 cm. longis; hypanthium obo- 
voideo-globosum 5 mm. altum, fere glabrum; sepala linearia 2 mm. 
longa serrata; corolla ca. 3 cm. longa, extus minutissime puberula, 
tubo supra calycem angustato, deinde sensim dilatato, ad faucem 6 
mm. lato, lobis lineari-lanceolatis attenuatis valde decurvis, superiori- 
bus ad 12 mm. longis; tubus staminalis longe exsertus villosus, 
antheris glabris, inferioribus apice dense albo-setosis. Ojo de Agua, 



1414 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

2,850 meters, H. E. Stork 3022 (type in Herb. Field Mus.). Cerro 
de Las Vueltas, Prov. San Jose", 2,700-3,000 meters, Standley & 
Valeria 43751 , 43951 . The leaves are the narrowest that I have noted 
among all the North American species. 

Centropogon Warscewiczii Vatke, Linnaea 38: 716. 1874. 
Type collected in Costa Rica or Panama by Warscewicz. Glabrous, 
the leaves oblong, rather acute, narrowed toward the base, the upper 
ones sessile; flowers arranged in short, umbelliform racemes; sepals 
linear-lanceolate, 2-3 times as long as the ovary; corolla pilose. 

Centropogon Wimmerii Standl., sp. nov. Herba metralis, 
caule pilis longis multilocularibus laxis sparse villoso; folia mediocria 
longipetiolata membranacea, petiolo gracili 1.5-3 cm. longo; lamina 
ovato-ovalis vel elliptico-ovata 8-9 cm. longa 4.5-5 cm. lata abrupte 
breviter acuminata, acumine anguste acute attenuate, basi late 
rotundata, supra in sicco fusca ad costam sparse villosa, subtus fere 
concolor praesertim ad venas pilis longis simplicibus patentibus 
villosa, margine irregulariter adpresso-crenato; flores axillares, ut 
videtur nutantes, pedicellis ad 9 cm. longis sparse villosis; hypan- 
thium hemisphaericum 5 mm. longum basi late rotundatum sparsis- 
sime villosum; sepala lineari-attenuata viridia 4 mm. longa ciliata 
erecta; corolla coccinea 3 cm. longa laxe villosa, tubo supra hypan- 
thium paullo constricto deinde sensim dilatato, ad faucem 6 mm. 
lato, lobis lineari-oblongis attenuatis, superioribus solemniter de- 
flexis; antherae glabrae, superiores apice dense albo-setosae. Las 
Nubes, Prov. San Jose", moist forest, 1,500-1,900 meters, Standley 
38739 (U. S. Nat. Herb., type). Probably related to C. costaricanus, 
but separated by its pubescence of simple, not branched hairs. 

HETEROTOMA Zucc. 

Annual or perennial herbs, the leaves membranaceous, alternate, 
dentate; flowers small or large, racemose; corolla prolonged at the 
base as a small or large spur. No other species are known from 
Central America. 

Heterotoma lobelioides Zucc. Forests of the central region, 
1,500-2,500 meters. Extending to Mexico. An herb 60 cm. or more 
in height, almost glabrous or pubescent; leaves ovate or lanceolate, 
acuminate, rounded at the base; flowers large, almost 4 cm. long, 
red and yellow. A very showy plant because of its large, vividly 
colored flowers. 



FLORA OF COSTA RICA 1415 

Heterotoma tenella Turcz. Reported from Costa Rica by 
Hemsley, collected by Endres, at 1,050 meters. Extending to 
Mexico. A small, annual herb, simple or branched, pubescent; 
leaves small, reniform or cordate; flowers very small, blue. 

ISOTOMA Lindl. 

A single species is found in Central America. 

Isotoma longiflora (L.) Presl. Jazmin de estrella, Jazmincillo. 
Occasional in the tierra caliente. A species of wide distribution. 
A somewhat pubescent herb with thick stems; leaves almost sessile, 
oblanceolate, coarsely dentate; flowers axillary, short-pedicellate, 
white, 10 cm. long; fruit capsular. Easy to recognize because of the 
large, white flowers, with a very long and slender tube. 

LAURENTIA Neck. 

Slender, small, annual herbs, the small, narrow leaves alternate, 
the peduncles axillary, 1-flowered; calyx 5-parted or 5-dentate; 
corolla straight or oblique, very small, the tube entire or shortly 
cleft; anthers 2, short-setose at the apex; fruit a small capsule, 
loculicidally bivalvate. Probably no other species are found in 
Central America. These small plants look much like some of the 
smaller species of Lobelia, such asL. xalapensis, but in the latter genus 
the corolla tube is cleft almost or quite to the base. 

Laurentia irazuensis (Planch. & Oerst.) Wimmer, comb. nov. 
Lobelia irazuensis Planch. & Oerst. Vid. Medd. Kjoebenhavn 153. 
1857. Irazu, Oersted. Frequent in meadows and pastures of the 
central region; regions of Dota and San Ramon; at 1,000-1,800 
meters. Endemic. Plants simple or somewhat fastigiately branched, 
usually 30 cm. high or less, the stems hispidulous or hirtellous, 
sometimes glabrate; leaves lance-oblong to almost linear, mostly 
less than 2 cm. long, acute to attenuate, dentate, sessile, glabrous 
or nearly so; pedicels filiform, mostly shorter than the subtending 
leaves, the flowers abruptly upcurved; sepals linear-attenuate, be- 
coming 5-6 mm. long, ciliate, equaling the capsule; corolla blue- 
purple, scarcely 5 mm. long. 

Laurentia micrantha (HBK.) Zahlbr. El Tablazo, region of 
San Ramon, and doubtless elsewhere, growing in pastures. Ranging 
to Mexico. Similar to the preceding species but often more slender, 
the stems glabrous or hirtellous; pedicels often much longer than 
the subtending leaves; sepals scarcely more than 3 mm. long; corolla 
violaceous. 



1416 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

LOBELIA L. 

Plants herbaceous or suffrutescent, annual or perennial; flowers 
small or large, axillary or in terminal racemes, pedicellate; corolla 
not spurred, cleft nearly to the base on one side; anthers glabrous, 
sometimes barbate at the apex. Other species are known from 
Central America. 

Lobelia irasuensis Planch. & Oerst. Vid. Medd. Kjoebenhavn 
153. 1857. L. Warscewiczii Vatke, Linnaea 38: 718. 1874 (Costa 
Rica or Veraguas, Warscewicz). Volcan de Irazu, Oersted. Meadows 
on the upper slopes of Irazu. Chiriqui region, Panama, at about 
3,000 meters. Plants perennial, almost glabrous, simple, mostly 
30 cm. high or less; leaves lance-linear, sessile, denticulate; racemes 
elongate, slender, few-flowered; corolla blue, about 8 mm. long. 

Lobelia laxiflora HBK. Yuquilla, Ceragallo. (l)Tupa costaricana 
Planch. & Oerst. Vid. Medd. Kjoebenhavn 154. 1857. L. costaricana 
Wimmer, Ann. Naturhist. Mus. Wien 46: 239. 1933. Frequent in 
thickets of the central region, 1,000-1,600 meters. Panama to 
Mexico. A branched herb about a meter high, more or less pubes- 
cent; leaves lanceolate, sessile or short-petiolate, denticulate; flowers 
axillary, red, about 3 cm. long. A conspicuous plant because of its 
large, bright red flowers, common in many regions of Central America. 
Wimmer considers (according to determinations in the herbarium) 
that the Costa Rican material of this relationship represents two or 
more species, but the characters by which they are to be separated 
are not apparent to the writer. It is probably more satisfactory 
to give the several forms varietal status, unless really good characters 
can be suggested for separating them as species. 

Lobelia poasensis Wimmer, Ann. Naturhist. Mus. Wien 46: 
239. 1933. Meadows of Volcan de Poas, 2,500 meters, Cufodontis 
605. Known, apparently, only from the type. Plants perennial (?), 
the stems erect or ascending, 15-25 cm. long, glabrous; leaves on 
very short petioles or the uppermost sessile, rhombic or rhombic- 
elliptic to lanceolate, acute, cuneate at the base, dentate, glabrous or 
nearly so; racemes 10-20 cm. long, the pedicels 7-10 mm. long; calyx 
tube 1 mm. long, the lobes subulate, entire, 1-2 mm. long, spreading; 
corolla lilac, glabrous, 6 mm. long, the tube 3 mm. long. 

Lobelia splendens Willd. Buenos Aires, 480 meters, and 
probably in other regions of the tierra caliente; region of San Ramon, 
800 meters. Widely distributed in tropical America. Plants stout 



FLORA OF COSTA RICA 1417 

and erect, growing usually on the banks of small streams, almost 
glabrous; leaves nearly linear, sessile, attenuate, entire; flowers red, 
3 cm. long, in short or elongate racemes. The plant is known in 
Panama by the name Flor de garza. 

Lobelia urticifolia Wimmer. L. neglecta Vatke. Frequent in 
thickets and other shady places of the central region, 1,000-2,300 
meters; El General. A species of wide distribution. A branched 
herb, erect or procumbent, pubescent or almost glabrous; leaves 
petiolate, ovate or broadly lanceolate, acuminate, dentate; flowers 
purplish white, axillary, 8 mm. long, the pedicels very long and 
slender. 

Lobelia xalapensis HBK. Pastures of the Meseta Central, 
descending to the Atlantic tierra caliente; region of San Ramon; 
at 1,600 meters or less. Widely distributed in tropical America. 
Plants annual, mostly less than 50 cm. high, usually branched, some- 
what pubescent with fine, short hairs; leaves petiolate, mostly deltoid- 
ovate, obtuse or acute, dentate; flowers blue or purplish, 4-5 mm. 
long, on very long, slender pedicels. The Costa Rican plant often 
has been referred to L. Cliff ortiana L. 

SIPHOGAMPYLUS Pohl 

Erect or somewhat scandent herbs, in most characters similar 
to Burmeistera and Centropogon, but differing from those genera by 
the capsular fruit. With flowering specimens it is difficult to distin- 
guish the two groups. 

Siphocampylus costaricae Vatke, Linnaea 38: 730. 1874. 
Volcan de Barba, Hoffmann 60 (photo, of type, Hoffmann 60, in 
Herb. Field Mus.). Known only from the original collection. Leaves 
petiolate, ovate, acuminate, cordate at the base, dentate, glabrous 
above, pubescent beneath; sepals linear, 4-5 mm. long; corolla red, 
5 cm. long. I have seen no specimens of this plant, which probably 
belongs to the genus Centropogon. The description is suggestive 
of C. palmanus. 

Siphocampylus nematosepalus (Donn. Smith) Wimmer, Re- 
pert. Sp. Nov. 38: 22. 1935. Centropogon nematosepalus Donn. 
Smith, Bot. Gaz. 44: 114. 1907. Cuesta de Los Borucas, Puntarenas, 
2,300 meters, Pittier 10533. Plants paleaceous-pubescent, the leaves 
membranaceous, lance-ovate, 7.5-10 cm. long, cordate at the base, 
denticulate; flowers axillary; sepals filiform, 9 mm. long; corolla 
purple-red, 4.5 cm. long, pubescent. 



1418 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Siphocampylus roseus Bonn. Smith, Bot. Gaz. 23: 249. 1897. 
Along the railroad line, Siquirres, Pittier 3165. Also at Puerto Viejo. 
Endemic. An almost glabrous herb; leaves ovate, acuminate, 8-13 
cm. long, dentate, puberulent on the nerves beneath; flowers axillary; 
sepals oblong, 7-8 mm. long; corolla puberulent, 4.5-5 cm. long; 
capsule globose, 2 cm. long. 

SPHENOCLEA Gaertn. 

A single species is found in America. 

Sphenoclea zeylanica Gaertn. Marshy places of the Atlantic 
coast. A species of wide distribution. A glabrous herb a meter high 
or less, branched; leaves petiolate, entire, elliptic to lanceolate, 
acute or obtuse, pale; flowers small, white, in very dense, short or 
elongate, pedunculate spikes. In appearance the plant is altogether 
unlike other representatives of the family. 

COMPOSITAE. Composite Family 

Herbs, shrubs, or trees, sometimes scandent or epiphytic; leaves 
opposite or alternate, entire to dissected, never truly compound 
although sometimes apparently so; flowers collected in a head (rarely 
1-flowered) on a receptacle, surrounded by an involucre of phyllaries 
(often called bracts); corolla gamopetalous, regular, tubular, and 
5-dentate (rarely 2-4-dentate), bilabiate, or ligulate (i.e., flattened, 
liguliform, and usually 2-5-dentate), rarely wanting in the pistillate 
flowers; stamens (in the perfect or staminate flowers) almost always 
5, united by the anthers or rarely free, inserted on the corolla; ovary 
inferior, 1-celled; style usually 2-branched, the branches stigmati- 
ferous inside, often with sterile appendages at the apex; fruit an 
achene, with a single seed, often bearing at the apex a pappus of 
setae, awns, or scales. The corollas are of four chief kinds: ligulate 
or strap-shaped, bilabiate, tubular, and filiform. Heads composed 
of one kind of flowers only are called homogamous; those of two or 
more kinds heterogamous; when heterogamous, the central flowers 
(disk) are always perfect, the peripheral pistillate or neutral (lacking 
the style). Homogamous heads in which all the flowers are perfect 
and have ligulate (in this case always 5-dentate) corollas are called 
ligulate; homogamous heads in which all the corollas are tubular 
and perfect or staminate, or filiform and pistillate, are called discoid. 
Heterogamous heads in which the peripheral corollas are ligulate 
are called radiate; those in which the peripheral flowers are pistillate, 
with tubular, filiform, or abortive corollas are called disciform. The 
receptacle may be naked, bristly, or paleaceous (bearing pales or 



FLORA OF COSTA RICA 1419 

chaff). The generic characters are drawn to a considerable extent 
from the character of the pappus, which may be of bristles, awns, 
scales, or teeth, reduced to a crown or cup, or entirely wanting. 
(From S. F. Blake, Contr. U. S. Nat. Herb. 23: 1401-1402. 1926.) 

The Compositae constitute the largest family of plants. Although 
well represented in Costa Rica, they are far less abundant, in general, 
in the tropics than in temperate areas. In preparation of the account 
of the Costa Rican species the writer is deeply indebted to Dr. S. F. 
Blake, who read most of the manuscript, and suggested changes and 
additions. He is, however, in no way responsible for any errors 
that may occur on the following pages. The account of the genus 
Senecio has been read by Dr. J. M. Greenman, who has suggested 
corrections, and supplied descriptions of several new species. 

Key to the tribes and genera 
Corollas all bilabiate or ligulate. 
Sap milky; corollas all ligulate; herbs XII. Cichorieae. 

Sap not milky; central corollas bilabiate, not ligulate; chiefly 
shrubs, sometimes herbs XL Mutisieae. 

Corollas not bilabiate, the central ones, at least, not ligulate but 
with regular, 4-5-fid corollas, the outer flowers often ligulate. 

Anthers caudate at the base. 

Anthers long-appendaged at the apex; plants prickly. 

X. Cynareae. 
Anthers not appendaged at the apex; plants never prickly. 

Pappus none or wool-like IX. Calenduleae. 

Pappus present, not wool-like IV. Inuleae. 

Anthers not caudate at the base. 

Style branches elongate, acute, hispidulous outside throughout 
their length. 

Heads yellow VIII. Senecioneae. 

Heads white or purplish, never yellow. 

Achenes 4-angulate, clavate; pappus consisting of paleae 
with a strong costa VI. Helenieae. 

Achenes not 4-angulate and clavate; pappus not consisting 
of paleae with a strong costa I. Vernonieae. 

Style branches not as above. 



1420 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Style branches usually elongate, linear or clavate, obtuse, not 
hispidulous outside at the apex, the stigmatic lines only 
below the middle and inconspicuous; heads never yellow. 

II. Eupatorieae. 

Style branches, when elongate, hispidulous outside or at the 
apex, the stigmatic lines reaching nearly to the apex; 
heads often yellow. 
Phyllaries scarious-marginate; heads often nodding; style 

tips truncate, hispidulous VII. Anthemideae. 

Phyllaries not scarious-marginate, or else the heads not 

nodding or the style tips acute. 

Receptacle paleaceous, at least in the staminate heads, 
or else the leaves all or mostly opposite; pappus never 

of bristles V. Heliantheae. 

Receptacle not paleaceous, rarely setose, or else the leaves 

alternate. 

Pappus present, chiefly or entirely of soft, capillary 
bristles; involucre not glandular-punctate. 

Style tips acute III. Astereae. 

Style tips truncate or subtruncate . VIII. Senecioneae. 
Pappus none, or of squamellae, awns, or stiff bristles, 
or else the involucre glandular-punctate. 

Leaves alternate III. Astereae. 

Leaves opposite, at least below, or else the involucre 
glandular-punctate VI. Helenieae. 

I. VERNONIEAE 
Heads distinct, not aggregated into secondary heads or glomerules. 

Pappus an entire, cartilaginous crown Struchium. 

Pappus of scales, bristles, or scales and bristles. 

Leaves lepidote Piptocarpha. 

Leaves not lepidote. 

Inner pappus of a few linear scales; heads 2-flowered. 

Oliganthes. 
Inner pappus of terete or slightly flattened bristles; heads 

with 3 to many flowers Vernonia. 

Heads aggregated into secondary heads or glomerules. 

Corolla 3-5-dentate; style branches short; phyllaries 2-5. 

Rolandra. 
Corolla 5-cleft; style deeply bilobate; phyllaries 8 or more. 



FLORA OF COSTA RICA 1421 

Pappus bristles dissimilar, 2 of them longer and conduplicate 

near the apex Pseudelephantopus. 

Pappus bristles all alike, straight. 
Pappus bristles numerous, not broadened at the base. 

Orthopappus. 

Pappus bristles about 5, dilated and flattened at the base. 

Elephantopus. 

II. EUPATORIEAE 

Pappus none or a very short crown. 

Heads 3-flowered; anthers not appendaged at the apex. .Piqueria. 
Heads several-many-flowered; anthers appendaged at the apex. 

Pappus none Alomia. 

Pappus a low, paleaceous crown Ageratum. 

Pappus present, well developed. 

Achenes prismatic and 8-10-costate Brickellia. 

Achenes 4-5-costate. 
Pappus of bristles only. 

Phyllaries 4; heads 4-flowered; plants scandent Mikania. 

Phyllaries and flowers more numerous; plants not scandent. 

Pappus bristles 5-8 Fleischmannia. 

Pappus bristles numerous Eupatorium. 

Pappus at least in part of squamellae or awns. 

Phyllaries 5-8, uniseriate, subequal Stevia. 

Phyllaries more numerous. 

Pappus of bristles and squamellae Fleischmannia. 

Pappus of squamellae only Ageratum. 

III. ASTEREAE 

Plants shrubs or at least conspicuously frutescent. 

Heads radiate Diplostephium. 

Heads discoid. 
Plants dioecious, the heads strictly staminate or pistillate. 

Baccharis. 

Plants polygamo-dioecious, the staminate heads as in Baccharis, 
the pistillate with 1-15 central perfect flowers. 

Archibaccharis. 
Plants herbaceous, low. 

Pappus none or reduced to a ring or collar. 



1422 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Heads discoid Laestadia. 

Heads radiate. 

Plants with branched, leafy stems Egletes. 

Plants with naked, scapiform stems Bellis. 

Pappus of numerous bristles. 
Phyllaries conspicuously unequal in length; style tips acute or 

acuminate Aster. 

Phyllaries all equal in length or nearly so; style tips short, 
obtuse Erigeron. 

IV. INULEAE 
Phyllaries dry but not scarious. 

Leaves dentate Pluchea. 

Leaves entire Tessaria. 

Phyllaries scarious, usually white or colored. 

Phyllaries in part raylike; heads large; cultivated plants. 

Helichrysum. 
Phyllaries not raylike; heads small; native plants. . . .Gnaphalium. 

V. HELIANTHEAE 
Heads with usually only 1 fertile flower. 

Involucre strongly compressed, the outermost phyllary cordate 

and bractlike Delilia. 

Involucre not compressed, the phyllaries not cordate. . . .Milleria. 
Heads with few to many fertile flowers. 
Ray flowers pistillate, fertile; disk flowers perfect, sterile. 

Leaves compound; plants scandent Hidalgoa. 

Leaves simple; plants not scandent. 
Pistillate corollas tubular. 
Pales of the disk enclosing the achenes; heads in small 

cymes Ichthyothere. 

Pales of the disk not enclosing the achenes; heads densely 
cymose-paniculate Clibadium. 

Pistillate corollas liguliform. 

Phyllaries subtending the ray achenes indurate and com- 
pletely enveloping the achenes Melampodium. 

Phyllaries not indurate and completely enveloping the 
achenes. 



FLORA OF COSTA RICA 1423 

Petioles dilated and clasping at the base; achenes not 
triquetrous Polymnia. 

Petioles not dilated at the base; achenes triquetrous. 

Plants shrubs Rensonia. 

Plants herbaceous Baltimora. 

Ray flowers, when present, pistillate or neutral; disk flowers 

perfect, fertile. 
Ray corollas sessile, persistent. 

Disk achenes strongly compressed Zinnia. 

Disk achenes quadrangular Heliopsis. 

Ray corollas with a distinct tube and deciduous, or wanting. 

Pappus of plumose squamellae Tridax. 

Pappus not of plumose squamellae, sometimes none. 
Achenes contracted into a conspicuous, stipitiform base. 

Podachaenium. 
Achenes not contracted into a stipitiform base. 

Involucre distinctly double, the outer phyllaries few, 

herbaceous, the inner submembranous. 
Trees or shrubs, the pubescence usually tomentose; 

rays white Montanoa. 

Herbs or, if shrublike, without tomentum; rays vari- 
ously colored. 

Achenes rostrate Cosmos. 

Achenes not rostrate. 
Pappus none. 

Plants without viscid pubescence Dahlia. 

Plants with abundant, viscid pubescence. 

Sigesbeckia. 
Pappus of awns. 

Pappus awns antrorse-ciliate or smooth ; achenes 
marginate or winged; cultivated plants. 

Coreopsis. 

Pappus awns retrorse-hispid or rarely smooth; 
achenes not marginate or winged; native 

plants Bidens. 

Involucre not distinctly double. 

Pappus of the disk achenes consisting of 4- many free, 
usually equal or subequal awns or squamellae. 



1424 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Heads white; plants herbaceous or chiefly so. 

Melanthera. 
Heads usually yellow; usually shrubs, rarely herbs. 

Achenes obovoid, somewhat compressed; phyl- 
laries not indurate Hymenostephium. 

Achenes subterete or 4-5-angulate; phyllaries 
indurate, or the outer ones sometimes herba- 
ceous Calea. 

Pappus of the disk achenes of 2-3 awns, with or with- 
out squamellae, or a crown of united squamellae, 
or wanting, or at least not as described above. 

Achenes strongly compressed, flat or flattish. 
Pappus of awns and squamellae. 

Rays neutral Oyedaea. 

Rays pistillate or none. 
Achenes marginate or winged, the wings not 

decurrent on the awns Zexmenia. 

Achenes winged, the wings decurrent on the 

awns Otopappus. 

Pappus various, but not of both awns and squa- 
mellae. 
Achenes, at least some of them, winged. 

Heads discoid; pappus annular; leaves alter- 
nate Garcilassa. 

Heads radiate. 

Outer achenes not winged Synedrella. 

Outer achenes winged. 
Pappus of 2-5 stout aristae united at the 

base; low herbs Calyptocarpus. 

Pappus of usually 2 slender, free aristae; 
shrubs or large, coarse herbs. 

Verbesina. 
Achenes not winged. 

Plants woody shrubs or vines Salmea. 

Plants herbaceous. 

Cauline leaves laciniate or lobate. 

Chrysanthellum. 

Cauline leaves merely dentate. 



FLORA OF COSTA RICA 1425 

Disk of the head strongly convex; flowers 

yellow or white; low herbs .Spilanthes. 

Disk not very convex; flowers yellow; 

tall, coarse herbs Simsia. 

Achenes thickened, not thin and compressed. 

Heads white, or the disk yellow and the rays white, 

pink, or red. 
Receptacle elongate, the disk very convex or 

elongate Isocarpha. 

Receptacle plane or merely convex. 
Disk yellow, the rays white, pink, or red. 

Galinsoga. 

Disk and rays white Eclipta. 

Heads yellow. 
Rays pistillate. 

Pappus of the disk achenes none . . Rumfordia. 
Pappus of the disk achenes present. 
Achenes acutely marginate or winged. 

Zexmenia. 

Achenes not marginate or winged. 
Achenes angulate; pappus none or of 

minute scales Jaegeria. 

Achenes not angulate; pappus annular 
or of more or less connate scales. 

,. Wedelia. 

Rays neutral or wanting. 

Heads discoid Eleuther anther a. 

Heads radiate. 

Rays white Montanoa. 

Rays yellow. 

Heads large, 5-14 cm. wide or broader. 
Peduncles fistulose; native plants. 

Tithonia. 
Peduncles not fistulose; cultivated 

plants Helianthus. 

Heads much smaller; peduncles not 

fistulose. 

Achenes each closely enclosed in a pale. 

Sclerocarpus. 

Achenes not closely enclosed in the 
pales Viguiera. 



1426 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

VI. HELENIEAE 

Heads discoid ; plants glabrous Porophyllum. 

Heads radiate; plants often pubescent. 
Phyllaries imbricate, numerous, in 2 or more series; disk globose; 

leaves alternate Helenium. 

Phyllaries in a single series or, if somewhat imbricate, only 5; 

disk not globose; leaves all or chiefly opposite. 
Ray achenes compressed and concave, dentate on the margins; 

pappus none Galeana. 

Ray and other achenes neither compressed nor concave, not 

dentate; pappus present. 

Pappus setose, the setae sometimes reduced to squamellae. 

Pectis. 
Pappus of paleae, these often dissected or awned. 

Pappus squamellae dissected into bristles; leaves not pin- 

natifid Dyssodia. 

Pappus squamellae not dissected; leaves usually pinnatifid. 

Tagetes. 

VII. ANTHEMIDEAE 

Receptacle paleaceous Achillea. 

Receptacle not paleaceous. 

Heads discoid, greenish Artemisia. 

Heads radiate, usually yellow with white rays, or the rays some- 
times colored. 

Achenes equally or irregularly 5-10-costate .... Chrysanthemum. 
Achenes 3-5-costate on the inner surface, ecostate dorsally. 

Matricaria. 

VIII. SENECIONEAE 
Phyllaries multiseriate, imbricate, the outer ones shorter. 

Leaves alternate, at least the lower ones lobate Neurolaena. 

Leaves opposite, not lobate. 

Receptacle naked; usually shrubs, often epiphytic; rays yellow. 

L/iabum. 

Receptacle paleaceous; herbs; rays white Schistocarpha. 

Phyllaries 1-seriate or nearly so, subequal, or graduated only in 

Gynura (cultivated). 

Pistillate flowers in several series; heads disciform; plants herba- 
ceous, with greenish or bright pink or purple flowers . Erechtites. 
Pistillate flowers in a single series. 



FLORA OF COSTA RICA 1427 

Style branches with long, subulate, hirtellous appendages; cul- 
tivated plants Gynura. 

Styles branches truncate or with short appendages. 

Heads red, pink, or orange; small, annual herbs Emilia. 

Heads yellow or white, rarely orange; perennial herbs, shrubs, 
or trees Senecio. 

IX. CALENDULEAE 
A single (cultivated) genus in Costa Rica Dimorphotheca. 

X. CYNAREAE 
A single genus in Costa Rica -. Cirsium. 

XL MUTISIEAE 
Plants acaulescent, the leaves all basal. 

Cultivated plants Gerbera. 

Native plants Chaptalia. 

Plants with leafy stems. 
Heads yellow or whitish. 

Heads yellow; involucre double; leaves pinnate-nerved. .Trixis. 
Heads whitish; involucre nearly 1-seriate; leaves palmate- 
nerved Jungia. 

Heads purple, crimson, or reddish. 

Leaves entire; shrubs Lycoseris. 

Leaves dentate and often lobate; large herbs Onoseris. 

XII. ClCHORIEAE 

Stems scapose, each bearing a single head; leaves all basal. 

Taraxacum. 
Stems more or less leafy, usually with several heads. 

Pappus setae plumose Hypochaeris. 

Pappus setae not plumose. 
Achenes compressed. 

Achenes truncate, not rostrate Sonchus. 

Achenes rostrate or narrowed at the apex Lactuca. 

Achenes terete or prismatic, not compressed. 

Achenes truncate at the apex Hieracium. 

Achenes narrowed at the apex Crepis. 



1428 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ACHILLEAL. Yarrow 

Perennial herbs with erect, leafy stems; leaves mostly pinnatifid 
and finely dissected, alternate; flower heads small, radiate, in terminal 
corymbs; involucre obovoid or campanulate, the phyllaries appressed, 
imbricate in few series, the outer ones shorter; receptacle flat or 
convex, paleaceous; ray flowers white or purple, pistillate, fertile; 
disk flowers perfect, fertile, the corollas white; achenes oblong or 
obovate, subcompressed; pappus none. No species are native in 
Central America. 

Achillea Millefolium L. Naturalized on banks in the potreros 
of Volcan de Turrialba; cultivated for ornament in gardens of the 
Meseta Central. Native of the Old World, but widely naturalized 
in temperate North America. Plants mostly 60 cm. high or less, 
generally simple below, abundantly pilose; leaves plumelike, divided 
into very numerous small, narrow segments; heads numerous, 4-6 
mm. broad, forming a compact corymb, the rays 4-6, scarcely 4 mm. 
long. Normally the rays are white, but especially in the cultivated 
forms they are frequently pink or purple. 



Achyrocline rufescens DC. was reported by Klatt from forests 
of Alto del Roble on the basis of Pittier 214- I have not seen this 
collection, and no material of the genus is available from Costa Rica. 
It is rather probable that the Costa Rican plant is a Gnaphalium, 
although the genus is represented both north and south of Costa 
Rica, and may well occur here. 

AGERATUM L. 

Reference: Robinson, Revision of the genus Ageratum, Proc. 
Amer. Acad. 49: 454. 1913. 

Annual or perennial herbs, rarely suffrutescent; leaves mostly 
opposite, broad, crenate or serrate; inflorescence usually terminal 
and cymose or corymbose, the heads discoid; involucre campanulate, 
the phyllaries narrow, subequal, 2-3-seriate, imbricate, mostly 2- 
costate; receptacle plane or convex, naked or paleaceous; achenes 
5-angulate, prismatic; pappus of distinct or connate, setiferous or 
muticous scales, sometimes an entire or dentate crown. Other 
species are known from Central America. Some species of the genus 
often are planted for ornament in the United States and other regions. 

Ageratum conyzoides L. Santa Lucia. Known in Costa Rica 
only from the region of Cartago, 1,200-1,400 meters, but to be ex- 
pected elsewhere. Widely distributed in the American tropics, and 



FLORA OF COSTA RICA 1429 

introduced in the Old World. Plants annual, commonly 60 cm. high 
or less, the stems usually purplish; leaves petiolate, obtuse, rounded 
to short-cuneate at the base, crenate, sparsely pilose; heads pedicel- 
late, 6 mm. broad, the phyllaries subabruptly acuminate, ciliate, 
usually erose below the apex, sparsely pilose or glabrate outside; 
corollas blue-purple; pappus scales 5, lanceolate, setiferous, distinct. 
A rather showy and handsome plant, in general appearance exactly 
like Alomia microcarpa, with which it often is confused by collectors. 
In Alomia the achenes bear no pappus. In some parts of Central 
America A. conyzoides is a common weed. 

Ageratum Houstonianum Mill. Reported from San Rafael 
de Cartago, and to be expected elsewhere. Extending to Mexico 
and West Indies. In general appearance very similar to A. cony- 
zoides; leaves hirsute; phyllaries entire, gradually long-attenuate, 
glandular-puberulent and hirsutulous. 

Ageratum latifolium Cav. Reported from hedges at Turrialba, 
200 meters, Tonduz 4139. Mexico to Peru and West Indies. Plants 
annual, erect or decumbent; leaves ovate or ovate-oblong, 2-5 cm. 
long, obtuse, rounded at the base, crenate-serrate; heads 5-8 in each 
corymb, 6 mm. broad, the phyllaries lance-oblong, acute, subgla- 
brate; corollas violaceous or white; pappus scales oblong or lanceolate, 
distinct, muticous. 

Ageratum Oerstedii Robinson, Proc. Amer. Acad. 49: 472. 
1913. Coelestina latifolia Benth. in Oerst. Vid. Medd. Kjoebenhavn 
71. 1852. Carelia latifolia Kuntze, Rev. Gen. 1: 325. 1891. Monte 
Aguacate, 600 meters, Oersted 251. Collected also near San Ramon. 
Endemic. Plants annual, 30-50 cm. high; leaves petiolate, ovate or 
ovate-oblong, 4-7 cm. long, obtuse or obtuse-acuminate, abruptly 
contracted at the base, crenate, sparsely pilose; heads few, 7.5 mm. 
broad, the phyllaries attenuate, sparsely hirtellous or glabrate; 
achenes glabrous, black, the pappus minute, coroniform, dentate. 

Ageratum riparium Robinson, Proc. Amer. Acad. 49: 473. 
1913. In sand along Rio Ceibo, Pittier 4914. Known only from 
the original collection. Leaves ovate, about 8 cm. long, obtuse- 
acuminate, narrowed and unequal at the base, undulate, scaberulous 
above, pilosulous beneath, especially on the veins; heads 4-8 in each 
corymb, 7 mm. long, the inner ones attenuate, the outer callose- 
obtuse, ciliate between the costae, otherwise glabrous; corollas 
white, glabrous; achenes glabrous or nearly so; pappus a 5-lobate 



1430 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

crown, the lobes muticous. Material of this species was referred 
incorrectly by Klatt to A. tomentosum Hemsl. 

Ageratum scabriusculum (Benth.) Hemsl. Region of San 
Ramon, in pastures or along roadsides, about 900 meters. Ranging 
to Mexico. Perennial, less than a meter high; leaves ovate, 2-5 cm. 
long, acuminate, serrate, lustrous above, scaberulous or minutely 
pilosulous, appearing glabrous; heads 3-7 in a corymb, 7 mm. long, 
the phyllaries very acute, crisp-puberulent or almost glabrous; 
corollas glabrous; achenes glabrous; pappus crateriform, usually 
5-dentate, scarcely 0.3 mm. long. The type locality is given by 
Robinson as Costa Rican, but it is actually Nicaraguan. 

Ageratum tomentosum (Benth.) Hemsl. Biol. Centr. Amer. 
Bot. 2: 82. 1881. Coelestina tomentosa Benth. in Oerst. Vid. Medd. 
Kjoebenhavn 71. 1852. Carelia tomentosa Kuntze, Rev. Gen. 1: 
325. 1891. Candelaria, 1,500 meters, Oersted. Extending to Mexico. 
Perennial, often suffrutescent; leaves ovate or deltoid, 2-3.5 cm. long, 
obtuse, crenate, rounded or truncate at the base and often short- 
decurrent, densely pale-tomentose beneath; corymbs long-peduncu- 
late, the heads 5 mm. long, the phyllaries narrowly lanceolate, 
tomentulose; corollas hispidulous toward the apex; achenes gla- 
brous; pappus coroniform. 

ALOMIA HBK. 

Reference; Robinson, Revision of the genus Alomia, Proc. Amer. 
Acad. 49: 438. 1913. 

Annual or perennial herbs, rarely suffrutescent, in general appear- 
ance exactly like Ageratum; leaves mostly ovate or rhombic and 
petiolate, thin, serrate or dentate; heads small, corymbose or panicu- 
late, discoid, many-flowered; involucre campanulate or subturbinate, 
the phyllaries subequal, biseriately imbricate, lance-linear, acute, 
1-3-costate, sometimes very unequal and in more numerous series; 
receptacle plane or conic, naked or paleiferous; corollas white to 
pink or purple; achenes prismatic, 5-angulate, glandular-atomiferous 
or hispidulous, sometimes glabrous, dark, the pappus very short and 
annular or none. At least three other species are known from Central 
America, in Guatemala and Panama. 

Alomia microcarpa (Benth.) Robinson, Proc. Amer. Acad. 
49: 452. 1913. Santa Lucia. Coelestina microcarpa Benth. in Oerst. 
Vid. Medd. Kjoebenhavn 72. 1852. Ageratum microcarpum Hemsl. 
Biol. Centr. Amer. Bot. 2: 82. 1881. In grassy places, Cartago, 
Oersted 10968. Abundant in fields and pastures of the central region, 



FLORA OF COSTA RICA 1431 

mostly at 1,000-1,500 meters; region of San Ramon; El General. 
Reported doubtfully from Venezuela. An erect or decumbent an- 
nual, usually about 50 cm. high, soft-pilose; leaves opposite, petiolate, 
deltoid-ovate, 3-6 cm. long, obtuse or subacute, cordate or sub- 
cordate at the base, crenate-serrate, pilose; heads numerous in each 
corymb, about 60-flowered, the phyllaries pilose; achenes black, 
glabrous, 1 mm. long; corollas blue-purple. Forma Torresii Standl. 
(Field Mus. Bot. 11: 274. 1936; La Pitahaya, Cartago, Ruben Torres 
Rojas 196) is a form with white flowers. Santa Lucia is one of the 
best known plants of central Costa Rica. It is especially abundant 
in the pastures around or above Cartago, some of which present 
solid masses of beautiful, clean color that are visible from a long 
distance. 

ARCHIBACCHARIS Heering 

Reference: Blake, Hemibaccharis, a new genus of Baccharidinae, 
Contr. U. S. Nat. Herb. 20: 543. 1924. 

Herbs or usually shrubs, often somewhat scandent, polygamo- 
dioecious; leaves alternate; heads small, whitish, discoid, cymose- 
paniculate, the pistillate bearing one or more central hermaphrodite 
flowers, the staminate often with a few marginal pistillate flowers; 
involucre 3-5-seriate, the phyllaries graduated, imbricate, linear to 
ovate, narrowly scarious-marginate; receptacle rather flat; achenes 
compressed and 2-nerved, hispidulous; pappus setose, 1-seriate. At 
least three other species occur in Central America. The genus is 
closely related to Baccharis. 

Archibaccharis irazuensis Blake, Journ. Wash. Acad. Sci. 
17: 60. 1927. Hemibaccharis irazuensis Blake, Contr. U. S. Nat. 
Herb. 20: 551. 1924. Laguna del Reventado, Volcan de Irazu, 2,300 
meters, Pittier 14079. Occasional in mountain forests of the central 
region, ascending to 3,000 meters; also in the region of San Ramon, 
1,000 meters. Endemic. A subscandent shrub, the branches short- 
pilose; leaves short-petiolate, lance-ovate or lance-elliptic, 4-8 cm. 
long, acuminate, cuneate at the base, mucronate-denticulate, pilosu- 
lous on both surfaces; panicles small and dense, rounded; heads 4.5-5 
mm. high, the phyllaries ciliate, the outer ones puberulous; achenes 
1.2mm. long. 

Archibaccharis torquis Blake, Contr. U. S. Nat. Herb. 23: 
1508. 1926. Hemibaccharis torquis Blake, op. cit. 20: 550. pi. 51. 
1924. San Jose", Tonduz 1535. Frequent in thickets and forests of 
the Meseta Central; regions of Dota and San Ramon, and widely 



1432 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

distributed on the Pacific slope, mostly at 900-1,400 meters. Extend- 
ing to southern Mexico. A shrub of 1-3 meters, often subscandent, 
the branches somewhat zigzag, puberulent or pilosulous; leaves short- 
petiolate, oval or ovate-oval, 3-8 cm. long, acute or short-acuminate, 
rounded or cuneate-rounded at the base, remotely dentate or denticu- 
late, glabrate but pilosulous beneath, chiefly on the nerves; panicles 
small and rounded, about 5 mm. wide, the heads dirty white, 4-5 mm. 
long; achenes 0.8 mm. long. Material of this species has been reported 
incorrectly under the name Conyza asperifolia Benth. & Hook. 

