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Botany

NEW SERIES. NO 28

FLORA COSTARICENSIS

William Burger, Editor

Family

#97 Krameriaceae

#98 Oxalidaceae

#99 Geraniaceae

#100 Tropaeolaceae

#101 Linaceae

#10 la Humiriaceae

#102 Erythroxylaceae

#103 Zygophyllaceae

October 31, 1991

Publication 1428

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

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FIELDIANA

Botany

NEW SERIES, NO. 28

William Burger, Editor

Curator

Department of Botany

Field Museum of Natural History

Chicago, Illinois 60605-2496

Family #97 Krameriaceae

William Burger and Beryl B. Simpson

Family #98 Oxalidaceae

William Burger

Family #99 Geraniaceae

William Burger

Family #100 Tropaeolaceae

William Burger

Accepted March 20, 1991

Published October 31, 1991

Publication 1428

Family #101 Linaceae

William Burger

Family #101a Humiriaceae

William Burger and Nelson Zamora

Family #102 Erythroxylaceae

Timothy Plowmanf

Family #103 Zgyophyllaceae

William Burger

JAN 2 7 1992

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

Library of Congress Catalog Card Number: 91-75909

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List of Illustrations

INTRODUCTION v

ACKNOWLEDGMENTS v

KRAMERIACEAE by William Burger and

Beryl B. Simpson 1

OXALIDACEAE by William Burger 2

GERANIACEAE by William Burger 16

TROPAEOLACEAE by William Burger 21

LINACEAE by William Burger 23

HUMIRIACEAE by William Burger and

Nelson Zamora 25

ERYTHROXYLACEAE by Timothy Plowman . . 30

ZYGOPHYLLACEAE by William Burger 36

LITERATURE CITED 41

INDEX . 42

1 . Krameriaceae: two species of Kramer ia . . 3

2. Oxalidaceae: two species of Biophytum . . 7

3. Oxalidaceae: four smaller species of

Oxalis 10

4. Oxalidaceae: four larger species of

Oxalis 13

5. Geraniaceae: two species of Geranium ... 19

6. Humiriaceae: four Costa Rican species . . 27

7. Erythroxylaceae: Erythroxylum hava-

nense 34

8. Zygophyllaceae: four Central American

species 38

111

Introduction

This is the seventh issue in the Flora Costari-

censis series. The first dealt with families num-

bered 40 and 41, Casuarinaceae and Piperaceae

(Fieldiana, Bot. 35, 1971). The second included

families 42 through 53, Chloranthaceae through

Urticaceae (Fieldiana, Bot. 40, 1977). The third

issue covered family 15, the Gramineae, authored

by Richard Pohl (Fieldiana, Bot., new series, No.

4, 1980). The fourth issue included families 54

through 70, Podostemonaceae through Cary-

ophyllaceae (Fieldiana, Bot., new series, No. 13,

1983). The fifth issue covered families 200 and

201, the Acanthaceae, authored by L. H. Durkee,

and Plantaginaceae (Fieldiana, Bot., new series,

No. 18, 1986). The sixth issue covered families 80

and 8 1 , the Lauraceae and Hernandiaceae (Field-

iana, Bot., new series, No. 23, 1990). The alpha-

betical listing and the sequence of families are giv-

en inside the front and back covers.

This issue includes the Geraniales and the fam-

ilies traditionally associated with them in the En-

gler sequence. Several recent collections repre-

senting new species for Costa Rica's flora are

included here. However, the most important con-

tribution in these pages is that for the Erythrox-

ylaceae. Timothy Plowman's untimely death cut

short his monographic work in Erythroxylum, but

he was able to complete a number of floristic treat-

ments, and we have the benefit of his insights in

these works. We were fortunate to have so gifted,

friendly, and conscientious a botanist on our staff

at the Field Museum, and we greatly mourn his

loss.

Acknowledgments

Beginning in 1961, fieldwork in Central Amer-

ica by Field Museum staff and their associates in

Honduras and Costa Rica was supported in part

by grants from the National Science Foundation.

The most recent of those grants was DEB-8 1 03 1 84.

Collections made with this support together with

many other herbarium holdings provide the basis

for much of our descriptive and geographic infor-

mation.

The staff and facilities of the Natural History

section of the Museo Nacional de Costa Rica have

been a primary resource for this project and have

been most helpful in carrying out field programs

for more than two decades. Recent work by col-

lectors associated with the Museo Nacional, Duke

University, and the Missouri Botanical Garden

have greatly expanded our knowledge of Costa

Rica's rich flora. Pablo Sanchez, Jorge Gomez-

Laurito, Nelson Zamora, Barry Hammel, and

Beryl Simpson have been especially helpful in pro-

viding information regarding the families included

in this issue of the Rora. An anonymous reviewer

made many helpful suggestions. Sadly, the pub-

lication of this group of families was initiated by

Timothy Plowman's contribution of the Erythrox-

ylaceae during the last year of his life.

FLORA COSTARICENSIS

Family #97 Krameriaceae

Family #98 Oxalidaceae

Family #99 Geraniaceae

Family #100 Tropaeolaceae

Family #101 Linaceae

Family #101a Humiriaceae

Family #102 Erythroxylaceae

Family #103 Zygophyllaceae

KRAMERIACEAE

By William Burger and Beryl B. Simpson

REFERENCE— B. B. Simpson, Krameriaceae.

Flora Neotropica Monogr. 49: 1-108. 1989.

Perennial herbs, shrubs, or rarely small trees, usually

vestitured with single-celled hairs, hemiparasitic on the

roots of other plants; stipules absent. Leaves alternate or

fasciculate, simple in Central America (rarely trifolio-

late), petiolate or sessile, the leaf blades with entire mar-

gins. Inflorescences of single axillary flowers, terminal

racemes (by condensation of internodes and loss of leaves)

or open panicles, peduncles with 2 opposite foliaceous

bracts, pedicels variable. Flowers bisexual, bilaterally

symmetrical (zygomorphic), hypogynous, colorful parts

pink, rose, or purple, rarely reddish brown to yellow,

imbricate, deciduous, sepals (4-)5, free and petaloid,

showy, unequal with the 3 outer usually larger than the

inner sepals and the petals; petals (4-)5, unequal and

dimorphic, small, the 3 upper (adaxial) long-clawed,

united near the base in some species, the petal-laminae

small or absent, the 2 lower (abaxial) petals smaller,

thick, sessile, laterally flanking the ovary, glandular on

the dorsal surfaces (and called elaiophores); stamens

(3-)4, usually alternating with the 3 upper petals and

angled downward and outward, filaments thick, free or

united near the base or adnate to the bases of the adaxial

petals, anthers basifixed, dehiscing near the apex by

membranous pores, pollen 3-porate (3-colporate), the

pores often expanded equatorially; pistil 1 , of 2 united

carpels with one developing and the other vestigial, ova-

ry with 1 locule with 2 pendulous ovules from an axial

placenta near the top of the locule, style obliquely ter-

minal, stigma simple. Fruit 1 -seeded, dry and indehis-

cent, usually covered with spines, the spines often with

retrorse barbs; seeds globose with a smooth seed coat,

with a straight embryo and 2 thick cotyledons, endo-

sperm absent.