ARTEMISIA L. 

Aromatic herbs or shrubs, the leaves alternate, often lobate or 
pinnatifid; heads small, discoid, racemose-spicate or paniculate, 
greenish or yellowish; involucre usually hemispheric, the phyllaries 
imbricate in few series, the outer gradually shorter; receptacle flat 
to hemispheric, naked or pubescent; achenes small, obovoid or oblong, 
2-costate; pappus none. No species are native in Central America, 
but several are found in Mexico. Artemisia Absinthium L. of 
Europe is the source of the liquor absinth. 

Artemisia vulgaris L. Ajenjo. Often planted in gardens of the 
temperate region. Native of Europe. An erect, perennial herb, less 
than a meter high; leaves deeply pinnatifid into linear or oblong 
segments, glabrate above, densely white-tomentose beneath. An 
infusion of the plant is used locally in domestic medicine as a tonic, 
febrifuge, anthelmintic, and emmenagogue. 

ASTER L. 

Perennial or rarely annual herbs, usually branched, with alternate 
leaves; heads radiate, corymbose or paniculate; involucre hemi- 
spheric to turbinate, the phyllaries imbricate in several series, the 
outer ones usually much shorter; receptacle flat or convex; rays 
mostly white, blue, or purple, pistillate, the disk flowers perfect, 
usually yellow; achenes commonly compressed and nerved; pap- 
pus bristles numerous, slender, scabrous or denticulate, normally 
1-seriate. Several other species are native in Central America. 

Aster exilis Ell. Common about San Jose", and probably in other 
places in the central region, at 1,100-1,300 meters; usually growing 
at the edges of streams. Widely distributed in tropical America. 
Plants annual, commonly 60 cm. high or less, often much branched, 
glabrous, somewhat succulent; leaves linear or linear-lanceolate, 3-10 
cm. long, entire or nearly so, sessile, acute or acuminate, the leaves 



FLORA OF COSTA RICA 1433 

of the branches subulate; heads numerous, paniculate; involucre 6 
mm. high, the phyllaries linear-subulate, appressed; rays purplish, 
4 mm. long; achenes pubescent. A common and rather weedy plant 
in many parts of Central America. Costa Rican material was 
referred by Klatt to A. pauciflorus Nutt. 

Aster laevis L. Lila. Cultivated for ornament in gardens of the 
Meseta Central and elsewhere. Native of temperate North America. 
Plants perennial, stout, erect, in cultivation about 50 cm. high, gla- 
brous; upper leaves sessile and usually cordate-amplexicaul, lanceo- 
late to ovate, the lower borne on winged petioles; rays blue or violet; 
heads as much as 2.5 cm. broad; achenes glabrous or nearly so. This 
is a common garden plant in many parts of Central America. Britton 
uses for the cultivated plant the name Aster versicolor Willd., based 
upon cultivated material, which, however, is believed to be a deriva- 
tive of the common A. laevis of the United States. 

Aster spinosus Benth. Manzanilla. Common on arenales 
(sandbars) about San Jose", and doubtless elsewhere on the Pacific 
slope. Ranging to southwestern United States. Plants perennial, 1 
meter high or less, usually forming dense, broad clumps, glabrous, 
the stems pale, angulate, armed with long, stout, stiff spines; flower 
heads small, the rays white; leaves mostly linear, often early deci- 
duous. Easily recognized in the genus by its abundant spines. The 
plant almost invariably, in Central America, grows along stream beds. 

BACCHARIS L. 

Dioecious shrubs or small trees; leaves alternate; heads discoid, 
small, paniculate or corymbose; involucre campanulate or oblong, 
the phyllaries numerous, imbricate in several series, the outer ones 
shorter; receptacle flat, naked; achenes somewhat compressed, 
costate; pappus of the fertile flowers copious, capillary, soft, that of 
the sterile flowers short. A few other species are found in northern 
Central America. The genus is a vast one, most abundantly devel- 
oped in South America, but with many species also in Mexico. 

Baccharis Braunii (Polak.) Standl., comb. nov. Eupatorium 
Braunii Polak. Linnaea 41 : 576. 1877. B. splendens Heering, Schr. 
Nat. Ver. Schleswig Hoist. 13: 48. 1906 (Rio Ceibo, near Buenos 
Aires, Pittier 1+932}. Sandy slopes, Angostura, Polakowsky 508. 
Occasional in mountain forests of the central region; region of San 
Ramon; mostly at 1,000-1,400 meters. Honduras to Panama. A 
shrub of 1.5-3 meters, glabrous throughout; leaves petiolate, oblong- 



1434 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

elliptic, mostly 3-4.5 cm. wide, acute at each end, entire, conspicu- 
ously 3-nerved, lustrous and very glutinous on the upper surface; 
heads small, mostly long-pedunculate, forming large, broad corymbs. 
Costa Rican material of this shrub was referred by Klatt to B. 
nervosa DC. 

Baccharis trinervis (Lam.) Pers. Alcotan, Santo Domingo. 
Frequent in thickets or hedges, Meseta Central and surrounding 
mountains, descending to the. upper Atlantic tierra caliente and to 
the Pacific coast; Guanacaste; sometimes ascending to 1,800 meters, 
but chiefly at much lower elevations. Widely distributed in tropical 
America. A much branched shrub of 3 meters or less, the branches 
usually recurved and sometimes subscandent; branches glabrous or 
nearly so; leaves on very short petioles, elliptic to lance-oblong, 
mostly 2 cm. wide or less, acute or acuminate, entire, conspicuously 
triplinerved ; flower heads about 4 mm. high, dirty white, in small 
or large panicles. B. trinervis var. rhexioides (HBK.) Baker is a 
form in which the branches and the veins of the lower leaf surface 
are tomentulose or puberulous. It grows in association with the 
more glabrate plants, and probably is scarcely worthy of special 
nomenclatorial designation. The shrub is a very common one in 
many parts of the Pacific slope, and is a most unattractive one in 
appearance. Pittier reports the TeYraba name as Zbin-kor-ga. The 
fresh leaves are applied as poultices to aid in the healing of sores. 

BALTIMORA L. 

Erect, usually much branched annuals, scabrous-pubescent; leaves 
opposite, petiolate, crenate-dentate; heads rather small, yellow, 
forming a broad panicle; involucre campanulate, the phyllaries few, 
biseriate; receptacle flat, paleaceous; disk flowers sterile, the ray 
flowers perfect, fertile; achenes triquetrous, truncate, the pappus 
short, cuplike, dentate, deciduous. No other species is found in 
Central America. 

Baltimora recta L. Collected at Jesus Maria, San Jose", and 
Bebedero (Guanacaste), and doubtless to be found in many other 
localities. Widely distributed in tropical America. Plants slender 
and low, or sometimes coarse and as much as 2.5 meters high; leaves 
broadly ovate, acute or acuminate; heads 5 mm. high, the phyllaries 
broad, acuminate, strigose, the rays long and narrow; achenes 
glabrous, smooth or somewhat tuberculate. This is said to be a 
good honey plant. It is a common weed in many parts of Central 
America. 



FLORA OF COSTA RICA 1435 

BELLIS L. Daisy 

Low, cespitose herbs, the stems usually scapose, the leaves 
mostly basal; heads radiate, the rays white or pink, the disk yellow; 
involucre hemispheric or broadly campanulate, the phyllaries narrow, 
herbaceous, subequal, in 1-2 series; receptacle naked; achenes com- 
pressed, obovate; pappus none or a ring of minute bristles. No 
species of the genus is native in Central America. 

Bellis perennis L. Common in meadows high on the slopes 
of Irazu and Turrialba, abundant in many localities. Native of 
Europe. A cespitose perennial, the scapose stems mostly 10 cm. 
high or less, pubescent; leaves basal, obovate, obtuse, obscurely 
dentate, 2.5-5 cm. long, pubescent; involucre about 7 mm. wide, 
the rays white or pink, the phyllaries usually purple. The English 
daisy, a highly esteemed garden plant, has become thoroughly 
naturalized in the potreros of the dairy districts of the central 
volcanoes. The cool, moist climate seems to be exactly suited to the 
plants, which are a beautiful sight in these meadows. The plants 
no doubt were introduced by accident with grass seed. 

BIDENS L. 

Reference: Sherff, The genus Bidens, Field Mus. Bot. 16. 1937. 

Annual or perennial herbs, sometimes scandent and somewhat 
suffrutescent; leaves chiefly opposite, serrate, simple or variously 
divided or dissected; heads rather large, radiate or discoid, the rays 
yellow or white; involucre campanulate to oblong, the phyllaries 
biseriate, distinct or slightly united at the base, the outer ones 
often foliaceous; receptacle flat or nearly so, paleaceous; disk flowers 
perfect and fertile; achenes flat, quadrangular, or almost terete, 
linear to cuneate or oblong; pappus of 2-6 teeth or subulate awns, 
the awns barbate or hispid. A few other species probably are found 
in Central America. 

Bidens ostruthioides (DC.) Schultz Bip. var. costaricensis 
(Benth.) Sherff, Bot. Gaz. 88: 298. pi. 21. 1929; Field Mus. Bot. 
16: pi. 132. B. costaricensis Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 94. 1852. B. irazuensis Calv. & Calv. Year C. R. Nat. Hist. 
137, 140. ill. 1917, nomen. Monte Aguacate, 600-1,500 meters, 
Oersted (photo, of type in Herb. Field Mus.). Also on the slopes of 
Irazu, chiefly at 2,000^4,000 meters, where apparently plentiful. 
Southern Mexico. Plants probably suffrutescent at the base; leaves 
bipinnatisect, the segments cuneate-lanceolate, incised-dentate, 
glandular-setulose above, pale beneath; outer phyllaries 57, folia- 



1436 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

ceous, lanceolate or broadly linear; rays yellow, 15-22 mm. long; 
achenes linear, glabrous, 7-9 mm. long, 3-aristate, the awns retrorse- 
barbate. The typical form of B. ostruthioides is found in Mexico 
and Guatemala. 

Bidens pilosa L. Mozote, Moriseco, Mozotillo. Occasional in 
waste ground or thickets, Meseta Central to the coasts. Widely 
distributed in the tropics of both hemispheres. An erect, branched 
annual, usually a meter high or less; leaves slender-petiolate, simple 
or mostly 3-5-parted, thin, serrate, pilose; heads discoid, at anthesis 
5-7 mm. long; outer phyllaries 7-9, linear or linear-spatulate, ciliate, 
4-5 mm. long; achenes linear, glabrous below, tuberculate-strigose 
above, 4-16 mm. long, the awns retrorse-barbate. A common and 
disagreeable weed in many parts of Central America. The achenes 
adhere tenaciously to clothing by means of the barbate awns. The 
species, as treated by Sherff at least, is a highly variable one. Repre- 
sented in Costa Rica are the following variants, of greater or minor 
importance: var. minor (Blume) Sherff, the heads radiate, the yellow- 
ish or whitish rays only 4-8 mm. long; var. radiata Schultz Bip., the 
heads radiate, the heads at anthesis 1.5-3 cm. broad, the rays white, 
larger; var. cakicola (Greenm.) Sherff, the heads radiate, in anthesis 
1-1.5 cm. broad. 

Bidens riparia HBK. Region of Nicoya. Southern Mexico to 
Peru and Brazil. An annual, 30-50 cm. high, the stems branched, 
glabrous; leaves bipinnate, pilosulous above, glabrous beneath, the 
segments ovate or lanceolate, dentate and incised-lobate; heads in 
anthesis 4-6 mm. long, the rays 3-5 mm. long, yellowish white; 
outer phyllaries 7-13, linear, 6-10 mm. long, glabrous or nearly so; 
achenes linear, curved, 10-15 mm. long, the short awns retrorse- 
barbate. 

Bidens squarrosa HBK. Mozotillo, Barbasco, Mozote. Fre- 
quent in thickets, Meseta Central to the Pacific coast; Guanacaste; 
regions of Dota and San Ramon. Mexico to Bolivia and Brazil. A 
suffrutescent or herbaceous vine, sometimes 3.5 meters long or more, 
the stems glabrous or tomentose; leaves pinnately 3-5-parted, the 
divisions ovate to lance-oblong, acute or acuminate, serrate, pubes- 
cent or glabrate; heads numerous, usually paniculate, radiate, at 
anthesis 1.5-4 cm. wide, 6-8 mm. long; involucre hispid or glabrate, 
the outer phyllaries 4-6, linear, 3-5 mm. long; rays bright yellow, 
8-15 mm. long; achenes linear, 6-9 mm. long, the awns retrorse- 
barbate or naked. A very showy and handsome plant when in 



FLORA OF COSTA RICA 1437 

flower. Material of this species has been referred by various authors 
to B. floribunda HBK. and B. rubifolia HBK. Soils 460, from Haci- 
enda Montecristo, was referred by Sherff to B. segetum Mart., this 
being the only record of its occurrence north of Brazil. As may be 
seen by inspection of SherfFs key to species, where B. segetum and B. 
squarrosa appear side by side, the differences, if any, between the two 
species are rather indefinite. It seems more in accordance with the 
probable facts, therefore, to refer the Soils collection to B. squarrosa. 

Bidens tenera 0. E. Schulz, Bot. Jahrb. 50: Suppl. 186. 1914; 
Sherff, Field Mus. Bot. 16: pi. 108, f. i-c. Forests of Boruca, Pittier 
4528. Pacific tierra caliente. Colombia to Brazil. A slender annual, 
35 cm. high or less, the stems glabrate, simple or sparsely branched; 
leaves simple or 3-foliolate, the segments thin, ovate or ovate- 
lanceolate, acuminate, serrate, sparsely short-pilose above; heads few, 
discoid, few-flowered ; involucre hispid at the base, the outer phyllaries 
about 4, linear or linear-spatulate, 4 mm. long; achenes linear, 
straight or slightly curved, glabrous, 12-15 mm. long, the awns 
retrorse-barbate. 

BRICKELLIA Ell. 

Reference: Robinson, A monograph of the genus Brickellia, Mem. 
Gray Herb. 1.1917. 

Herbs or shrubs; leaves opposite or alternate, sessile or petiolate; 
heads discoid, usually paniculate, sometimes racemose, cymose, or 
corymbose, white or reddish; involucre cylindric or campanulate, 
the phyllaries numerous, striate, imbricate in several series, the 
outer shorter; receptacle naked; achenes prismatic, commonly 10- 
costate, mostly hispidulous, at least on the costae; pappus bristles 
10-80, slender, almost smooth to subplumose. A few other species 
are known from Central America. 

Brickellia argyrolepis Robinson, Mem. Gray Herb. 1 : 90. /. 69. 
1917. In hedges, San Rafael, Pittier 1980. Frequent in thickets and 
hedges of the Meseta Central; region of Dota; at 1,100-1,800 meters. 
Endemic. A stout shrub 1-3 meters high; leaves opposite, short- 
petiolate, ovate, 4-6 cm. long, acute, rounded to subcordate at the 
base, serrulate, soft-tomentose beneath and pale; heads 2 cm. long, 
glomerate at the ends of the branches, greenish white, the phyllaries 
silvery-sericeous; achenes 5.5 mm. long. This plant was reported 
from Costa Rica by Klatt as B. Hartwegii Gray. 

Brickellia diffusa (Vahl) Gray. Occasional in waste ground 
or thickets, Meseta Central to the Pacific coast; Guanacaste. Widely 



1438 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

distributed in tropical America. A slender, usually much branched, 
erect annual, sometimes 2 meters high, sparsely puberulent or gla- 
brate; leaves slender-petiolate, orbicular-ovate or deltoid-ovate, 4-6 
cm. long, acuminate, serrate or crenate-dentate; pedicels filiform, the 
very numerous heads whitish, about 8-flowered, 8 mm. long; achenes 
1.5-2 mm. long, villous above. A common weed in places along the 
Pacific slope of Central America. 

CALEA L. 

Reference: Robinson & Greenman, Revision of the Mexican and 
Central American species of the genus Calea, Proc. Amer. Acad. 32: 
20-30. 1896. 

Usually densely branched, low shrubs; leaves opposite, mostly 
dentate or crenate; heads rather small, radiate or discoid, usually 
cymose or umbellate, the rays yellow or white; involucre several- 
seriate, graduate, the phyllaries dry or the outer sometimes herba- 
ceous; rays, when present, fertile; achenes sub terete or 4-5-angulate; 
pappus persistent, of 4-20 usually equal, paleaceous squamellae or 
awns, rarely none. A few other species grow in northern Central 
America. 

Calea dichotoma Standl., sp. nov. Frutex, ramis gracilibus 
brunnescentibus subteretibus ubique aequaliter repetite dichotomis, 
internodiis plerumque foliis brevioribus subsparse puberulis vel gla- 
bratis; folia parva breviter petiolata subcoriacea, petiolo puberulo 
vix ad 3 mm. longo; lamina ovata vel lanceolato-ovata 1.3-3 cm. 
longa 6-16 mm. lata anguste longiacuminata, basi longiuscule acute 
decurrens, grosse adpresso-serrata, supra sparse scaberula vel fere 
omnino glabra, subtus paullo pallidior dense impresso-puncticulata 
ad nervos hinc inde pilis paucis brevibus conspersa vel fere glabra, e 
basi solemniter trinervia, nervis valde elevatis; capitula discoidea ad 
apices ramorum umbellata, vulgo 3-5, pedicellis gracilibus puberulis 
vix ultra 6 mm. longis, capitulo e quaque axilla ramorum nascente 
solitario longius pedicellate; involucrum ca. 5 mm. longum anguste 
campanulatum basi rotundatum, phyllariis ca. 3-seriatis latis apice 
rotundatis minute ciliatis aliter glabris, pallidis, extimis brevissimis; 
achaenia immatura fusca ca. 2.5 mm. longa sparse hispidula, pappi 
squamis ca. 8 anguste oblongo-lanceolatis pallidis acutis imbricatis 
1.5 mm. longis. Maderal de San Mateo, October, 1922, Brenes 3665 
(type in Herb. Field Mus.). In appearance the plant is unique 
because of its regularly dichotomous branching. Noteworthy, too, 
are the very small leaves, which, however, may not furnish a con- 






FLORA OF COSTA RICA 1439 

stant character for separation. Except for its mode of branching, 
the plant suggests C. Zacatechichi. 

Calea Pittieri Rob. & Greenm. Proc. Boston Soc. Nat. Hist. 
29: 105. 1899. Rio Ceibo, near Buenos Aires, 200 meters, Pittier 
4913. Also at Boruca, and apparently frequent in the region of San 
Ramon, as well as elsewhere on the Pacific slope; El General; Car- 
tago; at 1,400 meters or less. Endemic. A shrub of 1-3 meters; 
leaves ovate to elliptic, mostly 3.5-6 cm. long, acute to obtuse, thick, 
crenate-serrate, rugose and very scabrous above, grayish-tomentose 
beneath or hispidulous, 5-nerved from near the base; heads discoid, 
slender-pedicellate, umbellate; pappus awns about 20, longer than 
the achene. Material of this species was referred by Klatt to 
C. prunifolia HBK., of Panama and South America. 

Calea urticifolia (Mill.) DC. Jalacate, Jaral. C. peUucidinerva 
Klatt, Bull. Soc. Bot. Belg. 31, pt. 1: 207. 1892 (TeYraba, 260 meters, 
Pittier 3707}. Frequent in pastures and thickets of the Meseta 
Central, often occurring in great abundance, also in many parts of 
the Pacific slope; region of San Ramon; at 1,400 meters or less. 
Panama to southern Mexico. A dense shrub, usually 1-1.5 meters 
high or lower; leaves ovate, 5-12 cm. long, acute, coarsely serrate, 
usually rugose, very scabrous above and often rough-pubescent 
beneath; heads almost 1 cm. long, the short rays yellow; achenes 
2.5 mm. long, the pappus 3.5 mm. long. The plant sometimes 
becomes a rather bad weed in pastures, especially in the Cartago 
region. It is a showy and rather handsome shrub when in flower. 
Costa Rican material has been referred to C. axillaris DC., which 
is now considered a variety of C. urticifolia. 

Calea Zacatechichi Schlecht. Region of San Ramon, 700-800 
meters. Ranging to Guatemala and Mexico. A much branched 
shrub, pubescent or almost glabrous; leaves ovate or triangular- 
ovate, 2-6 cm. long, coarsely dentate, acute or acuminate, somewhat 
rugose; heads small, about 12-flowered, forming dense, umbellate- 
cymose panicles, the pedicels mostly shorter than the heads; rays 
none; pappus shorter than the achene. Costa Rica has not been in- 
cluded previously in the range of C. Zacatechichi, but four collections 
from San Ramon agree very well with material from northern Central 
America, and probably are to be referred here. The species name 
is of Aztec origin, signifying "bitter grass." In Mexico the plant 
has long been used in domestic medicine. 



1440 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

CALYPTOCARPUS Less. 

Low annuals, prostrate or ascending, pubescent, branched; leaves 
small, opposite, dentate; heads small, yellow, radiate, axillary, 
usually short-pedunculate; phyllaries few, herbaceous; ray flowers 
pistillate, perfect, the disk flowers perfect and fertile; achenes of the 
ray and disk alike, or the ray achenes sometimes 3-angulate, not 
winged; pappus of 2-3 stout, hard, spinose awns. A single species is 
found in Central America. The generic name often is written in- 
correctly as Calyptrocarpus. 

Calyptocarpus vialis Less. Espinillo, Cabeza de vaca, Chiquisd. 
C. Wendlandii Schultz Bip. Bot. Zeit. 24: 165. 1866 (San Jose", 
Wendland 1078}. Synedrella vialis Gray. Waste or cultivated 
ground, Meseta Central. Widely distributed in tropical America. 
Plants branched from the base, the stems 30 cm. long or less; leaves 
ovate, 1-3 cm. long, acute, cuneate to subcordate at the base, 
thinly appressed-pilose, petiolate; involucre narrow, about 7 mm. 
long, the phyllaries green, imbricate, the rays short and incon- 
spicuous; achenes about 5 mm. long. 

CHAPTALIA Vent. 

Scapose, perennial herbs; leaves floccose-tomentose, entire to 
lobate; heads radiate, solitary at the end of a long, naked scape; 
involucre turbinate-campanulate, many-flowered, the phyllaries 
narrow, imbricate in few series, the outer shorter; rays pistillate, 
fertile, white to purple; disk flowers perfect, mostly neutral, their 
corollas somewhat bilabiate; achenes columnar or fusiform, 5-nerved; 
pappus of numerous long, soft bristles. A single species grows in 
Central America. 

Chaptalia nutans (L.) Polak. Common on open banks or in 
pastures, Meseta Central, and probably elsewhere. Widely distrib- 
uted in tropical America. Leaves mostly lyrate-lobate, sometimes 
entire, 8-25 cm. long, obtuse, green and almost glabrous above, 
densely white-tomentose beneath, thin; scapes 30-60 cm. high, 
slender, floccose-tomentose; heads 2-2.5 cm. long, purple or dark 
red; achenes 5 mm. long, glabrate or glandular, the filiform beak 
2-3 times as long; pappus yellowish, 10-13 mm. long. A common 
and rather weedy plant, generally distributed in Central America. 
Costa Rican material was referred by Klatt to C. albicans (Sw.) 
Northrop, a West Indian species. 



FLORA OF COSTA RICA 1441 

CHRYSANTHELLUM Rich. 

Glabrous annuals, usually much branched and diffuse or almost 
prostrate; basal leaves mostly incised-dentate, the cauline ones 
alternate and various divided; heads small, radiate, pedunculate, 
terminal or arising from the upper leaf axils; ray flowers yellow, 
pistillate, the disk flowers perfect; involucre campanulate, the 
phyllaries 1-2-seriate, membranaceous; receptacle plane, paleaceous; 
achenes linear-oblong, dorsally compressed, the outer ones with 
obtuse margins, the inner winged; pappus none. No other species 
are found in Central America. 

Chrysanthellum integrifolium Steetz. El Rodeo, Villa Colon, 
840 meters, M. Valeria 950. Widely distributed in tropical America, 
but uncommon in Central America; growing in savannas or pastures. 
Plants slender, often prostrate; leaves oblong to spatulate and serrate, 
somewhat succulent, or often pinnatifid; heads scarcely 1 cm. broad, 
terminating very long, slender peduncles; outer phyllaries with scari- 
ous margins. 

CHRYSANTHEMUM L. 

Annual or perennial herbs; leaves alternate, dentate to dissected; 
heads large, mostly long-pedunculate, radiate; involucre usually 
hemispheric or depressed, the phyllaries appressed, imbricate in 
several series, the outer ones shorter; receptacle flat to hemispheric, 
naked; ray flowers pistillate, fertile, the disk flowers perfect and 
fertile; achenes angulate or terete, 5-10-costate; pappus none or a 
short cup. No species of the genus are native in Central America. 

Chrysanthemum coronarium L. Conchita. Cultivated for 
ornament in gardens of the temperate region. Native of Europe. 
A tall, branched, almost glabrous annual with bipinnatifid leaves 
and pale yellow rays. 

Chrysanthemum indicum L. Crisantemo. The common 
chrysanthemum, native of China and Japan, is grown commonly in 
gardens of the temperate region, especially about San Jose", and in 
innumerable forms. The plants thrive here, and flowers grown out 
of doors during the dry season compare not unfavorably in size with 
those produced in greenhouses of the North. 

Chrysanthemum lacustre Brot. Margarita grande. Often 
grown for ornament in the temperate region. Said to be native of 
Portugal. A tall, much branched perennial; leaves ovate-lanceolate 
to linear-lanceolate, dentate; heads very large, with long, white rays. 



1442 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Chrysanthemum Leucanthemum L. Margarita. Frequent 
in pastures on the slopes of Turrialba, Irazu, and Barba. Naturalized 
from Europe. An almost glabrous perennial, usually about 60 cm. 
high ; basal leaves obovate or oblong, dentate or pinnatifid, the cauline 
leaves mostly amplexicaul, narrow, incised or dentate; heads long- 
pedunculate, 2.5-3 cm. broad, with 20-30 long, white rays. The 
ox-eye daisy seems to thrive in the pastures of the volcanoes, to which 
it has been introduced with European grass seed. It is especially 
abundant on the upper slopes of Irazu. 

Chrysanthemum Parthenium (L.) Pers. Altamisa. Feverfew. 
Often grown in gardens for ornament, and naturalized in many 
places, as at Escazu, in pastures, and about Cartago. Native of 
Europe. A cespitose perennial, 60 cm. high or less, strong-scented; 
leaves pinnately parted into ovate or oblong, incised or pinnatifid 
segments; heads numerous, small, forming dense, terminal corymbs, 
the disk scarcely 1 cm. broad, the short rays white. 

CIRSIUM Mill. Thistle 
Reference: Petrak, Beih. Bot. Centralbl. 27, pt. 2: 207. 1910. 

Coarse, perennial or biennial herbs, usually very prickly; leaves 
basal and cauline, the cauline ones alternate, variously lobate or 
pinnatifid; heads large, many-flowered, discoid, mostly white, pink, 
yellowish, or purple; involucre ovoid or globose, the phyllaries im- 
bricate in numerous series, often tipped with prickles; receptacle 
flat or convex, setaceous; achenes obovate or oblong, compressed or 
obtusely 4-angulate, glabrous; pappus of several series of long, 
slender, plumose bristles. A few other species are known from 
northern Central America. 

Cirsium costaricense (Polak.) Petrak, Beih. Bot. Centralbl. 
27, pt. 2: 219. 1910. Cardo. Cnicus costaricensis Polak. Linnaea 41: 
581. 1878. Waste ground, San Jose 1 , Polakowsky 64 (photo, of type 
in Herb. Field Mus.). Frequent in waste ground or thickets of the 
central region; region of San Ramon; chiefly at 1,000-2,000 meters. 
Extending to Honduras. Plants stout, mostly 1-1.5 meters high; 
cauline leaves not or scarcely decurrent, spinose-dentate, sparsely 
arachnoid-tomentose above and green, densely white-tomentose 
beneath; heads in flower about 2 cm. long, rose-purple, the bracts 
tipped with long and slender, rather weak prickles. 

Cirsium mexicanum DC. C. mexicanum var. bracteatum 
Petrak, Beih. Bot. Centralbl. 27, pt. 2: 231. 1910 (Tuis, 620 meters, 



FLORA OF COSTA RICA 1443 

Pittier 11239). Ascending to the Meseta Central, chiefly at 1,200 
meters or less. Ranging to Mexico. Plants usually tall and stout, 
branched; leaves long-decurrent on the stems; heads purple, similar 
to those of C. costaricense, usually glomerate. 

Cirsium pinnatisectum (Klatt) Petrak, Beih. Bot. Centralbl. 
27, pt. 2: 236. 1910. Cnicus pinnatisectus Klatt, Bull. Soc. Bot. Belg. 
35: 292. 1896. Paramos del Abejonal, 1,900 meters, Tonduz 7791. 
Collected also on Irazu, at 2,200 meters. Endemic. Plants about 
2 meters high ; leaves green above and arachnoid-tomentose, densely 
white- tomentose beneath, pinnatisect, very prickly; heads mostly 
solitary, declinate, about 7 cm. in diameter, yellowish; corolla lobes 
about equaling the tube. 

Cirsium subcoriaceum (Less.) Petrak. Cardo, Cardon. Fre- 
quent in the central region, chiefly in the mountains, often grow- 
ing in forest, mostly at 1,100-3,000 meters. Extending to Mexico. 
Plants 1-3 meters tall, branched; leaves not decurrent, green above, 
densely white-tomentose beneath, pinnatifid, very prickly; heads 
rose-red or yellowish, declinate, 4-6 cm. broad, the phyllaries tipped 
with short, stout prickles. A showy and rather handsome plant, 
plentiful at some places in the higher mountains, ascending to the 
upper slopes of Cerro de Las Vueltas. 

CLIBADIUM L. 

Reference: 0. E. Schulz, Beitrage zur Kenntnis der Gattung li- 
badium, Bot. Jahrb. 46: 613. 1912. 

Shrubs or small trees, usually with very rough pubescence; leaves 
opposite, mostly ovate, serrate, triplinerved ; heads very small, disci- 
form, few-many-flowered, greenish or whitish, disposed in a sub- 
corymbose panicle; involucre of 1-6 phyllaries, these ovate or rounded, 
concave, nerved; receptacle paleaceous near the margin, sometimes 
naked in the center; marginal flowers pistillate, fertile, the inner ones 
hermaphrodite, sterile; achenes obovoid, compressed, at maturity 
succulent and drupaceous; pappus none. A few other species are 
found in Central America. 

Clibadium acuminatum Benth. Bot. Voy. Sulph. 114. 1844. 
Cocos Island, Barclay. Endemic. Leaves petiolate, ovate, about 15 
cm. long and 7 cm. wide, long-acuminate, duplicate-serrate, very 
scabrous with minute hairs on both surfaces, cuneate at the base; 
panicles lax, corymbiform, the branches divaricate; heads scarcely 
4 mm. long, scabrous-puberulent ; fertile flowers about 5; achenes 
obscurely puberulent at the apex. 



1444 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Clibadium anceps Greenm. Proc. Amer. Acad. 39: 97. 1903. 
Forests of La Palma, 1,460 meters, Tonduz 12537. Also in the 
regions of San Ramon, Navarro, and El General, at 1,400 meters or 
less. Endemic. A small tree, the ultimate branches complanate, 
strigose; leaves ovate, 8-12 cm. long, 4-6 cm. wide, acuminate, 
cuneate at the base, remotely denticulate, hirtellous, above finally 
glabrate and somewhat lustrous; heads 5 mm. high, glomerate; 
phyllaries appressed-pubescent; fertile flowers usually 3; achenes 
2 mm. long, glabrous; sterile flowers 3-5. 

Clibadium glomeratum Greenm. Proc. Amer. Acad. 39: 98. 
1903. Forests of Tuis ("Luis" in the original publication), 650 meters, 
Tonduz 11508. Also on the slopes of Barba, at 2,000 meters or less. 
Endemic. A shrub of 1.5-2.5 meters, the branches pilose with short, 
spreading hairs; leaves broadly ovate, 10-20 cm. long, 5-15 cm. wide, 
acute, obtuse to subcordate at the base, remotely serrulate, rough- 
pubescent above, densely hirtous beneath; branches of the panicle 
ferruginous- tomentose, the heads glomerate, sessile, about 2 mm. 
long; phyllaries 2, orbicular, ciliate at the apex; fertile flowers 4, 
the sterile 5; achenes 2 mm. long, villous at the apex. 

Clibadium grande Blake, Contr. U. S. Nat. Herb. 22: 601. 
1924. La Florida, Atlantic tierra caliente, 80 meters, Pittier 11280. 
Leaves long-petiolate, broadly ovate, about 30 cm. long and wide, 
acute, broadly rounded at the base, dentate-serrate, scabrid-strigil- 
lose above, strigillose beneath; panicles sordid-tomentulose, 9 cm. 
long, the heads sessile, not glomerate, 5.5 mm. long; phyllaries 3, 
strigillose; pistillate flowers 6, the perfect flowers 8-9; achenes 2 mm. 
long, pilose at the apex. 

Clibadium grandifolium Blake, Contr. U. S. Nat. Herb. 22: 
599. 1924. Rio Pacuare, Llanuras de Santa Clara, 150 meters, J. 
D. Smith 16068. Atlantic tierra caliente. Endemic. Branches 
strigose; leaves long-petiolate, the blades broadly ovate or rounded- 
ovate, 15-28 cm. long, 12-23 cm. wide, short-pointed, truncate- 
rounded at the base and cuneately narrowed to the petiole, serrulate, 
scabrid above, strigillose beneath; panicles 17-24 cm. wide, the heads 
irregularly approximate, sessile, 5-6.5 mm. high; phyllaries 2-3, 
acute or acuminate, strigillose; fertile flowers 8-11, the sterile 9-10; 
achenes pilose at the apex. 

Clibadium leiocarpum Steetz. C. Pittieri Greenm. Proc. 
Amer. Acad. 39: 98. 1903 (La Florida, Atlantic tierra caliente, 80 
meters, Pittier 11290). C. terebinthaceum (Sw.) DC. var. Pittieri 



FLORA OF COSTA RICA 1445 

0. E. Schulz, Bot. Jahrb. 46: 626. 1912. Atlantic tierra caliente, 
ascending to the central region, at least on the lower slopes of Poas. 
Panama. A shrub of 1.5-2.5 meters, the branches strigose; leaves 
ovate, 10-15 cm. long, 4-8 cm. wide, acuminate, narrowed to the 
base, dentate above the middle, hirsute-pubescent on both surfaces; 
heads 5 mm. high, the phyllaries narrowly ovate, acute, strigose; 
fertile flowers 7-9; achenes pubescent at the apex. C. terebinthaceum, 
to which Schulz referred this plant, is a species of Jamaica and 
Colombia. C. leiocarpum var. strigosum Blake, Journ. Wash. Acad. 
27: 382. 1937 (Cerro de Piedra Blanca, above Escasu, Prov. San Jose", 
Standley 32593} is a form with strigose pubescence. The reduction 
of C. Pittieri to synonymy under C. leiocarpum is made at the sug- 
gestion of Dr. S. F. Blake. 

Clibadium polygynum Blake. C. Pittieri f. phrixium Greenm. 
Proc. Amer. Acad. 40: 38. 1904 (Puerto Viejo, Rio Sarapiqui, Biolley 
7399). Forests and thickets of the tierra caliente; region of San 
Ramon; Guanacaste. Extending to Guatemala. Branches densely 
hispidulous with spreading hairs; leaves ovate or ovate-lanceolate, 
7-9 cm. long, 2.5-4 cm. wide, acuminate, cuneate at the base, serrate, 
scabrous and hispidulous on both surfaces; panicles mostly small and 
dense, the heads short-pedicellate, 5 mm. high; phyllaries usually 2, 
subacute, strigose; fertile flowers about 29, the sterile about 8; 
achenes densely pilose at the apex. 

Clibadium Schulzii Blake, Contr. U. S. Nat. Herb. 22: 602. 
1924. Copey, 1,800 meters, Tonduz 11915. Frequent in thickets 
of the central region, and in Dota. Endemic. A shrub, or sometimes 
a tree of 7 meters, with a dense, rounded crown; branches densely 
pilose- tomentose with ascending hairs; leaves short-petiolate, lance- 
ovate, 6-9 cm. long, 2-3 cm. wide, acuminate, acutely cuneate at 
the base, serrulate, strigose above, densely pilose beneath with 
subappressed hairs; panicles 3-6 cm. wide, the heads on pedicels 2 
mm. long or less, 3.5-4 mm. high; phyllaries 3, obtuse or subacute, 
glabrate; fertile flowers 5-6, the sterile 12-14; achenes sparsely pilose 
at the apex. 

Clibadium surinamense L. Thickets of the Atlantic tierra 
caliente, ascending to the region of Cartago, at 1,200 meters or less; 
El General. Ranging to West Indies and the Guianas. A shrub of 
1-2 meters, the branches strigose; leaves short-petiolate, oblong- 
ovate, acute or acuminate, rounded at the base and cuneate-attenu- 
ate to the petiole, serrulate, 5-17 cm. long, 2-8 cm. wide, very 



1446 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

scabrous above, more softly pilose beneath; heads sessile, 5 mm. long; 
phyllaries 3-4, subacute, short-pilose; fertile flowers 3-6, the sterile 
8-13; achenes glabrous, black. 

Clibadium villosum Benth. Thickets of the Atlantic tierra 
caliente, ascending to the Meseta Central, at 1,200 meters or less. 
Colombia. Similar to C. surinamense; branches densely villous- 
tomentose; leaves scabrous above, villous-tomentose beneath; 
branches of the panicle densely tomentose; heads 4-4.5 mm. long; 
fertile flowers 3-4, the sterile 9-14. 

CONYZA L. 

Annual or perennial herbs, the leaves alternate, sometimes 
mostly basal, varying from entire to pinnatisect; heads small, 
usually numerous and paniculate or corymbose, disciform, the outer 
flowers pistillate and fertile, the inner ones perfect and all or mostly 
fertile; involucre campanulate, the phyllaries appressed, linear or 
lanceolate, the outer gradually shorter; receptacle plane or convex, 
not paleaceous; achenes compressed, truncate, sometimes costate; 
pappus of numerous slender, soft bristles. A few other species 
probably occur in Central America. 

Conyza chilensis Spreng. In pastures or thickets, Meseta 
Central and nearby mountain slopes; region of San Ramon. Widely 
distributed in tropical America. Stems erect, mostly about 40 cm. 
high or less, pilose, simple or sparsely branched; lower and basal 
leaved oblanceolate or linear-oblanceolate, coarsely crenate to lyrate- 
pinnatifid, the cauline leaves linear and entire, rough-pubescent; 
heads few, racemose or subcorymbose, long-pedunculate, the in- 
volucre about 1 cm. broad; achenes 1 mm. long, 1 -nerved on each face; 
pappus brownish. 

Conyza coronopifolia HBK. Pastures on the lower slopes of 
Irazu, perhaps introduced. Extending to Mexico. Plants probably 
perennial, the stems often several from the base, usually abundantly 
branched ; cauline leaves sessile, most of them deeply pinnate-lobate, 
the lobes entire, broad, obtuse, hispidulous; heads whitish, about 
8 mm. broad, rather closely clustered at the ends of the branches, 
the phyllaries linear, pilose; pappus more or less fulvous. 

Conyza lyrata HBK. Thickets of the Pacific coast. Widely 
distributed in tropical America. A coarse annual, sometimes a 
meter high, villous-pilose and very viscid; cauline leaves obovate, 
dentate and lyrate-lobate; heads greenish white, 7 mm. high, in 



FLORA OF COSTA RICA 1447 

broad, leafy panicles; pappus white or nearly so. A common weed 
along the Pacific coast of Central America. The plant has a strong 
and offensive odor. 

COREOPSIS L. 

Glabrous or pubescent herbs; cauline leaves opposite, most 
of the leaves often basal, entire to dissected; heads of medium or 
rather large size, yellow, solitary at the ends of the stems or laxly 
corymbose-paniculate radiate; phyllaries commonly 2-seriate, more or 
less connate at the base, the outer ones generally herbaceous, the 
inner membranaceous; ray flowers commonly neutral, the disk 
flowers mostly fertile; disk paleaceous; achenes obcompressed, mostly 
oblong, often 2- winged; pappus none or of 2 awns or teeth. One or 
more species are native in northern Central America. 

Coreopsis lanceolata L. Chispa, Margarita amarilla. Some- 
times planted for ornament in gardens of the temperate region. 
Native of the United States. An erect perennial, about 60 cm. high, 
glabrous or nearly so, slender; leaves mostly clustered at the base 
of the stems, long-petiolate, oblanceolate-linear; stems scapose, each 
bearing a single large, yellow head with long, spreading rays. 

Coreopsis tinctoria Nutt. Cultivated at Zarcero, and said to 
reproduce itself abundantly. Native of southwestern United States. 
A slender, erect annual, usually much branched; leaves parted into 
numerous narrow segments, the stems leafy; heads large and showy, 
with bright yellow rays. 

COSMOS Cav. 

Annual or perennial herbs; leaves opposite, mostly pinnatifid; 
heads usually rather large, long-pedunculate, radiate, the disk flowers 
yellow; involucre biseriate; ray flowers neutral, the disk flowers 
perfect and fertile; receptacle flat, paleaceous; achenes narrow, 
somewhat angulate, distinctly rostrate; pappus of 2-4 retrorsely 
barbate or hispid awns. 

Cosmos caudat us HBK. Occasional as a weed in waste ground 
of the tierra caliente; region of San Ramon; at 700 meters or less. 
Widely distributed in tropical America. Plants usually a meter high 
or less, branched, sparsely pilose or glabrate; leaves bipinnate, the 
divisions ovate to linear-lanceolate; involucre 12-15 mm. high, the 
outer phyllaries linear-lanceolate, ciliate; rays pink, 1-2 cm. long; 
achenes linear, 10-15 mm. long. 



1448 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Cosmos sulphureus Cav. Cambray. Sometimes planted for 
ornament in gardens of the temperate region. Native of Mexico. 
Somewhat similar to the preceding species, but the rays orange- 
yellow. 

CREPIS L. 

Herbs, the leaves alternate or basal, mostly dentate or pinnatifid ; 
sap milky; heads small or medium-sized, usually paniculate or corym- 
bose; involucre cylindric to campanulate, the principal phyllaries 
1-seriate and equal, some short ones present at the base of the 
involucre; receptacle generally flat, naked or short-fimbrillate; rays 
truncate and dentate at the apex; achenes linear-oblong, 10-20- 
costate, narrowed at the base and apex, not rostrate; pappus of 
slender, soft, white bristles. No species are native in Central 
America. 

Crepis capillaris (L.) Wallr. Naturalized in pastures, La 
Palma de San Ramon. Introduced from Europe. Plants annual, 
60 cm. high or less, leafy, corymbosely branched above, glabrous, 
or somewhat hirsute below; basal leaves spatulate, pinnatifid or 
dentate; cauline leaves lanceolate or oblong, amplexicaul; heads 
1-1.5 cm. wide, slender-pedunculate; involucre 6-8 mm. long, pubes- 
cent or glandular-pubescent, the phyllaries lanceolate; achenes 10- 
costate. 

Crepis heterophylla Klatt, Leopoldina Bot. Beibl. 8. 1895. 
San Rafael de Cartago, 1,500 meters, Pittier 6994- Plants nearly 
glabrous, the stems erect, striate, leafy, bearing 1 or 2 heads, 25-65 
cm. high; leaves 6 cm. long and 2.5 cm. wide, cordate-lanceolate, 
cordate-spatulate, or cordate, sinuate-dentate, mucronate-cuspidate, 
auriculate-amplexicaul, or petiolate; petioles broadly winged; 
peduncles glandular-hispid, bracteate at the base, the bracts lanceo- 
late, pectinate-dentate; involucre campanulate, the phyllaries lanceo- 
late, white-pilose, the inner ones 1 cm. long; achenes villous. I have 
summarized Klatt's description, but have seen no material of the 
plant he describes. In view of his many errors in referring Composi- 
tae to genera with which they had no close relationship, it is unwise 
to speculate upon the proper position of the present plant, whose 
description does not suggest to me any of the Cichorieae listed here 
for Costa Rica. 

DAHLIA Cav. 

Tall, coarse, perennial herbs, sometimes treelike and more or less 
ligneous, glabrous or pubescent; leaves opposite, variously pinnate- 



FLORA OF COSTA RICA 1449 

parted; heads large, long-pedunculate, radiate, the disk flowers 
yellow; ray flowers neutral, the disk flowers fertile; involucre biseri- 
ate, the outer phyllaries short, subfoliaceous, the inner membrana- 
ceous; receptacle plane, paleaceous; achenes glabrous, oblong or 
obovate, compressed, rounded at the apex; pappus none. One or 
more additional species are native in northern Central America. 

Dahlia rosea Cav. Dalia, Catarina, Catalina. D. dumicola Klatt, 
Bull. Soc. Bot. Belg. 35: 289. 1896 (in hedge, San Francisco de Guada- 
lupe, Pittier & Tonduz 8469). Common in the central region, especi- 
ally in hedges, often an abundant weed in cornfields; sometimes 
found in forest. Extending to Mexico. A tall, coarse herb or shrub, 
usually about 3 meters high, glabrous or nearly so, too well known to 
need special description here. In the Meseta Central the common 
dahlia behaves almost exactly like the sunflowers ( Helianthus annuus) 
of the central and western United States. It takes possession of the 
cornfields when the corn is well grown, and often is quite as con- 
spicuous as the corn itself. The wild plants have white or more often 
pink heads, which are either single or double-flowered. The wild 
plants with very large, white rays have been referred to D. Maxonii 
Safford, described from Guatemala, but I am inclined to believe that 
material so named, at least that from Costa Rica, is better referable 
to D. rosea (D. variabilis Desf.). Some of the better varieties of 
cultivated dahlias are, of course, to be found in Costa Rican gardens, 
and the common wild form likewise is common in village gardens. It 
is curious to observe that the country people give the name dalia to 
the garden plants, while the wild ones are called Catarina or Catalina. 
When asked what may be the differences between the plants receiving 
these two names, they always retort with some surprise that dalias 
are garden plants, while Catarina is a weed. 

DELILIA Spreng. 

Rather low and slender, annual herbs, branched ; leaves opposite, 
petiolate, dentate; heads numerous, small, pedicellate, aggregate at 
the ends of the branches, radiate; involucre compressed, the 2-4 
phyllaries membranaceous, the outer one large and broad, herbaceous, 
concealing the flowers; ray flowers 1-3, pistillate, fertile, the disk 
flowers 1-4, perfect, sterile; achenes obovate, compressed; pappus 
none. A single species is found in North America. In general 
appearance it is quite unlike most Compositae, and suggestive rather 
of some groups of Acanthaceae. 



1450 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Delilia biflora (L.) Kuntze. Elvira biflora Cass. Frequent in 
waste ground, thickets, or pastures, often in cultivated land, Meseta 
Central to the Pacific coast, and probably also in the Atlantic tierra 
caliente. Plants mostly 30-50 cm. high, much branched, the pubes- 
cence rather sparse or dense, of short, whitish, appressed or spreading 
hairs; leaves petiolate, mostly 3-5 cm. long, ovate to lanceolate, acute, 
3-nerved, incised-crenate; outer phyllaries green, rounded-cordate, 
4-6 mm. long, persistent; achenes glabrous, 2 mm. long. 

DIMORPHOTHECA Moench 

Annual or perennial herbs, glabrous or pubescent; leaves alternate 
or mostly basal, entire to incised, usually narrow; heads of medium 
or large size, long-pedunculate; disk flowers yellow to purplish, the 
rays of the same colors or white; heads radiate, the ray and outer 
disk flowers fertile; involucre broad, the phyllaries 1-2-seriate, sub- 
equal, linear, acuminate, scarious-margined; receptacle plane or sub- 
convex, naked; achenes glabrous, those of the ray subtrigonous, 
rugose or muricate, those of the disk compressed, the margin some- 
times winglike. All the species are natives of southern Africa. 

Dimorphotheca annua Less. Grown frequently for ornament 
in gardens of the temperate region, as elsewhere in Central America. 
Native of South Africa. A diffuse, simple or branched annual, gray- 
pubescent; leaves narrowly oblong or obovate-oblong, tapering to the 
base, with a few coarse, remote teeth; heads nodding, the flowers 
white and purple. 

DIPLOSTEPHIUM HBK. 

Reference: Blake, Key to the genus Diplostephium, with descrip- 
tions of new species, Contr. U. S. Nat. Herb. 24: 65. 1922. 

Low shrubs, more or less tomentose or lanate; leaves alternate, 
usually entire and coriaceous; heads small, solitary at the ends of the 
branches or corymbose-paniculate, radiate, the rays white to purple; 
involucre campanulate, the phyllaries several-seriate, graduated, 
with thin margins, usually somewhat recurved in age; receptacle flat 
or subconvex; ray flowers fertile, the disk flowers perfect, fertile or 
sterile; achenes 3-5-costate, glabrous, glandular, or hispid; pappus 
copious, mostly biseriate, the outer of short bristles or squamellae, 
the inner longer, setose. A single species is found in North America. 

Diplostephium costaricense Blake, Contr. U. S. Nat. Herb. 
24: 82. pi. 27. 1924. Cerro de La Muerte, 3,100 meters, Pittier 
10459. Collected also on Cerro de Las Vueltas, and in forest at 



FLORA OF COSTA RICA 1451 

La Palma de San Jose", 1,500 meters. Endemic. A dense shrub of 
4.5 meters or less; leaves oblanceolate or linear-oblanceolate, 2.5-4 
cm. long, subacute, narrowed to the sessile base, entire, grayish- 
tomentulose; heads 6-9 mm. high, in dense panicles, the rays white, 
the disk purple-red; achenes 1.5-2.5 mm. long, densely glandular. 
The Costa Rican plant has been referred to D. Schultzii Wedd., a 
Colombian species, and D. rupestre (HBK.) Wedd., of Colombia and 
Ecuador. The shrub is a characteristic one of the low, wet thickets 
at the upper limit of vegetation on the central volcanoes. 

DYSSODIA Cav. 

Herbs or low shrubs, usually with conspicuous oil glands; leaves 
opposite or alternate, entire to dissected; heads small or medium- 
sized, radiate or rarely discoid, yellow or orange; involucre usually 
campanulate, the principal phyllaries 1-seriate, narrow, equal, more 
or less connate, glandular-punctate, a few small ones often present 
at the base; achenes slender; pappus of 5-20 paleae, these often awned 
or aristate-dissected. Two or three other species are found in north- 
ern Central America. 

Dyssodia montana (Benth.) Gray. Girasolillo. Clomenocoma 
monlana Benth. Frequent in thickets or pastures, Meseta Central, de- 
scending on the Pacific slope almost or quite to the coast; region of 
San Ramon; Guanacaste; ascending rarely to 1,800 meters. Panama 
to Guatemala. A rather strong-scented, erect, perennial herb, a 
meter high or less, almost glabrous; leaves opposite, practically 
simple, the lateral divisions reduced to subulate lobes or bristles on 
the petiole, the blades ovate or lanceolate, 3-5 cm. long, serrate or 
laciniate, mostly acuminate, minutely strigillose beneath; heads 
solitary on long, bracteate peduncles; involucre 1.5 cm. high; rays 
orange, about 1 cm. long; achenes sericeous- villous, 3 mm. long, the 
pappus of 10 dissected squamellae 7-8 mm. long. Costa Rican mate- 
rial of this species has been referred incorrectly to D. grandiflora DC. 

ECLIPTA L. 

Erect or prostrate, branched, annual herbs, pubescent; leaves 
opposite, small; heads small, pedunculate, terminal and axillary, 
radiate, whitish; involucre broadly campanulate, the phyllaries 
imbricate, about 2-seriate, subequal or the outer longer; receptacle 
plane or convex, paleaceous; ray flowers pistillate and fertile, the 
disk flowers perfect and mostly fertile; achenes trigonous or com- 
pressed; pappus none or of a few small teeth. A single species is 
found in tropical America. 



1452 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Eclipta alba (L.) Hassk. Frequent in waste or cultivated 
ground, usually in moist places, Meseta Central to the coasts. Gen- 
erally distributed in tropics of both hemispheres, also in temperate 
regions. Plants usually prostrate or decumbent, the pubescence 
minute, appressed, rough; leaves lanceolate or linear-lanceolate, 
acute or acuminate, 2-10 cm. long, denticulate or entire, sessile, or 
the lower ones petiolate; heads 6-12 mm. broad, numerous but incon- 
spicuous, subsessile or slender-pedunculate, the very short rays 
easily overlooked. This is one of the most common and widely dis- 
tributed of all tropical weeds. 

EGLETES Cass. 

Low herbs, usually viscid and densely pubescent, branched; 
leaves alternate, dentate or lobate; heads small, pedunculate, radiate, 
terminal or opposite the leaves; involucre hemispheric, the phyllaries 
few-seriate, imbricate; receptacle ovoid or conic, naked; flowers per- 
fect; achenes oblong, compressed, smooth; pappus reduced to a den- 
tate or ciliate, cartilaginous ring. A single species is known from 
Central America. 

Egletes viscosa (L.) Less. Guanacaste (Bebedero), collected 
by Brenes. Rather widely distributed in Central America, but 
apparently of infrequent occurrence. Plants commonly 50 cm. 
high or less and densely dichotomous-branched, the abundant pubes- 
cence of short, spreading, viscid hairs; leaves small, broadly obovate, 
coarsely dentate or sublobate, toward the base often lyrate-lobate; 
heads about 5 mm. broad, yellow, the minute rays white. 

ELEPHANTOPUS L. 

Reference: Gleason, N. Amer. Fl. 33: 106. 1922. 

Perennial herbs, the stems sometimes scapiform; leaves alternate, 
but often chiefly basal, entire or dentate; heads small, mostly in 
glomerules that are pedunculate and corymbose, subtended by 1-3 
foliaceous bracts, 1-5-flowered; involucre of 2 decussate pairs of 
phyllaries; achenes truncate, usually 10-costate; pappus of 5-8 rigid 
scales, these prolonged into setae. No other species are known from 
Central America. 

Elephantopus hypomalacus Blake, Contr. Gray Herb. 52: 
20. 1917. Orotina, 180 meters, Holway 314- Frequent in pastures 
or thickets of the tierra caliente; region of San Ramon; Guanacaste; 
at 900 meters or less. Extending to Mexico. Plants with leafy, 
branched stems, mostly 60 cm. high or less, pilose with chiefly long, 



FLORA OF COSTA RICA 1453 

spreading hairs; cauline leaves broadly obovate to oblong-obovate, 
10-20 cm. long, acute, inconspicuously serrate, densely and softly 
pilose beneath; glomerules of heads numerous, 1 cm. long; involucre 
8 mm. long; achenes 3.5 mm. long, pilose; pappus bristles 4-5 mm. 
long, gradually dilated below to a narrowly triangular base. Some 
of the Costa Rican material has been referred to E. mollis HBK., a 
species ranging from Mexico into South America, but all the speci- 
mens from Costa Rica which I have examined seem to be referable 
rather to E. hypomalacus, if this species is really distinct from E. 
mollis, which is rather doubtful. 

Elephantopus scaber L. Hierba de San Antonio, Lechuguilla, 
Escobilla, Achicoria. Common in pastures and thickets of the central 
region, extending into the upper part of the tierra caliente; Aguacate; 
region of San Ramon. Believed to have been introduced from the 
East Indies; established also in Guatemala. Plants mostly 30-40 
cm. high, simple or branched; leaves all or chiefly basal, oblong-obo- 
vate, 8-12 cm. long, obtuse, densely pilose beneath; achenes 2.5 
mm. long. The plant has every appearance of being perfectly at 
home in central Costa Rica, and it is hard to believe that it is an intro- 
duced species. The flower heads are white, as in the preceding species. 
A decoction made from the leaves of the plant is one of the local 
remedies for dysentery. 

ELEUTHERANTHERA Poit. 

Low, more or less hirsute, perennial herbs; leaves thin, opposite, 
petiolate, dentate; heads small, usually discoid, terminal and axillary, 
the flowers all perfect, or a few neutral flowers with very small rays 
sometimes present; involucre broadly campanulate, the phyllaries 
few, unequal; receptacle convex, paleaceous; achenes thick, oblong- 
obovate, somewhat compressed, rounded at the apex; pappus cupu- 
lar, ciliate-dentate or with 2-3 short awns. The genus consists of a 
single species. 

Eleutheranthera ruderalis (Swartz) Schultz Bip. Waste 
ground or thickets of the Atlantic tierra caliente (Changuinola Val- 
ley). Widely distributed in tropical America. Plants 60 cm. high 
or less, often much branched, sometimes diffusely spreading; leaves 
ovate to oblong-lanceolate, 6 cm. long or less, 3-nerved, acute, 
rounded or narrowed at the base; peduncles 2-12 mm. long; involu- 
cre 5-6 mm. high, the phyllaries oblong or ovate-oblong, obtuse; 
achenes 2-3 mm. long. 



1454 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

EMILIA Cass. 
Reference: Garabedian, A revision of Emilia, Kew Bull. 137. 1924. 

Somewhat succulent herbs; leaves alternate and basal, the cauline 
ones often amplexicaul; heads small, long-pedunculate, discoid, pink 
to purple, orange, red, or white, solitary or laxly corymbose; involu- 
cre cylindric or campanulate, the phyllaries linear, uniseriate, equal ; 
receptacle plane, naked; achenes sub terete or 5-angulate; pappus of 
numerous soft, white, capillary bristles. The species are natives of 
the Old World. 

Emilia sagittata (Vahl) DC. E. coccinea Sweet. Sometimes 
grown in gardens of the Meseta Central, and perhaps escaping from 
cultivation. Probably native of tropical Africa, but occasionally 
naturalized in tropical America. Plants slender, erect, sparsely vil- 
lous; cauline leaves dentate or entire; heads dark red, the flowers 
conspicuously longer than the involucre. 

Emilia sonchifolia (L.) DC. Naturalized in pastures or waste 
ground of the tierra caliente, but apparently infrequent. Native 
of the Old World tropics; widely naturalized in tropical America. 
A slender annual, glabrous or sparsely pubescent, pale, branched; 
cauline leaves repand-dentate to lyrate-pinnatifid; heads cylindric, 
the involucre 8-12 mm. long, the flowers pale purple or pink, scarcely 
exceeding the involucre. 

ERECHTITES Raf. 

Coarse, erect, simple or branched herbs; leaves alternate, coarsely 
dentate to deeply pinnatifid; heads rather small, disciform, whitish 
or purple, corymbose-paniculate; involucre cylindric, the principal 
phyllaries 1-seriate, linear, subequal, sometimes with a few very 
short ones at the base; receptacle concave, naked; flowers all fertile; 
achenes linear-oblong, angulate or striate; pappus capillary, of 
abundant, long, soft, capillary bristles. No other species are 
known from Central America. 

Erechtites hieracifolia (L.) Raf. E. carduifolia DC.(?). Fre- 
quent in waste or cultivated ground, sometimes in forest, Meseta 
Central and upper part of the Pacific tierra caliente; region of San 
Ramon; ascending rarely to 1,800 meters. Widely distributed in 
tropical and temperate America. Plants pale green, slightly suc- 
culent, usually a meter high or less, simple or sparsely branched, 
hirsute or glabrate; leaves mostly lanceolate or ovate-lanceolate, 
coarsely dentate and the lower ones sometimes incised, 5-20 cm. 



FLORA OF COSTA RICA 1455 

long, the upper sessile and auriculate-clasping, the lower narrowed 
to slender petioles; heads 12-20 mm. long, whitish; pappus white. 
The Central American plants referred to this species are somewhat 
variable and may represent two or more species or varieties. The 
proper name to be used for the tropical plant is somewhat uncertain. 
Probably it is not true E. hieracifolia, which is properly a northern 
species. 

Erechtites valerianaefolia (Wolf) DC. Eupatorium angosturae 
Polak. Linnaea 41: 575. 1877 (Angostura, Polakowsky 479). Fre- 
quent in thickets or forest, Meseta Central and especially on the 
nearby forested mountain slopes, chiefly at 1,200-2,450 meters; 
Guanacaste. Widely distributed in tropical America. Plants 
mostly 1-2 meters high, thinly villous or glabrate, simple or branched ; 
leaves mostly pinnate-lobate, the lobes often numerous and narrow, 
incised; heads numerous, bright purple; pappus pink or purple. 
This species is much more showy than E. hieracifolia, but both are 
decidedly unattractive weeds. 

ERIGERON L. 

Annual or perennial herbs, simple or branched; leaves alternate, 
or sometimes chiefly basal, entire to pinnatifid; heads very small to 
medium-sized, paniculate or solitary, radiate (rays sometimes very 
small); involucre campanulate, the phyllaries linear, subequal, 1-3- 
seriate; receptacle almost plane, naked; rays white, pink, or purple, 
pistillate, the disk flowers yellow, perfect; achenes compressed, 1-2- 
nerved; pappus of numerous slender bristles in 1 or 2 series. Other 
species are found in Central America. 

Erigeron adenophorus Greenm. Proc. Amer. Acad. 39: 94. 
1902. Senecio eriocephalus Klatt, Bull. Soc. Belg. 31: 212. 1892. Be- 
tween Rancho del Jaboncillal and summit of Cerro de La Muerte, 
3,100 meters, Pittier 3425. Known only from the original collection. 
Plants suffruticose, branched, the branches hirsute, glandular- 
pubescent; leaves sessile, amplexicaul, at first erect or spreading, 
later reflexed, linear-lanceolate, 1-2 cm. long, acute, sparsely dentate, 
densely glandular-pubescent and with long, flaccid hairs; heads 
forming dense cymes, about 1 cm. high, the outer phyllaries covered 
with long, matted hairs; achenes puberulent. 

Erigeron bonariensis L. Frequent in thickets or waste ground 
of the Meseta Central, and doubtless at lower elevations; reported 
from El General. Widely distributed in tropical America. Plants 



1456 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

rather slender, erect, often a meter high, branched or almost simple, 
hispidulous or glabrate; lower leaves lanceolate or oblanceolate, 
acute, sometimes 10 cm. long and 1.5 cm. wide, but usually much 
smaller, acute or acuminate, dentate or somewhat pinnate-lobate, 
the upper leaves lanceolate or linear, usually entire; heads in large 
but rather narrow panicles, very numerous, 5 mm. high; phyllaries 
acuminate, hispidulous; pappus fulvous or brownish. A common 
weed in many regions of Central America. This species has been 
reported from Costa Rica under the name Conyza floribunda HBK. 

Erigeron irazuensis Greenm. Proc. Amer. Acad. 40: 36. 1904. 
La Playita, Volcan de Irazu, 3,300 meters, Pittier 14075. Endemic, 
and known only from the slopes of Irazu, growing usually on wet 
banks, at 1,800 meters or more. Plants somewhat ligneous at the 
base, the stems erect or ascending, 10-20 cm. long, pilose; leaves 
spatulate to oblanceolate, 1-3.5 cm. long, apiculate-acute, somewhat 
dentate, gradually narrowed to a marginate petiole, pilose-hirsute; 
heads few, 10-12 mm. high, terminating slender peduncles; involucre 
pubescent; rays pink or rose-red; achenes pubescent. The plant 
was reported from Costa Rica by Klatt as E. nevadensis Wedd. 

Erigeron jamaicensis L. Collected at Las Concavas, south 
of Cartago, and in the region of San Ramon at 1,050 meters. Hon- 
duras to Panama; Greater Antilles. Plants very slender, the stems 
suberect or decumbent, simple or sparsely branched, pubescent; 
leaves mostly basal, spatulate, usually entire or subentire, some- 
times deeply lobate, mostly 2 cm. long or less, those of the stems 
chiefly linear and bractlike, usually entire; heads few, 4-5 mm. high, 
the bracts narrowly scarious-marginate; rays white, small. This 
is presumably the plant listed by Klatt as E. cuneifolius DC. 

Erigeron Karvinskianus DC. E. mucronatus DC. Frequent 
in forest or on brushy slopes about Escazu, at 1,800 meters, and in 
the region of Cartago. Panama to Mexico. Plants perennial, 
erect or ascending, branched, the stiff branches strigose or almost 
glabrous, densely leafy; leaves lanceolate to narrowly spatulate, 
mostly 2.5 cm. long or less, short-petiolate, acute, remotely lobate- 
dentate or entire, sparsely strigose or glabrate; heads few, long- 
pedunculate, the involucre scarcely 5 mm. long, the numerous 
slender rays white or pinkish. 

Erigeron pusillus Nutt. Frequent in thickets or waste ground 
about the Meseta Central, and probably also in the tierra caliente; 



FLORA OF COSTA RICA 1457 

often in cultivated places. Widely distributed in tropical and tem- 
perate America. Plants slender, usually a meter high or less, simple 
or branched, the stems glabrate; lower leaves spatulate, dentate or 
entire, the upper ones linear, entire, ciliate; heads very numerous, 
forming a narrow panicle, greenish white; involucre only 2-3 mm. 
high; rays very short and inconspicuous, often overlooked. The 
plant is closely similar to E. canadensis L., a common species of 
North America, with which it is commonly confused, but is nearly 
or quite glabrous, and the phyllaries have purplish tips. 

Erigeron spathulatus Vahl. Common in waste ground or 
thickets, often in cultivated land, Meseta Central to the coasts; 
rarely ascending to 1,800 meters or more. Widely distributed in 
tropical America, and in the tropics of the Old World. Plants erect, 
rather stout, usually a meter high or less, often much branched, 
abundantly short-pilose; lower leaves spatulate to obovate, coarsely 
dentate, 10 cm. long or less, on marginate petioles, the upper leaves 
smaller, mostly obtuse, sometimes entire; heads whitish, in open, 
terminal corymbs, 4-5 mm. high. 

Erigeron subspicatus Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 82. 1853. Volcan de Irazu, 2,400 meters, Oersted. Slopes of 
Volcan de Irazu, and in the region of Dota, usually in pastures or on 
open banks, at 2,400-3,400 meters. A variety of the species has 
been described from Guatemala. Plants erect, simple, about 30 cm. 
high, scabrous-puberulent and pilose-lanate, whitish; leaves dilated 
at the base and more or less decurrent, linear-oblong or lanceolate, 
2 cm. long or less, with a few coarse teeth; heads small, numerous, 
forming a dense spike along the upper part of the stem, the spike 
leafy-bracted ; rays minute and inconspicuous, white. 

EUPATORIUM L. 

Shrubs or small trees, sometimes annual or perennial herbs; 
leaves mostly opposite, very variable in form; heads discoid, red to 
purple, blue, or white, with few to many flowers, usually arranged 
in corymbose or thyrsoid panicles; achenes columnar to obovoid, 
5-costate or 5-angulate; pappus of numerous slender bristles. A 
vast genus in tropical America, with many other species in Central 
America. 

Eupatorium Allenii Standl., sp. nov. Herba erecta metralis, 
caule subtereti vel obtuse hexangulari infra inflorescentiam glabro, 
internodiis superioribus elongatis foliis subaequalibus; folia modica 



1458 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

opposite petiolata membranacea, petiolo usque 1 cm. longo cras- 
siusculo sparse pilosulo vel fere omnino glabro; lamina anguste 
rhombico-ovata vel oblongo-lanceolata ca. 8 cm. longa atque 3 cm. 
lata, longe sensim attenuato-acuminata, basi cuneata, in toto mar- 
gine subgrosse crenata, in sicco fusca, supra viridis tantum ad 
costam venasque minute puberula, subtus fere concolor, ad costam 
nervosque minute adpresso-pilosula vel fere glabra; inflorescentia 
ampla foliosa corymbosa ca. 15 cm. longa atque aequilata, ramis 
valde adscendentibus oppositis sparse sordido-puberulis, capitulis 
numerosis parvis in cymulas sat densas subglobosas aggregatis, 
pedicellis gracilibus plerumque 3-4 mm. longis, bracteis parvis 
linearibus; involucrum campanulatum ca. 4 mm. altum, phyl- 
lariis subbiseriatis linearibus, interioribus angustioribus apicem 
versus attenuatis subacutis glabratis, extimis paullo brevioribus 
sparse adpresso-pilosulis vel fere glabris; flores 20 vel paullo ultra 
albi; achaenia linearia nigra fere 2 mm. longa 4-angulata ad angulos 
hirtella; pappi setae numerosae albae 3.5 mm. longae. In thickets, 
summit of Volcan de Poas, 2,575 meters, P. H. Allen 597 (type in 
Herb. Field Mus.). The present plant has much the general appear- 
ance of Archibaccharis asperifolia (Benth.) Blake, with which I at 
first confused it. Although it has no particularly conspicuous 
distinguishing characters, I have been unable to place it with any 
of the species previously recorded for Costa Rica. There is avail- 
able only the upper part of the stem, consequently it is probable 
that the lower leaves are substantially larger than those described 
above. Their venation is rather curious and distinctive: from the 
cuneate part of the blade, in the lower third of its length, there issue 
3-4 pairs of very oblique lateral nerves, while just above them 2 
pairs of similar nerves issue from a common point, making the 
venation there practically palmate; in the upper part of the blade 
only about 2 pairs of nerves issue from the costa. 

Eupatorium amygdalinum Lam. Thickets and pastures of 
the Pacific tierra caliente; region of El General. Extending to 
Venezuela. Plants erect, stiff, herbaceous or suffrutescent, usually 
60 cm. high or less, pubescent or almost glabrous; leaves sessile, 
lance-oblong or ovate-oblong, coriaceous, obtuse, narrowed to the 
base, crenate-serrate to entire, 5-10 cm. long, conspicuously venose; 
heads purple-pink, 7-8 mm. high, about 40-flowered, usually pedicel- 
late and glomerate, forming open, terminal panicles; phyllaries linear, 
purple-tinged. The plant is a characteristic species of the savannas. 



FLORA OF COSTA RICA 1459 

Eupatorium angulare Robinson, Contr. Gray Herb. 96: 19. 
1931. E. fistulosum Robinson, Proc. Bost. Soc. Nat. Hist. 31: 249. 
1904, non Barratt, 1847. Forests of La Palma de San Jose", 1,550 
meters, Pittier 10167. Frequent at the edge of forest, slopes of 
Irazu and Barba, especially in the region of La Palma; region 
of San Ramon; at 1,000-1,600 meters. Endemic. A stout shrub of 
1.5-4 meters, sparsely branched, the stems fistulose, tomentulose 
when young; leaves long-petiolate, very large, suborbicular, often 
20 cm. long or larger, cordate at the base, 3-12-lobate, dentate, 
palmately 3-9-nerved, pubescent above, tomentose beneath; heads 
rose-purple, 1 cm. long, 4-5-flowered, very numerous, pedicellate, 
forming a very large and broad, dense panicle; phyllaries very un- 
equal, several-seriate, obtuse, ciliate; achenes glabrous, the pappus 
stramineous. A showy and handsome plant, abundant in some 
places on the middle or upper slopes of the central volcanoes, often 
forming dense thickets. In appearance it is most unlike all other 
Costa Rican species. 

Eupatorium anisochromum Klatt, Bull. Soc. Bot. Belg. 31: 
186. 1892. E. adspersum Klatt, Leopoldina Bot. Beibl. 1. 1895; Bull. 
Soc. Bot. Belg. 35: 279. 1895 (bridge of Cariblanco, road to Sara- 
piqui, Biolley 7^22}. E. polanthum Klatt, Leopoldina Bot. Beibl. 3. 
1895; Bull. Soc. Bot. Belg. 35: 281. 1895 (Cuesta de Tarrazu, 1,900 
meters, Tonduz 7797). Laguna de Barba, 2,755 meters, Pittier 1940. 
Frequent in thickets or meadows on the upper slopes of the central 
mountains, chiefly at 1,500-3,400 meters, but also in the region of 
San Ramon, at about 1,200 meters. Endemic. An herb or shrub, 
sometimes 2 meters high, but usually much lower, often much 
branched, the stems terete, villosulous or glabrate; leaves thick, 
short-petiolate, oblong-lanceolate to ovate-lanceolate, mostly 4-10 
cm. long, acute to acuminate, cuneate or obtuse at the base, serrate, 
green above, pale beneath, more or less villosulous, conspicuously 
punctate; heads white or lavender, 5 mm. high, pedicellate, 12- 
flowered, forming small, very dense, rounded, corymbose panicles; 
phyllaries biseriate, villosulous, the outer acute, purplish at the apex, 
the inner obtuse; achenes 1 mm. long, glabrous, the pappus white. 

Eupatorium araliaefolium Less. Tapanti; region of San 
Ramon; mountains of Guanacaste; at 1,300 meters or less. Extend- 
ing to Mexico. An epiphytic shrub, glabrous except in the inflores- 
cence, there more or less sordid-puberulent; stems terete or somewhat 
6-angulate; leaves oblong-elliptic to rhombic-lanceolate, acute or 
acuminate, acute or cuneate at the base, short-petiolate, entire, 



1460 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

penninerved, somewhat fleshy, drying fuscous, 11-19 cm. long, 
heads in small or large panicles, about 25-flowered, pedicellate, 1 cm; 
long or less; phyllaries long and narrow, subequal, caducous, the 
outer much shorter and persistent. 

Eupatorium arthrodes Robinson, Contr. Gray Herb. 68: 8. 
1923. In pasture, La Palma de San Jose", 1,550 meters, Tonduz 
12430. Apparently frequent in the region of San Ramon, about 
1,300 meters. Endemic. An epiphytic, branched shrub, 1-4 meters 
long, glabrous except in the somewhat puberulent inflorescence; 
petioles mostly 2-3.5 cm. long; blades oblong to elliptic-oblong or 
elliptic-ovate, usually 7-10 cm. long, acuminate, acute to almost 
rounded at the base, fleshy, drying fuscous, entire or remotely and 
obscurely dentate, penninerved; heads white or pink, pedicellate, 
in large or small, terminal corymbs, about 12-flowered, 1 cm. long 
or less; phyllaries about 10, ciliolate, otherwise glabrous, very 
unequal, broad, rounded at the apex, the outer ones broadly ovate, 
the inner oblong; achenes scaberulous on the angles, the pappus 
whitish. 

Eupatorium Aschenbornianum Schauer. Frequent in forest 
and meadows of the central mountains, chiefly at 900-2,800 meters; 
regions of Dota and San Ramon. Panama to Mexico. An herb or 
shrub, sometimes 1.5 meters high but usually lower, branched, 
densely or sparsely villosulous with purplish or brownish hairs, the 
stems terete; leaves rounded-ovate or ovate, mostly 3-8 cm. long, 
subacute to acuminate, rounded or cordate at the base, finely crenate, 
slender-petiolate; heads white, 20-40-flowered, 5 mm. long, campanu- 
late; phyllaries linear, subscarious, lacerate-ciliate toward the apex; 
pappus dirty white. 

Eupatorium badium Klatt, Bull. Soc. Bot. Belg. 31: 186. 1892. 
E. chlorophyllum Klatt, op. cit. 187 (between La Division and Alto 
del Palmital, valley of Rio General, 1,100-2,160 meters, Pittier 3429). 
Along the road from Cerro de La Muerte to La Division, 2,160-2,900 
meters, Pittier 3407. Known in Costa Rica, apparently, only from 
the type region. Perhaps also in southern Mexico. Plants frutes- 
cent, the branches terete, pilosulous; leaves long-petiolate, rhombic, 
acuminate, cuneate at the base, 5-nerved, cartilaginous-serrate, about 
8 cm. long and 4 cm. wide, pilosulous on both surfaces; petioles 1-1.5 
cm. long; panicles axillary and terminal, the branches bearing 2-3 
heads, the pedicels 2-6 mm. long; heads about 20-flowered, the phyl- 
laries biseriate, lanceolate, 3 mm. long, cuspidate, ciliate and pubes- 
cent; achenes pilose on the angles, the pappus white. 



FLORA OF COSTA RICA 1461 

Eupatorium Brenesii Standl., sp. nov. Frutex epiphyticus 
vel interdum terrestris atque scandens, vulgo 1.5-2.5 m. longus, 
ramis crassis in sicco subteretibus striatis glabris vel in statu juvenili 
tomentulosis, internodiis elongatis; folia majuscula longipetiolata 
subcarnosa, petiolo gracili 4-7 cm. longo fere glabro; lamina ovata 
vel oblongo-ovata plerumque 13-22 cm. longa atque 7-9 cm. lata 
acuminata, basi acuta vel subrotundata penninervia, grosse arcteque 
dentata vel remote denticulata vel serrata, in sicco fusca, supra gla- 
bra, subtus sparse vel subdense praesertim ad venas sordido-villo- 
sula; inflorescentia laxissima grandis multicapitata, vulgo 20-30 cm. 
longa atque aequilata, nuda, ramis sordide puberulis vel villosulis, 
pedicellis gracilibus plerumque 3-5 mm. longis, capitulis laxe cymose 
dispositis; capitula 5-9-flora ca. 7 mm. alta, corollis lilacinis; phyl- 
laria biseriata valde inaequalia, exteriora oblonga obtusa ciliolata 
dorso sparse puberula, interiora glabrata lineari-oblonga apicem 
obtusum versus paullo attenuata; achaenia angusta glabra fere 
3 mm. longa, pappi setis stramineis. Alto de La Palma de San 
Ramon, 1,260 meters, February, 1923, Brenes 3842 (type in Herb. 
Field Mus.), 3834. Viento Fresco, Prov. Alajuela, 1,600-1,900 
meters, Standley 47955. Yerba Buena, northeast of San Isidro, Prov. 
Heredia, 2,000 meters, Standley & Valeria 50067. PANAMA: Bajo 
Chorro, Prov. Chiriqui, 1,800 meters, Mrs. M. E. Davidson 318. 
Considering the number of species of Eupatorium described from 
Costa Rica, it is somewhat remarkable that no name has been found 
for this plant, represented by several collections from separated locali- 
ties. It is closely related to E. araliaefolium, which differs conspicu- 
ously in the form of its entire leaves, which usually are quite glabrous. 
The specimens referred to E. Brenesii exhibit a good deal of variation 
in dentation of the leaves, but all probably are conspecific. 

Eupatorium carnosum Kuntze, Rev. Gen. 1: 337. 1891. Type 
collected by Kuntze on the Atlantic slope at 700 meters. Known 
definitely only from the original collection. A glabrous shrub of 2-3 
meters, presumably epiphytic, the branches terete; leaves broadly 
lanceolate, fleshy, penninerved, entire or subentire, 6-8 cm. long, the 
petioles about 1 cm. long; heads in terminal corymbs, 1 cm. long; 
phyllaries caducous, about 4-seriate, the inner ones oblong, the outer- 
most suborbicular, obtuse, multistriate, whitish; flowers 20-25, pink; 
pappus stramineous. I have seen no material that seemed referable 
to this species, which apparently is too closely related to E. araliae- 
folium, differing chiefly in its smaller leaves. 