A monogeneric New World family with 1 7 spe-

cies ranging from the southwestern United States

(with disjunct populations in Florida and Geor-

gia), Mexico, and the West Indies southward to

northern Chile and Argentina. Major centers of

species diversity are in northern Mexico and cen-

tral and eastern Brazil. All the species are found

on sandy or rocky soils in arid or seasonally dry

climates. The family is not now of commercial

importance, but the roots have been used medic-

inally and as a source of yellow and reddish brown

dyes. TJie Krameriaceae were once thought to be

closely related to the Leguminosae, but a relation-

ship with the Polygalales is more likely.

Krameria Loefling

With the characters of the family (see above). The

genus is quite distinctive because of its zygomorphic

flowers, the unique configuration of calyx and corolla,

and its unusual fruit. The flowers are usually held erect

with the longitudinal plane of the open perianth vertical,

somewhat like those of Senna or Cassia (Leguminosae).

This orientation and the unusual morphology of the

flowers are part of an interesting pollination syndrome

in which the lower petals secrete lipids that are collected

by female Centris bees. The flora parts were misinter-

preted in the past; see the article by B. B. Simpson (1982)

and the monograph cited above.

Key to the Species of Krameria

1 a. Leaves with distinct petioles, leaf blades narrowly elliptic to lanceolate and gradually narrowed at

the base; flowers pinkish fading to white K. ixine

Ib. Leaves sessile or subsessile, leaf blades linear to linear-lanceolate, abruptly narrowed at the base;

flowers yellow and dusky rose K. revoluta

FIELDIANA: BOTANY, N.S., NO. 28, OCTOBER 31, 1991, PP. 1-43

Krameria ixine Loefl., Iter hispan. 195. 1758. K.

cuspidata K. Presl, Reliq. haenk. 2: 103. 1835.

Figure 1.

Small shrubs, erect to 1(-1.5) m tall, often many

branched and to 1 m broad, internodes l-15(-25) mm

long, leafy stems l-3(-4) mm thick, densely strigillose

with thin whitish or grayish hairs 0.2-0.8 mm long. Leaves

simple, petioles 3-7(-9) mm long, merging gradually with

the decurrent lamina-base, strigillose; leaf blades 8-20

(-33) mm long, 3-8 mm broad, narrowly oblong or nar-

rowly elliptic to narrowly obovate or lanceolate, acute

with a spinose tip 0.5-1.5 mm long, acute at the base

and decurrent on the petiole, obscurely palmately

3-veined, densely strigillose. Inflorescence in lateral or

terminal racemes, peduncles usually shorter than the

subtending leaves, with a pair of opposite bracts in the

middle, densely strigillose. Flowers ca. 8 mm long and

1 2 mm broad when dry, perianth rose-pink to deep red,

magenta or reddish brown, turning white with age, sepals

4-10 mm long, broadly oblong, densely whitish sericeous

on the outer surfaces; upper petals united in the lower

half; stamens didynamous with anthers ca. 1 mm long.

Fruit globose, body of the fruit 4-7 mm in diameter,

covered with short ( 1 mm) whitish hairs and longer (1.6-

4.6 mm) orange or reddish spines with thin retrorse bar-

bules distally.

Plants of seasonally deciduous vegetation along

the Pacific slope of Mexico and Central America.

Growing from near sea level to 400 m elevation

in Costa Rica (to 1000 m in Honduras and 1500

m in Guatemala). Flowers and fruits have been

collected during the wet season in Costa Rica (late

May to December). This species ranges from west-

ern Mexico (Sonora) to northwestern Costa Rica,

through most of the Greater and Lesser Antilles,

and into South America in northeastern Colom-

bia, northern Venezuela, and eastern Guyana.

Krameria ixine is recognized by its short shrub-

by habit, grayish puberulence, small, petiolate,

narrowly elliptic and spine-tipped leaves, unusual

pink flowers with the sepals turning white with

age, and distinctive rounded fruits with dense

whitish hairs and longer barbed spines. The spe-

cies is known only from northern Guanacaste

Province in Costa Rica, where it ranges over a

wider area than its congener, K. revoluta.

Krameria revoluta Berg, Bot. Zeitung (Berlin) 14:

751. 1856. Figure 1.

Small shrubs or herbs with woody bases, to 50 cm tall,

with many branches, woody stems 2-5 mm thick, dark

gray to blackish and glabrescent, with longitudinal fis-

sures or smooth, leafy internodes l-4(-8) mm long, 0.6-

2 mm thick, grayish white with appressed ascending

strigulose hairs 0.9-1.5 mm long. Leaves alternate, pale

grayish or grayish white, sessile or subsessile (petioles

less than 2 mm long if present); leaf blades 6-22 mm

long, 1 .2-2.8(-3.5) mm broad, linear to linear-lanceolate

(narrowly lanceolate), tapering gradually to the acute apex,

with a small gland I i kc tip or spine that dries dark, abrupt-

ly narrowed at the base, margins usually revolute when

dried, covered with white or pale grayish appressed-as-

cending strigose hairs ca. 0.5 mm long. Inflorescences

terminal racemes (but flowers may appear to be solitary

in distal leaf axis in early stages because of the leaflike

bracts), flowers borne on slender strigulose peduncles 4-

1 2 mm long, usually with 2 opposite leaflike linear bracts

in the middle, larger outer sepals 5-9 mm long and ca.

3 mm broad, appressed whitish sericeous on the outside,

petaloid (upper) petals ca. 6 mm long and 2.5 mm broad,

basally connate for 3.4—4 mm, blades yellow or dull red

to brownish red, glandular petals orbicular, 2-3 mm in

diameter; stamens ca. 4 mm long. Fruits 4-6 mm in

diameter (to 1 1 mm measuring the spines), globose to

ovoid, covered by short dense whitish hairs and slender

red and yellow spines to 3.5 mm long, spines with re-

trorse barbs.