1462 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Eupatorium collinum DC. Common in hedges or thickets, 
Meseta Central; regions of Dota and San Ramon; chiefly at 1,000- 
1,400 meters. Extending to Mexico. A shrub of 1-4.5 meters, the 
slender branches terete; leaves long-petiolate, deltoid-ovate, acumi- 
nate, cuneate to rounded or subcordate at the base, crenate to serrate 
or sometimes almost entire, obscurely puberulent to spreading- 
pubescent, 5-10 cm. long, glandular-punctate beneath; heads 24-46- 
flowered, 8 mm. high, white, fastigiate-grouped into small, dense, 
round-topped corymbs, usually slender-pedicellate; phyllaries lanceo- 
late to oblong, mostly obtuse. Called Vara blanca in Salvador. 
Costa Rican material of this species was listed in error by Klatt under 
the name E. plectranthifolium Benth. 

Eupatorium costaricense Kuntze, Rev. Gen. 1: 337. 1891. 
Angostura, 1,600 meters, Kuntze. Ecuador. A scandent shrub, the 
upper branches scabrous by the bases of deciduous hairs; leaves short- 
petiolate, ovate, acuminate, rounded at the base, 5-8 cm. long, scabe- 
rulous, especially beneath, remotely dentate, triplinerved ; inflores- 
cences axillary, corymbose; heads 10-20-flowered, campanulate, 8-10 
mm. long; phyllaries pilose, few-striate, about 3-seriate, the inner 
ones subacute, linear, the outer ovate; achenes glabrous, the pappus 
stramineous. Known to the writer only from description. Kuntze 
states that it is similar to Gynoxis in habit, and related to the Mexi- 
can E. albicaule Schultz Bip. 

Eupatorium daleoides (DC.) Hemsl. In thickets and forest, 
Meseta Central and nearby mountains; regions of Dota and San 
Ramon; chiefly at 1,000-1,500 meters, but ascending to probably 
2,000 meters. Panama to Mexico. A shrub or tree of 2.5-6 meters, 
somewhat fulvous-villosulous, at least on the young parts, the 
branches terete; leaves short-petiolate, thin, oblong to lance-oblong 
or ovate-oblong, 10-20 cm. long, acute or acuminate, cuneate at the 
base, serrate, pinnate-nerved, pellucid-punctate; heads white or 
greenish white, 5-flowered, sessile, forming large, rounded or ovoid 
panicles; phyllaries stramineous, ovate to narrowly oblong, obtuse, 
the involucre 5-6 mm. high. 

Eupatorium Durandii Klatt, Bull. Soc. Bot. Belg. 31: 189. 
1892. Between Potrero del Alto and the crater of Volcan Poas, 
2,000 meters, Pittier 387. In wet thickets, upper slopes of Poas and 
apparently also of Turrialba, at 2,000-3,000 meters. Endemic. A 
densely branched shrub, 1.5 meters high or less, the branches slender, 
villosulous, rather densely leafy; leaves small, ovate or lance-ovate, 



FLORA OF COSTA RICA 1463 

mostly 1-3 cm. long, obtuse or acute, cuneate or obtuse at the base, 
nearly sessile, 3-plinerved, almost glabrous above, pale beneath, 
almost or quite glabrous, conspicuously punctate; heads dull white, 
4 mm. long, pedicellate, 10-flowered, forming small, compact, 
corymbose panicles; phyllaries biseriate, oblong-ovate, purplish, 
erose-dentate, villosulous; achenes puberulent on the angles, the 
pappus dirty white. Closely related to E. anisochromum, and it is 
not altogether clear to the writer that the species are really distinct. 

Eupatorium elatum Steetz. Thickets and forest of the tierra 
caliente; region of San Ramon. Panama. Plants slender, herba- 
ceous, much branched, 1-2 meters high, sparsely puberulent or short- 
villosulous throughout, the stems terete, striate; leaves thin, pinnate- 
nerved, sessile, lanceolate or oblanceolate, the larger as much as 25 
cm. long, acuminate, entire or undulate, attenuate to the narrow 
base, the very base somewhat dilated and amplexicaul ; heads green- 
ish white, 5 mm. high, 20-25-flowered, short-pedicellate, in small 
clusters at the ends of the branches of a large, very open panicle; 
phyllaries 2-3-seriate, obtuse, sparsely villosulous; achenes sparsely 
hirtellous on the angles, the pappus dirty white. 

Eupatorium eximium Robinson, Contr. Gray Herb. 73: 11. 
1924. Near La Palma de San Jose", on the road to La Hondura, 
1,500-1,700 meters, Maxon & Harvey 7941. Frequent on the 
mountain slopes near La Palma; slopes of Barba; mountains south 
of Cartago; 1,500-2,000 meters. Endemic. An epiphytic shrub, 
often forming dense clumps, about 1 meter long, glabrous except 
in the somewhat puberulent inflorescence, the branches terete; 
leaves short-petiolate, oblong, about 5-6 cm. long, acuminate, acute 
at the base, fleshy, entire or slightly undulate, penninerved; pani- 
cles large or small, terminal, corymbiform; heads slender-pedicellate, 
about 9-flowered, 8-9 mm. high, pale purple; phyllaries about 10, 
unequal, 3-seriate, the outer ones lanceolate, subacute, the inner 
oblong, ciliolate, rounded at the apex, 4 mm. long; achenes glabrous, 
the pappus yellowish white. A showy and handsome plant, like 
other related species of epiphytic habit. 

Eupatorium glandulosum HBK. is reported, doubtless in error, 
by Klatt from several localities. The proper reference of the mate- 
rial so reported is unknown to the present writer. E. glandulosum 
is a synonym of E. adenophorum Spreng., a species unknown at 
present from Costa Rica. 



1464 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Eupatorium hebebotryum (DC.) Hemsl. Occasional in 
forest of the central region, at about 1,300 meters. Ranging to 
Mexico. A large shrub or small tree, the slender branches slightly 
angulate, puberulent; leaves rhombic-lanceolate to elliptic-oblong, 
10-20 cm. long, acuminate or long-acuminate, acute at the base, 
crenate-serrate, penninerved, conspicuously pellucid-punctate and 
lineolate, short-petiolate; inflorescence dense, large, terminal, 
rounded or thyrsiform, the branches tomentellous; heads about 5- 
flowered, white, 8 mm. long, short-pedicellate; phyllaries very 
unequal, pale green or stramineous, the inner oblong, obtuse or 
rounded at the apex, glabrous or nearly so; pappus yellowish white. 
Easy of recognition because of the closely pellucid-punctate leaves. 

Eupatorium hygrohylaeum Robinson, Contr. Gray Herb. 
77: 19. 1926. Wet forest, La Hondura, Prov. San Jose", 1,200-1,500 
meters, Standley & Valeria 51933. Known only from the original 
material. An erect, perennial herb, the stems terete, obscurely 
tomentulose; leaves petiolate, ovate, about 8 cm. long and 5 cm. 
wide, acuminate, rounded at the base, serrate, pinnate-nerved, 
glabrous above, somewhat puberulent beneath on the nerves and 
veins; heads in large, terminal corymbs, pedicellate, about 30- 
flowered, 7 mm. high; phyllaries about 4-seriate, rounded at the 
apex, white-striate, puberulent, lacerate-ciliolate, the outer ones 
ovate, the inner ovate-oblong; corollas white; achenes sparsely hir- 
tellous on the angles, the pappus white. 

Eupatorium hylonomum Robinson, Proc. Bost. Soc. Nat. 
Hist. 31: 250. 1904. In forest, Las Vueltas, Tucurrique, 650-700 
meters, Tonduz 12882. Region of San Ramon, at about 1,000 meters. 
Endemic. A shrub or tree of 3-5 meters, the branches slender, sub- 
terete, puberulent or minutely tomentulose; leaves short-petiolate, 
oblong or oblong-lanceolate, 10-18 cm. long, narrowly long-acumin- 
ate, acute or attenuate at the base, undulate-serrulate or almost 
entire, glabrous or nearly so; panicles large, pyramidal, sordid- 
tomentulose; heads 4 mm. long, about 7-flowered, very numerous, 
arranged in subglobose glomerules; phyllaries about 9, very unequal, 
the outer ones short, ovate, subacute, puberulent, the inner broadly 
oblong, subacute, glabrous, 2 mm. long; achenes puberulent, yellow- 
ish white. This species is too close to E. Pittieri, and its claim to 
specific status is decidedly dubious. 

Eupatorium hymenophyllum Klatt, Bull. Soc. Bot. Belg. 31: 
190. 1892. Rio San Pedro, between El General and Buenos Aires, 



FLORA OF COSTA RICA 1465 

Pittier 3709. Forests of the mountains south of Cartago, extending to 
the upper part of the Atlantic tierra caliente; region of Dota; Guana- 
caste; at 1,600 meters or less. Endemic. Plants herbaceous or 
suffrutescent, about a meter high, branched, the branches terete, 
striate, pilose; leaves thin, petiolate, rhombic-lanceolate, about 10 
cm. long and 3 cm. wide, acuminate, cuneate at the base and attenu- 
ate to the petiole, incised-serrate, 7-nerved, glabrous; branches of 
the panicle hirsute; heads long-pedicellate, 20-25-flowered, the 
flowers yellowish white to purple; phyllaries glabrous, 3-seriate, 
lanceolate, acute, stramineous, scarious-margined, 2 mm. long; 
achenes scabrous on the angles, the pappus whitish. 

Eupatorium ixiocladon Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 77. 1853. Chirrite bianco. Southern slope of Volcan de 
Irazu, 2,700 meters, Oersted 9597 (photo, of type in Herb. Field 
Mus.). Perhaps also in the region of San Ramon. Endemic. 
Plants herbaceous or suffrutescent, almost glabrous, the branches 
terete, glutinous; leaves long-petiolate, ovate or deltoid-ovate, 7-13 
cm. long, 5-7 cm. wide, long-acuminate, truncate to rounded- 
cuneate at the base, rather coarsely crenate; heads numerous, 
white, 4-5 mm. long, forming small, terminal, corymbose panicles, 
the branches minutely puberulent; heads about 20-flowered, the 
phyllaries linear, subbiseriate, scarcely striate, the inner ones 
acuminate; achenes scabrous on the angles, the pappus dirty white. 

Eupatorium laevigatum Lam. Savannas of the Pacific tierra 
caliente; thickets at Pejivalle; at 900 meters or less. Mexico to 
Argentina. A shrub of 1-2 meters, very viscid but otherwise glabrous 
or nearly so, the branches angulate; leaves short-petiolate, rather 
thick, rhombic-ovate to ovate-oblong, acute, cuneate at the base, 
3-nerved from the base, coarsely serrate; heads about 12 mm. long, 
sessile or pedicellate, pale lavender, numerous, forming very dense, 
convex corymbs; involucre cylindric, the phyllaries indurate, pale, 
closely appressed, obtuse or rounded at the apex, striate. 

Eupatorium laurifolium Robinson, Proc. Bost. Soc. Nat. Hist. 
31: 251. 1904. Hills near Rio Chirripo, Pittier 16065. Known, 
apparently, only from the original material. A scandent shrub, the 
branches slender, striate, glabrate; leaves oval, about 12 cm. long 
and 5 cm. wide, caudate-acuminate, acute at the base, 5-nerved, 
undulate-dentate or entire, glabrous, obscurely pellucid-punctate, 
the petioles 1.5 cm. long; panicles pedunculate, sordid-pubescent; 
phyllaries about 4-seriate, oval-oblong, striate, concave, ciliate but 



1466 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

otherwise glabrous; corollas 5 mm. long; achenes pubescent, the 
pappus sordid yellowish. 

Eupatorium ligustrinum DC. E. semialatum Benth. Fre- 
quent in forest and thickets of the Meseta Central, sometimes form- 
ing almost pure stands, extending to the upper part of the Pacific 
tierra caliente; regions of Dota and San Ramon. Ranging to Mexico. 
A shrub or tree of 1-6 meters; leaves oblong, 4-9 cm. long, acuminate, 
cuneate at the base, petiolate, pinnate-nerved, remotely serrate- 
dentate, glabrous, glandular-dotted beneath; heads 4-8-flowered, 
7 mm. long, white or pinkish, fragrant, forming small, dense, termi- 
nal corymbs; phyllaries linear, gland-dotted, half as long as the 
flowers; pappus often deep pink. 

Eupatorium macrophyllum L. Frequent in thickets or weedy 
fields of the tierra caliente; region of San Ramon; Guanacaste. 
Widely distributed in tropical America. Plants herbaceous, simple 
or branched, 1-2 meters high, pale-tomentulose, especially on the 
lower leaf surface, the stems terete; leaves thin, large, long-petiolate, 
broadly ovate, acuminate, broadly cordate at the base, crenate; 
heads greenish white, 50-75-flowered, 7 mm. high; phyllaries gradu- 
ated, many-seriate, lanceolate, acute, pale-nerved; pappus dirty 
white. A common, weedy plant in many regions of the Central 
American tierra caliente. 

Eupatorium microstemon Cass. E. guadalupense Spreng. In 
thickets or forest, Meseta Central; region of Dota; Guanacaste. 
Mexico to West Indies and Bolivia. A slender, weak, usually 
branched annual, a meter high or commonly lower, puberulent; 
leaves slender-petiolate, deltoid-ovate, 2.5-7 cm. long, thin, nar- 
rowed to an acute or obtuse apex, subtruncate at the base, crenate- 
serrate; heads 4-5 mm. high, purple or white, slender-pedicellate, 
forming a lax panicle; phyllaries 3-4-seriate, 3-nerved, the outer ones 
lanceolate, acute, the inner linear-oblong, rounded at the apex. Var. 
lilacinum Kuntze (Rev. Gen. 1: 338. 1891; described from Trinidad 
and Costa Rica) is a form with purple or lilac corollas, those of the 
typical form of the species being white. 

Eupatorium morifolium Mill. E. populifolium HBK. In 
thickets or hedges, region of San Jose" to the coasts; region of San 
Ramon; at 1,1 00 meters or less. Mexico to Brazil. Plants herbaceous 
or suffrutescent, simple or sparsely branched, 1.5-3 meters high, the 
stout stems subterete, glabrous or nearly so; leaves large, broadly 
ovate to ovate-oblong, 12-20 cm. long or larger, acute or short-acu- 



FLORA OF COSTA RICA 1467 

minate, acute to cordate at the base, coarsely serrate, rather thick, 
triplinerved or pinnate-nerved; heads greenish white, 8-12-flowered, 
sessile or short-pedicellate, about 7 mm. high, forming large, thyrsoid 
panicles; phyllaries stramineous, ovate, obtuse or rounded at the 
apex, arachnoid- tomentose or glabrate, about 5-seriate; pappus ful- 
vous whitish. Material of this species was reported from Costa Rica 
by Klatt under the name E. hebebotryum Benth. 

Eupatorium Neaeanum DC. is reported by Klatt from San Pedro 
de La Calabaza, presumably in error. The species is known from 
Mexico and Guatemala. 

Eupatorium nemorosum Klatt. In fields or thickets, some- 
times in wet soil, regions of San Ramon and of La Palma de San Jose", 
1,000-1,400 meters. Extending to Bolivia. Plants annual or peren- 
nial, herbaceous, stout, erect, 1.5 meters high or less, the stem usually 
simple, terete, villosulous and glandular-pilose; leaves oblong, 7-15 
cm. long, acuminate, serrate, attenuate into a broadly winged petiole, 
pinnate-nerved, rough-pilose on both surfaces; heads numerous, 
greenish white, many-flowered, 1 cm. high, forming a narrow panicle, 
pedicellate; involucre campanulate, about 3-seriate, the phyllaries 
lanceolate, acute; achenes glabrous or nearly so, the abundant pappus 
dirty white. In general appearance this plant is quite unlike other 
local species. In habit and form of the heads it reminds one some- 
what of Erechtites hieraciifolia. 

Eupatorium odoratum L. E. conyzoides Mill. Common in 
thickets, Meseta Central to the coasts; region of San Ramon; Guana- 
caste. Generally distributed in tropical America. A shrub of 1-2 
meters or more, often subscandent, the branches commonly recurved 
or pendent, the stems terete, villous; leaves petiolate, deltoid-ovate 
or rhombic-ovate, 7-10 cm. long, long-acuminate, abruptly narrowed 
and acute at the base, subentire to coarsely dentate, often subhastate, 
sparsely villous above, pubescent or velutinous beneath; heads laven- 
der, about 1 cm. long, cylindric, 20-35-flowered, in small, dense, 
trichotomous, convex corymbs; phyllaries several-seriate, rounded 
at the apex, closely appressed, indurate, striate. Probably the most 
common Eupatorium species of tropical America, often a weed in 
abandoned or cut-over land. Rather showy when covered with its 
abundant flower clusters. E. odoratum var. mallotophyllum Robinson, 
originally described from Colombia, is reported by Blake (in litt.) 
to be represented in Costa Rica, according to material at the Gray 
Herbarium. It is distinguished by having the leaves densely griseous- 
tomentose on both surfaces. 



1468 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Eupatorium Oerstedianum Benth. in Oerst. Vid. Medd. Kjoe- 
benhavn 1852: 74. 1853. Cerro de Aguacate, 600 meters, Oersted 
9616 (photo, of type in Herb. Field Mus.); also Cartago and Cerros 
de Candelaria. Thickets and forest of the Meseta Central, and upper 
part of the Pacific tierra caliente; region of San Ramon; Guanacaste. 
Ranging to Guatemala and British Honduras. A stiff shrub, 1-2.5 
meters high, the branches terete, glabrate; leaves sessile or short- 
petiolate, oblong or lance-oblong, mostly 15-20 cm. long, acuminate, 
acute or obtuse at the base, serrate, pinnate-nerved, glabrous above, 
puberulent beneath on the veins; heads cylindric, about 40-flowered, 
white, 1 cm. long; phyllaries indurate, closely appressed, stramineous, 
obtuse or rounded at the apex, ciliate; achenes puberulent. 

Eupatorium pacacanum Klatt, Leopoldina Bot. Beibl. 3. 
1895. E. roseum Klatt, Bull. Soc. Bot. Belg. 31: 194. 1892 (Rodeo de 
Pacaca, Pittier 3321+}. Both names were based upon the same col- 
lection. Known, apparently, only from the original material. An 
almost glabrous shrub, the branches terete; leaves long-petiolate, 
ovate, 3-5 cm. long, 3 cm. wide, acute, rounded at the base, mem- 
branaceous, crenate, 3-nerved, ferruginous-pilose beneath along the 
nerves; heads in dense corymbs, 40-flowered, pedicellate, the pedi- 
cels pilose; phyllaries biseriate, spatulate, 3 mm. long, pink at the 
apex, 2-striate, scarious-margined; achenes hirtellous on the angles, 
the pappus white. 

Eupatorium parasiticum Klatt, Ann. Naturhist. Hofmus. 
Wien 9: 357. 1894. Type Endres 147 from Costa Rica, without 
definite locality. Apparently common in the region of San Ramon; 
also at La Hondura de San Jose"; 1,000-1,300 meters. Endemic. A 
small, epiphytic shrub, less than a meter long, often only 20 cm. 
high, glabrous except for the inconspicuously puberulent inflores- 
cence; leaves small, short-petiolate, oblong or oblong-elliptic, about 
4 cm. long, obtuse or subacute, acute at the base, fleshy, coriaceous 
when dried, remotely and coarsely dentate or subentire; panicles 
small and lax, corymbiform; heads slender-pedicellate, pale purple, 
5-flowered; phyllaries about 6, lanceolate, 5 mm. long, ciliolate, 
puberulent near the apex; achenes minutely pilose on the angles, 
the pappus white. 

Eupatorium pithecobium Robinson, Contr. Gray Herb. 77: 
28. 1926. Wet forest, Laguna de La Chonta, northeast of Santa 
Maria de Dota, Prov. San Jose", 2,000 meters, Standley 4-21 81 . Known 
only from the region of Dota, 2,000-2,400 meters. Endemic. An 



FLORA OF COSTA RICA 1469 

epiphytic shrub, pendent, 1.5 meters long or less, the branches 
obscurely puberulent; leaves petiolate, oblong or oblong-lanceolate, 
5-12 cm. long, acuminate, cuneate at the base, closely serrate- 
dentate, penninerved, hirtellous beneath, especially on the costa; 
petioles 1-1.5 cm. long; corymbs terminal, of about 10 heads; heads 
about 22-flowered, 11 mm. long; involucre campanulate, the phyl- 
laries 3-4-seriate, the outermost lanceolate, acute, the inner ones 
oblong or spatulate, obtuse, pink; corollas pink; achenes scaberulous 
on the angles, the pappus white. 

Eupatorium Pittieri Klatt, Bull. Soc. Bot. Belg. 31: 192. 1892. 
Quitirri chico. Along Rio Segundo, Pittier 1698. Frequent in 
forest of the central mountains, ascending to about 2,000 meters; 
upper part of the Atlantic tierra caliente, and mountains of Guana- 
caste, at 600 meters or more. Panama to southern Mexico. A 
shrub or tree, 3-6 meters high, the slender branches minutely 
puberulent or almost glabrous; leaves short-petiolate, elliptic-oblong 
to oblong-lanceolate, 9-20 cm. long, narrowly long-acuminate, 
attenuate to the base, coarsely serrate to subentire, almost glabrous; 
heads very numerous, 5-6 mm. long, whitish, pedicellate, forming 
large panicles, about 10-flowered, campanulate; phyllaries 3-seriate, 
ovate, obtuse, very unequal, the inner 3 mm. long, striate, ciliate; 
achenes hirtellous on the angles, the pappus dirty white. 

Eupatorium platyphyllum Robinson. Region of San Ramon; 
lower slopes of Poas; El General; at 1,400 meters or less. Mexico 
and Guatemala. A branched shrub of 3-4 meters, the branches 
terete or nearly so, at first tomentulose-puberulent, becoming 
glabrate; leaves slender-petiolate, broadly ovate or rounded-ovate, 
sometimes shallowly hastate-lobate, acute or acuminate, rounded 
or very broadly cuneate at the base, thin, 3-nerved from the base, 
undulate or dentate, almost glabrous; heads numerous, large, 60-75- 
flowered, 1 cm. high or larger, broad, slender-pedicellate, forming a 
large, corymbiform panicle; phyllaries pluriseriate, laxly imbricate, 
scarious, very unequal in length, stramineous, scarious and erose at 
the rounded apex, glabrous, many-nerved; corollas white; achenes 
glabrous, 2 mm. long, the pappus white. 

Eupatorium plectranthifolium Benth. in Oerst. Vid. Medd. 
Kjoebenhavn 1852: 76. 1853. Naranjo, 1,200 meters, Oersted. 
Birris, on the lower slopes of Irazu; region of Dota; slopes of Barba, 
1,600 meters. Endemic. A branched herb about a meter high, the 
branches terete, tomentulose or glabrate; leaves long-petiolate, ovate, 



1470 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

acuminate, cordate or subcordate at the base, 8-11 cm. long, crenate- 
serrulate, thin, 7-nerved, glabrate above, tomentulose beneath on 
the reticulate veins; heads rose-purple, 4-5 mm. long, 30-40-flowered, 
short-pedicellate, forming dense, leafy, corymbose panicles; phyl- 
laries ovate-lanceolate or lance-linear, acute, very unequal, the 
outer ones short, tomentulose, the inner 3-5-striate, tomentulose 
at the apex; achenes glabrous. 

Eupatorium prionophyllum Robinson, Proc. Bost. Soc. Nat. 
Hist. 36: 484. 1901. Banks of Rio Toros, Pittier 1705. Collected 
also at Rancho Flores, 2,030 meters, and at La Ventolera, slopes of 
Poas. Panama. A shrub or small tree, appearing glabrous but with 
traces of a short, close tomentum on the branchlets, petioles, leaf 
veins, and pedicels; leaves slender-petiolate, broadly ovate, acu- 
minate, usually obtuse at the base, incisely serrate-dentate, thin, 
pinnate-nerved, 7.5-9 cm. long; heads 25-30-flowered, in terminal, 
rounded or thyrsoid panicles; phyllaries 3-seriate, the outer short, 
ovate, acute, the inner oblong, subobtuse, striate, ciliolate, stra- 
mineous; pappus white. The plant was listed incorrectly by Klatt 
as E. ixiocladon Benth. A variety has been described from Guate- 
mala. The flowers of the La Ventolera plant were pale purple. 

Eupatorium psoraleum Robinson, Proc. Bost. Soc. Nat. Hist. 
31: 253. 1904. La Palma de San Jose", 1,460 meters, Tonduz 12589. 
Apparently known only from the original material. A shrub, 
apparently epiphytic, the branchlets slightly 6-angulate, striate, 
covered with minute, transverse scales; leaves petiolate, broadly 
lanceolate, 4-9 cm. long, acuminate at each end, entire, pinnate- 
nerved, glabrous; heads 5-flowered, short-pedicellate, forming a 
spreading panicle; phyllaries scarious, stramineous or pink, ciliolate, 
otherwise glabrous, very unequal, 3-4-seriate, the inner ovate- 
oblong, obtuse, slightly ampliate at the apex; corollas 4-5 mm. long, 
pink; achenes glabrous, 2 mm. long, the pappus sordid. According to 
the author of the species, it bears much resemblance to E. daleoides. 

Eupatorium pycnocephalum Less. Chirrite, Santa Lucia. 
E. pratense Klatt, Bull. Soc. Bot. Belg. 31: 193. 1892 (pastures at 
Turrialba, 200 meters, Pittier 4128). Frequent in thickets or forest 
of the Meseta Central, descending into at least the upper part of 
the tierra caliente, ascending the slopes of the mountains to about 
2,300 meters; regions of Dota and San Ramon. Mexico to Venezuela. 
A herbaceous perennial, 1.5 meters high or usually much lower, 
finely incurved-pubescent; leaves slender-petiolate, deltoid-ovate, 



FLORA OF COSTA RICA 1471 

4-6 cm. long, acuminate, cuneate to truncate or shallowly cordate 
at the base, crenate-serrate, 3-nerved from the base, at least sparsely 
pilose on both surfaces; inflorescence with spreading or ascending 
branches, each bearing a rounded, terminal cluster of 7-20 or more 
short-pedicellate heads, these purple, about 25-flowered, 3-5 mm. 
high; involucre campanulate, the phyllaries about 3-seriate, the 
inner oblong, obtuse, 3-nerved, the outer ones acute, pubescent or 
almost glabrous; achenes Somewhat pubescent. 

Eupatorium quadrangulare DC. is cited by Klatt from the Pacific 
tierra caliente. This is probably an error of determination, since 
Robinson gives the range of the species as extending from Mexico 
only as far southward as Salvador. 

Eupatorium Schultzii Schnittspahn. Occasional in forest and 
clearings of the central mountains; region of San Ramon; usually 
at 1,000-1,200 meters. Extending to southern Mexico. A branched 
shrub or coarse herb about a meter high, the slender branches terete, 
viscid-pilosulous; leaves large, thin, rounded-ovate, acuminate, rather 
deeply cordate at the base, often as broad as long as broader, undu- 
late or coarsely dentate, somewhat puberulent, beneath densely and 
conspicuously glandular-punctate; heads white, campanulate, about 
7 mm. high, many-flowered, slender-pedicellate, forming a large, 
broad panicle, the pedicels densely glandular-puberulent; phyllaries 
scarious, whitish, oblong or elliptic-oblong, rounded at the apex, 
glabrous; pappus white. 

Eupatorium sexangulare (Klatt) Robinson, Proc. Amef. Acad. 
43: 35. 1907. Piptocarpha sexangularis Klatt, Leopoldina Bot. Beibl. 
1. 1895. "Descente de la Ardilla," 900 meters, Tonduz 7760. Also 
in the region of San Ramon, at about 1,100 meters. Endemic. A 
glabrous shrub of 1.5-2 meters, the stems sharply 6-angulate and 
sulcate, the internodes elongate; leaves large, short-petiolate, ovate- 
lanceolate, 10-26 cm. long or larger, acute or acuminate, acute and 
long-decurrent at the base, thick, very lustrous, sinuate-dentate or 
almost entire, penninnerved ; heads white, 5-flowered, 7-8 mm. long, 
cylindric, forming large, terminal, open, corymbose panicles, the 
individual heads sessile or nearly so and densely glomerate; phyllaries 
multiseriate, stramineous, the inner caducous, ovate-lanceolate, 
striate, obtuse, the outer ones short, broadly ovate; achenes glabrous, 
3 mm. long, the pappus yellowish white. 

Eupatorium Sideritidis Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 77. 1853. Ujarras, Oersted 9656 (photo, of type in Herb. 



1472 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Field Mus.). Frequent about the Meseta Central, descending to the 
tierra caliente, ascending to about 1,500 meters; region of San Ramon. 
Extending to Guatemala. Plants herbaceous, perennial, 30-60 cm. 
high, often forming dense clumps, very slender, rather sparsely pube- 
rulent; leaves narrowly lanceolate or almost linear, small, remotely 
serrate, 3-nerved, attenuate to the short petiole, glabrate; heads 4 
mm. high, bright purple, 20-25-flowered, pedicellate, forming small, 
lax, terminal clusters; phyllaries 2-3-seriate-linear-lanceolate, acumi- 
nate, striate; pappus white or dirty white. Easily recognized among 
Costa Rican species by the very narrow leaves. The plant is highly 
specialized as to habitat, growing practically always at the edge of 
streams, on rocks or in mud. 

Eupatorium silvicola Robinson, Proc. Bost. Soc. Nat. Hist. 31 : 
254. 1904. Forests of Copey, 1,800 meters, Tonduz 11694. Known, 
apparently, only from the original locality. Endemic. Plants fruti- 
cose, the branches sordid-tomentose; leaves large, ovate, 12-17 cm. 
long, 8-12 cm. wide, acuminate, obtuse at the base, serrate or suben- 
tire, thin, almost glabrous except beneath on the sordid-tomentose 
veins; petioles 3-7 cm. long; heads 7-10-flowered, pedicellate, 7 mm. 
long, arranged in a leafy panicle; phyllaries about 3-seriate, very 
unequal, the outer ones short, ovate, subacute, viscid-tomentose, 
the inner lance-linear, acute, subscarious, green-striate, glabrous or 
glabrate; achenes sparsely glandular. 

Eupatorium Sinclairii Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 79. 1853. Cartago, Oersted. Thickets and fields, Meseta 
Central to the tierra caliente; region of San Ramon; Guanacaste. 
Panama to Guatemala, and probably ranging even more widely. 
Closely related to E. microstemon, and like that species in general 
appearance and in most details; heads smaller, only 3 mm. high, usu- 
ally purple, forming large and lax panicles. 

Eupatorium solidaginoides HBK. E. decussatum Klatt, Bull. 
Soc. Bot. Belg. 35: 295. 1896 (Rio Virilla, near San Juan, Tonduz 
9869). Thickets, regions of San Jose", San Ramon, and El General. 
Widely distributed in tropical America. Plants erect, slender, much 
branched, tall, herbaceous or suffrutescent, sometimes subscandent; 
leaves slender-petiolate, ovate, 5-12 cm. long, caudate-acuminate, 
truncate to cordate at the base and more or less hastate, dentate, 
thin, puberulent to subtomentulose beneath on the nerves; heads 
10-15-flowered, slender-pedicellate, subracemose on the spreading 
branches of a large, leafy-bracted panicle, 5-6 mm. long, white; phyl- 
laries lanceolate, acute, laxly imbricate, usually purple-tinged. 



FLORA OF COSTA RICA 1473 

Eupatorium Standleyi Robinson, Contr. Gray Herb. 77: 40. 
1926. Brushy slope, Alto de La Estrella, Prov. Cartago, Standley 
39288. Also at La Hondura de San Jose", wet forest, 1,300-1,700 
meters. Endemic. Plants herbaceous, erect or sometimes 3 meters 
long and scandent, glabrous or nearly so except in the inflorescence; 
leaves petiolate, ovate-oblong, 12-16 cm. long, subacute or obtuse, 
penninerved, coarsely dentate, somewhat fleshy; panicles large, pube- 
rulent, the heads 8 mm. high, 6-flowered, pedicellate; phyllaries about 
8, very unequal, acute or subobtuse, ciliolate, the outer ones ovate; 
corollas pink; achenes glabrous, 2.5 mm. long, the pappus yellowish. 

Eupatorium subcordatum Benth. in Oerst. Vid. Medd. Kjoe- 
benhavn 1852: 77. 1853. E. splendens Klotzsch ex Polak. Linnaea 
41: 575. 1877 (Angostura, Polakowsky). Southern slope of Volcan 
de Irazu, 2,400 meters, Oersted 9663 (photo, of type in Herb. Field 
Mus.). Slopes of Irazu and Turrialba, ascending to 3,400 meters. 
Endemic. Herbaceous or frutescent, sometimes 1.5 meters high, the 
branches minutely brown-puberulent or almost glabrous; leaves long- 
petiolate, ovate or broadly deltoid-ovate, 3.5-6 cm. long, acuminate, 
truncate or cordate at the base, crenate, rufous-pubescent beneath 
on the nerves and veins; heads 4-5 mm. long, pinkish white or pale 
lavender, about 20-flowered, forming dense, leafy corymbs, pedicel- 
late; phyllaries linear, subbiseriate, puberulent, very unequal; achenes 
scabrous on the angles, the pappus dirty white. Material of this 
species (Pittier 3432) was referred by Klatt toE. Dombeyanum DC., 
a South American species. 

Eupatorium Thieleanum Klatt, Bull. Soc. Bot. Belg. 31: 191. 
1892. E. myrianthum Klatt, Leopoldina Bot. Beibl. 3. 1895 (based 
on the same material as E. Thieleanum). Rodeo de Pacaca, 900 
meters, Pittier 1603. Mountains of the central region, descending to 
the Pacific tierra caliente; El General; region of San Ramon; at 1,800 
meters or less, in forest. Panama. Plants branched, herbaceous or 
suffrutescent, 1-3 meters high, the stems slender, terete, sordid- 
tomentulose; leaves apparently all alternate, slender-petiolate, 
broadly ovate or rounded-ovate, often as broad as long, acuminate, 
rounded to deeply cordate at the base, crenate-dentate, often some- 
what trilobate or hastate-lobate, thin, scaberulous or glabrate above, 
short- villosulous or tomentulose beneath on the nerves and veins; 
panicles small, dense, terminal and axillary, the heads glomerate, 
sessile or short-pedicellate, 4 mm. long, white, about 25-flowered; 
phyllaries biseriate, lanceolate, obtuse or subacute, minutely villo- 



1474 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

sulous; receptacle pilose; achenes 1 mm. long, glabrous or nearly so, 
the pappus white. 

Eupatorium Tonduzii Klatt, Leopoldina Bot. Beibl. 4. 1895; 
Bull. Soc. Bot. Belg. 35: 282. 1896. Cuesta de Tarrazu, 1,900 meters, 
Tonduz 7799. Oak forest, region of Dota, 1,500-2,100 meters. 
Endemic. A shrub of 1-2 meters, puberulent; leaves petiolate, 
lanceolate, 10-14 cm. long, acuminate at each end, crenate or 
crenate-serrate, penninerved, obscurely puberulent on the costa, 
elsewhere glabrous; heads numerous, in terminal corymbs, pinkish 
white, pedicellate, about 28-flowered, 7 mm. high; phyllaries about 
16, linear-lanceolate, attenuate, often purple-tinged at the apex, 4 
mm. long; achenes 2 mm. long, scaberulous on the angles, the 
pappus white. The species is related to E. ligustrinum DC. 

Eupatorium triptychum Robinson, Contr. Gray Herb. 77: 43. 
1926. Wet forest, El Mufieco, south of Navarro, Prov. Cartago, 
1,400 meters, Standley 33458. Known only from the original locality. 
A large shrub, almost glabrous except in the inflorescence; leaves 
trifoliolate, long-petiolate; leaflets petiolulate, oblong, 6-10 cm. 
long, acuminate, rounded to subacute at the base, obsoletely crenate- 
undulate; heads 12-flowered, 9 mm. high, slender-pedicellate, numer- 
ous, forming large, terminal corymbs; phyllaries linear-oblong, 4-4.5 
mm. long, rounded at the apex, ciliolate, glabrate; achenes puberulent 
on the costae, the pappus dirty white. Easy of recognition because 
of its compound, 3-foliolate leaves. 

Eupatorium Valerianum Standl., sp. nov. Herbacea vel 
frutescens ut videtur elata, ramis gracilibus teretibus brunnescenti- 
bus minute adpresse sordido-tomentulosis vel glabratis, internodiis 
vulgo elongatis; folia opposita graciliter longipetiolata membranacea, 
petiolo 1.5-3.5 cm. longo sordido-puberulo vel glabrato; lamina ovata 
usque lanceolata 6.5-11.5 cm. longa 2.5-7 cm. lata longe attenuato- 
acuminata, basi rotundata vel breviter abrupte decurrens, in toto 
margine grosse crenato-dentata, 5-plinervia, nervis superioribus alte 
supra basin laminae nascentibus, supra viridis, ad nervos villosulo- 
puberula, subtus fere concolor, ad nervos venasque dense tomentulosa, 
aliter glabrata; inflorescentia terminalis irregulariter corymboso- 
paniculata foliosa, capitulis numerosissimis parvis breviter pedicel- 
latis ut videtur albis, inflorescentia densa, capitulis cymose dispositis; 
capitula ca. 4 mm. alta campanulata ca. 10-flora; phyllaria ca. 10 
subbiseriata, valde inaequalia, exteriora brevia oblongo-lanceolata 
acuminata minute puberula, interiora 2.5 mm. longa spathulato- 



FLORA OF COSTA RICA 1475 

oblonga scariosa viridescentia 2-costata apice subdilatato late 
rotundata vel subtruncata glabrata; corolla glabra ca. 2.5 mm. longa; 
achaenia ca. 1.5 mm. longa ad angulos minute scaberula, pappi setis 
albis 2 mm. longis scaberulis. Colinas de San Pedro de San Ramon, 
March, 1935, Brenes 20498 (type in Herb. Field Mus.). Cataratas 
de San Ramon, February, 1931, Brenes 13428, 13455. Similar to E. 
prionophyllum Robinson, but conspicuously different in the crenate- 
dentate rather than sharply serrate leaves, and in the tiny heads, 
which are about the smallest found in any local species of the genus. 
The species is named in honor of Professor Juvenal Valeric Rodriguez, 
Director of the Museo Nacional de Costa Rica. 

Eupatorium Vitalbae DC. Occasional in forest or thickets 
of the central region, extending to the tierra caliente of both coasts; 
at 1,500 meters or less. Central America to Peru and Brazil. A 
shrub of 1-2.5 meters, sometimes scandent, the stems and inflores- 
cence finely puberulent; leaves petiolate, coriaceous, elliptic-ovate 
or ovate-lanceolate, acuminate, rounded at the base, 3-5-nerved, 
remotely serrate; heads bright pink, 12 mm. high, 45-70-flowered, 
on long, stout pedicels, forming large, open panicles; outer phyllaries 
ovate-oblong, acute, multistriate, the inner narrowly lance-oblong; 
achenes minutely hispid on the angles. A handsome plant because 
of the bright-colored flower heads. 

Eupatorium vulcanicum Benth. in Oerst. Vid. Medd. Kjoe- 
benhavn 1852: 78. 1853. Chirrite bianco. Southern slope of Volcan 
de Irazu, 2,700 meters, Oersted (photo, of type in Herb. Field Mus.). 
Occasional in thickets and forest of the central mountains, chiefly 
at 1,400-2,700 meters. Endemic. Plants herbaceous or suffrutes- 
cent, about a meter high, the branches rufous- villous; leaves petio- 
late, rounded-ovate, obtuse or acute, subcordate at the base, crenate, 
3-5-nerved, pubescent above, rufous-villous beneath; heads numer- 
ous, in dense, terminal corymbs, about 20-flowered, white; phyllaries 
subbiseriate, linear-lanceolate, acuminate, the outer ones densely 
pubescent; achenes hispidulous on the angles. 

FLEISCHMANNIA Schultz Bip. 

Plants herbaceous or suffrutescent; leaves opposite or alternate, 
dentate to biternately parted; heads small, discoid, white or purplish, 
long-pedunculate, solitary or subcorymbose; phyllaries many-seriate, 
graduated, dry, acuminate, striate; receptacle naked; achenes linear, 
5-angulate; pappus of 5-8 bristles, with minute, setulose squamellae 



1476 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

between them, or the squamellae united into a low crown, or obsolete. 
Probably no other species are known in Central America. 

Fleischmannia arguta (HBK.) Robinson. F. rhodostyla Schultz 
Bip. Eupatorium quinquesetum Benth. Reported by Klatt from 
Quebrada de Tapshi, near Boruca, Pittier 7344- Nicaragua to south- 
ern Mexico. An herb of 20-40 cm., in general appearance suggestive 
ofErigeron, puberulent, slender, branched; lower leaves opposite, the 
upper alternate, petiolate, rhombic to lanceolate, 4 cm. long or less, 
acute or acuminate, cuneate at the base, coarsely dentate, 3-nerved; 
heads few, about 8 mm. broad, the phyllaries 2-3-seriate, 4 mm. 
long, puberulent; achenes obscurely scaberulous on the angles, the 
pappus of 5 bristles. 

GALEANA Llave 

Reference: Rydberg, N. Amer. Fl. 34: 42. 1914. 

Slender, dichotomously branched annuals, more or less viscid- 
pubescent, the leaves glandular-punctate, petiolate, mostly opposite; 
heads small, radiate, in leafy cymes; phyllaries 5, broad, concave, 
oval; ray flowers 3, pistillate and fertile, the disk flowers partly 
fertile and partly neutral; achenes pyriform, triquetrous, some of 
the angles somewhat corky-thickened, some with corky, incurved, 
crenate wings. No other species are known from Central America. 

Galeana pratensis (HBK.) Rydb. Villanova pratensis Benth. & 
Hook. Occasional in pastures or thickets of the Meseta Central and 
the upper part of the Pacific slope. Ranging to southern Mexico. 
Plants much branched, bushy, usually 20-30 cm. high, glandular- 
pubescent; leaves deltoid-ovate to oblong, 2-3 cm. long, acute or 
obtuse, truncate or acute at the base, coarsely serrate, 3-nerved; 
involucres 3 mm. high, the bracts glandular-pubescent; rays whitish, 
scarcely 3 mm. long; achenes black, glabrate, 3 mm. long. 

GALINSOGA Ruiz & Pavon 

Reference: St. John & White, The genus Galinsoga in North 
America, Rhodora 22: 97. 1920. 

Branched, annual herbs, pubescent; leaves opposite, usually 
petiolate, dentate; flower heads very small, radiate, white, pink, or 
purple, inserted in the upper leaf axils; involucre hemispheric, the 
phyllaries broad, few, biseriate, striate, subequal, the outer slightly 
shorter; receptacle conic, paleaceous; ray flowers pistillate, fertile, 
the rays 4-5, short; disk flowers yellow, perfect; achenes angulate or 
the outer ones compressed; pappus of the disk flowers of several short, 



FLORA OF COSTA RICA 1477 

laciniate or fimbriate scales, that of the ray flowers of few short, 
slender bristles, or none. Probably no other species are to be found 
in Central America. 

Galinsoga bicolorata St. John & White. Open banks, regions 
of San Jos and Cartago. Ranging to Mexico. Plants 20-30 cm. 
high, the stems hispid or glabrate; leaves ovate-cordate or lanceolate, 
1.5-4 cm. long, acuminate, serrate, hispid; heads 3-4 mm. high, the 
outer phyllaries sparsely glandular-pilose; rays purple or pink; ray 
achenes hispid, the pappus scales linear, long-aristate, longer than 
the corolla tube; disk pappus of linear, fimbriate, long-aristate scales 
equaling or exceeding the corolla tube. All the species of this genus 
are alike in general appearance, the specific differences being found in 
the color of the rays and in the pubescence and pappus of the achenes. 

Galinsoga caracasana (DC.) Schultz Bip. Frequent in mead- 
ows of the central region, especially on the mountain slopes, ascending 
to perhaps 3,000 meters. Widely distributed in tropical America. 
Rays pink or dark red; pappus of the disk flowers of linear-lanceolate, 
fimbriate scales half as long as the corolla, the corolla longer than 
the achene. 

Galinsoga ciliata (Raf.) Blake. G. aristulata Bicknell. Occa- 
sional in fields of the central region; Dota. Widely distributed in 
tropical America. Rays white; pappus of the disk flowers aristate, 
fimbriate; pappus of the ray flowers of linear, minutely fimbriate 
scales equaling the tube of the corolla. 

Galinsoga parviflora Cav. Frequent in fields and waste ground 
of the central region, often in cultivated land, descending to the upper 
part of the tierra caliente. Widely distributed in tropical America. 
Rays white; pappus of the disk flowers not aristate; pappus of the 
ray flowers none or of a few bristles. This species, like some of 
the others listed, although essentially a tropical plant, was intro- 
duced long ago into the eastern and northern United States, where it 
is perfectly hardy and thrives, especially in vacant ground about 
cities. It is particularly plentiful in Chicago in the courts of apart- 
ment buildings. 

GARCILASSA Poepp. & Endl. 

A slender annual, branched, erect or ascending, scabrous or his- 
pid; leaves alternate, 3-nerved, serrate; heads small, discoid, aggre- 
gate at the tips of the branches into globose glomerules, 4-5-flowered ; 
involucre very short, the phyllaries few; receptacle paleaceous; 



1478 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

achenes oblong, laterally compressed, short-pilose, sometimes nar- 
rowly winged; pappus very short, annular, ciliate-fimbrillate. The 
genus consists of a single species. 

Garcilassa rivularis Poepp. & Endl. In forest or thickets of the 
Atlantic tierra caliente; Guanacaste; at 700 meters or less. Ranging 
to Peru. Leaves petiolate, ovate or lanceolate, the pubescence rough; 
heads small, green, about 5 mm. long. An inconspicuous and rather 
weedy plant, apparently of rare occurrence in the regions where it 
is found. 

GERBERA L. 

Scapose, perennial herbs; leaves basal, entire or sinuate-pinnatifid, 
tomentose beneath; heads solitary at the end of the scape, erect, of 
medium or large size, radiate, yellow, reddish, or white; involucre 
turbinate or broadly campanulate, the phyllaries narrow, imbricate 
in few or numerous series, the outer gradually shorter; receptacle 
plane, naked; achenes somewhat compressed, 5-costate, villous, 
often rostrate; pappus of numerous slender setae. The species are 
natives of Africa and Asia. 

Gerbera Jamesoni Bolus ex Hook. f. Sometimes planted for 
ornament in gardens of the temperate region. Native of South 
Africa. Leaves greatly elongate, more or less pinnatifid, narrow, 
lanate or glabrate beneath; heads large, the narrow rays orange or 
flame-colored. 

GNAPHALIUM L. 

Annual or perennial herbs, sometimes suffrutescent, the pubes- 
cence whitish, lanate; leaves alternate, entire, often decurrent; heads 
small, disciform and glomerate, corymbose, spicate, or racemose; 
phyllaries scarious; receptacle plane to conic, naked; pistillate flowers 
in several series, their corollas filiform; central flowers few, perfect, 
tubular; achenes oblong or obovate, terete or slightly compressed, 
ecostate; pappus a single series of capillary bristles, coherent at the 
base or separately deciduous. A few other species are known from 
Central America. The specific limits in this group are poorly under- 
stood, and the genus is seriously in need of careful revision. Some 
of the names used below are of doubtful application to the Costa 
Rican plants. 

Gnaphalium attenuatum DC. Ajenjillo cimarron. Common 
in pastures and thickets of the Meseta Central, and on the slopes of 
the neighboring mountains; region of Dota; chiefly at 1,100-2,400 
meters. Panama to Mexico, and probably extending into South 



FLORA OF COSTA RICA 1479 

America. Plants annual or probably persisting for more than one 
year, erect, branched, mostly 50-90 cm. high, the stems densely 
white-lanate; leaves linear or lance-linear, often 7 mm. wide, long- 
attenuate at each end, sessile or nearly so, not decurrent, densely 
lanate beneath, usually green and glabrate above; heads white, aggre- 
gate in small, rounded clusters, or the clusters sometimes rather lax 
and open, usually disposed in broad panicles. It is by no means cer- 
tain that the Costa Rican material is the same as the Mexican plant 
that typifies this species, but G. attemtatum is the conventional name 
used by most botanists in naming any Gnaphalium from Mexico or 
Central America without any outstanding characters to force its 
reference to some different species. 

Gnaphalium indicum L. I have seen one specimen of this 
species from San Jose", at 1,160 meters, M. Valeria 204; it is probable 
that other collections that have been determined as G. purpureum L. 
may be referable here. Native probably of the Old World tropics, 
but naturalized in the West Indies, and in South America. A slender 
annual, the stems 30 cm. long or less, very weak and decumbent; 
leaves spatulate to narrowly obovate, 1-4 cm. long, obtuse, densely 
lanate on both surfaces, narrowed to a somewhat amplexicaul base, 
thin; heads small, yellowish, forming dense, leafy spikes; involucre 
2 mm. long. 

Gnaphalium lavandulaefolium (HBK.) Blake. Chionolaena 
Lavandulaceum Benth. & Hook, ex Hemsl.; G. lavandulaceum DC. 
Thickets of the upper slopes of Volcan de Irazu, and Cerro de La 
Muerte, 3,000-3,300 meters. Also in Mexico. Plants densely tomen- 
tose, suffrutescent, much branched, about 30 cm. high, the branches 
densely leafy; leaves ascending, or the older ones reflexed, about 8 
mm. long; heads densely clustered at the tips of the branches, the 
phyllaries white at the tip, dark brown at the base. 

Gnaphalium rhodarum Blake, Journ. Wash. Acad. Sci. 17: 61. 
1927. In paramo, Cerro de Las Vueltas, Prov. San Jose", 2,700- 
3,000 meters, Standley & Valeria 43623. Also on Cerro de La Muerte. 
Endemic. Plants annual, single-stemmed, about 30 cm. high, erect, 
branched only above; leaves almost uniform, 2.5-5 cm. long, lanceo- 
late, acuminate, broadest near the base and decurrent, green and 
glandular above, arachnoid-tomentose beneath; heads pink, in dense 
clusters crowded in a small panicle; involucre 4 mm. high. 

Gnaphalium roseum HBK. G. roseum var. hololeucum Benth. 
in Oerst. Vid. Medd. Kjoebenhavn 1852: 105. 1853 (summit of Vol- 



1480 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

can de Irazu, 3,300 meters, Oersted (duplicate of type, Oersted 10577, 
in Herb. Field Mus.);G. roseum var. angustifolium Benth. loc. cit. 
(near Cartago, 1,500 meters, Oersted); G. roseum var. sordescens 
Benth. loc. cit. (near Cartago, Oersted 10576; duplicate of the type 
in Herb. Field Mus.). G. roseum var. stramineum Kuntze, Rev. Gen. 
1: 340. 1891 (type from Costa Rica, Kuntze). Frequent in meadows 
or thickets on the slopes of Irazu and Poas, 2,500-3,400 meters, and 
probably elsewhere. Ranging to Mexico. Plants perennial, with 1 or 
several stems from the base, often branched above, densely white- 
lanate almost throughout; leaves lance-linear, acute or acuminate, 
somewhat dilated, amplexicaul, and slightly decurrent at the base, 
rarely glabrate on the upper surface; heads small, pink to white or 
yellowish, densely glomerate, the clusters grouped in small, rounded, 
headlike panicles. The plants referred here are especially variable 
in color of the bracts, but all seem to represent a single species, which, 
however, may not really be G. roseum of Mexico. Material placed by 
Klatt under G. Sprengelii Hook. & Arn. may be referable here. 
Most Costa Rican collections named G. oxyphyllum DC. likewise 
may represent this same species. 

Gnaphalium spicatum Lam. G. purpureum var. macrophyllum 
Greenm. Proc. Amer. Acad. 39: 97. 1903 (Copey, 1,800 meters, 
Tonduz 1 1 771 ) . Frequent in meadows or other grassy places, Meseta 
Central, ascending the slopes of the mountains to 2,500 meters or 
more; region of San Ramon, 1,000 meters; Dota. Widely distributed 
in tropical America. Plants probably annual, perhaps sometimes 
perennial, simple or often branched from the base, erect or ascending, 
rather stout, commonly about 30 cm. high; leaves spatulate or linear- 
spatulate, 2-6 cm. long, obtuse or rounded at the apex, green and 
glabrate above, white-lanate beneath, narrowed to the sessile base; 
heads small, usually rather dark brown, forming a dense, leafy, 
terminal spike, the phyllaries glabrous. 

Gnaphalium subsericeum Blake, Journ. Wash. Acad. Sci. 17: 
61. 1927. Wet meadow, La Palma, Prov. San Jose 1 , 1,600 meters, 
Standley 32941 Collected also at Cartago, and apparently frequent 
in the region of San Ramon, descending to 1,000 meters. Endemic. 
Plants annual or perennial, about 30 cm. high, the stems mostly 
solitary, branched above; leaves linear, very narrow, 4-6 cm. long, 
green above, densely white-sericeous or lanate beneath, narrowed to 
each end, not decurrent, the margins revolute; heads 4 mm. high, 
white, densely glomerate, the clusters few, forming a small panicle; 
phyllaries sparsely arachnoid-tomentose below. The Costa Rican 



FLORA OF COSTA RICA 1481 

species is compared by its author with G. salicifolium (Bertol.) 
Schultz Bip., of Guatemala and Mexico, to which it is perhaps too 
closely related for convenient separation. 

GYNURA Cass. 

Herbs, the leaves alternate, entire to pinnate-lobate or dissected; 
heads medium-sized, discoid, solitary or corymbose at the ends of the 
branches, yellow or purplish, the flowers all perfect; involucre cylin- 
dric or campanulate, the phyllaries narrow, equal, 1-seriate, smaller 
ones sometimes present at the base of the involucre; receptacle plane, 
naked or short-fimbrillate; achenes 5-10-striate, glabrous; pappus 
of numerous many-seriate, soft, white bristles. The species are 
natives of the Old World. 

Gynura aurantiaca DC. Tunica del Nazareno, Tornasol. 
Planted for ornament in gardens of the Meseta Central. Native of 
Java. Plants a meter high or less, stout, branched, densely covered 
with violet or purple hairs; leaves ovate, large and soft, irregularly 
dentate, the upper ones amplexicaul; heads yellow or orange, 1.5 
cm. high, forming a lax, terminal corymb. 

HELENIUM L. 

Reference: Rydberg, N. Amer. Fl. 34: 120. 1915. 

Annual or perennial herbs; leaves alternate, impressed-punctate, 
usually decurrent and forming wings on the stems; heads medium- 
sized, solitary or corymbose, radiate; involucre at first rotate, the 
phyllaries narrow, 2-3-seriate, subequal; receptacle convex to conic, 
naked; ray flowers yellow, pistillate and fertile, the disk flowers per- 
fect and fertile, yellow, brown, or purple; achenes obpyramidal, 4-5- 
angulate, with the same number of intermediate costae; pappus of 
4-8 scarious squamellae. A few other species are known from 
Central America. 

Helenium centrale Rydb. Near Cartago, 1,270 meters, col- 
lected only by Cooper; perhaps introduced here. Ranging to Guate- 
mala. A puberulent annual, probably 60 cm. high or more, the 
stems rather broadly winged; lowest leaves oblanceolate, 5-10 cm. 
long, entire or denticulate, the cauline leaves linear-lanceolate, 
attenuate; heads corymbose, long-pedunculate; phyllaries 4 mm. 
long; rays 4 mm. long; disk brown, globose, 1 cm. broad; achenes 1.5 
mm. long, hispid on the angles. The Costa Rican collection was 
referred by Klatt to H. mexicanum HBK. 



1482 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

HELIANTHUSL. Sunflower 

Coarse, annual or perennial herbs; leaves opposite or alternate, 
simple; heads large, radiate, usually long-pedunculate, corymbose 
or solitary, the rays yellow, the disk yellow, brown, or purple; invo- 
lucre hemispheric or depressed, the phyllaries imbricate in several 
series; receptacle flat to conic, paleaceous; ray flowers neutral, the 
disk flowers perfect, fertile; achenes thick, oblong or obovate, com- 
pressed or slightly 4-angulate; pappus of 2 scales or awns, deciduous. 
Probably no species are native in Central America. 

Helianthus annuus L. GirasoL Cultivated occasionally for 
ornament. Native of the western United States. A well known 
plant, often becoming a pernicious and extremely abundant weed in 
many parts of the United States. Cultivated plants usually have 
much larger heads than wild ones, the disk of the head in some forms 
becoming almost 30 cm. broad. The seeds are edible, and in Russia, 
for example, the plant is much grown as a source of human food and 
of oil. The seeds are used commonly as food for parrots and other 
cage birds. 

HELICHRYSUM Gaertn. 

Herbs or shrubs, usually tomentose or lanate; leaves alternate, 
entire; heads large or small, said to have sometimes as many as 
1,000 flowers, solitary or corymbose, disciform; involucre scarious, 
usually white or brightly colored in yellow or red, imbricate in 
numerous series, erect or spreading; achenes small, 5-angulate or 
terete; pappus of numerous slender bristles. All the species are 
natives of the Old World. 

Helichrysum bracteatum (Vent.) Andr. Oropel. Straw-flower. 
A stout, branched annual, a meter high or less, the stems glabrous 
or nearly so; leaves oblong-lanceolate, short-petiolate, green; heads 
2.5-5 cm. broad, yellow or orange, the phyllaries short, obtuse. 
There are numerous color varieties with white or red heads. The 
dried flower heads are very ornamental, and may be preserved for 
a long time in bouquets. They are often used for preparing funeral 
wreaths. 

HELIOPSIS Pers. 

Annual or perennial herbs; leaves opposite, 3-nerved, dentate; 
heads usually large, long-pedunculate, terminal and axillary, radiate, 
yellow; involucre hemispheric, the phyllaries oblong or lanceolate, 
imbricate in 2-3 series; receptacle convex or conic, paleaceous; ray 
flowers pistillate, fertile, the large rays firm and persistent upon the 



FLORA OF COSTA RICA 1483 

achene; disk flowers perfect; achenes obtusely 3-4-angulate, trun- 
cate; pappus none or of 2-4 teeth, or a coroniform border. Probably 
no other species reach Central America. 

Heliopsis buphthalmoides (Jacq.) Dunal. Slopes of Volcan 
de Irazu, above Cartago, at 1,400 meters or less. Mexico to Peru. 
Plants about a meter high, branched, rather slender, sparsely pubes- 
cent with short and rather soft hairs, or almost glabrous; leaves long- 
petiolate, thin, green, ovate or broadly ovate, acute or acuminate, 
truncate or rounded at the base, rather coarsely dentate; heads 
showy, on long, naked peduncles, the disk more than 1 cm. broad, 
the spreading, rather broad rays 1.5 cm. long. 

HIDALGOA Llave & Lexarza 

Plants scandent, herbaceous or suffrutescent, slender, glabrous 
or nearly so; leaves opposite, compound, ternate or biternate, the 
leaflets petiolulate, broad, serrate, the lateral ones often parted; 
heads axillary, solitary, radiate, long-pedunculate, the rays red or 
orange; ray flowers pistillate or neutral, the disk flowers perfect, 
fertile; involucre biseriate, the inner phyllaries few and broad, sub- 
equal, connate at the base, the outer ones smaller, foliaceous; recep- 
tacle plane, paleaceous; achenes oblong or obovate, dorsally 
compressed, rounded at the apex, naked or obsoletely 2-dentate. 
The genus consists of a single species. The group is noteworthy for 
its scandent habit and apparently compound leaves. 

Hidalgoa ternata Llave & Lex. H. Wercklei Hook. f. Bot. Mag. 
125: pi. 7684- 1899 (described from cultivated material said to have 
been obtained in Costa Rica by Werckle"). Childsia Wercklei Childs 
ex Hook. f. loc. cit. in syn. Frequent in thickets or forest in some 
places in the mountains of Guanacaste, 700 meters; region of La 
Palma de San Jose". Mexico to Ecuador. A large, chiefly herbaceous 
vine; leaves ternate and with 3 leaflets, or biternate, the leaflets ovate 
or broadly ovate, sometimes elliptic, thin, acute or acuminate, 
coarsely serrate; pseudostipules orbicular, herbaceous; disk about 13 
mm. long; rays 2 cm. long or more, orange-red. 

HIERACIUM L. 

Perennial herbs, usually pilose; leaves alternate or chiefly basal, 
entire to pinnatifid; heads medium-sized or rather large, commonly 
yellow, solitary, corymbose, or paniculate; involucre cylindric to 
hemispheric, the principal phyllaries 1-3-seriate, the outer ones 
shorter; receptacle flat, naked or short-fimbrillate; corollas all alike, 



1484 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

liguliform; achenes oblong or fusiform, not rostrate, 10-15-costate, 
terete or 4-5-angulate; pappus of 1-2 rows of rather stiff, brownish 
bristles. A few other species have been collected in Central America. 

Hieracium abscissum Less. Oak forest in the region of Dota, 
and doubtless elsewhere. Extending to Mexico. Plants slender, 
erect, about 60 cm. high or less, minutely glandular, deflexed-pilose 
near the base with long, white hairs, leafy below; leaves setulose and 
pilose, borne chiefly at or near the base of the stem, 25 cm. long and 
2.5 cm. wide or usually much smaller, subobtuse, remotely repand- 
denticulate; heads paniculate, numerous; involucre 5-7 mm. long, mi- 
nutely glandular, floccose; achenes black, 2-2.5 mm. long, columnar. 

Hieracium frigidum Wedd. is reported by Klatt from Cerro de 
Buena Vista, but the record needs verification. Probably the plant 
so reported is one of the following species. 

Hieracium irasuense Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 113. 1853. Papelillo macho. Volcan de Irazu, 2,850 
meters, Oersted. In thickets or meadows, upper slopes of Irazu and 
Turrialba; also near Aserri; chiefly at 2,400-3,400 meters. Endemic. 
Plants about 30 cm. high, short-pubescent and glandular-pilose, the 
cauline leaves mostly reduced and bractlike; basal leaves lanceolate, 
7-10 cm. long, entire or glandular-serrate, long-pilose above, densely 
lanate beneath with long, white hairs; heads corymbose, yellow, the 
involucre 6 mm. long, rather densely covered with intermixed simple 
and glandular hairs; achenes subfusiform, 10-costate. 

Hieracium sphagnicola Blake, Journ. Wash. Acad. Sci. 17: 
62. 1927. Open sphagnum bog, Laguna de La Chonta, northeast 
of Santa Maria de Dota, Prov. San Jose", 2,000 meters, Standley 
42139. Known certainly only from the original locality, but a collec- 
tion from Cerro Gallito is probably conspecific. Basal leaves 
oblanceolate, 17 cm. long or less, sparsely long-pilose; stems sparsely 
long-pilose below, with 2 or fewer leaves below the middle; heads 
5-8, cymose, the involucre 8-11 mm. long, sparsely pilose and stipi- 
tate-glandular; achenes slightly tapering above or almost columnar, 
3-4 mm. long. 

Hieracium Standleyi Blake, Journ. Wash. Acad. Sci. 17: 64. 
1927. In paramo, Cerro de Las Vueltas, Prov. San Jose', 3,000 
meters, Standley & Valeria 43624. Known only from the original 
collection. Plants lanate at the base, the stems scapose; basal leaves 
obovate, 3.5-4 cm. long, rounded and apiculate at the apex, ob- 



FLORA OF COSTA RICA 1485 

scurely denticulate, sparsely long-pilose; heads few, the involucre 
9-11 mm. long, pilose at the base, not glandular; achenes deep 
purple-brown, slightly tapering above, 2.5-3.8 mm. long. 



Hinterhubera Laseguei Wedd. was reported from Costa Rica by 
Klatt, on the basis of Pittier 3431 from Cerro de Buena Vista. 
The collection is actually Alchemilla ocreata Donn. Smith (see p. 478; 
Rosaceae). 

HYMENOSTEPHIUM Benth. 

Tall, coarse herbs, or the plants often suffrutescent; leaves chiefly 
opposite, petiolate, ovate or lanceolate, dentate; heads small or 
medium-sized, cymose or cymose-paniculate, radiate, yellow; in- 
volucre 2-3-seriate, the bracts subequal or the outer ones gradually 
shorter, ovate or lance-ovate, usually narrow and not foliaceous- 
appendaged; achenes obovoid, somewhat compressed, pubescent or 
glabrous; pappus of a few unequal squamellae without awns, or 
none. One or two other species grow in northern Central America. 

Hymenostephium cordatum (Hook. & Arn.) Blake. Gym- 
nopsis costaricensis Benth. in Oerst. Vid. Medd. Kjoebenhavn 1852: 
90. 1853 (Aguacate and Ujarras, Oersted). Aspilia costaricensis Klatt, 
Bull. Soc. Bot. Belg. 31: 201. 1892. Common in thickets or forest, 
Meseta Central to the Atlantic coast, and in the upper part of the 
Pacific tierra caliente; region of San Ramon; mostly at 1,300 meters 
or less. Mexico to Colombia. Plants herbaceous or somewhat 
shrubby, usually 1-2 meters high, occasionally subscandent, the 
branches sparsely strigillose or glabrate; leaves slender-petiolate, 
mostly 6-10 cm. long, broadly ovate to lance-ovate, acuminate, 
usually abruptly narrowed at the broad base, coarsely serrate, 
strigillose beneath; heads numerous, in small cymes, the involucre 
5 mm. high; achenes glabrous or pubescent, the pappus none or of 
a few unequal squamellae. Material reported by Klatt as Gymno- 
lomia rudbeckioides HBK,, a South American species of Hymeno- 
stephium, may be referable here. 

Hymenostephium guatemalense (Rob. & Greenm.) Blake. 
H. pilosulum Blake. Thickets in the regions of San Jose" and Cartago, 
1,100-1,300 meters. Panama to southern Mexico. Similar to the 
preceding species, to which it is rather too closely related, but differ- 
ing in the more abundant and spreading pubescence of the branches 
and lower leaf surface. 



1486 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

HYPOCHAERIS L. 

Low, perennial herbs, the stems usually naked, often branched; 
leaves all or chiefly basal, entire to pinnatifid; heads rather large, 
long-pedunculate, yellow, the corollas all liguliform; involucre 
cylindric to campanulate, the phyllaries green, narrow, imbricate 
in several series, the outer shorter; receptacle flat, paleaceous; 
achenes oblong to linear, 10-costate, rostrate, the pappus of plumose 
bristles. No species are native in North America. 

Hypochaeris radicata L. Margarita amarilla. Common in 
meadows of the middle and upper slopes of the central volcanoes; 
region of San Ramon; at 1,200-2,700 meters. Native of Europe, 
but rather widely naturalized in America. Stems glabrous, 30-60 
cm. high, usually branched; leaves oblanceolate to obovate, dentate 
to pinnatifid, 5-15 cm. long, hirsute; heads about 2.5 cm. high, 
the phyllaries glabrous or sparsely pilose. 

ICHTHYOTHERE Mart. 

Herbs or somewhat suffrutescent plants, glabrous or pubescent; 
leaves opposite, sessile or petiolate, entire or somewhat serrate; 
heads small, disciform, whitish, sessile or short-pedicellate, in small, 
dense cymes; outer phyllaries 1-2-seriate, small, the inner ones and 
the outer paleae of the disk broad, subtending the perfect flowers; 
receptacle oblong or linear; achenes obovoid, dorsally compressed, 
smooth or costate; pappus none. The other species of the genus 
are South American. 

Ichthyothere scandens Blake. Regions of Guapiles and El 
General, in the tierra caliente. Venezuela to Peru. A suffrutescent 
vine(?), the branches appressed-pilose; leaves slender-petiolate, ovate 
or lance-ovate, 7-11 cm. long, falcate-acuminate, acute or acuminate 
at the base, serrulate, sparsely sordid-pubescent on the veins, 3-5- 
plinerved; heads pedicellate, 4.5 mm. high; outer phyllaries subulate- 
triangular, the inner obovoid; achenes about 8-costate, truncate, 
glabrous, 2-3 mm. long. 

ISOCARPHA R. Br. 

Erect, slender herbs; leaves opposite or alternate, petiolate or 
sessile, entire or dentate; heads rather small, discoid, whitish, mostly 
solitary at the end of a long peduncle; phyllaries narrow, small, 
2-3-seriate, rigid-tipped; receptacle conic, paleaceous; achenes 4-5- 
angulate, truncate; pappus none. No other species are found in 
Central America. 



FLORA OF COSTA RICA 1487 

Isocarpha atriplicifolia (L.) R. Br. Nicoya; Patarra. West 
Indies and northern South America. Rather similar to the more 
common /. oppositifolia, but more branched, the leaves broader, 
chiefly ovate, usually conspicuously dentate; petioles auriculate- 
dilated at the base; heads shorter, very obtuse at the base; achenes 
pubescent. 

Isocarpha oppositifolia (L.) R. Br. Isla de Chira, Pacific 
coast, M. Valeria. Texas and Mexico to Venezuela and West Indies. 
Plants branched, erect, pubescent, usually less than a meter high; 
leaves sessile or short-petiolate, lanceolate or linear-lanceolate, obtuse 
to attenuate, acute at the base, entire or nearly so; petioles not auric- 
ulate at the base; phyllaries glandular and appressed-pilosulous; 
achenes 2 mm. long, glabrous. 



Iva annua L., a South American plant, was reported by Klatt 
from "Bolenak (Haut Zhorquin)" on the basis of Pittier 8656. I 
have not seen this collection, and have no idea of what it may be. 

JAEGERIA HBK. 

Slender, branched annuals, the pubescence usually abundant; 
leaves opposite, dentate; heads very small, radiate, yellow, terminal 
or axillary, long-pedunculate, solitary or cymose, the rays minute 
and inconspicuous; involucre campanulate, the phyllaries 1-2-seriate, 
subequal; receptacle convex or conic, paleaceous; achenes obscurely 
5-angulate, glabrous; pappus none. Probably a single species occurs 
in Central America. 

Jaegeria hirta (Lag.) Less. Miekilla. Common in meadows 
of the central region, especially on the mountain slopes; mostly at 
1,000-2,600 meters. Widely distributed in tropical America. Plants 
slender and rather weak, usually 30 cm. high or less, rather densely 
pilose throughout; leaves sessile, oblong or ovate-oblong, 2-4 cm. 
long, obtuse or subacute, 3-plinerved, often entire; heads 3-4 mm. 
long. In general appearance the plant is much like Galinsoga, but 
differs at first glance in its yellow rather than white or pink rays. 
The rays, however, are so small that they are easily overlooked. 

JUNGIA L. f. 

Herbs or shrubs, erect or subscandent; leaves alternate, usually 
rounded-cordate, dentate or lobate, most often tomentose beneath; 
heads small or medium-sized, paniculate at the ends of the branches, 
the flowers all alike or nearly so, the heads appearing discoid; 



1488 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

involucre campanulate or cylindric, the phyllaries subequal, 1-2- 
seriate, the outer sometimes shorter; receptacle paleaceous; achenes 
oblong or linear, sub terete, 5-costate, contracted above and rostrate; 
pappus bristles 1-seriate, plumose to denticulate. No other species 
are known from North America. 

Jungia ferruginea L. f. Occasional in forests of the higher 
mountains, 1,600-2,600 meters. Colombia. Plants scandent, the 
stout branches more or less brownish-tomentose; leaves slender- 
petiolate, cordate-orbicular, mostly 5-9 cm. wide, shallowly or deeply 
cordate at the base, shallowly lobate and coarsely crenate, densely 
villous above, densely brownish-tomentose beneath; heads white 
or yellow, in small, dense cymes, the involucre 4 mm. long, densely 
pubescent; achenes glabrous, scarcely more than 2 mm. long. 
Material of this genus at hand for study is altogether inadequate. 
There is some probability that the plant of Central America repre- 
sents a distinct and endemic species. Part of the material was 
referred by Klatt to J. spectabilis Don. 

LACTUCA L. 

Tall herbs with milky sap; leaves alternate, entire to pinnatifid; 
heads small, paniculate, yellow, blue or white; involucre cylindric, 
the phyllaries imbricate in several series, the outer ones shorter; 
receptacle plane, naked; heads appearing radiate, the corollas liguli- 
form; achenes oval to linear, compressed, 3-5-costate, contracted 
into a slender beak; pappus of numerous soft, capillary, white or 
brown bristles. One species is native in northern Central America. 

Lactuca sativa L. Lechuga. Lettuce. The garden lettuce, 
believed to be a native of Asia, is cultivated as commonly in Costa 
Rica as in other parts of the earth for use as a salad plant. It 
thrives at all elevations. 

LAESTADIA Kunth 

Perennial herbs, sometimes suffrutescent, cespitose or prostrate, 
glabrous or pubescent; leaves alternate or subopposite, small, 
crowded, entire; heads small, discoid, solitary on elongate, terminal 
and axillary peduncles, violaceous; involucre hemispheric, the 
phyllaries about 2-seriate, narrow, subequal; receptacle plane or 
subconvex, naked; achenes scarcely compressed, 7-10-costate, gla- 
brous, contracted at the apex into a short beak; pappus none. 
No other species are known in North America. 



FLORA OF COSTA RICA 1489 

Laestadia costaricensis Blake, Contr. U. S. Nat. Herb. 22: 
593. 1924. Cerro de Las Vueltas, 3,000 meters, Pittier 10500. 
Known only from the original locality, where it was collected also 
by Juvenal Valerio and the present writer. Endemic. A low plant, 
forming dense tufts, the stems 25 cm. long or less, rooting at the 
nodes, very slender, puberulent; leaves narrowly spatulate or linear- 
oblanceolate, 6-11 mm. long, acute, entire, sparsely pubescent; 
heads 3 mm. high, the phyllaries glandular and ciliate; achenes 1.5 
mm. long. The same plant was reported by Klatt from Cerro de 
La Muerte (Cerro de Buena Vista) as L. Lechleri Schultz Bip., a 
South American species. 

LIABUM Adans. 
Reference: Rydberg, N. Amer. Fl. 34: 289-301. 1927. 

Perennial herbs or shrubs, often epiphytic and more or less 
scandent; leaves opposite, entire or dentate; heads radiate or discoid, 
yellow, the phyllaries imbricate in 3 or more series; receptacle naked 
or often hirsute or paleaceous; ray flowers pistillate and fertile, the 
disk flowers perfect and fertile; achenes cylindric, 10-15-costate; 
pappus of numerous capillary, scabrous bristles in a single series, 
a few short ones sometimes present. A few additional species 
exist in Central America. 

Liabum Bourgeaui Hieron. Occasional in thickets or forest 
of the central region, at Cartago and on the mountain slopes at 
middle elevations; region of San Ramon; extending into the upper 
part of the tierra caliente. Panama to Mexico. A shrub, terrestrial 
or epiphytic, 1.5-2.5 meters high, the branches white-tomentose; 
petioles broadly winged, conspicuously dilated and clasping at the 
base; leaf blades ovate to elliptic, 8-20 cm. long, acuminate, con- 
tracted and decurrent at the base, denticulate, glabrate and green 
above, densely white-tomentose beneath, triplinerved; involucres 
8-9 mm. high, the phyllaries all acute or attenuate; achenes 1 mm. 
long, hispidulous. 

Liabum discolor (Hook. & Arn.) Benth. & Hook. In forest, 
slopes of Barba and Poas, mostly at 2,200 meters or less. Ranging 
to Mexico. A shrub of 3-4.5 meters, the stout branches glabrous; 
leaves slender-petiolate, broadly ovate to lance-ovate, 8-15 cm. 
long, acute or acuminate, more or less narrowed at the base, denticu- 
late, glabrous above, white-tomentose beneath; involucre 8 mm. 
high, the inner phyllaries rounded at the apex; achenes glabrous. 



1490 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Liabum igniarium (Humb. & Bonpl.) Less. Reported from 
the region of TeYraba and Boruca by Klatt, and from Costa Rica 
by Rydberg, but I have seen no Costa Rican specimens. Extending 
to Peru. Similar in most characters to L. Bourgeaui, but the petioles 
naked, abruptly dilated at the base into large, green, rounded 
pseudostipules. 

Liabum polyanthum Klatt, Bull. Soc. Bot. Belg. 31: 209- 
1892. Sinclairia polyantha Rydb. N. Amer. Fl. 34: 299. 1927. S. 
Pittieri Rydb. op. cit. 300 (near Alajuelita, Prov. San Jose", Pittier 
9093) . El General, Pittier 431 9. Frequent in forests of central region, 
descending into the upper part of the tierra caliente, mostly at 500- 
1,500 meters; region of San Ramon; Guanacaste. Endemic. 
Branches laxly floccose-tomentose; leaves slender-petiolate, oval 
to rounded, 10-15 cm. long, acute or acutish, rounded at the base, 
glabrate above, white- tomentose beneath; heads numerous, radiate, 
the involucre 7 mm. high, the inner phyllaries rounded at the apex; 
achenes strigose. 

Liabum Tonduzii Robinson, Proc. Bost. Soc. Nat. Hist. 31: 
270. 1904. Sinclairia Tonduzii Rydb. N. Amer. Fl. 34: 298. 1927. 
Rio Virilla near San Jose", 1,100 meters, Tonduz 7064- Collected 
also at Santa Maria de Dota. Endemic. A shrub of 3-4.5 meters, 
the young branches tomentose and somewhat brown-hirsute; leaves 
petiolate, rhombic-ovate, 7-15 cm. long, acuminate, rounded to 
subacute at the base, green and scaberulous above, densely white- 
tomentose beneath; involucres 6-7 mm. high, the inner phyllaries 
rounded at the apex; achenes hirsutulous, 2.5 mm. long. 

Liabum Valerii Standl., sp. nov. Frutex scandens, ramis 
gracilibus primo tomento adpresso albido dense obtectis, internodiis 
elongatis; folia inter minora petiolata membranacea, petiolo gracili 
15-18 mm. longo dense tomentoso basi non vel vix dilatato; lamina 
lanceolato-oblonga 9.5-13 cm. longa 2.5-4.5 cm. lata longe sensim 
attenuato-acuminata, basi rotundata usque subacuta, remote obscure 
serrulata, supra primo laxe floccosa cito glabrata, subtus ubique 
tomento denso pallide brunnescente vel fere albo obtecta; capitula 
in cymas axillares parvas breviter pedunculatas disposita, graciliter 
usque 8 mm. longe pedicellata; phyllaria 3-4-seriata lineari-lanceo- 
lata vel lanceolata, omnia acuta vel interiora acute attenuata, dorso 
glabrata, apicem versus ciliata, involucro ca. 5 mm. longo; capitula 
ut videtur discoidea, floribus non visis; achaenia immatura paullo 
ultra 2 mm. longa breviter hispidula, pappi setis fulvis 6 mm. longis. 



FLORA OF COSTA RICA 1491 

In wet forest, near Finca La Cima, above Los Lotes, north of 
Copey, Prov. San Jose', 2,100-2,400 meters, Standley 42555. Dis- 
tinguished from all other Costa Rican species by the definitely 
scandent habit and narrow leaves. 