Plants of seasonally very dry deciduous forest

and shrub communities, from 50 to 250 m ele-

vation in Costa Rica (to 1400 m elsewhere). Flow-

ers and fruits have been collected in September

and October in Costa Rica. The species flowers

from July through December elsewhere. The spe-

cies ranges along the Pacific side of middle Amer-

ica, from southern Mexico to northwesternmost

Costa Rica.

Krameria revoluta is recognized by its almost

linear alternate leaves covered with whitish hairs,

unusual yellow and dusky rose flowers, puberulent

fruit with barbate spines, and restriction to very

dry habitats. In Costa Rica the species is known

only from two collections made in Santa Rosa

National Park, Guanacaste (Salas & Poveda s.n.

[1974] CR, Callaway 485 CR).

OXALIDACEAE

By William Burger

REFERENCE— Alicia Lourteig, Oxalidaceae, in R.

E. Woodson, R. W. Schery et al., Flora of Panama,

Ann. Missouri Bot. Gard. 67: 823-850. 1980.

Herbs, rarely shrubs or small trees; stipules present

and adnate to the petiole or absent. Leaves alternate,

subopposite or closely clustered (sometimes all basal),

pinnately or palmately compound, often (in Oxalis) tri-

foliolate, rarely unifoliolate, often folding up at night,

petiolate, the leaf blades usually entire. Inflorescences

axillary or cauliflorous, usually cymes, sometimes in

compound racemiform or umbelliform panicles, some-

FIELDIANA: BOTANY

Krameria ixine

2 mm

5 mm

Krameria revoluta

FIG. 1 . Krameriaceae: two species of Krameria.

BURGER ET AL.: FLORA COSTARICENSIS

times densely clustered or reduced to solitary flowers.

Flowers bisexual, radially symmetrical (plant rarely with

apetalous and minute cleistogamous flowers), hypogy-

nous, often with trimorphic heterostyly, sepals 5, united

only near the base, imbricate (valvate), persisting in fruit;

petals 5, free or connate near the middle, contorted or

imbricate in bud, narrowed at the base (clawed), nectar

glands often present between petals and stamens and

alternating with the petals; stamens basically 10 in 2

whorls of 5 ( 1 5 in Hypseocharis), the outer stamens usu-

ally shorter and opposite the petals, all the stamens func-

tional or 2-5 not bearing anthers and staminodial, fila-

ments slender and united in the lower half or only at the

base, anthers ovoid, versatile, 2-thecous, dehiscing lon-

gitudinally and introrse; pistil 1, (3-)5-carpellate, ovary

(3-)5-locular and lobed, each locule with (l-)2-l 5 ovules

pendulous from axile placentae, styles 5 and free (1 in

Hypseocharis), persisting, stigmas bifid or bicapitate. Fruit

a 5-lobed capsule (rarely a berry), usually 5-locular, calyx

and style persisting, dehiscence loculicidal, seeds often

ejected by a basal aril (the elastic outer integument); seed

with fleshy endosperm (rarely endosperm absent), em-

bryo straight.

A family of six genera and about 950 species

found throughout the world in cold climates and

warm. The largest genus of the family is Oxalis,

with over 800 species worldwide. Averrhoa is

widely planted in the tropics for its fruit, and

Biophytum is pantropical. Hypseocharis is endem-

ic to the tropical Andes, but Boesewinkel has re-

cently suggested this genus be transferred to Ge-

raniaceae (see Acta Bot. Neerl. 37: 1 1 1-120. 1988).

The family is recognized by the compound leaves,

5 -parted flowers, obdiplostemonous stamens, pis-

til with usually five styles and five locules, and

loculicidal capsule or berry. The following treat-

ment is based largely on the herbarium annota-

tions and publications of Alicia Lourteig.

Key to the Genera of Oxalidaceae

1 a. Trees, leaves pinnate; fruit over 5 cm long, fleshy and edible; plants grown in gardens and parks . .

Averrhoa

1 b. Herbs or small (to 3 m) subshrubs; fruit a small dehiscent capsule; ornamental and wild plants . .

2a

2a. Leaves usually 3-foliolate; capsule dehiscing by longitudinal slits, carpels (mericarps) remaining

attached to the central axis of the fruit Oxalis

2b. Leaves long-pinnate with many small leaflets; capsules loculicidal in a radial form, carpels remaining

attached only at the base Biophytum

Averrhoa Linnaeus

Trees, usually small and widely branching, puberulent;

stipules minute or absent. Leaves alternate, compound

and imparipinnate, petiolate, leaflets short-petiolulate,

alternate or subopposite on the rachis, margins entire.

Inflorescences solitary and axillary or cauliflorous, of

cymes in racemiform panicles, bracts and bracteoles su-

bulate and caducous. Flowers bisexual and regular, het-

erostylous, sepals 5, imbricate, united near the base; pet-

als 5, free or united near the middle, contorted in bud;

stamens 10 in 2 series or 5 fertile and 5 staminodial;

ovary 5-lobed, each lobe with 1 locule and 2-7 ovules.

Fruit an elongate fleshy indehiscent berry, pendant at

maturity, usually with 5 longitudinal ridges or lobes;

seeds 2-several in each locule, endosperm fleshy.

A genus of only two species, probably native to

Southeast Asia. Both species are widely planted in

the tropics and subtropics. The fleshy, usually yel-

low or orange fruits have a pleasant but acidic

flavor and are used to make drinks or preserves

or are eaten raw.

Key to the Species of Averrhoa

la. Leaves with 6-15 leaflets, leaves evenly spaced along the distal branchlets; petals to 9 mm long,

fertile stamens 5; fruit longitudinally deeply 5-lobed; commonly planted A. carambola

1 b. Leaves with 14-40 leaflets, leaves clustered near the tips of branchlets; petals 10-20 mm long, fertile

stamens 10; fruit smooth or slightly lobed; less common . . .A. bilimbi

Averrhoa bilimbi L., Sp. PI. 428. 1753.

Trees to 1 5 m tall, trunks to 30 cm thick, branches

usually ascending, leafy branchlets ca. 7 mm thick, densely

brownish or yellowish puberulent. Leaves clustered at

the tips of branchlets, to 65 cm long, petioles 4-13(-17)

cm long, ca. 2.5 mm thick and expanded at the base,

densely puberulent; leaf blades 2.5-8(-15) cm long, 1.2-

3(-5) cm broad, oblong to ovate-oblong, asymmetrical,

FIELDIANA: BOTANY

acuminate at the apex, rounded to truncate at the base,

minutely puberulent above and below. Inflorescences

axillary or cauliflorous on aborted branchlets, racemi-

form or paniculate, 1 5-20-flowered, puberulent and

glandular, bracts ca. 4 mm long, bracteoles 1.5-2 mm

long, subulate, pedicels 4-20 mm long, articulate near

the middle. Flowers small, sepals 1.5-2.5 mm long, 1.5-

3 mm wide, ovate to elliptic (1 internal sepal oblong),

pubescence appressed and glandular on both surfaces;

petals 10-20 mm long, 2-5 mm wide, narrowly spa tu-

la to. reddish purple, caducous; stamens 10, all fertile,

biseriate with longer (10 mm) and shorter (4 mm) fila-

ments; pistil 7.5-12 mm long, ovary 4-7.5 mm long.

Fruits 5-10 cm long, ca. 3.5 cm in diameter, smooth or

longitudinally 5-lobed (circular to pentagonal in cross

section).

Averrhoa bilimbi has smaller, more sour fruits

than its congener, and the fruits are not so prom-

inently ridged. The fruits are used for preserves or

for flavoring. This species is not commonly plant-

ed in Central America. It is called mimbro and

tiriguro in Costa Rica.

Averrhoa carambola L., Sp. PI. 428. 1753.

Trees to 10(-25) m tall, much branched, leafy branch-

lets 2-3 mm thick and minutely (0.1 mm) puberulent

with ascending thin whitish hairs, becoming glabrous

and brownish. Leaves separate along the distal twigs,

10-20 cm long, petioles 15-35 mm long, thickened near

the base and minutely puberulent; distal leaf blades (leaf-

lets) 5-8 cm long, 2-3 cm broad, ovate to ovate-oblong,

short-acuminate at the apex, truncate or rounded at the

base, proximal laminae 1.5-3 x 1-2 cm, minutely pu-

berulent on the major veins above and below. Inflores-

cences axillary or cauliflorous, to 8 cm long, of cymes

borne on racemiform panicles or fasciculate, peduncle

to 1 cm long, pedicels 3-4 mm long. Flowers ca. 8 mm

long, sepals 2.5-3.5 mm long, 1-2 mm wide, oblong;

petals 6-9 mm long, 1 .5-3 mm broad, spatulate, connate

at the middle, papillate-glandular within, violet to purple

and white; fertile stamens 5 with longer (2-2.5 mm) fil-

aments, 5 inner stamens/staminodes with shorter fila-

ments and usually without anthers; pistil 3-4 mm long,

cylindrical-ellipsoid. Fruits 8-13 cm long, 5-6 cm thick,

elliptic-oblong to ovoid, with 5 prominent longitudinal

ridges and grooves (strongly pentagonal or star-shaped

in cross section), yellowish translucent, mildly acidic to

sweet.

Averrhoa carambola is widely planted, both as

an ornamental and for its value as a fruit tree. The

acidic fruits are eaten fresh and are used in salads

and desserts. The glabrous, 5-ribbed, lustrous yel-

low fruits are distinctive. The fruit is often called

carambola, both in Spanish and English.

Biophytum DeCandolle

Herbs or subshrubs, sometimes woody at the base,

erect or decumbent, stems usually bearing a ring of stiff

retrorse hairs at the apex; stipules obscure. Leaves fas-

ciculate at the apex of the stem or at ground level on the

rootstock, pi 11 na ic I > compound with many leaflets, short-

petiolate, paripinnate (the terminal leaflet has been re-

duced to a bristle), the leaf blades (leaflets) opposite, very

short petiolulate and articulate at the rachis, gradually

differing in shape from base to apex (along the rachis)

with the lower pairs reniform to broadly ovate and small-

er than the more distal leaflets, puberulent. Inflores-

cences solitary and axillary, cymose or few-flowered,

pseudo-umbellate, peduncles long, pedicels subtended

by the spiral and imbricate bracteoles. Flowers bisexual

and regular, 5-parted, sepals essentially free, imbricate,

scarious; petals coherent near the middle, white to yel-

low, orange, pink, or red; stamens 10, biseriate with 5

shorter and 5 longer filaments, connate near the base,

all fertile or sometimes the shorter without anthers; ovary

5-lobed and 5-locular, each locule with 2-6 ovules al-

ternating along 2 rows within each locule, styles 5, free,

each with an enlarged bifid subcapitate or papillate stig-

ma. Fruits capsular and covered by the persisting sepals,

obovoid to subglobose or ellipsoid, each valve (carpel)

1-6-seeded, loculicidally dehiscent, the valves opening

upward over the sepals in a 5-pointed star pattern and

remaining attached at their base (centrally); seeds ex-

pelled explosively from the outer integument, usually

tuberculate.

Biophytum is a pantropical genus of about 75

species, with one species in northern Central

America and five additional species in Panama.

Only one species has been recorded in Costa Rica,

but it seems likely that a second will be found in

Costa Rica. The plants are quite distinctive be-

cause of the relatively long, narrow leaves clus-

tered at the ends of unbranched stems, and the

many opposite little leaflets that change their shape

along the length of the leaf. The plants look like

miniature palms or tree ferns. The following ac-

count is based on the treatment for the Flora of

Panama by Lourteig (Ann. Missouri Bot. Gard.

67: 825-834. 1980).

Key to the Species of Biophytum

la. Middle leaflets usually oblong to slightly falcate in outline, the larger leaves usually with less than

22 pairs of leaflets (range: 9-26 pairs); cymes condensed and 2-4 mm broad B. dendroides

1 b. Middle leaflets usually slightly falcate in outline, the larger usually with more than 22 pairs of leaflets

(range: 13-34 pairs); cymes ca. 5 mm broad B. falcifolium

BURGER ET AL.: FLORA COSTARICENSIS

Biophytum dendroides (H.B.K.) DC., Prodr. 1 : 690.

1824. Oxalis dendroides H.B.K., Nov. gen. &

sp. 5: 194. 1822. Figure 2.