Liabum vulcanicum Klatt. La Hondura de San Jose", in wet 
forest. Colombia. Plants about 2 meters high, chiefly herbaceous, 
the stems fistulous, branched; leaves long-petiolate, broadly ovate- 
cordate, about 20 cm. long and wide, sinuate-lobate and dentate, 
green but somewhat pilose on both surfaces; heads radiate, long- 
pedicellate and subumbellate, forming a large panicle; phyllaries 
obtuse, broad; achenes glabrous. Very different in appearance 
from other Central American species because of the lobate leaves, 
green on the lower surface. 

LYCOSERIS Cass. 

Erect or subscandent shrubs, the branches usually at least 
recurved; leaves alternate, sessile or short-petiolate, commonly 
entire, glabrous above, more or less tomentose beneath, 3-5-pli- 
nerved; heads dioecious, very large, solitary or few at the ends of 
the branches, the pistillate usually larger than the staminate, the 
corollas orange-yellow, the outer ones elongate and liguliform; 
involucre campanulate or subglobose, the phyllaries multiseriate, 
imbricate, narrow, very numerous; receptacle plane, fimbrilliferous; 
achenes oblong or elongate, 5-costate and usually multistriate, 
glabrous; pappus of numerous slender bristles. One other species 
occurs in Panama. 

Lycoseris crocata (Bertol.) Blake. L. squarrosa Benth. Bot. 
Voy. Sulph. 121. 1844 (based in part upon material from Nicoya). 
L. macrocephala Greenm. Proc. Amer. Acad. 39: 119. 1903 (forests 
of the hills of Nicoya, Tonduz 13602}. Region of Nicoya. Panama 
to Guatemala. A shrub with recurved and striate branches; leaves 
lanceolate or lance-oblong, 3-5-plinerved, glabrous above, thinly 
white-lanate beneath; heads 2.5 cm. or more in diameter; phyllaries 
narrowly lanceolate, attenuate, rather densely arachnoid-tomentose. 

Lycoseris grandis Benth. in Oerst. Vid. Medd. Kjoebenhavn 
1852: 11. 1853. Onoseris grandis Kuntze, Rev. Gen. 1: 354. 1891. 
San Mateo, Oersted. Region of San Mateo and at various places 
along the Pacific coast; Nicoya. Endemic(?). Similar to the preced- 
ing species, but the heads usually larger, the bracts broad, obtuse 
or merely apiculate, glabrous or nearly so. 



1492 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

MATRICARIA L. 

Annual herbs, glabrous or somewhat pubescent, strong-scented; 
leaves alternate, once to thrice pinnatifid into narrow divisions; 
heads solitary or corymbose, radiate or discoid; involucre depressed 
to hemispheric, the phyllaries 2-4-seriate, somewhat imbricate, 
subequal, obtuse, scarious-marginate; receptacle conic to subglobose, 
naked; rays usually white, the disk yellow; achenes somewhat 
asymmetric, 3-5-costate on the inner side; pappus a more or less 
evident corona or margin. No species are native in Central America. 

Matricaria Chamomilla L. Manzanilla. Planted commonly in 
gardens for use in domestic medicine; sometimes escaping to waste 
ground. Native of Europe and Asia. Plants 50 cm. high or less, 
usually much branched, glabrous or nearly so; heads solitary at the 
ends of the branches, the rays white; involucre about 3 mm. high, 
the ligules 5-8 mm. long. A decoction of the plant is one of the 
local remedies for pains in the stomach. 

MELAMPODIUM L. 

Reference: B. L. Robinson, Synopsis of the genus Melampodium, 
Proc. Amer. Acad. 36: 455-466. 1901. 

Annual or perennial herbs, pubescent or glabrate, usually dichoto- 
mous-branched ; leaves opposite, entire to pinnate-lobate; heads 
small or medium-sized, radiate, yellow, pedunculate, inserted in 
the axils of the branches or of the leaves; involucre double, the 
outer phyllaries 4-5, broad, often connate at the base and accrescent 
after anthesis, the inner ones closely investing the ray flowers and 
closed above the achene after anthesis; receptacle convex or conic, 
paleaceous; achenes obovoid, without pappus. A few other species 
are found in Central America. 

Melampodium brachyglossum Bonn. Smith. In thickets or 
waste ground, vicinity of San Jose" and Cartago. Extending to 
Guatemala. Plants erect, sparsely branched, 90 cm. high or less, 
the stems pilose with weak, spreading hairs; leaves long-petiolate, 
deltoid-ovate, acute or acuminate, cuneate at the base and long- 
decurrent, remotely and inconspicuously dentate, rough-pilose; heads 
long-pedicellate, 4-5 mm. broad, the pedicels densely spreading- 
pilose, the outer phyllaries broad, obtuse, the rays minute, hidden 
by the phyllaries. 

Melampodium divaricatum (Rich.) DC. M. paludosum HBK. 
Common in waste and cultivated ground of the Pacific tierra caliente; 



FLORA OF COSTA RICA 1493 

Guanacaste; the species should be found also in the Atlantic tierra 
caliente, but I find no specimens or reports for that region ; reported 
from the region of Dota. Mexico to West Indies and northern 
South America. Plants erect, rather stout, often much branched, 
a meter high or less, the stems usually glabrous or nearly so; leaves 
on long or short petioles, rather large, broadly ovate to lance-ovate, 
acute or acuminate, acute at the base and decurrent, rather coarsely 
dentate, rough-pubescent or glabrate; heads mostly long-peduncu- 
late, about 8 mm. broad, the outer phyllaries broad, green, obtuse, 
the rays short but conspicuous. One of the most common weeds 
of the tierra caliente of Central America. 

Melampodipm flaccidum Benth. Waste ground or thickets, 
often in cultivated fields, Meseta Central to the tierra caliente. 
Extending to Mexico. Similar to M. divaricatum, but often lower, 
rather weak, frequently decumbent; heads mostly on short pedicels, 
or almost sessile, but sometimes long-pedicellate, about 7 mm. 
broad, the rays very small, hidden by the inner phyllaries. 

Melampodium oblongifolium DC. Mielilla. In thickets or 
waste ground, regions of San Jose" and Palmira, 1,000-1,800 meters. 
Extending to Mexico. Plants very slender, usually densely branched, 
sometimes a meter high, the stems pilose with long, spreading hairs; 
leaves small, ovate to narrowly lanceolate, acuminate, usually taper- 
ing to the base of the petiole, more or less serrate-dentate, rough- 
pubescent; heads very numerous, slender-pedicellate, 3-4 mm. broad, 
the outer phyllaries ovate or lanceolate, acute; rays minute. This 
has much smaller flower heads than any of the other species, and 
differs also in the small, narrow, acute phyllaries. 

Melampodium perfoliatum (Cav.) HBK. Collected only at 
La Caja, San Jose", probably introduced, M. Valerio 1345. Mexico 
and Guatemala. Plants stout, erect, a meter high or less, branched, 
the stems glabrous; leaves broadly ovate, their bases enlarged and 
united about the stem, rough-pubescent; heads numerous, on long 
or short peduncles, larger than in the other species, the broad, green 
outer phyllaries 1-1.5 cm. long, the rays inconspicuous. 

MELANTHERA Rohr 

Perennial herbs, usually large and coarse, with rough pubescence; 
leaves opposite, petiolate, dentate, often hastate-lobate; heads small, 
discoid, whitish, pedunculate; involucre hemispheric, the phyllaries 
few, ovate or lanceolate, subequal, imbricate in 2-3 series; receptacle 



1494 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

convex, paleaceous; achenes 4-angulate, the pappus of caducous 
awns. At least one other species is known from Central America. 

Melanthera nivea (L.) Small. Paira. M. aspera (Jacq.) Rich.; 
M. deltoidea Rich.; M. lanceolata Benth.; M. oxylepis DC. Common 
in thickets or waste ground, often in cultivated fields, Meseta Central 
to the coasts; region of San Ramon; Guanacaste. Generally distrib- 
uted in tropical America. Plants erect, much branched, sometimes 
3 meters high and then suffrutescent, rarely subscandent; leaves 
mostly ovate-oblong or lanceolate, hastate-lobate, serrate, 3-pli- 
nerved, very rough; heads about 1 cm. broad, the black anthers 
conspicuous against the general white of the head; pappus of 2-4 
short bristles. The plant is a common weed throughout a great 
part of Central America. There is probably no sadder spectacle in 
the field of systematic botany than the seriousness with which a 
large number of botanists have labored for a century or more, shuffling 
and reshuffling the plants, or rather specimens, of this small genus, 
painfully describing their own new "species," while reducing to 
synonymy those of earlier workers. As more and more material 
accumulates in herbaria, it becomes more apparent that the so-called 
species have no systematic value whatever, except for a very few 
localized plants that really do have good differential characters. 
Any one who has examined even casually the variation of leaf form 
in a colony of this plant, or even upon one individual, will have 
scant regard for most "species" of Melanthera. It is scarcely neces- 
sary to add that characters supposed to exist in the phyllaries and 
disk scales have no greater merit as specific characters. 

MIKANIA Willd. 

Reference: Robinson, Synopsis of the Mikanias of Costa Rica, 
Proc. Bost. Soc. Nat. Hist. 31: 254-257. 1904. 

Plants scandent, herbaceous or woody; leaves opposite, petiolate; 
heads discoid, white or pink, spicate, racemose, corymbose, or 
glomerate and often paniculate; involucre narrow, the phyllaries 4, 
narrow, subequal; flowers 4; receptacle naked; achenes truncate, 
5-angulate; pappus of numerous capillary bristles in 1-2 series. 
A few other species are known from Central America. The Costa 
Rican ones are none too well known, several of them being based 
upon single collections. It seems rather probable, too, that the 
available material may represent a greater number of forms than are 
named below. 



FLORA OF COSTA RICA 1495 

Mikania bogotensis Benth. Reported by Robinson from 
Palmares del General, Diquis Valley, Pittier 12080. Colombia. 
Plants slender, nearly or quite glabrous; leaves ovate, acuminate, 
subcordate at the base, thin, 3-nerved from the base; panicle large 
and open; heads sessile, spicate, 5 mm. high; phyllaries subobtuse; 
pappus rufous. 

Mikania cordifolia (L. f.) Willd. Frequent in forest and 
thickets of the central mountains, ascending to the upper part of the 
tierra caliente; region of San Ramon; Guanacaste. Widely distrib- 
uted in tropical America. A small or large vine, herbaceous or 
somewhat woody, the stems tomentose or tomentulose; leaves ovate, 
acute or acuminate, cordate at the base, undulate-dentate to almost 
entire, 3-5-nerved from the base, villous to sericeous, usually pale; 
heads dirty white, fragrant, corymbose, pedicellate; phyllaries 6-8 
mm. long, pubescent or glabrate; achenes glabrous, the pappus dull 
white, becoming rufescent. 

Mikania cristata Robinson, Proc. Amer. Acad. 47: 195. 1911. 
La Palma de San Jose", 1,460 meters, Tonduz 12583. Known, appar- 
ently, only from the original material. Stems puberulent-tomentu- 
lose; leaves long-petiolate, broadly ovate, acuminate, deeply cordate 
at the base, subentire, velutinous-tomentose, 3-5-nerved; petioles 
dilated at the base and connected by a cristate, appendaged stipular 
margin; heads corymbose, pedicellate; phyllaries subobtuse, 6 mm. 
long, puberulent dorsally; achenes glabrous, the pappus rufous. I 
have seen no material of this species. 

Mikania eupatorioides Blake, Journ. Wash. Acad. Sci. 28: 1938. 
Edge of forest, El General, Prov. San Jose", 1,525 meters, Skutch 
3041. Endemic, and known only from the region of the type. A 
woody vine, the branches pilosulous; leaves slender-petiolate, ovate 
or lance-ovate, 11-15 cm. long, long-acuminate, cuneate or rounded- 
cuneate at the base, entire or very obscurely denticulate, 5-plinerved, 
subglandular-puberulous above and sparsely pilosulous, similarly 
pubescent beneath; individual panicles convex, forming an open, 
leafy, thyrsoid panicle; pedicels 2 mm. long or less; heads 7-9 mm. 
high, the phyllaries puberulous toward the tip; corollas 4.5 mm. long; 
achenes 3 mm. long, the pappus flesh-colored, 4.2 mm. long. 

Mikania Guaco Humb. & Bonpl. Guaco. M. olivacea Klatt, 
Bull. Soc. Bot. Belg. 31: 195. 1892 (forests of Buenos Aires, 200-300 
meters, Pittier 4433). Occasional in thickets or forest of the tierra 
caliente, also in the central mountains, ascending to 1,500 meters; 



1496 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

region of San Ramon. Widely distributed in tropical America. 
Plants herbaceous or frequently woody, the stems glabrate; leaves 
ovate, often very large, acute or acuminate, cuneate-decurrent at 
the base, undulate or denticulate to entire, puberulent or glabrate 
above, tomentulose beneath, especially on the nerves, or glabrate, 
pinnate-nerved; heads dirty white, about 1 cm. long, mostly sessile 
and glomerate, forming a large, ovoid panicle; phyllaries puberulent, 
rounded at the apex; pappus buff or fulvous. In Costa Rica, as 
in other Central American countries, this plant is famous as a remedy 
for fevers and snake bites. 

Mikania Holwayana Robinson. Regions of Orosi and El 
Muneco. Ecuador. Stems puberulent or glabrate; leaves broadly 
ovate, short-acuminate, cordate at the base, denticulate or sinuate, 
puberulent above, whitish-tomentulose beneath, 3-5-nerved from 
the base; heads about 1 cm. long, pedicellate, forming a large, pyram- 
idal panicle; phyllaries obovate-elliptic, 6.5 mm. long, obscurely puber- 
ulent; corollas creamy white; achenes glabrous, the bristles white. 

Mikania Houstoniana (L.) Robinson. Forests and thickets 
of the tierra caliente; region of San Ramon; Guanacaste. Mexico to 
Bolivia. A woody or herbaceous vine, the stems puberulent to pilose; 
leaves broadly ovate, acuminate, rounded at the base, entire, glabrous 
or nearly so, 5-nerved from near the base; heads 4-6 mm. high, race- 
mose, short-pedicellate, white; phyllaries oblong, subacute, sparsely 
puberulent; pappus whitish. 

Mikania hylibates Robinson, Contr. Gray Herb. 77: 52. 1926. 
Wet forest, El Muneco, south of Navarro, Prov. Cartago, 1,400 
meters, Standley 33848. Known only from the original material. A 
large vine, glabrous except in the puberulent inflorescence; leaves 
oblong, long-acuminate, rounded to subacute at the base, entire, 
coriaceous, 3-5-nerved from the base; heads 8 mm. long, sessile, the 
phyllaries oblong, obtuse, puberulent; corollas dirty white; pappus 
dirty white. 

Mikania leiostachya Benth. Thickets and forest of the Atlantic 
tierra caliente; region of San Ramon. Honduras to Ecuador. A 
slender vine, almost glabrous; leaves ovate, acuminate, rounded at 
the base, subcoriaceous, entire; heads white, 7 mm. long, sessile, 
spicate, forming large, lax panicles; phyllaries ovate-oblong, obtuse, 
pubescent or glabrate; corollas white; achenes glabrous, the pap- 
pus white. 



FLORA OF COSTA RICA 1497 

Mikania micrantha HBK. In thickets or forest, often forming 
dense tangles over shrubbery, Meseta Central to the upper part of 
the Pacific tierra caliente; region of San Ramon; Guanacaste. Widely 
distributed in tropical America. Plants chiefly herbaceous, almost 
glabrous or pubescent; leaves thin, ovate or broadly ovate, often 
sagittate or hastate, with a deep, open basal sinus, acuminate, cre- 
nate-undulate or dentate, mostly 3-nerved from the base, usually 
glabrate; heads dirty white, only 4-5.5 mm. long, pedicellate, 
corymbose; phyllaries obovate-oblong, acute or short-acuminate; 
pappus white at first, becoming fulvous. Probably the most common 
species of Central America. It has been reported from Costa Rica 
under the name M. scandens Willd. 

Mikania nubigena Robinson, Contr. Gray Herb. 77: 57. 1926. 
Wet forest, Las Nubes, Prov. San Jose*, 1,500-1,900 meters, Standley 
38509. Material from the Rio Navarro is perhaps referable here. 
Endemic. Plants glabrous except for the puberulent inflorescence; 
leaves ovate-oblong, caudate-acuminate, rounded to subacute at the 
base, remotely cuspidate-denticulate, pinnately 5-nerved, 8-10 cm. 
long; heads white, 7 mm. long, sessile, forming a terminal, compound 
panicle; phyllaries narrowly oblong, rounded at the apex, ciliolate, 
dorsally glabrous; pappus yellowish white. 

Mikania Pittieri Robinson, Proc. Bost. Soc. Nat. Hist. 31: 255. 
1904. Cuesta de Los Borucas, Pittier 10540. Reported by Cufo- 
dontis from Guayabillos, Volcan de Irazu, 2,250 meters. Endemic. 
Plants glabrous except in the puberulent inflorescence; leaves ovate, 
caudate-acuminate, rounded at the base, subcoriaceous, entire, pin- 
nate-nerved; heads sessile, forming a dense, thyrsoid panicle; phyl- 
laries narrowly oblong, somewhat dilated and obtuse at the apex, 
glabrate, 5-6 mm. long; achenes glabrous, 4 mm. long, longer than 
the corolla. 

Mikania riparia Greenm. ex Robinson, Proc. Bost. Soc. Nat. 
Hist. 31: 255. 1904. In thickets, Rio de Las Vueltas, Tucurrique, 
635 meters, Tonduz 13163. Thickets and forest of the tierra caliente, 
ascending to the region of Dota and the mountains south of Cartago; 
El General; region of San Ramon. Endemic. Plants slender, herba- 
ceous, glabrous except in the puberulent inflorescence; leaves thin, 
broadly ovate, usually small, caudate-acuminate, cordate at the base, 
near the base coarsely dentate or undulate and often hastate; heads 
dirty white, short-pedicellate, racemose, forming a large, open, leafy 



1498 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

panicle; phyllaries narrowly oblong, obtuse or subacute, glabrous, 
or puberulent at the apex; achenes glabrous, the pappus pale 
reddish brown. 

Mikania Skutchii Blake, Brittonia 2: 333. 1937. In forest, 
vicinity of El General, Prov. San Jose", 1,130 meters, Skutch 2514- 
Frequent in thickets and forest of the central region; regions of Dota 
and San Ramon; chiefly at 1,000-1,800 meters, common in many 
localities. Endemic. A large or small vine, herbaceous or more or less 
woody, the stems densely fulvous- or rufous-pilose with soft, spreading 
hairs; leaves ovate or lance-ovate, mostly 8-20 cm. long, acuminate, 
rounded or subcordate at the base, densely pilose, remotely denticu- 
late or subentire, pinnate-nerved; heads small, dirty white, sessile 
or nearly so, glomerate, forming a large, thyrsiform panicle; phyl- 
laries oblong, obtuse, 5 cm. long, pilose; achenes sparsely pilose 
toward the apex, the pappus whitish, becoming fulvous. This was 
reported from Costa Rica by Robinson as M. hirsutissima DC., a 
species of Brazil and Paraguay. 

Mikania Standleyi Robinson, Contr. Gray Herb. 77: 59. 1926. 
Wet forest, La Estrella, Prov. Cartago, Standley 39155. Collected 
also at La Palma de San Jos and El General. Endemic. A large, 
more or less woody vine, the stems fulvous-puberulent; leaves ovate, 
acuminate, cordate at the base, denticulate, glabrous above, puberu- 
lent beneath, 3-5-nerved from the base; heads white or greenish 
yellow, pedicellate, forming a large, pyramidal panicle; phyllaries 
6.5 mm. long, puberulent toward the apex, the outer ones subacute, 
the inner rounded at the apex; achenes glabrous, the pappus fulvous. 

Mikania Tonduzii Robinson, Proc. Bost. Soc. Nat. Hist. 31: 
256. 1904. Tucurrique, 635 meters, Tonduz 13274. Forests of 
Guanacaste. Extending to southern Mexico. A large, herbaceous 
vine, glabrous or nearly so except in the puberulent inflorescence; 
leaves ovate, coriaceous, acuminate, rounded-cordate at the base, 
entire, glabrous and lustrous above, puberulent or glabrate beneath; 
heads white, sessile, forming globose glomerules, these disposed in 
large, leafy panicles; phyllaries oblong, 3 mm. long, griseous-tomen- 
tulose, rounded at the apex; achenes glabrous, 2 mm. long, the pappus 
more or less fulvous. 

Mikania vitifolia DC. M. punctata Klatt, Bull. Soc. Bot. Belg. 
31: 195. 1892. El General, 600 meters, Pittier 3434. Frequent in 
thickets and forest of the tierra caliente, ascending to about 1,000 
meters; region of San Ramon; Guanacaste. Southern Mexico to 



FLORA OF COSTA RICA 1499 

Brazil. A large vine, the stems somewhat pilose or almost glabrous; 
leaves ovate to triangular, acuminate, cordate at the base, 5-7- 
nerved from the base, undulate-dentate and sometimes subhastate, 
punctate, rusty-pubescent beneath along the nerves and veins; 
upper leaves triangular-ovate, rounded to truncate at the base; 
heads white, sessile or nearly so, glomerate, forming a compound 
panicle; phyllaries linear-oblong, 5 mm. long, pubescent toward the 
apex; pappus dull white, becoming reddish brown. 

MILLERIA L. 

Tall, rather slender, annual herbs, somewhat villosulous or 
glabrate; leaves opposite, large, thin, petiolate; heads very small, 
yellow, radiate, arranged in open, paniculate cymes; involucre 
oblique-subglobose, somewhat enlarged in fruit and closed above the 
single achene, the bracts few, subherbaceous, one much larger than 
the others, cucullate; receptacle naked; ray flower 1, fertile, the disk 
flowers 4 or fewer, sterile; achene obovoid, compressed, glabrous, 
without pappus. The genus consists of a single species. 

Milleria quinqueflora L. Thickets or waste ground of the 
Pacific tierra caliente; region of San Ramon; Guanacaste. Widely 
distributed in tropical America. Plants often a meter high or more; 
leaves broadly ovate, acuminate, usually long-decurrent upon the 
petiole, the upper leaves often sessile; flower heads in fruit about 
5 mm. broad. The plant is a weedy and unattractive one, common 
in some parts of Central America. 

MONTANOA Cervantes 

Reference: Robinson & Greenm. Proc. Amer. Acad. 34: 508. 1899. 

Shrubs or trees; leaves mostly opposite, dentate to lobate or 
pinnatifid; heads medium-sized, radiate, usually very numerous, 
the rays white, the disk yellow; outer phyllaries 5-7, linear or oblong; 
rays neutral; paleae of the disk accrescent in fruit, usually scarious, 
often spinulose-tipped, greatly exceeding the obovoid, epappose 
achenes. A few other species are native in Central America. 

Montanoa dumicola Klatt, Bull. Soc. Bot. Belg. 31: 200. 1892. 
In hedge, Alajuelita, 1,000 meters, Pittier 1454. Frequent in hedges 
and forest about San Jose"; region of San Ramon. Endemic. A 
shrub or tree, sometimes 9 meters high, with a broad, open crown, 
the bark gray, shredded; leaves slender-petiolate, the petioles not 
appendaged; blades ovate to broadly ovate, usually not lobate, 
obscurely dentate or subentire, sometimes shallowly trilobate, green 



1500 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

and glabrate above, pale beneath and very densely and minutely 
pubescent; heads rather large, the phyllaries 4 mm. long, the rays 
about 10 and 1.5 cm. long; pales of the disk somewhat pubescent, 
spinulose- tipped. 

Montanoa hibiscifolia Benth. Tora. M. Wercklei Berger, 
Gard. Chron. III. 50; 122. 1911 (based upon cultivated plants, 
grown from seeds sent to Italy by Werckle"). In thickets or forest, 
Meseta Central, ascending the slopes of Barba to 2,000 meters or 
more; region of San Ramon; Pejivalle; at 900 meters or more. 
Extending to southern Mexico. A shrub or small tree; leaves large, 
palmately 5-7-lobate, 20-30 cm. wide or smaller, often cordate at 
the base, the lobes acuminate, green and glabrate above, pale 
beneath and densely pubescent; petiole bearing near its apex 2 large, 
leafy auricles; heads numerous, in a large corymb; rays 8-10, about 
1 cm. long; pales short-mucronate, somewhat pubescent at the apex. 
Like other species of this genus, this is a remarkably handsome and 
showy tree or shrub when in flower, and it deserves wide cultivation 
as an ornamental plant. 

Montanoa Pittieri Rob. & Greenm. Proc. Amer. Acad. 34: 517. 
1899. Alajuelita, Pittier 1455. Known, apparently, only from the 
original collection. Leaves green and glabrate above, pale beneath 
and appressed-puberulent, obsoletely trilobate, caudate-acuminate, 
auriculate-appendaged at the base; heads few, the phyllaries 4 mm. 
long, the rays about 8 and 1.5 cm. long; pales pubescent above, 
with a short, incurved point at the apex. 



Montanoa xanthiifolia Schultz Bip. is reported by Klatt from 
El Rodeo, 750 meters, Pittier, but the determination is probably 
incorrect, the species ranging farther north. 

NEUROLAENA R. Br. 

Tall herbs, the leaves alternate, dentate, the lower ones often 
trilobate; heads paniculate, usually discoid, the involucre campan- 
ulate; phyllaries imbricate, 3-4-seriate, the outer ones gradually 
shorter, obtuse, membranous; receptacle plane, paleaceous; achenes 
oblong, glabrous or pubescent; pappus abundant, of slender, per- 
sistent, scabrous bristles in 1-2 series. Two other species have been 
described from Central America. 

Neurolaena lobata (L.) R. Br. Gavilana, Capitana. Eupa- 
torium Valverdeanum Klatt, Bull. Soc. Bot. Belg. 31: 188. 1892 
(Piedra del Convento, between El General and Buenos Aires, Pittier 



FLORA OF COSTA RICA 1501 

3710}. E. chrysocephalum Klatt, Leopoldina Bot. Beibl. 2. 1895 
(also based on Pittier 3710). Common in thickets and waste ground 
of the tierra caliente; region of San Ramon; ascending to 1,050 
meters. Widely distributed in tropical America. Plants stout, 
sometimes 3 meters high, simple or sparsely branched, the stems 
angulate, dense and finely pubescent; leaves mostly sessile, chiefly 
lanceolate, often very large, serrate, the lower ones mostly trilobate, 
rough-pubescent; heads forming large panicles, the involucre 5-6 
mm. high; phyllaries 3-nerved, puberulent; flowers yellow; achenes 
black, 1.5 mm. long. The heads are much like those of the genus 
Eupatorium. The plant is used in domestic medicine in Central 
America. 

OLIGANTHES Cass. 
Reference: Gleason, N. Amer. Fl. 33: 102. 1922. 

Shrubs or small trees; leaves broad, alternate, petiolate, more 
or less tomentose; heads discoid, 1-8-flowered, aggregate in dense, 
corymbiform panicles at the ends of the branches; involucre cylindric 
or narrowly campanulate, the phyllaries few, imbricate in several 
series; receptacle subconvex, naked; achenes striate or costate; 
pappus usually 2-seriate, of linear scales, these twisted toward the 
apex. No other species are known from Central America. 

Oliganthes discolor (HBK.) Schultz Bip. Reported from El 
General and from forests of Buenos Aires. Panama and Colombia. 
A shrub or small tree; leaves elliptic, 10-18 cm. long, acuminate, 
narrowed to the base, entire, glabrous above or nearly so, tomentose 
beneath; heads 2-flowered, 5 mm. high. Pittier lists the TeYraba 
name as Shkur. 

Oliganthes ferruginea Gleason, N. Amer. Fl. 33: 102. 1922. 
Forests of Alto de Mano de Tigre, Diquis Valley, 700 meters, Pittier 
12138. Endemic. A tree of 10-12 meters; leaf blades 10-15 cm. 
long, acuminate, entire, narrowed to the acute base, glabrous above 
or nearly so, almost glabrous beneath; heads 2-flowered, 7 mm. high. 

ONOSERIS DC. 

Herbs or shrubs; leaves mostly basal, or alternate, white-tomen- 
tose beneath; heads large, appearing radiate, solitary or paniculate, 
the phyllaries grading into the bracts of the peduncles, the flowers 
mostly purple or red ; involucre hemispheric or radiate, the phyllaries 
narrow, multiseriate, imbricate, the outer gradually shorter; recep- 
tacle naked or fimbrillose; achenes subterete, 5-costate, glabrous or 



1502 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

pubescent; pappus bristles numerous, rather stiff, in 2 or more 
series. At least one other species is known from Central America. 

Onoseris silvatica Greenm. Proc. Amer. Acad. 49: 514. 1913. 
Hoja quema. Forests of the hills of Nicoya, Tonduz 13597. El 
Rodeo; Tabarcia; region of San Ramon. Probably endemic. Plants 
large and coarse, 1 meter high or more, simple or branched; leaves 
crowded near the base of the stem, large, runcinate-pinnatifid, acute 
or short-acuminate, the terminal segment more or less triangular- 
ovate and much larger than the others, rounded or cordate at the 
base, very coarsely dentate, green and glabrate above, densely 
white-tomentose beneath; heads numerous, purple-red, forming a 
large panicle, 2-2.5 cm. high; phyllaries densely white-tomentose; 
achenes sericeous. There is some doubt as to whether this is distinct 
from 0. onoseroides (HBK.) Robinson. 



The genus Orthopappus almost surely is represented on the 
Pacific slope of Costa Rica by 0. angustifolius (Swartz) Gleason, 
but I have found no evidence of its having been collected. 

OTOPAPPUS Benth. 

Reference: Blake, Journ. Bot. 53: 229-235. 1915. 

Shrubs, often scandent; leaves opposite, short-petiolate, dentate; 
heads radiate or discoid, yellow; involucre campanulate, the phyl- 
laries graduate, 3-6-seriate, indurate, often herbaceous-tipped; 
achenes compressed, 2-winged, the wings decurrent on the awns; 
pappus of 2 awns and a lacerate corona of connate squamellae. 
A few other species grow in Central America. 

Otopappus verbesinoides Benth. In thickets or forest, upper 
part of the Pacific tierra caliente; region of San Ramon; Atlantic 
tierra caliente; Guanacaste. Extending to southern Mexico. A 
shrub of 3 meters or less, with recurved branches, or often scandent, 
the branches strigillose; leaves ovate to lance-oblong, 7-15 cm. long, 
acuminate, obtuse or acute at the base, somewhat serrulate; heads 
radiate, about 1 cm. high, in panicles of 3-5 at the ends of the 
branches; outer phyllaries with reflexed, linear to spatulate, her- 
baceous tips. 

OYEDAEA DC. 

Reference: Blake, Revision of the genus Oyedaea, Contr. U. S. 
Nat. Herb. 20: 411-422. 1921. 

Chiefly shrubs; leaves opposite, entire or serrate; heads radiate, 
yellow, medium-sized; ray flowers neutral; involucre campanulate, 



FLORA OF COSTA RICA 1503 

the phyllaries graduated, 2-5-seriate, subequal or the outer ones 
longer, usually with herbaceous tips, the outer sometimes foliaceous; 
receptacle flat or subconvex, paleaceous; achenes compressed, oblong 
to cuneate-obovate, 2- winged; pappus of 2 slender, persistent awns 
and a corona of 6-12 narrow, fimbriate or lacerate, more or less 
united squamellae. A single species is known from Central America. 

Oyedaea verbesinoides DC. Viguiera acuminata Benth. in 
Oerst. Vid. Medd. Kjoebenhavn 1852: 91. 1853 (Irazu, 2,745 meters, 
Oersted 96, 106). V. macrophylla Benth. in Oerst. loc. cit. (Cerro 
de Aguacate, 610 meters, Oersted 104)- 0. acuminata Benth. & 
Hook, ex Hemsl. Biol. Centr. Amer. Bot. 2: 176. 1881. 0. macro- 
phylla Benth. & Hook, ex Hemsl. loc. cit. V. drymonia Klatt, 
Leopoldina Bot. Beibl. 5. 1895 (Canas Gordas, at 1,100 meters, 
Pittier 7351). Thickets and forest of the central region, descending 
to the Pacific tierra caliente, and ascending high on the slopes of the 
central mountains. Ranging south to Venezuela. A shrub or tree, 
sometimes 6 meters high, the branches pilose or hispid; leaves short- 
petiolate, thick, ovate or ovate-lanceolate, 8-20 cm. long, long- 
acuminate, cuneate at the base, serrate, densely rough-pubescent; 
heads large and showy, the involucre 1-1.5 cm. high; rays 1.5-2 cm. 
long, bright yellow; achenes 3.5-6 mm. long. The tree or shrub 
is abundant in some localities, as at Santa Maria de Dota and on 
the hills near Agua Caliente. It is showy and handsome because of 
the abundance of large, brightly colored flower heads, suggestive 
of sunflowers (Helianthus). 

PEGTIS L. 

Reference: Rydberg, N. Amer. Fl. 34: 194. 1916. 

Low, strong-scented, annual or perennial herbs, usually glabrous 
or nearly so; leaves opposite, narrow, gland-dotted, generally entire, 
commonly with bristles on the margins near the base; heads small, 
solitary or cymose, radiate, yellow; involucre cylindric to turbinate, 
the phyllaries 3-12, uniseriate, gland-dotted; receptacle naked; 
achenes linear, terete or angulate, pubescent or glabrate; pappus of 
few or many squamellae, awns, or bristles. Several other species 
have been collected in Central America. 

Pectis capillipes (Benth.) Hemsl. Reported from Guanacaste 
by Klatt. Nicaragua. Perennial, the stems branched, erect, 7-12 
cm. high; leaves linear, 1.5-2 cm. long; heads solitary in the forks 
of the branches, on slender pedicels 3-5 cm. long; involucre turbinate, 
the 5 phyllaries oblong, obtuse; pappus of the disk flowers of 5-6 



1504 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

bristles somewhat dilated at the base, and of many shorter ones. 
Rydberg in the North American Flora gives the original locality of 
this species as Guatemalan, but in error, for the type was collected 
in Nicaragua. 

Pectis diffusa Hook. & Arn. was reported from the Pacific slope 
by Klatt, but' the species is not known from Central America. 

Pectis elongata HBK. Coronillo. In pastures or savannas, 
regions of San Ramon and Buenos Aires, Pacific tierra caliente. 
Southern Mexico to West Indies and Peru. An erect annual, 60 
cm. high or less, simple or sparsely branched, almost glabrous; leaves 
linear, 2-6 cm. long; heads cymose-paniculate, slender-pedicellate, 
the involucre 5-6 mm. high; phyllaries 5, acuminate, glabrous; 
achenes 2-2.5 mm. long, hirsutulous or glabrate; pappus bristles 5-20. 

Pectis febrifuga van Hall. Reported by Rydberg from Costa 
Rica, the locality not indicated. Extending to Venezuela and West 
Indies. An erect annual, diffusely branched, 20 cm. high or less, 
the stems scabrous-pubescent; leaves linear or lance-linear, 2 cm. 
long or less; heads solitary in the forks of the branches, the pedicels 
1.5-3 cm. long; involucre 4-4.5 mm. long, the 5 phyllaries acuminate 
or apiculate; achenes 2-2.5 mm. long; pappus in the disk flowers of 
5 awned squamellae. 

Pectis grandiflora Klatt, Leopoldina Bot. Beibl. 6. 1895. 
P. falcata Cufodontis, Archivio Bot. 9: 202. 1933 (Golfo Dulce, 
near Puerto Jime'nez, Cufodontis 177; duplicate type in Herb. Field 
Mus.). Type from Hato Viejo. Collected several times in the 
Golfo Dulce region. Panama. Plants coarse, perennial, procumbent, 
the stems 1 meter long or less, glabrous or puberulent, very densely 
leafy; leaves linear, 2-4 cm. long; heads terminal, solitary, on 
peduncles 2-3 cm. long; involucre 1 cm. high, the phyllaries broad, 
obtuse; achenes 6 mm. long; pappus of 12-15 subulate squamellae. 
Differing from other local species in the very large heads. 

Pectis Oerstediana Rydb. Anisillo. Collected at Alajuela, 
and doubtless elsewhere on the Pacific slope. Nicaragua. Plants 
erect, densely branched, 30-60 cm. high, glabrous; leaves linear, 
5 cm. long or less; heads axillary and terminal; involucre 5 mm. 
high, the phyllaries linear, acuminate; achenes 2.5 mm. long, hispidu- 
lous; pappus of 3-5 bristles. 

Pectis polyantha Rydb. Reported from Costa Rica, doubtless 
from the lower Pacific slope. Nicaragua. A diffuse perennial, the 



FLORA OF COSTA RICA 1505 

stems 10-20 cm. long, puberulent; leaves linear, 1-2.5 cm. long; heads 
solitary in the upper leaf axils, the peduncles 4-6 cm. long; involucre 
7 mm. high, the 8-12 phyllaries linear; achenes 3 mm. long, pubes- 
cent; pappus of about 20 hirsutulous bristles. 

Pectis prostrata Cav. Near seashore, San Lucas, Pacific coast, 
J. M. Orozco 230. Widely distributed in tropical America. A much 
branched annual, usually prostrate, short-pilose; leaves oblanceolate- 
linear, 1-3 cm. long, acute, often mucronate, setose-ciliate near the 
base; heads axillary and terminal, the lower solitary, the upper 
glomerate, sessile or nearly so; involucre 5-6 mm. long, the 5 phyl- 
laries oblong, rounded or truncate at the apex; achenes 3-4 mm. long; 
squamellae of the ray flowers 2, narrowly lanceolate, with 1-3 smaller 
setae; squamellae of the disk flowers 5, unequal. 

PIPTOCARPHA R. Br. 

Shrubs, often with long, recurved branches, or subscandent, the 
pubescence chiefly stellate or lepidote; leaves alternate, petiolate, 
penninerved; heads small, discoid, densely clustered in the leaf 
axils, whitish, few-flowered; involucre ovoid or campanulate, the 
phyllaries multiseriate, imbricate, dry, the inner ones usually 
caducous, the outer shorter; receptacle convex, naked; achenes 10- 
costate, truncate; pappus setae more or less biseriate, stiff, the 
outer ones often shorter. No other species are known from Central 
America. 

Piptocarpha chontalensis Baker. Barbudo. P. costaricensis 
Klatt, Bull. Soc. Bot. Belg! 31: 184. 1892 (in forest, Te'rraba, 260 
meters, Pittier 3704)- Forests of the tierra caliente. Ranging to 
British Honduras. A shrub, sometimes 6 meters high and more or 
less scandent, the branches appressed-tomentose or glabrate; leaves 
coriaceous, short-petiolate, ovate to oblong, 10-16 cm. long, acute 
or subobtuse, somewhat oblique at the base and obtuse to sub- 
cordate, lustrous and almost glabrous above, pale beneath and 
densely appressed-tomentose; involucre 4-5 mm. long; heads 6- 
flowered. It is possible that P. costaricensis may be a distinct 
species, but I find no characters for separating it. 

PIQUERIA Cav. 

Reference: Robinson, Revision of the genus Piqueria, Proc. 
Amer. Acad. Sci. 42: 4. 1906. 

Annual or perennial herbs, sometimes shrubs; leaves opposite 
or alternate, petiolate, usually serrate or dentate; heads small, 



1506 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

white, discoid; involucre cylindric to ovoid or campanulate; phyl- 
laries few, subequal, imbricate or almost 1-seriate; receptacle plane, 
naked; achenes prismatic, 5-angulate, rounded at the apex; pappus 
none, or a deciduous, annular disk, rarely of a few short setae. 
No other species have been found in Central America. 