R hi/omatous herbs, with leaves from the apex of an

underground stem or from an erect or decumbent stem

to 15(-35) cm long and 0.5-3(-5) mm thick, stems un-

branched (rarely with 2-4 branches), puberulent with

retrorse appressed hairs but glabrescent, roots to 10 cm

long, fibrous. Leaves pseudoverticillate at the ends of

stems, pinnate with 9-18(-26) pairs of leaflets, petioles

1-4 cm long, leaflets sometimes overlapping, rachis 2-

10 cm long and hirsutulous, petiolules ca. 0.2 mm long;

proximal leaf blades (leaflets) 1-2.5 mm long, 1-2.5 mm

wide, asymetrically ovate to triangular, middle leaflets

5-10 mm long and 1.5-4 mm wide, subrectangular to

subfalcate, distal leaflets 6-1 1 mm long and 2.5-5 mm

wide, asymmetrically obovate, glabrous to sparsely hir-

sutulous on 1 or both surfaces, midvein with 8-18 as-

cending secondary veins on each side, a submarginal

vein present but obscure. Inflorescences from the axils

of the fasciculate leaves, with a distal cluster of flowers

but only 1 flower blooming at a time, peduncles 1-8 cm

long, minutely puberulent to hirsutulous, pilose beneath

the flowers, bracts subulate to 6 mm long, bracteoles 2-

7 mm long, 0.3-1.5 mm wide, linear to lanceolate, inner

bracteoles equaling or exceeding the pedicels, pedicels

1-5 mm long, articulate in the lower half, glabrous or

finely puberulent. Flowers whitish pink or lilac, sepals

5-7 mm long, 1-1.5 mm wide, lanceolate and acuminate

with a slender tip, glabrous or with ascending hairs,

densely hirsutulous at the base, with parallel raised ve-

nation; petals 6-10 mm long, spatulate to obovate, nar-

rowed at the base, slightly retuse at the apex; stamens

with filaments enlarged at the base, longer stamens 4.5-

5.5 mm long, shorter stamens 3-4 mm long; pistil mi-

crostylous or macrostylous, puberulent apically, stigmas

small and bifid. Fruits capsular, 2.5-4 mm long, oblong

to subglobose, yellowish to violet, puberulent near the

apex, valves 1-6-seeded, short pubescent within; seed

ca. 1.5 mm long, semiovoid to triquetrous, planar on

one side and convex on the other, longitudinally tuber-

culate.

Herbs of evergreen formations from near sea

level to 1 200 m elevation. This species ranges from

Mexico through Guatemala, Honduras, Nicara-

gua, and Panama to Ecuador; it has not been re-

corded from Costa Rica.

Biophytum dendroides is recognized by its un-

usual habit (resembling miniature palms or tree

ferns) with the verticillate long-pinnate leaves aris-

ing from the apex of a small, usually unbranched,

erect woody stem. The opposite asymmetrical sub-

sessile leaflets that change their size and shape

along the length of the rachis, and the long, thin,

few-flowered peduncles also help make these plants

distinctive. This species has fewer leaflets on the

mature leaves than does B. falcifolium. The ap-

parent absence of B. dendroides in Costa Rica is

puzzling.

Biophytum falcifolium Lourteig, Ann. Missouri

Bot. Gard. 67: 829. 1980. Figure 2.

Erect or decumbent herbs, erect stems 5-12 cm long

and branched only near the base, ca. 2 mm thick, puber-

ulent in early stages and drying blackish. Leaves pseu-

doverticillate at the end of the stem, 5-1 1(-15) cm long

and 7-14 mm broad, pinnate with 13-34 pairs of op-

posite or subopposite leaflets, the leaves linear-cuneate

in outline, petioles 4-11 mm long, ca. 0.5 mm thick,

rachis minutely puberulent above and hirsutulous be-

neath, petiolules 0-0.2 mm long, thick; proximal leaf

blades (leaflets) 3-6 mm long and 1.5-3 mm wide, ob-

long to rhombic-oblong and slightly falcate (curved for-

ward), central leaflets 5-1 1 mm long and 2-4 mm wide,

falcate and oblong-rhombic, the distal leaflets equal to

or shorter than the middle leaflets, usually with a short

(1-2 mm) filamentous mucro at the apex, broadly obtuse

to subtruncate at the base, glabrous to finely puberulent

above, puberulent beneath. Inflorescences 1-3, subses-

sile and subcapitate with the condensed cymes globose

to cylindrical, with 1-5 flowers in various stages of de-

velopment, pedicels 8-16 mm long, minutely puberu-

lent, bracts ca. 2.5 mm long, subulate, purplish, brac-

teoles 1.5-3 mm long, 1 -veined, keeled, spirally

overlapping and green, puberulent and glandular. Flow-

ers whitish, sepals 4.5-7 mm long, 0.7-1.3 mm wide,

linear-elliptic, 5-7 veined, densely glandular; petals 6-

10 mm long, oblong-spatulate, cream-white with pink

veins and yellowish near the base, narrowed at the base;

longer stamens 3-3.5 mm long and unequal, anthers ovoid

and cordate at the base, shorter stamens 1-1.5 mm long,

with sterile anthers or reduced to glands; pistil 3-4 mm

long, ovary pilose near the apex and 5-lobed, each locule

with 3-4 ovules, styles glandular-puberulent, styles ca.

1.5 mm long, stigma bifid. Fruits capsular, globose, ca.

3-5 mm long, shorter than the persisting calyx, glandular

and puberulent at the apex; mature seeds unknown.

Plants of wet evergreen forest formations from

about 300 to 1 200 m elevation. The species is only

known from Costa Rica and Panama.

Biophytum falcifolium is recognized by the larg-

er number of leaflets on each leaf, the narrower,

more falcate leaflets, and the whitish petals. The

species has only recently been collected in Costa

Rica (L. D. Gomez et al. 23843 CR, MO) from Rio

Uren, Limon Province, at an altitude of ca. 1000

m. This species is closely related to B. panamense

Lourteig, but that species has more oblong leaflets.

Oxalis Linnaeus

REFERENCES— Melinda Denton, a monograph of

Oxalis, section lonoxalis (Oxalidaceae) in North

America. Publ. Mus. Michigan State Univ. 4(10):

459-615. 1973. Alicia Lourteig, Oxalidaceae ex-

tra-austroamericanae. I. Oxalis L. Sectio Tham-

noxys Planchon. Phytologia 29: 449^71. 1975;

FIELDIANA: BOTANY

5mm

FIG. 2. Oxalidaceae: two species of Biophytwn.

BURGER ET AL.: FLORA COSTARICENSIS

II. Oxalis L. Sectio Corniculatae DC, Phytologia

42: 57-198. 1979.

Perennial herbs or subshrubs (rarely vines), stems rhi-

zomatous, bulbous, or aerial, herbaceous or woody, pu-

berulent or rarely glabrous; stipules present or absent,

sometimes glandlike or interpetiolar. Leaves alternate or

subopposite to pseudoverticillate, basal or cauline, pin-

nately or palmately 3-foliolate (rarely 1-foliolate or

4-foliolate), petiolate, leaf blades (leaflets) usually entire,

often obcordate. Inflorescences axillary, umbelliform

cymes or solitary flowers, bracts small, bracteoles 2.

Flowers usually showy, often heterostylous (rarely cleis-

togamous and reduced), sepals 5 (4), free or united near

the base, imbricate, persisting in fruit; petals 5 (4), free,

narrowed at the base, contorted in bud, caducous; sta-

mens 1 0, the outer shorter stamens opposite the petals

and alternating with the sepals, filaments slender, united

near the base; pistils heterostylous and bi- or trimorphic,

ovary 5-locular, with 1-15 ovules in each locule, styles

5 and free, stigmas capitate. Fruits cylindrical to oblong

or globose capsules, glabrous or puberulent, loculicidally

dehiscent, each locule with (l-)2-15 seeds, valves con-

nate to the central axis and persisting; seeds usually ovoid,

flattened on the sides, oblong to apiculate, testa char-

taceous, longitudinally ribbed to transversely striate or

sculptured and densely verrucate, external integument

fleshy and ariliform, breaking elastically and expelling

the ripe seed, endosperm fleshy.

A worldwide genus of more than 800 species in

both warm and cold environments. The trifolio-

late leaves that are narrowed to the base and, to-

gether, form an almost circular outline character-

ize many species. The leaves usually fold together

at night and in inclement weather; they are often

marked with red or purple. Species of the genus

are often cultivated as ornamentals. Oxalis tub-

erosa (Oca) is an important tuber crop in the high

Andes. Some species are called acedera and vin-

agrillo in Central America. The following work is

based on Lourteig's treatment in the Flora of Pan-

ama (Ann. Missouri Bot. Gard. 67: 835-850. 1980)

and includes three species not yet known to occur

in Costa Rica (O. dombei, O. microcarpa, and O.

tetraphylld).

Key to the Species of Oxalis

1 a. Leaves pinnately 3-foliolate, the distal leaflet borne on a petiole-like extension of the rachis, laminae

rounded apically, never deeply emarginate or obcordate distally 2a

Ib. Leaves digitately (palmately) 3- or 4-foliolate, all the leaflets arising from the apex of the petiole,

laminae usually emarginate to obcordate distally (except in O. rhombifolid) 5a

2a. Petals pinkish distally (white or yellowish at the base); terminal leaflets ovate to ovate-oblong

O. barrelieri

2b. Petals yellowish throughout 3a

3a. Leaflets with small white punctate cystoliths, leaflets broadly ovate to oblong-orbicular;

inflorescences often with foliaceous bracts; plants not yet collected in Costa Rica

O. dombei

3b. Leaflets lacking small white punctate cystoliths; inflorescences with small linear bracts; plants

known to grow in Costa Roca 4a

4a. Inflorescences umbelliform with 2-7 (-15) flowers; capsule ovoid to oblong; terminal leaflets

often narrowly ovate to narrowly triangular (lanceolate), bluntly obtuse to slightly emarginate

at the apex; sepals 2.5-5 mm long; a common species O. frutescens

4b. Inflorescences elongate, with 5-1 2(— 40) flowers; capsule globose; leaflets usually ovate-elliptic

to rhombic, rounded at the apex; sepals 2-3 mm long; a wide-ranging species, but not yet

collected in Costa Rica O. microcarpa

5a. Leaves and inflorescences arising from a single bulbous or woody rootstock, elongate woody or

herbaceous internodes absent, petioles and peduncles usually over 10 cm long; flowers pink to

lavender 6a

5b. Leaves and inflorescences arising from aerial or rhizomatous stems, elongate internodes present;

flowers yellow 9a

6a. Leaves and inflorescences arising from a short, rhizomatous woody stem covered with the

bases of old leaves; leaflets broad distally, rounded and deeply notched O. articulata

6b. Leaves and inflorescences arising from a basal bulb, bulb with many subulate scales and

fibrous roots at the base 7a

7a. Leaflets 4, usually with rounded divergent distal lobes O. tetraphylla

7b. Leaflets 3, deeply notched to slightly emarginate distally 8a

FIELDIANA: BOTANY

8a. Leaflets often broadly triangular with relatively straight margins, usually retuse to slightly

obcordate distally; sepals with distal brownish calli 0.3-1 mm long O. latifolia

8b. Leaflets rounded, broadly obovate in general outline; sepals with distal brownish calli 0.3-

0.5 mm long o. debilis

9a. Petiole base without adnate stipules; stems erect and often climbing, clambering, or shrublike, to

2.5 m tall; distal leaflets often rhombic to ovate-triangular and acute at the apex O. rhombifolia

9b. Petiole base with adnate stipules (these sometimes difficult to see among the hairs); herbaceous

plants rarely more than 50 cm tall; leaflets usually obovate with incised (notched) apex lOa

lOa. Stems slender and decumbent, outer cortex often splitting off in age; leaflets usually less than 1

cm long; capsules densely and minutely puberulent, sepals usually more than half the length of

the capsule; petals 8-14 mm long o. filiformis

lOb. Stems slender and decumbent to erect, outer cortex not splitting off; leaflets (0.5-)l-3 cm long

• • 1 la

11 a. Stipules with an abruptly truncated apex or the apex gradually merging with the petiole; distal

leaflets broadly obovate to triangular; capsules covered with minute hairs, persisting calyx ^ -Vi

as long as the capsule; petals 5-1 3 mm long O. corniculata

lib. Stipules with a short acute apex (often difficult to see among the hairs); distal leaflets narrowly

obovate to obovate-oblong with a notch (ca. 3 mm deep in large leaflets); capsule with few erect

multicellular hairs, persisting sepals equaling the capsule in length; petals 1 1-18 mm long

O. spiralis

Oxalis articulate Savigny, in Lamarck, Encycl. 4:

686-687. 1797.