Piqueria trinervia Cav. In forest, Meseta Central, chiefly at 
1,100-1,500 meters. Extending to Mexico, and in Haiti. An erect, 
perennial herb, a meter high or less, branched, the stems puberulent 
in lines; leaves opposite, lanceolate or narrowly ovate, acuminate, 
acute at the base, serrate, glabrate, 3-nerved; heads 4-flowered, 
3 mm. long, cymose-corymbose; phyllaries rounded at the apex. 
Costa Rican material has been referred incorrectly to P. pilosa 
HBK. and P. trinervia var. pilosa Kuntze. P. trinervia var. luxurians 
Kuntze (Rev. Gen. 1: 355. 1891; P. luxurians Robinson ex Volkens, 
Verh. Bot. Ver. Brandenb. 65: 118. 1923; type from Irazu, Kuntze) is a 
form in which the leaves are somewhat larger, 5-nerved, and rounded 
at the base. It has been collected several times on Volcan de Irazu. 



Pluchea odorata (L.) Cass. must surely occur in Costa Rica, 
but I find neither specimens nor records of its occurrence. 

PODACHAENIUM Benth. 

Tall shrubs or small trees; leaves large, opposite, petiolate, 
angulate-lobate; heads small, radiate, forming large, corymbose 
panicles, the disk yellow, the rays white; involucre small, the phyl- 
laries 2-3-seriate, narrow, herbaceous, the outer ones shorter; 
receptacle convex, the whole head becoming globose in fruit; achenes 
compressed, pubescent, the margin ciliate or cartilaginous-winged, 
the base contracted into a 2-winged stipe; pappus of 2 dilated awns 
at each angle, with 2 small squamellae on each side. The genus 
consists of a single species. 

Podachaenium eminens (Lag.) Schultz Bip. Tora, Tora 
blanca. P. paniculatum Benth. in Oerst. Vid. Medd. Kjoebenhavn 
1852: 99. 1853 (Volcan de Irazu, 2,400 meters, Oersted; photo, of 
type in Herb. Field Mus.). Frequent in hedges, thickets, or forest 
of the central region; Dota; descending into the upper part of the 
Atlantic tierra caliente; at 600-2,400 meters. Ranging to Mexico. 
A shrub or tree of 6 meters or less, sometimes wholly herbaceous; 
leaves slender-petiolate, ovate to orbicular, often 30 cm. long, 
sometimes entire but usually lobate or coarsely dentate, green and 



FLORA OF COSTA RICA 1507 

somewhat rough above, rather densely sordid-tomentose beneath; 
heads about 1 cm. broad; achenes 2 mm. long. 

POLYMNIA L. 

Coarse, perennial herbs, or shrubs, glabrous or variously pubes- 
cent, often viscid; leaves chiefly opposite, broad, entire to angulate 
or lobate, the petioles often auriculate-appendaged; heads usually 
large, corymbose-paniculate, yellow, radiate, the ray flowers fertile, 
the disk flowers sterile; involucre hemispheric or broader, the 
phyllaries biseriate, membranaceous or herbaceous, the outer ones 
sometimes large and green; receptacle plane, paleaceous; achenes 
glabrous, thick, obovoid; pappus none. A few other species have 
been recorded from Central America. 

Polymnia latisquama Blake, Journ. Wash. Acad. Sci. 16: 421. 
1926. Along stream, southern slope of Volcan de Turrialba, near 
the Finca Volcan de Turrialba, 2,000-2,400 meters, Standley 35340. 
Collected also on Irazu, at 2,850 meters. Endemic. A coarse herb, 
1.5-3 meters high, the stems glabrous or nearly so; leaves ovate or 
triangular-ovate, 10-20 cm. long, coarsely serrate, almost glabrous; 
heads solitary, long-pedunculate, about 6 cm. wide, the rays 3 cm. 
long; outer phyllaries 4, rounded-ovate, 18 mm. wide, glabrous; 
achenes 6 mm. long. 

Polymnia maculata Cav. Tora, Purca. Frequent in thickets 
or forest of the Meseta Central, chiefly at 1,000-1,300 meters. 
Ranging to Mexico. A coarse, rough-pubescent herb, 1-1.5 meters 
high, usually viscid; petioles broadly dilated at the base and clasping; 
leaf blades deltoid to ovate-rounded, usually lobate and coarsely 
dentate, very rough; outer phyllaries lanceolate, villous or hirsute; 
heads, excluding the rays, about 1.5 cm. broad. Blake (Contr. 
Gray Herb. 52: 33. 1917) divides the Costa Rican material into 
two varieties: var. vulgaris Blake with hispid-pilose, sparsely glan- 
dular branches, the leaves green beneath, densely or sparsely hispid- 
pilosulous and hispid-pilose; and var. adenotricha Blake, with the 
branches densely stipitate-glandular and sparsely hispid-pilose, the 
leaves glandular-roughened and rather sparsely pubescent beneath. 
Polymnia sonchifolia Poepp. & Endl. as reported from Costa Rica 
by Klatt is P. maculata var. vulgaris. 

POROPHYLLUM Adans. 
Reference: Rydberg, N. Amer. Fl. 34: 181. 1916. 
Annual or perennial herbs, rarely shrubs, usually glabrous; 
leaves alternate or opposite, petiolate, broad or narrow, entire, 



1508 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

with pellucid glands, at least along the margins; heads small or 
medium-sized, discoid, mostly yellowish or purplish; involucre 
cylindric or campanulate; phyllaries 5-9, linear to obovate, 1-seriate, 
commonly with 2 rows of dorsal glands; receptacle naked; achenes 
slender, striate, usually hispidulous; pappus of numerous scabrous 
or hispidulous bristles. One or two other species are found in 
Central America. 

Porophyllum punctatum (Mill.) Blake. Thickets of the 
plains of Guanacaste. Extending to Mexico. Plants much branched, 
perhaps sometimes suffrutescent; leaves oval or elliptic, 1.5-3.5 cm. 
long; involucre 1-1.5 cm. high; corollas cream-colored; achenes 
6 mm. long. 

Porophyllum ruderale (Jacq.) Cass. P. ellipticum Cass. 
Occasional in thickets or fields of the tierra caliente. Widely dis- 
tributed in tropical America. An erect, sparsely branched annual; 
leaves elliptic, 1-6 cm. long, obtuse; involucre 2 cm. high; achenes 
8 mm. long. 

PSEUDELEPHANTOPUS Rohr 

Erect, branched, perennial herbs with leafy stems; leaves alter- 
nate, entire or crenate; heads small, 4-flowered, white or lavender, 
discoid, glomerate and disposed in panicled spikes, each glomerule 
subtended by 2-3 bracts; involucre of 4 pairs of phyllaries; achenes 
compressed, 10-striate; pappus a single series of 5-15 setae, the 
2 lateral setae longer than the others and contorted at the tip. 
A single species is found in North America. 

Pseudelephantopus spicatus (Juss.) Rohr. Escobilla, Escoba 
real, Lechuguilla. Elephantopus spicatus Juss.; Distreptus spicatus 
Cass. Elephantopus spicatus var. roseus Klatt, Bull. Soc. Bot. Belg. 35 : 
279. 1896 (pastures, San Jose", Pittier 7237}. A common weed in waste 
or cultivated ground, often in pastures, Meseta Central to the 
coasts. Generally distributed in tropical America. Plants usually 
a meter high or less, the branches pilose with long, appressed, 
whitish hairs; leaves spatulate to obovate, 6-15 cm. long, the upper 
ones almost linear and much smaller; involucres 8-12 mm. long, 
the phyllaries lanceolate, aristate-acuminate, with whitish margins. 
One of the most common weeds found in Central America. Pittier 
reports the TeYraba name as Tshicordias, evidently a corruption of 
Achicoria. 

RENSONIA Blake 

Shrubs, erect or ascending, sometimes more or less scandent, 
rarely epiphytic; leaves opposite, ovate, slender-petiolate, serrate, 



FLORA OF COSTA RICA 1509 

scabrous; heads small, yellow, radiate, in terminal, cymose panicles; 
phyllaries 8-9, oblong-obovate, indurate below, thick-herbaceous 
above; receptacle flat, paleaceous; achenes of the fertile flowers 
obovate, obcompressed, epappose, 2-winged, the wings prolonged 
above the achene into 2 triangular, lacerate teeth. The genus 
consists of a single species. 

Rensonia salvadorica Blake. In thickets or forest, mountains 
of Guanacaste, 600-700 meters. Salvador. A slender shrub of 
1-5 meters, the branches strigillose; leaf blades 10-25 cm. long, 
acuminate, acutely cuneate at the base; heads 6-7 mm. high, the 
phyllaries acute, 3-5-nerved; achenes 5.5 mm. long, blackish, his- 
pidulous on the outer surface. Illustrated, Journ. Wash. Acad. 
Sci. 17: 63. 1927. 

ROLANDRA Rottb. 

Erect, perennial herbs, branched; leaves alternate, entire, short- 
petiolate, closely and densely white- tomentose beneath; heads dis- 
coid, 1-flowered, small, forming axillary and terminal, globose 
clusters; involucre of 2 aristate phyllaries, the outer one larger; 
achenes angulate, truncate; pappus a lacerate crown. The genus 
consists of a single species. 

Rolandra fruticosa (L.) Kuntze. Thickets of the Atlantic 
tierra caliente. Widely distributed in tropical America, but in 
Central America known definitely only from Costa Rica and Panama. 
Plants a meter high or less, the branches sparsely or densely strigose; 
leaves oblong-lanceolate to elliptic, 5-10 cm. long, acute at each 
end, green above and rough; glomerules of heads about 1.5 cm. 
in diameter, green, the phyllary tips uncinate; corolla white. 

RUMFORDIA DC. 

Reference: Robinson, A revision of the genus Rumfordia, Proc. 
Amer. Acad. 44: 592. 1909. 

Shrubs or tall herbs, the stems branched, often fistulose; leaves 
opposite, chiefly ovate or lanceolate, often connate at the base and 
perfoliate, serrate or denticulate; heads of medium or large size, 
paniculate, the rays yellowish or in age white; ray flowers pistillate 
and fertile, the disk flowers perfect and fertile; achenes obovoid, 
somewhat compressed, glabrous, without pappus; involucre biseriate, 
the outer phyllaries "herbaceous, ovate or elliptic, spreading, the 
inner ones much smaller, ovate or lanceolate, paleiform, enclosing 
the outer achenes. A few other species are known in northern 
Central America. 



1510 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Rumfordia aragonensis Greenm. Proc. Amer. Acad. 40: 38. 
1904. Thickets near Aragon, Turrialba, 630 meters, Pittier 13246. 
Known only from the original collection. Stems fistulose; leaves 
rhombic-ovate, about 12 cm. long, denticulate, glabrate above, 
sparsely pubescent beneath, 1-lobate or angulate near the base on 
each side, the petioles winged ; outer phyllaries 5-6, ovate, acuminate, 
1.5 cm. long, inconspicuously puberulent, the inner ones ovate, 
acuminate, dorsally glandular-hispidulous, 5 mm. long; rays 5 mm. 
long; achenes 2 mm. long. 

Rumfordia polymnioides Greenm. Proc. Amer. Acad. 39: 
99. 1903. In maize fields, Copey, Prov. San Jose", 1,800 meters, 
Tonduz 11947. Collected also at Santa Maria de Dota in the same 
region. A coarse herb 1-1.5 meters high, the stems whitish-pubes- 
cent; leaves broadly ovate, acuminate, 3-nerved, green above and 
short-pubescent, paler beneath and griseous-tomentulose, narrowed 
to the winged petiole; outer phyllaries 5, broadly ovate, acute, 
griseous-hirsute, the inner ones linear, hispidulous. 

SALMEA DC. 

Reference: Blake, Journ. Bot. 53: 196. 1915. 

Shrubs or woody vines, the stems terete or nearly so; leaves 
opposite, petiolate, more or less coriaceous and dentate; heads 
small, dirty white, discoid, in small, terminal and axillary corymbs; 
involucre campanulate or turbinate, the phyllaries appressed, imbri- 
cate in few series; receptacle conic, paleaceous; achenes laterally 
compressed, marginate or narrowly winged; pappus of 2 awns. 
No other species are known from Central America. 

Salmea scandens (L.) DC. In thickets, region of San Jos4, 
descending to the Pacific coast. Widely distributed in tropical 
America. A scandent shrub, sometimes 10 meters long; leaves 
ovate to oblong-ovate, 5-12 cm. long, remotely serrate or almost 
entire, acute to acuminate, lustrous, almost glabrous; heads 4.5-7 
mm. high, the phyllaries ovate or ovate-lanceolate, acute or acumi- 
nate. This shrub is rather well known in Salvador, where it is 
called "duerme-boca." It is said that if the root is chewed, it 
deadens sensation in the tongue. In Puerto Rico the plant is called 
"bejuco de muela," presumably because it is chewed as a remedy 
for toothache. In addition, it receives in Salvador the name "salta- 
afuera," an allusion to its use as a barbasco or fish poison, as which 
it is said to be unusually efficient. These properties well deserve 



FLORA OF COSTA RICA 1511 

scientific investigation, since they might possibly have some eco- 
nomic application. 

SCHISTOCARPHA Less. 
Reference: Rydberg, N. Amer. Fl. 34: 303. 1927. 

Coarse, annual or perennial herbs; leaves opposite, petiolate, 
broad, thin, dentate, usually decurrent into the petiole, the petioles 
generally connate-amplexicaul at the base; heads radiate, rather 
small, cymose-paniculate; involucre campanulate, the phyllaries 
3-4-seriate, imbricate, the outer shorter, striate, membranaceous; 
receptacle convex, paleaceous; ray flowers pistillate, the disk flowers 
perfect; achenes oblong, glabrous, striate or smooth, black, lustrous; 
pappus bristles about 20, uniseriate, caducous. Two or three other 
species are recorded for Central America. In general appearance 
the plants resemble the genus Eupatorium, which is not at all closely 
related, but they are distinguished easily enough in the growing 
state by their yellow heads. 

Schistocarpha bicolor Less, is reported by Cufodontis from the 
Atlantic coast, on the basis of too mature specimens. Probably 
the record is referable rather to S. oppositifolia. 

Schistocarpha oppositifolia (Kuntze) Rydb. S. Hoffmannii 
Kuntze. Frequent in thickets or cutover land of the Atlantic tierra 
caliente, ascending sometimes in the central mountains to 2,400 
meters; Atlantic slope of the mountains of Guanacaste. Mexico to 
Bolivia. A coarse herb, 1-1.5 meters high, branched, short-hirsute; 
leaves long-petiolate, broadly ovate, 10-20 cm. long, acuminate, 
abruptly contracted and cuneate-decurrent at the base, dentate, 
sparsely hirsutulous; heads 6-9 mm. high, in small or large panicles; 
phyllaries obtuse or subacute, ciliolate; rays none or minute and 
inconspicuous; achenes 1.5 mm. long, glabrous. A weedy plant, 
abundant in many parts of the Central American tierra caliente. 

Schistocarpha paniculata Klatt, Bull. Soc. Bot. Belg. 31: 
210. 1892. Southeastern slope of Volcan de Irazu, Pittier 866. 
Occasional in thickets or forest of the upper part of the Atlantic 
tierra caliente, and ascending the slopes of the central mountains 
to about 2,800 meters; El General. Ranging to Colombia. A 
shrub of 2-4.5 meters, the branches hirsute and viscid; leaves ovate 
or ovate-deltoid, acuminate, dentate, pilose; involucre 7 mm. high 
or more, the phyllaries hirsute; rays conspicuous, 6-8 mm. long; 
achenes glabrous. 



1512 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

SCLEROCARPUS Jacq. 

Branched, more or less pubescent herbs; leaves chiefly alternate, 
the lowest sometimes opposite; heads rather large, yellow, radiate, 
mostly solitary and terminal or opposite the leaves; involucre 
campanulate, the phyllaries few, about 2-seriate, usually herbaceous; 
receptacle convex or conic, paleaceous; ray flowers neutral, the 
disk flowers perfect; achenes smooth, deciduous with the enveloping 
palea, the pappus none or a short crown. One or two other species 
may occur in Central America. 

Sclerocarpus divaricatus (Benth.) Benth. & Hook. Fields 
and waste ground about San Jose', descending into the upper part 
of the Pacific tierra caliente; region of San Ramon. Mexico and 
Central America. Plants rather slender, erect to prostrate, rarely 
a meter high, rough-pubescent, branched; leaves slender-petiolate, 
mostly 6 cm. long or less, rhombic-ovate or deltoid-ovate, acuminate, 
abruptly decurrent at the base, coarsely serrate; phyllaries green, 
the outer ones foliaceous, about 7 mm. long; disk corollas 1 cm. 
long or more, very slender; rays broad and conspicuous. The proper 
specific name for this plant is doubtful, to the writer at least, and 
the whole genus seems to be in a chaotic condition. So far as I 
know, there is available no treatment that brings order out of the 
apparent chaos exhibited by the usual herbarium material. 

SENECIO L. 

Herbs, shrubs, or small trees, sometimes epiphytic; leaves 
alternate, very variable in form; heads mostly yellow, sometimes 
whitish or orange, radiate or discoid; involucre cylindric to cam- 
panulate, the principal phyllaries narrow and uniseriate, usually with 
a few much smaller ones at their base; ray flowers, when present, 
fertile; achenes sub terete, usually costate, glabrous or pubescent; 
pappus of numerous, soft, white bristles. The largest genus of the 
Compositae, with other species in Central America, although, in 
general, the group is rather sparsely represented in this region. 
It is usually regarded as the largest of all genera of the phanerogams. 
For assistance in preparing the account of this genus the writer is 
greatly indebted to Dr. J. M. Greenman, who has suggested cor- 
rections in the manuscript, and has supplied descriptions of three 
new species. 

Senecio andicola Turcz. Paramos of Cerro de Las Vueltas 
and Cerro de La Muerte, about 3,000 meters. Colombia. A densely 
branched shrub of 1-4.5 meters, the young branches lanate-tomen- 



FLORA OF COSTA RICA 1513 

tose, soon glabrate; leaves almost sessile, oblong or narrowly elliptic- 
oblong, 3-4.5 cm. long, acute or obtuse, acute at the base, entire, 
glabrous above and lustrous, ochraceous-tomentose beneath, coria- 
ceous; heads few, discoid, corymbose at the apices of the branches; 
involucre campanulate, 8 mm. high; principal phyllaries 8, oblong- 
ovate, broadly marginate, sparsely tomentulose or glabrate; achenes 
glabrous. Collections cited by Klatt under the name of S. ledifolius 
DC., a South American species, are referable here, according to 
information supplied by Dr. Greenman. 

Senecio Brenesii Greenm. & Standl., sp. nov. Frutex epiphy- 
ticus scandens vel dependens, ramis stramineis vel ochraceis saltern 
in sicco angulatis, novellis sat dense lanato-tomentosis, internodiis 
foliis duplo vel triple brevioribus; folia inter minora petiolata sub- 
rigide coriacea, petiolo crasso 1-1.5 cm. longo tomentoso vel glabrato; 
lamina elliptica vel ovato-elliptica 6-8 cm. longa 2.5-4 cm. lata 
obtusa vel breviter acutata, basi obtusa vel subacuta, margine vulgo 
plus minusve revoluto, supra glabra vel tan turn ad costam impressam 
sparse tomentosa, evenosa, subtus ubique tomento laxo brunnescente 
induta, costa saliente, nervis venisque occultis; capitula discoidea 
pauciflora thyrsoideo-paniculata, paniculis terminalibus atque axilla- 
ribus usque 10 cm. longis, basi interdum foliis paucis reductis bractea- 
tis, capitulis breviter pedicellatis vel subsessilibus dense aggregatis; 
involucrum ca. 8 mm. longum, phyllariis principalibus vulgo 8 dorso 
laxe tomentosis vel glabratis obtusis linearibus apice subinflexis; 
achaenia glabra. In forest, La Palma de San Ramon, 1,100 meters, 
January, 1927, Brenes 5342 (type in Herb. Field Mus.); February, 
1927, Brenes 5406. Among Costa Rican species this is easily recog- 
nized by the dense tomentum of the lower surface of the entire leaves. 

Senecio calyculatus Greenm. in Donn. Smith, Bot. Gaz. 37: 
419. 1904. Volcan de Turrialba, 2,700 meters, Pittier 7503 (13242}. 
Upper slopes of Turrialba and Irazu, ascending to at least 2,700 
meters. Endemic. An epiphytic shrub, the branches pendent, 
about a meter long, tomentose at first, soon glabrate; leaves slender- 
petiolate, elliptic-lanceolate to elliptic-obovate, mostly 3-6 cm. long, 
obtuse or acute, acute at the base, coriaceous, entire; inflorescence 
covered with a dense, white tomentum; heads discoid, yellow, 8-9 
mm. high, very numerous and densely crowded, sessile; phyllaries 
8; achenes glabrous. This species is easy of recognition because 
of the combination of entire, glabrous leaves and densely white- 
tomentose inflorescence. 



1514 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Senecio candelariae Benth. in Oerst. Vid. Medd. Kjoebenhavn 
1852: 108. 1853. Vara blanca. Candelaria, at 1,800 meters, Oersted. 
Frequent in forest of the central region, ascending on Poas to 2,600 
meters, but chiefly at 1,000-2,000 meters; region of San Ramon. 
Endemic. An epiphytic shrub, usually about a meter long, almost 
glabrous except in the inflorescence and there sordid-puberulent or 
sparsely tomentulose; leaves petiolate, lanceolate or lance-oblong 
to elliptic, mostly less than 10 cm. long, acuminate, acute or obtuse 
at the base, subcoriaceous, entire, the lateral nerves inconspicuous 
or obsolete; heads discoid, whitish, on short or elongate pedicels, 
forming a compact or rather lax, corymbiform inflorescence; involucre 
4-6 mm. high, the phyllaries usually 8, glabrate; achenes glabrous. 
One sheet referred here for the present has been indicated by Green- 
man as a new species, still unpublished. The rather numerous sheets 
at hand exhibit considerable variation, and it is possible that they 
represent as many as three species, although I am inclined to believe 
that all are forms of a single variable entity. I have not seen type 
material of S. candelariae, and am uncertain as to which of the varia- 
tions is nearest the type. 

Senecio Cineraria DC. Sometimes grown for ornament in 
gardens of the central region. Native of the Old World. A low per- 
ennial, very densely white-tomentose throughout; leaves deeply 
pinnatifid, the numerous divisions narrow, obtuse; heads yellow, 
radiate, in small, dense cymes. The plant is called "dusty miller" 
in the United States. 

Senecio Cooperi Greenm. Field Mus. Bot. 2: 284. 1907. Car- 
tago, 1,310 meters, Cooper 5803. Slopes of Irazu and Poas, ascending 
to at least 1,800 meters; El General. Also in Chiriqui, Panama. 
A rather succulent herb, or sometimes suffrutescent, as much as 4 
meters high, simple or branched, the stems and inflorescence abun- 
dantly brownish-hirtellous, densely leafy; leaves very large, oblong- 
ovate, 50 cm. long or less, obtuse to acuminate, often lyrate-pinna- 
tifid toward the base, irregularly dentate, puberulent or sparsely 
tomentulose on both surfaces or glabrate, petiolate, the petiole naked 
or winged to the base; heads very numerous, forming a large, corym- 
biform panicle; heads radiate, yellow, 1.5 cm. high; phyllaries 8, 
linear, glabrous except at the apex; achenes pubescent. 

Senecio copeyensis Greenm. Field Mus. Bot. 2: 285. 1907. 
Forests of Copey, 1,800 meters, Tonduz 11663. Collected also at 
Agua Caliente, 1,600 meters, and probably also on Irazu and Poas. 



FLORA OF COSTA RICA 1515 

Endemic. A shrub or small tree, about 4.5 meters high, sparsely 
branched, the branches stout, tomentose; leaves large, on naked 
petioles, 20-30 cm. long or more, oblong-ovate or ovate, pinnate- 
lobate, acute at the base, tomentulose beneath at first but soon 
glabrate, the lobes broad or narrow, acute; heads yellow, very numer- 
ous, forming dense and broad, tomentose, corymbose panicles; 
phyllaries usually 8, only 4-5 mm. long; achenes glabrous. Costa 
Rican material that has been referred to S. arbor escens Steetz, a 
species of Mexico and northern Central America, is presumably 
referable here. 

Senecio Durandii Klatt, Bull. Soc. Bot. Belg. 31: 211. 1892. 
Rio de Las Vueltas, 2,100 meters, Pittier 220. Forests of Rancho 
Flores; slopes of Volcan de Barba; Cerro Gallito; at 2,000-2,400 
meters. Endemic. A slender, glabrous herb, or often somewhat 
woody below, erect or decumbent, 30-60 cm. high, the stems dark 
purple-red; leaves dark green, linear-lanceolate, herbaceous, 10 cm. 
long or less, 1-1.5 cm. wide, narrowly long-attenuate, attenuate to 
the slender petiole, sharply and conspicuously serrate; heads few, 
terminal, subcorymbose or cymose, long-pedicellate, the pedicels with 
numerous linear bracts; involucre 1 cm. long, the linear phyllaries 
about 13, scarious-marginate; rays large, orange-yellow. I have 
no doubt that I have determined correctly the material placed here, 
although Klatt describes the achenes as pilose, while in the collec- 
tions now referred here they are glabrous. The species is a most 
distinct one, altogether unlike any other with which I am familiar. 
Although referred by Greenman to his section Streptothamni, it cer- 
tainly has little if anything in common with other species grouped 
in that section. Where I have seen the plant, which appears to be 
rare, it grew on rocks at the edge of streams in deep, dark forests, 
sometimes in the spray of waterfalls. The crushed leaves have the 
peculiar odor of Eryngium foetidum. 

Senecio firmipes Greenm. Proc. Amer. Acad. 39: 119. 1903. 
Cerro de La Muerte, 3,100 meters, Pittier 10472. Also on Cerro de 
Las Vueltas, at about the same elevation. Endemic; growing in wet 
thickets at the edge of the paramos. A dense, heathlike shrub, 
2-4.5 meters high, the branches glabrous; leaves lanceolate, 2-3 cm. 
long, 4-6 mm. wide, acute, somewhat pungent- tipped, slightly 
hirtellous on the margins, narrowed to the base, glabrous, somewhat 
glaucous beneath; heads discoid, 5-6 mm. high, forming terminal, 
corymbiform cymes; phyllaries 8, oblong-ovate, glabrous, thick, 
obtuse; achenes glabrous. 



1516 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Senecio heterogamus (Benth.) Hemsl. Biol. Centr. Amer. Bot. 
2: 242. 1881. Cacalia heterogama Benth. in Oerst. Vid. Medd. 
Kjoebenhavn 1852: 107. 1853. Southern slope of Volcan de Irazu, 
2,400 meters, Oersted 8858 (photo, of type in Herb. Field Mus.). 
In forest, slopes of Irazu and Cerro de Las Vueltas, 2,400-3,000 
meters. Chiriqui, Panama; a variety is described from Guatemala. 
Plants herbaceous or suffrutescent, 1-1.5 meters high, rather copi- 
ously scabrous-pubescent; leaves long-petiolate, peltate, ovate- 
rounded, large, angulate-lobate; heads discoid, pale yellow, the 
involucre 8 mm. long; phyllaries about 12; achenes glabrous. This 
differs from all other Costa Rican species in its peltate leaves, 
but other species with peltate leaves are found in other parts of 
Central America. 

Senecio Hoffmannii Klatt, Leopoldina 25: 106. 1889. San 
Rafael. S. Berlandieri Auct. San Jose", Hoffmann 497. Common in 
thickets of the Meseta Central, descending to the coasts, at 1,500 
meters or less; region of San Ramon; Guanacaste. Panama. A 
rather coarse, terrestrial vine, herbaceous or suffrutescent, with 
sparse, rough pubescence; leaves petiolate, ovate to lance-oblong, 
acuminate, obtuse to subcordate at the base, coarsely or finely den- 
tate, somewhat succulent; heads few, large, long-pedunculate, subcy- 
mose at the ends of the branches; involucre about 1 cm. long, the 
linear phyllaries about 20, puberulent and somewhat tomentulose; 
flowers all orange or sometimes dark red, the rays numerous and con- 
spicuous; achenes puberulent. The plant is a rather handsome and 
showy one, plentiful in many places about San Jose", often growing 
over hedges. It is easily recognized by its scandent habit, other 
scandent Senecios of Costa Rica being small, epiphytic vines. The 
flowers are much visited by bees, and the vine is said to be a prolific 
source of honey. 

Senecio megaphyllus Greenm. Field Mus. Bot. 2: 284. 1907. 
S. multivenius var. oliganthus Greenm. loc. cit. in syn. "Bords du 
Paraita Grande," Copey, 1,800 meters, Tonduz 11700. Wet forest, 
La Hondura de San Jose", and slopes of Volcan de Barba, 1,200-2,000 
meters. Endemic. Plants herbaceous or somewhat frutescent, 
1.5-3 meters high, simple or branched, the stout stems arachnoid- 
tomentose above; leaves large, oblong-obovate, often 50 cm. long, 
acute or acuminate, coarsely dentate, narrowed below to the petiole 
and decurrent upon it to its amplexicaul base, somewhat arachnoid- 
tomentose or soon glabrate; heads numerous, bright yellow, radiate, 
forming a dense, corymbose panicle; phyllaries 8, linear, 1 cm. long, 



FLORA OF COSTA RICA 1517 

glabrous; achenes glabrous. A showy and handsome plant, like other 
related species. 

Senecio minis Klatt, Bull. Soc. Bot. Belg. 31: 213. 1892. La 
Division, Valle de El General, 2,160 meters, Pittier 3405. Appar- 
ently known only from the original locality. Plants herbaceous, the 
stem purple, glabrous, winged; lower leaves petiolate, large, cordate, 
serrate, violaceous beneath, pilose on the costa, pinnate-lobate below, 
the petiole dilated at the base, the upper leaves entire, adnate- 
decurrent; heads terminal, radiate, long-pedicellate, laxly corymbose, 
the pedicels bearing numerous filiform bracts; phyllaries 21, lanceo- 
late, barbate at the apex; achenes glabrous. I have seen no repre- 
sentation of this species. 

Senecio multivenius Benth. in Oerst. Vid. Medd. Kjoebenhavn 
1852: 109. 1853. Quiebrahacha (Copey), Tabaquillo. Southern slope 
of Volcan de Irazu, 2,400 meters, Oersted. Frequent in forest or in 
clearings, slopes of the mountains of the central region, and in the 
region of Dota, 2,000-3,000 meters. Endemic. A simple or branched 
herb or shrub, 1-4.5 meters high, almost glabrous or when young laxly 
and sparsely tomentulose; leaves ovate or lanceolate, often 40 cm. 
long, coarsely dentate and usually more or less pinnate-lobate but 
shallowly so; petioles long and slender, naked; heads very numerous, 
bright yellow, radiate, forming a large, dense, corymbose panicle; 
involucre 6 mm. long, the 8 phyllaries linear; achenes glabrous. 

Senecio Oerstedianus Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 109. 1853. Papelillo. Crater of Reventado, Volcan de 
Irazu, 2,700 meters, Oersted. Forests and pastures of the upper 
slopes of Irazu and Turrialba, 2,700-3,450 meters; Cerro de La 
Muerte. Endemic. A stout herb, 1-2 meters high, densely white- 
tomentose almost throughout, glabrate only on the upper leaf sur- 
face; lower leaves petiolate, ovate, the upper ones sessile and amplexi- 
caul, large, rounded to acuminate, closely and rather coarsely 
dentate, the lowest ones usually cordate at the base; heads numerous, 
bright yellow, radiate, forming a large, dense, corymbose panicle; 
involucre 7 mm. long, the phyllaries about 20, linear, densely white- 
lanate; achenes glabrous. 

Senecio phanerandrus Cufodontis, Archivio Bot. 9: 203. 1933. 
Volcan Poas, 2,400 meters, Cufodontis 544- Upper slopes of Poas, 
2,400-2,600 meters. Endemic. A scandent shrub, almost glabrous; 
leaves petiolate, subcoriaceous, ovate-elliptic, 5 cm. long or less, 
obtuse or acute, truncate to acute at the base, obtusely crenate- 



1518 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

dentate; heads discoid, small, very numerous, forming a somewhat 
leafy, rather thyrsiform panicle; phyllaries 8, linear, 5 mm. long, 
glabrous except at the apex; achenes glabrous. A very distinct 
species, apparently local in distribution. A portion of the type 
material has been forwarded through the courtesy of the author of 
the species. To this are referred for the present Standley & Valeria 
50544 from Cerros de Zurqui and Manuel Valeria 1098 from Cerro 
Gallito. In inflorescence characters they seem to be exactly like S. 
phanerandrus, and on the stems there are traces of the curious 
tuberculate-scabrous indument that characterizes S. phanerandrus. 
However, their leaves are narrower and less evidently dentate. It 
may be that they represent a distinct species, but this can be decided 
only when more ample material is available for comparison. 

Senecio schizotrichus Greenm., sp. nov. Frutex scandens 
ubique cum pilis albo-ramosis obtectis; caulibus ramisque subangu- 
latis, striatis; foliis alternis, petiolatis, elliptico-lanceolatis, 3.5-8.5 
cm. longis, 1.5-4.5 cm. latis, acutis, integris, basi obtusis, supra 
viridibus, subtus pallidioribus et prominenter venosis, utrinque 
pubescentibus; petiolis 10-12 mm. longis; inflorescentiis terminalibus 
paniculatis, inflorescentiae bracteis plus minusve foliaceis; capitulis 
numerosis, homogamis; involucris anguste campanulatis, brevius 
calyculatis, paulo pubescentibus; involucri squamis 8, lineari-lanceo- 
latis, 7-8 mm. longis; floribus tubularibus vel tubuloso-campanulatis, 
flavis, exterioribus paululo minoribus; achaeniis glabris. Vicinity 
of El General, Province of San Jose", alt. 975 meters, Jan., 1936, 
Alexander F. Skutch 2502 (type in Mo. Bot. Gard. herb.). This 
species is well marked and easily distinguished from all known species 
of the genus by the white, branched hairs on all vegetative parts of 
the plant. 

Senecio streptothamnus Greenm., sp. nov. Frutex scandens; 
caule tereti in sicco cortice cinereo-brunneo tecto; foliis alternis, 
petiolatis, elliptico-lanceolatis, acutis vel breve acuminatis acutisque, 
integris, utrinque glabris subtus paululo pallidioribus; inflorescentiis 
terminalibus, paniculato-cymosis; capitulis heterogamis, ca. 8-10 mm. 
altis; involucris campanulatis, paulo calyculatis; involucri bracteis 
plerumque 8, lineari-lanceolatis, acutis, 5-6 mm. longis, glabris; 
floribus femineis 3-5, ligulis anguste lineari-oblongis; floribus disci 
8-10; achaeniis glabris vel in angulis minute hirtellis. Forests 
of Las Vueltas, Tucurrique, alt. 635-700 meters, March, 1889, 
Adolfo Tonduz 13275 (Gray Herb., type). 



FLORA OF COSTA RICA 1519 

Senecio Tonduzii Greenm. Bot. Jahrb. 32: 22. 1901, nomen. 
Frutex scandens; caulibus lignosis vel juventate aliquanto carnosis, 
glabris; foliis alternis, brevipetiolatis, ellipticis vel ovatis, 6-14 cm. 
longis, 3.5-6 cm. latis ad apicem acutis vel subrotundatis, integris 
vel leviter sinuato-denticulatis, utrinque glabris, inconspicue venosis, 
basi in petiolum usque ad 1 cm. longum angustatis; foliis superioribus 
fere sessilibus; inflorescentiis paniculatis, inflorescentiae bracteis 
subfoliaceis; involucris anguste campanulatis, breve calyculatis; capi- 
tulis heterogamis, radiatis; involucri squamis plerumque 8, lanceo- 
latis, 6-7 mm. longis, glabris; floribus ligulatis 5-8, ligulis anguste 
oblongis, 3-4 mm. longis, flavis; floribus disci 12-15; pappi setis 
albidis; achaeniis glabris. Costa Rica, "sur les troncs d'arbres des 
paturages de La Palma," alt. 1,460 meters, August, 1898, Adolf o 
Tonduz 12542 (U. S. Nat. Herb., type; Gray Herb., Kew Herb., 
Berlin Herb., British Mus. Herb.). Cascajal, in 1920, C. H. Lankes- 
ter K146 (Kew Herb.). 

SIGESBECKIA L. 

Annual herbs, erect or ascending, branched, viscid-pubescent; 
leaves broad, opposite, dentate; heads small, subradiate, yellow, the 
flowers few, the ray flowers pistillate, fertile, the disk flowers perfect, 
fertile; phyllaries few, herbaceous, glandular, the outer ones usually 
5, linear-cuneate, spreading, the inner small, erect, concave; achenes 
glabrous, obovoid-oblong, turgid, usually incurved, obtuse, without 
pappus. Probably a single species is native in North America. 

Sigesbeckia jorullensis HBK. Mielilla. In forest or thickets, 
Meseta Central and region of Dota, ascending high on the slopes of 
the mountains, at 1,100-3,000 meters. Mexico to Panama, and per- 
haps extending farther southward. A coarse annual, a meter high or 
less, usually rather freely branched, very viscid-pubescent, the stems 
usually dark red; leaves thin, ovate to lance-ovate, often cordate 
at the base, acute or acuminate, coarsely serrate, the long petioles 
broadly winged, broadly dilated at the base and amplexicaul; heads 
very numerous, 5-7 mm. broad, the outer phyllaries conspicuous, 
often greatly elongate. An inconspicuous, weedy plant of unattrac- 
tive appearance. The very viscid flower heads adhere in an annoying 
fashion to clothing. 

SI MSI A Pers. 

Reference: Blake, Proc. Amer. Acad. 49: 376. 1913. 
Coarse, annual or perennial herbs; leaves alternate, or the lower 
usually opposite; heads rather large, radiate or discoid, the flowers 



1520 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

yellow or purple; phyllaries 3-4-seriate, subequal or slightly gradu- 
ated, narrow; receptacle slightly convex, paleaceous; rays usually 
yellow; disk achenes flat, obovate or oblong, glabrous or sericeous, 
the edges thin, not marginate; pappus none or of 2 awns. A few 
other species are found in Central America. 

Sinisia grandiflora Benth. Thickets of the Pacific coast; region 
of Nicoya. Nicaragua to Panama, and perhaps ranging even more 
widely. A coarse annual, a meter high or often much taller, branched, 
viscid-pilose; lower leaves broadly ovate, acute, truncate at the base, 
crenate or serrate, glandular-puberulent and setose, long-petiolate, 
the upper leaves sessile or nearly so; heads few, axillary and terminal, 
the disk 2-3 cm. broad; phyllaries lance-ovate to ovate-oblong, sub- 
acute, glandular-hispid; rays about 1 cm. long; achenes 5-6 mm. long. 
In some regions of the Pacific coast of Central America this plant is 
a common weed in cultivated fields. 

SONCHUS L. 

Annual or perennial herbs with milky sap, somewhat succulent; 
leaves alternate, auriculate-amplexicaul and sessile, entire to dentate 
or pinnatifid, prickly-margined; heads medium-sized, corymbose 
or paniculate, yellow, the flowers all with liguliform corollas; involu- 
cre ovoid or campanulate, the phyllaries herbaceous, imbricate in 
several series, the outer shorter; receptacle flat, naked; achenes oval 
to linear, compressed, costate, truncate; pappus of numerous soft, 
white, capillary bristles. No other species are known from Central 
America, and none are native in North America. 

Sonchus oleraceus L. Lechuguilla, Serraja, Serrajilla. Fre- 
quent in waste or cultivated ground, especially about dwellings, 
Meseta Central and elsewhere, ascending to 2,400 meters. Native 
of the Old World, but abundantly naturalized in temperate and 
tropical America. An annual, commonly about 60 cm. high, simple 
or sparsely branched; basal and lower cauline leaves lyrate-pinnatifid, 
the terminal segment usually larger and deltoid; heads pale yellow, 
2-3 cm. broad. 