Herbs, to 40 cm high, stem rhizomatous, to 15 cm

long and (l-)2 cm in diameter, covered distally with the

short bases of leaves and their adnate stipules, roots

tuberous, cylindrical, to more than 10 cm long; stipules

4-16 mm long and adnate to the leaf-base. Leaves fas-

ciculate from the apex of the stem, 5-15(-30) cm long,

erect or ascending, petioles (4-)9-18(-30) cm long, slen-

der, pubescent to glabrate with thin hairs ca. 0.5 mm

long, petiolules 0.5-1 mm long, thick, pubescent; leaf

blades (leaflets) 5-20(-30) mm long, 7-35(-45) mm wide,

obcordate and widely obovate in outline, rounded dis-

tally and emarginate to retuse at the apex, cuneate at the

base, usually with appressed ascending thin whitish hairs

ca. 0.7 mm long on both surfaces, with dark or orange

punctations or lines (calli) 0.2-0.5 mm long near the

margins (less often scattered over the surface). Inflores-

cences umbelliform and cymose with 6-20 flowers, pe-

duncles usually longer than the leaves (2.5-40 cm), pu-

bescence similar to the petioles, bracts and bracteoles

0.5-3 mm long, lanceolate, pedicels l-3(-5.5) cm long,

slender and articulate at the base. Flowers violaceous to

pink, sepals 2-5(-6.5) mm long, 1-2 mm broad, narrowly

elliptic to lanceolate, appressed puberulent with thin

straight ascending hairs; petals 5-15(-20) mm long, ob-

ovate, connate near the middle, areas exposed in bud

covered with appressed whitish hairs; longer stamens ca.

3.5 mm long, shorter stamens ca. 2 mm long, united for

more than half their length; pistil ca. 5.5 mm long, ovary

usually pilose distally, each locule with 4-8 ovules, styles

puberulent, the flower with long, medium or short styles,

stigma small and capitate. Fruits cylindrical to oblong,

8-1 1 mm long, acute at the apex, with ascending ap-

pressed hairs distally on the carpels or all over, the per-

sisting calyx ca. lh as long; seeds 1-1.3 mm long, ovoid

to ellipsoid and acute at both ends, with 8 or 9 ribs and

4—8 pits, pale brown.

A cultivated species originally from Uruguay

and adjacent areas, escaping and becoming natu-

ralized in some areas. Oxalis articulata is recog-

nized by the short, thick stems bearing the closely

clustered leaves and inflorescences, the pink to

violaceous flowers, and the rounded leaflets punc-

tate along the margin.

Oxalis barrelieri L., Sp. PI. ed. 2, 624. 1762. Fig-

ure 4.

Herbs or subshrubs, to 1 m tall, leafy internodes

(0-)0.5-8 cm long, 0.5-3 mm thick, sparsely puberulent

with thin whitish hairs, glabrescent, roots fibrous; stip-

ules linear and ca. 3 mm long or absent. Leaves alternate

to subopposite, petioles 1-4 cm long, ascending, cana-

liculate, sparsely puberulent with thin hairs, more dense-

ly puberulent and articulate near the base, the rachis 3-

15 mm long, petiolules 0.5-1 mm long, with thin hairs;

leaf blades (leaflets) 10-55 mm long, 8-25 mm broad,

ovate to elliptic-oblong (rarely suborbicular), bluntly ob-

tuse to acute at the apex, obtuse to rounded at the base,

asymmetric at the base in lateral leaflets, sparsely pu-

berulent above and below. Inflorescences usually axillary

cymes longer than the leaves, (l-)3-l l(-30)-flowered,

peduncles to 6.5 cm long, 0.5-0.8 mm thick (dry), gla-

brescent, terminated by a flower and with 2 unbranched

secondary branches to 3 cm long, bracts and bracteoles

1-2 mm long, pedicels 1.5-3.5(-5) mm long, articulate

above the base. Flowers pink, sepals greenish, 3—4 mm

long, 1-2 mm wide, linear to narrowly oblong, acute and

minutely mucronate, sparsely and minutely puberulent;

petals 7-9 mm long, ca. 3.5 mm broad, retuse, obovate

to spatulate, pink distally, white or yellow in the lower

half; longer stamens ca. 3 mm long, I igulate at the middle,

BURGER ET AL.: FLORA COSTARICENSIS

O. corniculata

O. latifolia

5mm

FIG. 3. Oxalidaceae: four smaller species of Oxalis.

10

FIELDIANA: BOTANY

shorter stamens 1.5-2 mm long; pistil 3.5-4 mm long,

ovary glabrous and ovoid, each locule with 3 or 4 ovules,

styles puberulent, stigma bifid and subcapitate. Fruits 5-

9 mm long, to 4 mm thick, ovoid, persisting calyx half

as long; seeds 1.5-2 mm long, ca. 1 mm broad, ovoid

with slightly flattened sides, with 6-8 prominent trans-

verse ridges/grooves, and minutely tuberculate, brown.

Plants of the evergreen lowland Caribbean forest

formations from near sea level to 1 000 m elevation

(rarely from as high as 2000 m in the Chiriqui

highlands); flowering collections have been made

throughout the year. This species is native to trop-

ical and subtropical America and has been intro-

duced into the Old World tropics.

Oxalis barrelieri is recognized by its erect, slight-

ly woody stems, pinnately trifoliolate leaves (ter-

minal leaflet borne on a slender rachis), leaflets

acute to rounded distally (neither retuse nor emar-

ginate), and pink flowers.

Oxalis corniculata L., Sp. PI. 435. 1753. Figure 3.