SPILANTHES Jacq. 

Reference: A. H. Moore, Revision of the genus Spilanthes, Proc. 
Amer. Acad. 42: 521. 1907. 

Annual or perennial herbs, erect to prostrate; leaves opposite, 
sessile or petiolate; heads small, discoid or radiate, yellow or white, 
the rays usually small; phyllaries few or numerous, obtuse or acute, 



FLORA OF COSTA RICA 1521 

ovate or lanceolate; achenes compressed, the margin ciliate or naked; 
pappus none or of 2 short awns. A few other species are recorded 
from Central America. 

Spilanthes americana (Mutis) Hieron. Calabaza. S. Mutisii 
HBK. S. parvifolia Benth. in Oerst. Vid. Medd. Kjoebenhavn 1852: 
100. 1853 (Santa Rosa, Guanacaste, Oersted). Moist banks or fields, 
Meseta Central to the Pacific coast. Widely distributed in tropical 
America. Plants slender, prostrate or procumbent, rooting at the 
nodes, somewhat pubescent or glabrate; leaves ovate or ovate-lanceo- 
late, mostly 3 cm. long or less, serrate or subentire, obtuse or acute; 
heads radiate, the disk about 7 mm. broad, the rays short but con- 
spicuous, yellow. Moore reported for Costa Rica S. americana var. 
parvula (Robinson) Moore and S. americana var. parvula f. parvifolia 
(Benth.) Moore (based on S. parvifolia Benth.), both of which seem 
to be forms of no particular systematic importance. The plants often 
form dense carpets of yellow along country roads. 

Spilanthes macrophylla Greenm. Proc. Amer. Acad. 39: 109. 
1903. Rio del Volcan, Pittier 3717. Alto del Zacatal; San Marcos; 
along streams, region of San Ramon. Honduras to Panama. Plants 
much larger than in the other species, apparently usually a meter 
high or more and said to reach almost 2 meters, the stems sparsely 
hirsute; leaves ovate-lanceolate, 5-15 cm. long, acute or acuminate, 
entire or serrate, short-petiolate, glabrous or sparsely hirsute beneath; 
heads radiate, yellow, 8-10 mm. high, axillary and terminal; phyl- 
laries biseriate, acute, 4-7 mm. long. 

Spilanthes ocymifolia (Lam.) A. H. Moore. Frequent in moist 
fields and waste ground, Meseta Central to the Pacific tierra caliente; 
region of Dota; Guanacaste; region of San Ramon. Widely dis- 
tributed in tropical America. Plants usually erect, 60 cm. high or 
less, pubescent or glabrate; leaves ovate, dentate or subentire; heads 
whitish, discoid, in fruit about 1 cm. long. Easy of recognition 
because of the whitish, discoid heads, which very rarely have short 
and inconspicuous, whitish rays. Var. acutiserrata Moore (Proc. 
Amer. Acad. 42: 533. 1907; Cartago, Cooper 5807) is a form of slight 
importance with acuminate, acutely serrate leaves. 

Spilanthes poliolepidica A. H. Moore, Proc. Amer. Acad. 42: 
540. 1907. Chilamate, near Rio Sarapiqui, Biolley 7420. Honduras. 
Plants grayish-pilose; leaves lanceolate, 3-5 cm. long, short-petiolate 
or almost sessile, entire or nearly so; heads yellow, 6-8 mm. long, 
radiate; phyllaries pilosulous; achenes ciliate. 



1522 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Spilanthes uliginosa Swartz is reported from Costa Rica by 
Klatt and may occur here, since it is found in Panama. The speci- 
mens cited by Klatt do not seem to be mentioned by Moore in his 
account of the genus, and their proper specific position must remain 
uncertain until they are re-examined. 

STEVIA Cav. 

Reference: Robinson, Observations on the genus Stevia, Contr. 
Gray Herb. 90: 36. 1930; The Stevias of North America, op. cit. 
90. 1930. 

Shrubs or herbs, often viscid; leaves chiefly opposite, petiolate 
or sessile, dentate or entire; heads small, discoid, few-flowered 
(chiefly 5-8-flowered), corymbose-paniculate; phyllaries 5-8, sub- 
equal, rather rigid, narrow, subherbaceous; corolla white, pink, or 
purple; achenes slender, commonly 5-costate, glabrous or hispidulous; 
pappus of 2-12 short squamellae or slender awns, or of both. Other 
species are known from Central America. The genus is much more 
abundantly represented farther north, especially in Mexico, and in 
the South American Andes. 

Stevia elatior HBK. Las Concavas, south of Cartago, Lankester 
266. Mexico to Venezuela and Ecuador. An erect perennial, about 
60 cm. high, the abundant pubescence of spreading, gland-tipped 
hairs, the stems usually simple; leaves broadly deltoid-ovate to ovate- 
oblong, obtuse, crenate-serrate, coarsely pubescent on both surfaces; 
heads about 1 cm. long, long-pedicellate, forming a lax, open panicle; 
phyllaries acute, glandular-pubescent. 

Stevia elongata HBK. Reported from Heredia and the region 
of San Jose". Mexico to Venezuela. An erect perennial, the stems 
glandular-puberulent; leaves ovate or rhombic-ovate to lance-oblong, 
the upper sessile, the middle and lower ones petiolate, acuminate to 
rounded, narrowed to the base, serrate or crenate, pubescent chiefly 
on the nerves and veins; heads 9-11 mm. long, sessile or nearly so, 
in dense, paniculate glomerules; phyllaries oblong or lanceolate, 
acute, glandular-pubescent; corollas pale pink or lilac. The Costa 
Rican material is referred by Robinson to var. caracasana (DC.) 
Robinson. The species seems to be none too well differentiated 
from S. rhombifolia, and so far as Costa Rican material is concerned, 
I rather doubt that there is any constant difference. 

Stevia lucida Lag. S. nitida Walp. Reported by Robinson 
from Volcan de Irazu, at 2,300 meters. Mexico to Venezuela. A 



FLORA OF COSTA RICA 1523 

glutinous shrub a meter high or less; leaves slender-petiolate, ovate- 
oblong or lance-oblong, crenate-serrate or dentate, acuminate or 
acute, glabrous, 4-7 cm. long; heads 1 cm. long, sessile or nearly so, 
in dense, corymbose glomerules; phyllaries linear, acute, very viscid; 
corollas white or pinkish; achenes bearing a short crown of some- 
what dentate, connate squamellae. 

Stevia rhombifolia HBK. S. compacta Benth. Frequent in 
thickets or forest of the central region, descending into the upper 
part of the Pacific tierra caliente; regions of Dota and San Ramon; 
mostly at 800-1,300 meters. Mexico to Peru. An erect, perennial 
herb, a meter high or less, simple or sparsely branched, the stems 
puberulent; leaves ovate or rhombic-ovate to lance-oblong, mostly 
3-6 cm. long, acute or acuminate, narrowed at the base, crenate- 
serrate, sessile or nearly so, pubescent on both surfaces; heads 6-9 
mm. long, white or pale pink, sessile or nearly so, in compact glom- 
erules arranged in a convex panicle; phyllaries narrowly oblong, 
acute, puberulent and glandular; achenes bearing a crown of short, 
erose squamellae. The Costa Rican material is referred by Robinson 
to var. stephanocoma Schultz Bip. 

STRUCHIUM P. Br. 

Somewhat succulent, erect or ascending herbs; leaves alternate, 
thin, petiolate, serrate; heads small, axillary, sessile, discoid, white 
or purplish; involucre hemispheric, the phyllaries imbricate in several 
series; receptacle convex; achenes 4-angulate, truncate, glabrous or 
minutely glandular; pappus a cartilaginous crown. The genus con- 
sists of a single species. 

Struchium sparganophorum (L.) Kuntze. Sparganophorus 
Vaillantii Crantz. Thickets or wet ground, Atlantic tierra caliente. 
British Honduras to Panama and West Indies; tropical Africa. 
Plants a meter high or less, sparsely appressed-pilose on the young 
parts; leaves lanceolate to elliptic, acute at each end, 5-12 cm. long, 
often almost entire; heads 6-9 mm. broad, densely aggregate in 
the leaf axils; phyllaries acuminate, scarious-marginate; achenes 
1.5 mm. long. 

Syncephalantha sanguinea Klatt was based upon material col- 
lected by Warscewicz in "Guatemala et Costa Rica," and it is 
reported from Costa Rica by Rydberg (N. Amer. Fl. 34: 160. 1915), 
probably upon the basis of Klatt's record. The plant is frequent 
in Guatemala, but has not been found recently in Costa Rica, there- 
fore the record for the latter country probably should be deleted. 



1524 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

SYNEDRELLA Gaertn. 

Annual herbs, erect or ascending; leaves opposite, petiolate, 
dentate; heads small, axillary, usually sessile and glomerate, radiate, 
yellow; involucre oblong, the phyllaries few, imbricate, the outer 
1-2 foliaceous, the inner paleaceous; receptacle small, paleaceous; 
ray flowers pistillate and fertile, the disk flowers perfect; ray achenes 
compressed, 2-winged, the wings produced into awns, the disk 
achenes narrower, 2-3-aristate. No other species are found in 
Central America. 

Synedrella nodiflora (L.) Gaertn. Waste ground or thickets 
of the tierra caliente, ascending rather high on the Pacific slope. 
Widely distributed in tropical America. Plants appressed-pilose, 
erect or sometimes weak and reclining, much branched ; leaves ovate 
or elliptic, 3-10 cm. long, triplinerved, acute, narrowed at the base to 
the short petiole, serrate; heads several together in dense, sessile 
clusters; involucre 8 mm. high, the outer phyllaries oblong-lanceolate, 
pubescent; achenes 4-5 mm. long, those of the ray flowers with 
marginal bristles and 2 short awns. 

TAGETES L. 

Reference: Rydberg, N. Amer. Fl. 34: 148. 1915. 

Annual or perennial, strong-scented herbs; leaves opposite, or 
the upper alternate, commonly pinnate or pinnatifid, conspicuously 
gland-dotted; heads radiate, yellow or orange, pedunculate, solitary 
at the ends of the branches or in leafy cymes; involucre fusiform to 
campanulate; phyllaries 3-10, narrow, uniseriate, united almost to 
the apex, glandular; ray flowers pistillate, fertile, the disk flowers 
perfect and fertile; achenes narrow and slender, angulate or terete, 
glabrous or pubescent; pappus squamellae 3-10, very unequal, 1 or 2 
of them much longer than the others. A few other species occur 
in Central America. 

Tagetes erecta L. Flor de muerto. Marigold. Cultivated com- 
monly in gardens for ornament, sometimes escaping. Perhaps native 
of Mexico. A tall, stout, branched annual, the stems glabrous; leaves 
pinnate, the 11-17 leaflets lanceolate or linear-lanceolate, 1-5 cm. 
long, serrate; involucre 1.5-2 cm. long, glabrous, the phyllaries 5-8; 
disk corollas glabrous; achenes 7-8 mm. long. A well known orna- 
mental plant, cultivated in almost all parts of the earth. 

Tagetes filifolia Lag. Anis, Manzanilla, Anisillo. T. congesta 
Hook. & Arn. Frequent in fields of the central region; regions of 



FLORA OF COSTA RICA 1525 

Dota and San Ramon. Extending to Mexico. A slender annual, 
erect, 20-40 cm. high, simple below, branched above, glabrous or 
nearly so; leaves pinnatifid or bipinnatifid into linear-filiform, short 
divisions; heads mostly pedicellate; involucre 7 mm. long, not angu- 
late at the base, the phyllaries 5; ray flowers 1-3, the ligules 1.5 mm. 
long; disk corollas glabrous; achenes 4.5 mm. long. In domestic 
medicine, an infusion of the plant is employed as a diuretic. 

Tagetes foetidissima DC. Region of Santa Maria de Dota; 
Guanacaste. Extending to Mexico. A glabrous, branched annual, 
a meter high or less; leaflets 11-29, oblong or lance-oblong, 1-2.5 
cm. long; heads long-pedunculate, the involucre 1.5 cm. long, the 
phyllaries 5; ray flowers usually 5, the ligules 2 mm. long; disk 
corollas viscid-hispid; achenes hirsute, 6 mm. long. 

Tagetes microglossa Benth. Flor de muerto; Rudillo, Man- 
zanilla. T. macroglossa Polak. Linnaea 41 : 580. 1878 (near San Jose", 
Polakowsky 372; photo, in Herb. Field Mus.). Common in fields 
and thickets of the Meseta Central, descending to the Pacific coast; 
regions of Dota and San Ramon. Guatemala to Ecuador. A slender, 
glabrous annual, a meter high or less, often densely branched; leaf- 
lets 7-11, oblanceolate, 1-2 cm. long, serrate; heads numerous, long- 
pedunculate, the involucre 1.5 cm. long; phyllaries 5; ray flowers 
3-5, the ligules small; disk corollas glabrous; achenes 8-9 mm. long, 
hispidulous. 

Tagetes pusilla HBK. Flor de muerto. Frequent in fields or 
thickets of the Meseta Central; region of San Ramon. Guatemala 
to Bolivia. Closely related to T. filifolia, and probably only a form 
of that species, if at all distinct; plants usually low and diffusely 
branched, about 10 cm. high; heads sessile or nearly so, the involucre 
conspicuously angulate at the base. 

Tagetes remotiflora Kunze. Flor de muerto. Fields and thickets 
of the Meseta Central. Extending to Mexico. A glabrous annual, 
70 cm. high or less; leaflets 9-25, lanceolate or linear-lanceolate, 1-3 
cm. long, serrate; heads long-pedunculate, the involucre 13-14 mm. 
long; phyllaries usually 5; ray flowers 5, the ligules obovate, almost 
1 cm. long; disk flowers puberulent. 

Tagetes subulata Cervantes. In pastures, region of San Ramon, 
1,050 meters. Mexico to Colombia. A slender, glabrous annual, 
50 cm. high or less, usually openly branched; leaflets 5-13, 1 cm. 
long or less, pectinately dissected into narrow lobes; heads long- 



1526 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

pedunculate, the involucre 1.5 cm. long, the phyllaries 4-5; ray 
flowers 3-5, the ligules 2 mm. long; achenes 5 mm. long. 

Tagetes tenuifolia Cav., which ranges from Mexico to Colom- 
bia, has been reported from Costa Rica and may occur here, but I 
have seen no specimens certainly referable to it. 

TARAXACUM Hall. Dandelion 

Acaulescent, perennial herbs with thick roots, the sap milky; 
leaves all basal, forming a rosette, pinnatifid or sinuate-dentate; 
heads large, yellow, solitary at the ends of hollow scapes, the corollas 
all liguliform; inner phyllaries 1-seriate, subequal, slightly united at 
the base, with several series of short, spreading outer ones at the 
base; receptacle flat, naked; achenes oblong or linear-fusiform, 5-10- 
nerved, roughened above, tapering into a slender beak; pappus of 
numerous soft, .slender bristles. No species are native in Central 
America. 

Taraxacum officinale Weber. Diente de Icon, Amargon, Arnica. 
Frequent in fields and on open banks about San Jose" and Cartago; 
pastures of Volcan de Turrialba. Native of Europe, but widely ad- 
ventive in North America. Leaves oblong to spatulate, pinnatifid to 
almost entire, sparsely pubescent when young; scapes 5-20 cm. high, 
the showy heads 2.5-5 cm. broad. 

TESSARIA Ruiz & Pavon 

Grayish-pubescent shrubs; leaves alternate, usually entire; heads 
small, discoid, whitish or purplish, in terminal cymes or corymbs, 
involucre ovoid, the phyllaries multiseriate, imbricate, scarious; 
graduated; receptacle plane, naked; achenes small, 4-5-angulate, 
glabrous or pubescent; pappus of numerous slender bristles. A 
single species is found in North America. 

Tessaria mucronata DC. Usually along sandy stream beds, 
Pacific tierra caliente and even at higher elevations on the Pacific 
slope, ascending to about 1,200 meters; region of San Ramon. 
Extending to Colombia and perhaps farther southward. A shrub 
of 1.5-3.5 meters, the branches sericeous or glabrate; leaves mostly 
oblong-oblanceolate, acute, attenuate to the base, entire, densely 
grayish- or brownish-sericeous on both surfaces; heads about 5 mm. 
long, sessile and densely aggregate, the outer phyllaries somewhat 
tomentose, the inner ones glabrous. The Costa Rican plant has been 
referred to T. legitima DC. The taxonomy of the genus is apparently 
in a chaotic condition, and the proper name for the Central American 
species is questionable. 



FLORA OF COSTA RICA 1527 

TITHONIA Desf. 

Reference: Blake, Revision of the genus Tithonia, Contr. U. S. 
Nat. Herb. 20: 423-436. 1921. 

Tall, coarse herbs, or shrubs; leaves alternate, or the lowest oppo- 
site, petiolate, dentate or lobate; heads very large, radiate, yellow, 
the long peduncles fistulose; involucre broad, the phyllaries about 
4-seriate, broad, graduate; rays neutral; achenes turgid; pappus of 
free or connate squamellae, with or without awns, or absent. At 
least one other species is native in Central America. 

Tithonia diversifolia (Hemsl.) Gray. Mirasol. Frequent in 
thickets or fields of the Meseta Central; region of Dota; chiefly at 
1,000-1,400 meters. Ranging to Mexico. Plants herbaceous or 
more or less woody, usually 1.5-3 meters high, branched, the stems 
hispid-pilose; leaves large, ovate or deltoid-ovate, usually 3-5-1 obate, 
cuneate at the base, the lobes acuminate to attenuate, tuberculate- 
hispidulous above, soft-pilose beneath; disk of the head 1.5-4 cm. 
broad; phyllaries broadly oval or oval-ovate, rounded at the apex, 
sparsely pilose or almost glabrous; rays 6 cm. long or less; pappus of 
2 awns and 6-10 squamellae. In many regions this is a common weed 
of cornfields. The Costa Rican plant was referred by Klatt in error 
to T. speciosa Hook. 

Tithonia longeradiata (Bertol.) Blake. T. scaberrima Benth. 
Rio Tiliri, near San Jose*. Ranging to southern Mexico. A stout, 
coarse herb, sometimes 5 meters high, the stems densely pilose or 
hispid-pilose; leaves ovate-lanceolate to broadly ovate, acuminate, 
narrowed to the petiole, crenate-serrate, scabrous above, densely 
pilose beneath; disk 1.5-3.5 cm. broad; phyllaries 4-seriate, oval or 
obovate, broadly rounded at the apex, hispid-pilose or glabrate; 
achenes glabrous; pappus none. 

Tithonia Pittieri (Greenm.) Blake, Contr. Gray Herb. 54: 9. 
1918. Gymnolomia Pittieri Greenm. Proc. Amer. Acad. 39: 101. 1903. 
Rio Ceiba, Buenos Aires, Pittier 3735. Regions of Buenos Aires 
and San Jose", at 1,200 meters or less. Ranging to Guatemala. 
Plants tall, herbaceous, the stems hispid-pilose; leaves lanceolate, 
acuminate, tapering to the short petiole, crenate-serrulate, hispidu- 
lous above, pilosulous beneath; disk 1-2 cm. broad; phyllaries 4-seri- 
ate, the outermost oblong to obovate-oval, obtuse or subacute, the 
innermost obovate, rounded at the apex; rays about 2 cm. long; 
pappus none. 



1528 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Tithonia rotundifolia (Mill.) Blake. T. aristata Oerst. Vid. 
Medd. Kjoebenhavn 1852: 114. 1853 (Cerro de Aguacate, Oersted). 
T. speciosa Hook. Urbanisol aristatus Kuntze, Rev. Gen. 1: 371. 
1891. In thickets and fields, often along streams, Meseta Central 
to the Pacific coast; Guanacaste; region of San Ramon. Panama to 
Mexico. A stout annual, usually 1-2 meters high, the younger 
parts densely grayish-pilosulous; leaves large, broad, mostly trilobate, 
the lobes caudate-acuminate, hispid-pilose on both sides; disk 2-3 
cm. broad; involucre 2-seriate, the phyllaries ovate to ovate-oblong, 
acute or acuminate, the outer ones with herbaceous tips, pilosulous 
or glabrate; rays 2-2.5 cm. long; achenes pilose, the pappus of squa- 
mellae and 2 awns. 

TRIDAX L. 

Annual or perennial herbs, usually low, often prostrate or pro- 
cumbent; leaves opposite, dentate or incised; heads radiate, long- 
pedunculate, the rays often trilobate; involucre ovoid to hemispheric, 
the phyllaries subequal, few-seriate; receptacle flat or convex, palea- 
ceous; achenes sericeous- villous; pappus of numerous aristate, 
plumose scales. One or two other species may reach northern 
Central America. 

Tridax procumbens L. Frequent in waste ground or thickets, 
Pacific tierra caliente; probably also in the Atlantic tierra caliente. 
Widely distributed in tropical America. Plants hirsute, branched 
from the base, the branches prostrate or ascending; leaves ovate to 
ovate-lanceolate, 2-6 cm. long, short-petiolate, acute or acuminate, 
incised-dentate, cuneate at the base; peduncles solitary, terminal, 
10-30 cm. long; involucre campanulate, 6 mm. high, hispid, the inner 
phyllaries elliptic, the outer lanceolate; rays almost white; achenes 
2 mm. long. One of the most common weeds in many parts of the 
Central American tierra caliente. 

TRIXIS P. Br. 

Reference: Robinson & Greenman, Revision of the Mexican and 
Central American species of Trixis, Proc. Amer. Acad. 40: 6-14. 1904. 

Shrubs, often somewhat scandent; leaves alternate, entire or 
dentate; heads yellow, cymose or paniculate; involucre double, the 
outer phyllaries few, usually shorter, linear to ovate, herbaceous, 
the inner ones 5-10, equal, subherbaceous; flowers all perfect and 
fertile, the corollas all bilabiate; achenes subcylindric, papillose; 
pappus of numerous bristles, usually brownish. A few other species 
are known to occur in Central America. 



FLORA OF COSTA RICA 1529 

Trixis divaricata (HBK.) Spreng. Reported from El General, 
the determination by Blake. Unknown elsewhere in North America, 
but extending southward to Argentina. A slender shrub, probably 
more or less scandent; leaves sessile or nearly so, oblong-lanceolate 
or linear-lanceolate, subentire, attenuate-acuminate, acute at the 
base, green above but sericeous, densely pale-tomentose beneath; 
branches of the inflorescence flexuous, the cymes open, the heads 
mostly pedicellate, 1 cm. long. 

Trixis radial is (L.) Kuntze. T. frutescens P. Br. In thickets 
and hedges, Meseta Central to the Pacific coast; region of San 
Ramon; Guanacaste. Widely distributed in tropical America. A 
shrub of 1-2.5 meters, the branches often recurved or subscandent, 
glabrous or sparsely appressed-pilose; leaves narrowly lanceolate 
to lance-ovate, 3-10 cm. long, acuminate, subentire, glabrous or 
glabrate; inflorescence dense and leafy, the heads sessile or short- 
pedicellate, 1.5-2 cm. high. One of the common, weedy shrubs in the 
Pacific tierra caliente of Central America, rather showy and hand- 
some when in full bloom. 

VERBESINA L. 

Reference: Robinson & Greenman, Synopsis of the genus Verbe- 
sina, with an analytical key to the species, Proc. Amer. Acad. 34: 
534-566. 1899. 

Herbs, shrubs, or small trees; leaves opposite or alternate, dentate 
to pinnate-lobate; heads small or medium-sized, radiate or discoid, 
solitary to numerous and paniculate, yellow, orange, or white; phyl- 
laries about 2-seriate, usually unequal and more or less herbaceous; 
rays usually fertile; achenes strongly compressed, 2- winged; pappus of 
2 awns, rarely none. Other species are found in Central America. 

Verbesina Fraseri Hemsl. Hacienda La Argentina, Grecia, 
825 meters, M. Valeria 580. Extending to Guatemala. Plants appar- 
ently somewhat scandent, the stems broadly winged, glabrous or 
glabrate; leaves scabrous, opposite, deeply pinnate-lobate; heads 
large, subglobose, discoid; outer phyllaries obovate or oblanceolate, 
obtuse. From Costa Rica I have seen a single sterile specimen that 
seems to belong here, but it may be that it is merely an abnormal 
form of V. Tonduzii. 

Verbesina myriocephala Schultz Bip. Thickets, fields, and 
hedges of the Meseta Central; region of San Ramon. Panama to 
southern Mexico. A coarse, branched herb, 1.5-3 meters high, the 
stems glabrous, usually dark purple; leaves alternate, large, deeply 



1530 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

pinnatifid, scabrous above, soft-pilose or rarely glabrate beneath; 
heads dirty white, discoid, 8 mm. high, forming large, dense, 
terminal corymbs; phyllaries narrowly oblong; achenes obscurely 
winged. V. costaricensis Robinson, Proc. Amer. Acad. 47: 212. 1911 
(Rio Virilla, San Jose", Tonduz 9833}, seems to be doubtfully if at all 
distinct from V. myriocephala. Costa Rican material of this species 
has been referred also to V. nicaraguensis Benth. and V. gigantea Jacq. 

Verbesina Oerstediana Benth. in Oerst. Vid. Medd. Kjoeben- 
havn 1852: 96. 1853. Torilla. Southern slope of Volcan de Irazu, 
2,400-2,850 meters, Oersted 8729 (photo, and duplicate of the type 
in Herb. Field Mus.). Frequent in thickets and forest of the central 
region, especially on the middle and higher slopes of the mountains; 
region of San Ramon; at 1,000-2,900 meters. Endemic. A shrub 
or tree, sometimes 9 meters high, with a trunk 25 cm. in diameter, 
the branches pilose or subtomentose; leaves short-petiolate, lance- 
oblong or oblong-oblanceolate, acuminate, attenuate to the base, 
pilose or glabrate above, densely or sparsely pilose beneath, rather 
inconspicuously denticulate; heads bright yellow, radiate, forming 
large or small, compact corymbs at the tips of the branches; in- 
volucre 5 mm. high; achenes narrowly winged. Var. glabrior Blake 
(Field Mus. Bot. 17: 404. 1938; oak forest, Santa Maria de Dota, 
Standley 41619) is a form with less abundant and shorter pubescence 
than the type. It occurs chiefly at lower elevations than typical 
V. Oerstediana. 

Verbesina sublobata Benth. Tora. Thickets and fields of 
the Meseta Central; region of Dota; San Ramon; 1,000-1,800 
meters. Extending to southern Mexico. An herb or shrub of 
1-4.5 meters, the branches densely sordid-tomentose or pilose, not 
winged; leaves alternate, large, deeply sinuate-lobate, very rough 
above, pilose-tomentose beneath; heads very numerous, dirty white, 
with short and inconspicuous rays; involucre 3 mm. high, the 
phyllaries linear or oblanceolate; achenes narrowly winged. 

Verbesina Tonduzii Greenm. Proc. Amer. Acad. 40: 42. 1904. 
Las Vueltas, Tucurrique, 635 meters, Tonduz 12765. In thickets 
or open, dry forest, tierra caliente, ascending to about 1,000 meters 
on the Pacific slope; region of San Ramon; Guanacaste. Nicaragua. 
A small, suffrutescent vine, the stems narrowly winged, glabrous 
or nearly so; leaves opposite, ovate, 5-14 cm. long, acuminate, 
unequally sinuate-dentate, narrowed to the base, rough-pubescent; 
heads few, solitary at the ends of long, naked peduncles, discoid, 



FLORA OF COSTA RICA 1531 

bright orange, 1.5-2.5 cm. broad; phyllaries very numerous, densely 
puberulent; achenes broadly winged, 5-7 mm. long. Called Capi- 
taneja in Nicaragua. Costa Rican collections have been referred 
to V. Fraseri Hemsl. and V. crocata (Cav.) Less. The species is 
very close to V. crocata, differing principally in leaf characters. 

Verbesina turbacensis HBK. V. nicaraguensis Benth.; V. 
microcephala Benth. Thickets and fields of the Meseta Central; 
regions of Dota and El General; 600-1,300 meters. Mexico to 
Colombia. Plants tall and coarse, herbaceous or somewhat woody, 
commonly 1-3 meters high, the branches densely sordid-tomentose, 
winged by the decurrent leaf bases; leaves large, alternate, deeply 
pinnate-lobate, rough above, puberulent or tomentose beneath; 
heads very numerous, white, forming large, corymbose panicles, 
the white rays very small; involucre 3 mm. high. 

VERNONIA Schreb. 
Reference: Gleason, N. Amer. Fl. 33: 52-101. 1922. 

Annual or perennial herbs, shrubs, trees, or woody vines; leaves 
alternate, entire or dentate; heads discoid, white, pink, or blue, 
mostly in scorpioid cymes, these usually panicled, the heads some- 
times solitary or corymbose, 5-many-flowered ; involucre cylindric 
to campanulate, the phyllaries laxly or closely imbricate in few or 
numerous series, the outer ones shorter; receptacle flat or subconvex; 
achenes costate or smooth, truncate; pappus biseriate, the outer 
short, of scales or bristles, the inner of stiff bristles. Other species 
occur in Central America. 

Vernonia argyropappa Buek. Occasional in thickets and 
forest of the central region; apparently common in the region of 
San Ramon; 1,000-1,800 meters. Southern Mexico to South Amer- 
ica. Plants herbaceous or suffrutescent, 1-1.5 meters high, sparsely 
branched, the branches brown-pilose; leaves oblong-lanceolate, 8-12 
cm. long, acuminate, entire or nearly so, pilose above, strigose- 
hirsute and resinous beneath; inflorescence leafy-bracted, the 
branches long and slender; heads 20-30-flowered, the involucre 
8-10 mm. high, the flowers bright purple; phyllaries subulate, 
erect; pappus white; achenes 3 mm. long, hirsute. Costa Rican 
material of this species was listed by Klatt as V. remotiflora var. 
tricholepis Baker. 

Vernonia brachiata Benth. in Oerst. Vid. Medd. Kjoebenhavn 
1852: 67. 1853. Cana de danto, Tabaquillo. Cacalia brachiata Kuntze, 



1532 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Rev. Gen. 969. 1891. Cerro de Aguacate, 600 meters, Oersted (photo. 
of type in Herb. Field Mus.). Frequent in thickets and open forest 
of the tierra caliente, especially the Atlantic, ascending in the 
central mountains to about 1,500 meters, but rare at higher eleva- 
tions; region of San Ramon; Atlantic slope of Guanacaste. Also in 
northern South America. Plants herbaceous or suffrutescent, 1-2.5 
meters high, the stems glabrous or nearly so; leaves thin, elliptic, 
10-20 cm. long, acute or acuminate, narrowed to the base, entire 
or serrulate, almost glabrous; branches of the inflorescence very 
long and slender, leafless; heads 21-flowered, purple, the involucre 
3-4 mm. high; phyllaries ciliate, acute or acuminate; achenes 
sparsely hirsute; pappus brownish. 

Vernonia canescens HBK. V. buttata Benth. in Oerst. Vid. 
Medd. Kjoebenhavn 67. 1853 (mountains south of Cartago, 1,500 
meters, Oersted 8540; photo, of type in Herb. Field Mus.). Cacalia 
bullata Kuntze, Rev. Gen. 969. 1891. Common in thickets of the 
Meseta Central, ascending to 1,800 meters or more; region of San 
Ramon; Guanacaste. Mexico to South America. A slender shrub 
of 1-3 meters, sometimes somewhat scandent, the branches pubes- 
cent or glabrate; leaves rather rigid, often bullate, oblong-lanceolate 
to broadly ovate, 5-10 cm. long, acuminate, rounded or obtuse at 
the base, entire, rough above, densely strigose and resinous beneath; 
inflorescence large and usually much branched, the bracts small 
and inconspicuous; heads pink, 21-flowered, the involucre 4-5 mm. 
high; phyllaries laxly villous, triangular-subulate or lanceolate, 
acuminate; achenes 2 mm. long, hirsute; pappus white. A common, 
weedy shrub in the central region, where it often forms extensive 
thickets. Material referred by Klatt to V. mollis HBK. probably 
is to be placed here. 

Vernonia Lankesteri Blake, Journ. Wash. Acad. Sci. 15: 106. 
1925. Santa Clara de Cartago, Lankester 712. Occasional in forests 
of the central region, especially in the mountains south of Cartago; 
La Palma de San Jose"; chiefly at 1,500-2,500 meters. Endemic. 
Plants suffrutescent, 1-3 meters high, the young branches tomentose; 
leaves obovate, 12-21 cm. long, acuminate, long-attenuate to the 
base, serrate, glabrate in age; heads in panicles of 4-8, about 2.5 
cm. broad, rose-purple; phyllaries triangular to oblong-obovate, 
with broad, obtuse or rounded, herbaceous, spreading tips; achenes 
glabrous; pappus yellowish white. A showy and unusually hand- 
some plant, very different in general appearance from other local 
species. The heads are much larger than in other species occurring 



FLORA OF COSTA RICA 1533 

in Costa Rica. Vernonia Salvinae Hemsl. was reported from Costa 
Rica by Gleason on the basis of a specimen really referable to 
V. Lankesteri. 

Vernonia patens HBK. Tuete, Tuete bianco. V. pacchensis 
Benth. In thickets of the tierra caliente, ascending to the region 
of La Palma, about 1,400 meters. Southern Mexico to South 
America. A stiff shrub of 2-3 meters, the stems puberulent or 
glabrate; leaves thick and stiff, oblong-lanceolate, 8-15 cm. long, 
often rugose, acuminate, narrowed to the base, entire or serrulate, 
puberulent, the hairs of the lower surface very short and conic; 
inflorescence large but dense, the branches short; heads 21-flowered, 
the flowers white or pale pink, fragrant; involucre 4-5 mm. high, the 
phyllaries appressed, subacuminate to rounded at the apex, tomen- 
tose-ciliate; achenes minutely pubescent, the pappus pale brown. 

Vernonia Seemanniana Steetz is reported from Costa Rica by 
Gleason, but probably in error. The type locality is Volcan de 
Chiriqui, Panama (not Costa Rica, as stated by Gleason), and very 
likely the species is endemic there. 

Vernonia stellaris Llave & Lexarza. Tuete. V. Deppeana Less. 
Common in thickets or forest of the Meseta Central; region of San 
Ramon; chiefly at 900-1,300 meters. Ranging to Mexico. A shrub 
of 3-6 meters, the branches pubescent or tomentose; leaves oblong 
to narrowly elliptic, 8-10 cm. long, acute, narrowed to rounded at 
the base, entire or nearly so, rough above, tomentose beneath; 
inflorescence large and much branched, the branches short; heads 
white or pale pink, the flowers fragrant; involucre 3-4 mm. high, 
the phyllaries erect or appressed, ovate to oblong, acute or acumi- 
nate, arachnoid-ciliate; achenes pubescent, the pappus fulvous. 

Vernonia triflosculosa HBK. Quitirri. Eremosis triflosculosa 
Gleason. Frequent in thickets or forest about San Jos and in the 
region of San Ramon, 1,000-2,000 meters. Panama to southern 
Mexico. A shrub or tree, sometimes 6 meters high or more, the 
trunk as much as 15 cm. in diameter, the crown rounded, the branches 
pubescent or glabrate; leaves thin, oblanceolate to elliptic, 8-12 cm. 
long, acute or acuminate, attenuate to the base, entire or nearly so, 
nearly or quite glabrous; heads 3-flowered, white, in small, dense 
clusters arranged in leafy panicles; involucre cylindric, pale brown, 
4-5 mm. high, the phyllaries subrotund to oblong, acute, glabrous 
or nearly so, minutely ciliate; achenes pubescent, the pappus white. 



1534 FIELD MUSEUM OF NATURAL HISTORY BOTANY, VOL. XVIII 

Vernonia vernicosa Klatt, Bull. Soc. Bot. Belg. 35: 294. 1896. 
Tuete. V. vernicosa var. comosa Greenm. Proc. Amer. Acad. 30: 
93. 1903 (Matamber, near Nicoya, Tonduz 13607). Banks of Rio 
Virilla, Tonduz 7065 (9860). Frequent in thickets, region of San 
Jos to the Pacific coast; region of San Ramon; Guanacaste. En- 
demic. A stout shrub of 1-2.5 meters, the branches sparsely pubes- 
cent or glabrate; leaves coriaceous, oblong-lanceolate to ovate, 
6-10 cm. long, acute or acuminate, rounded at the base, entire, 
strigose-hirsute or glabrate; inflorescence lax and sparsely branched, 
very leafy; heads about 50-flowered, white or pale pink, fragrant; 
involucre 1 cm. high, the phyllaries very numerous, appressed, 
narrow, sparsely villous or glabrous; achenes hirsute; pappus white. 
This species usually has been confused with V. tortuosa (L.) Blake 
(V. Schiedeana Less.), a plant of Mexico and northern Central 
America. 

VIGUIERA HBK. 

Reference: Blake, A revision of the genus Viguiera, Contr. 
Gray Herb. 54: 1-205. pi 1-3. 1918. 

Herbs or shrubs, usually with rough pubescence; leaves opposite, 
at least the lower ones; heads small or large, yellow, radiate; invo- 
lucre 2-7-seriate, the phyllaries graduated or subequal, usually with 
indurate base and herbaceous tip; rays neutral; achenes thick, the 
pappus persistent, of 2 awns and several free or united squamellae, 
or wanting. A few other species are known in Central America. 

Viguiera strigosa Klatt, Bull. Soc. Bot. Belg. 31: 204. 1892. 
El Rodeo, Rio Tiribi, 700 meters, Pittier 1604- Collected also at 
Libano, Guanacaste, in dry forest. Endemic. Herbaceous, the 
stems sparsely pilose and puberulous; leaves rhombic-ovate or lance- 
ovate, acuminate, cuneate at the base, crenate-serrate, hispid- 
strigose above, beneath sparsely strigillose and somewhat gland- 
dotted, petiolate; heads long-pedunculate, the disk 6 mm. high, 
in age 9-11 mm. thick; phyllaries 2-seriate, obovate-oblong, sparsely 
strigillose, acute; achenes appressed-pilose, 3 mm. long; pappus of 
2 awns and about 6 unequal squamellae. 

Viguiera sylvatica Klatt, Bull. Soc. Bot. Belg. 31: 204. 1892. 
"Massif du Barba," Carrizal, 1,900 meters, Pittier 779. Collected 
also at San Ramon and in El General; Cerro de La Carpintera; region 
of Dota. Panama. Herbaceous, the stems hispid-pilose; leaves 
ovate, acuminate, cuneate to rounded at the base, serrate, strigillose 
and short-hispid above, hispidulous-pilosulous and hispid beneath; 
heads numerous, irregularly paniculate; involucre 2-seriate, 5-7 mm. 



FLORA OF COSTA RICA 1535 

high; phyllaries oblong-lanceolate, short-acuminate, hispid-pilose; 
achenes black and white, appressed-pilose, 2.6 mm. long; pappus 
of 2 awns and 6 lacerate squamellae. 

Viguiera tenuis Gray. Region of Boruca. Panama to Mexico. 
A slender annual, 30-50 cm. high; stems strigose or strigose-hispid ; 
leaves mostly opposite, sessile, lanceolate or lance-ovate, 3-6 cm. 
long, acuminate, rounded or cuneate at the base, entire or serrulate, 
strigose-pilose; disk 8-9 mm. high; involucre 2-seriate, the phyllaries 
narrowly linear-lanceolate, attenuate almost from the base, canes- 
cent-strigillose; achenes subsericeous, 2.5 mm. long; pappus of 2 
awns and about 6 lacerate squamellae. 

WEDELIA Jacq. 

Erect or prostrate herbs; leaves opposite, entire or dentate, 
sometimes lobate; heads small, radiate, yellow, long-pedunculate; 
phyllari