Annual herbs, much branched and with prostrate or

decumbent stems to 50 cm long (rarely with erect stems

to 40 cm tall), often rooting at the nodes, internodes

variable in length (0-)l-4 cm long, 0.3-1.3 mm thick

(dry), usually with a few slender whitish hairs ca. 0.5-1

mm long, roots usually fibrous; stipules 0.8-3 mm long

and adnate to the petiole-base, l-2(-3) mm wide and

truncated at the apex, ciliate along the margin. Leaves

alternate to subopposite or often fasciculate, petioles 1-

5(-7) cm long, slender and ascending, with thin ascend-

ing whitish hairs, petiolules ca. 1 mm long, thick and

pubescent; leaf blades (leaflets) 4-12(-20) mm long, 5-

20 mm broad, broadly obovate and usually wider than

long, rounded and retuse to deeply emarginate distally

(obcordate) with a notch to 6 mm deep in large leaflets,

cuneate to obtuse at the base, margin entire and ciliate,

the pubescence variable with sparse to dense slender

ascending or appressed hairs 0.5-1 mm long (often gla-

brous above). Inflorescences cymose or umbelliform,

( l-)2-7-flowered, peduncles to 2-7(-2 1 ) cm long, slender

with thin appressed hairs, bracts and bracteoles 0.5-4

mm long, linear to lanceolate, pedicels slender, ca. 10

mm long (lengthening in fruit). Flowers yellowish, sepals

2.5-5.5 mm long, elliptic-oblong to narrowly ovate (lan-

ceolate), acute at the apex, translucent, lacking calli; pet-

als 5-13 mm long, 1-3 mm broad, often pinkish or

whitish within, yellow distally; longer stamens 4-6 mm

long, shorter stamens 3.5-4 mm long and united in the

lower third; pistils micro-, meso-, or macrostylous, to 8

mm long, ovary oblong and acute, each locule with 2-15

ovules, stigma 2-lobed and papillose. Fruits 5-25 mm

long, 1-3 mm thick, subcylindrical and 5-ribbed, acute

apically, minutely puberulent or with multicellular glan-

dular hairs, persisting sepals !4-'/3 the length of the fruit;

seeds ca. 1 mm long, ovoid but flattened on the sides,

apiculate at the apex, with 6 longitudinal ribs (3 on each

side) and prominent transverse ridges, brown or reddish

brown.

A species originating in the Mediterranean area

but now distributed worldwide. In Central Amer-

ica it is primarily found between 1000 and 2000

m elevation in open secondary vegetation, flow-

ering throughout the year. The species is weedy

and capable of hybridizing with indigenous spe-

cies.

Oxalis corniculata is recognized by its small

stature, trifoliolate leaves with laminae rounded

and notched distally, unusual stipules, and yellow

petals. The flowers usually open in the morning

and close by the middle of the day. This species

represents a complex of forms that have been

treated as three subspecies and several varieties

by Lourteig. It has been called acederilla in Costa

Rica.

Oxalis debilis H.B.K., Nov. gen. sp. 5: 236, t. 466.

1822. 0. martianaZuccarini, Denkschr. Konigl.

Akad. Wiss. Munchen 9: 144. 1825. Figure 3.

Herbs, arising from a globose, ovoid, or oblong bulb

1-2 cm long, the bulb covered with scales and leaf- bases,

the scales 7-13 mm long, 3-5 mm broad near the base,

lanceolate and with 3 longitudinal ribs, roots fibrous;

stipules to 13 mm long and 2 mm wide, completely

connate to the leaf-bases (difficult to see). Leaves trifo-

liolate, petioles to 30 cm long, 0.4-1.5 mm thick, with

thin transparent hairs 0.8-1 .5 mm long or glabrous, pet-

iolules 0.5-1 mm long; leaf blades (leaflets) 1.2-5.5 cm

long, 1.5-6 cm broad, broadly obovate to suborbicular

in outline, obcordate, broadly rounded distally and

emarginate to retuse with a notch 2-7 mm deep, glabrous

to sparsely puberulent above, more densely puberulent

beneath with thin hairs ca. 1 mm long, minutely (0.1

mm) dark punctate along the margin and more sparsely

scattered over the surface. Inflorescences bifid cymes

(sometimes twice bifid), with 6-15 flowers, to 30(-45)

cm long, peduncles similar to the petioles, bracteoles (at

the base of the pedicels) 1-3 mm long, often with brown-

ish calli, pedicels to 3 cm long, slender. Flowers pinkish

to red-violet, sepals 4-7 mm long, 0.7-1.5 mm wide,

narrowly ovate-elliptic or oblong, glabrescent, usually

with 2 brown or orange antherlike calli ca. 0.5 mm long

on the outside of the sepal apex; petals 2-3 times the

length of the sepals, obovate and narrowed at the base;

longer stamens 4.5 mm long, puberulent near the apex,

shorter stamens ca. 3 mm long and glabrous, connate in

the lower '/,; pistil ca. 7.5 mm long, ovary glabrous,

macro- or mesostylous (rarely microstylous), with scat-

tered thin hairs and glandular hairs, each locule with 2-

1 2 ovules. Fruits ca. 1 8 mm long with styles 2 mm long,

cylindric, calyx V4-Y^ the length of the capsule; seeds ovoid

with flattened sides, with 12 longitudinal ribs and 12

transverse ridges, brown.

A weedy species found in evergreen areas from near

sea level to 1 500 m elevation in our area. The species

is originally South American, but it is now widespread.

BURGER ET AL.: FLORA COSTARICENSIS

11

Oxalis debilis is recognized by its scaly bulbs, lack of

visible internodes, rounded trifoliolate leaves with shal-

low distal notches, pinkish flowers, and unusual brown-

ish calli at the tips of the sepals. These plants fruit in-

frequently, reproducing by bulblets.

Oxalis dombeii St. Hilaire, Fl. bras, merid. 1: 111.

1825. O. darienensis Woodson, Ann. Missouri

Bot. Gard. 27: 312-313. 1940.

Herbs, stems erect, to 60 cm tall, leafy internodes to

10 cm long, to 4(-6) mm thick and woody near the base,

glabrous to sparsely puberulent (sometimes more dense-

ly viscous-puberulent near the base), rhizome cylindri-

cal, roots fibrous; stipules obscure. Leaves alternate, sub-

opposite or pseudoverticillate, pinnately trifoliolate,

petioles 1-4 cm long, 0.2-0.4 mm thick (dry), articulate

near the base, glabrescent, canaliculate above, rachis 3-

6 mm long, petiolules ca. 0.5 mm long and drying dark;

leaf blades (leaflets) 9-25 mm long, 5-22 mm broad,

obovate to suborbicular, rounded at the apex, cuneate

to the base, lateral leaflets often asymmetric, essentially

glabrous above and below (fine ciliate on the margin),

with whitish minute (0. 1 mm) cystoliths usually visible

on the dried leaf. Inflorescences bifid-cymose (the pe-

duncle terminated by a flower and with 2 racemose

branches), 10-25-flowered, peduncle to 7 cm long, 0.5-

1.1 mm thick, glabrous, bracteoles 1-2 mm long, linear-

lanceolate, pedicels about 3 mm long, articulate near the

base, usually glabrous. Flowers yellow, sepals 4.5-7 mm

long, 1-2.5 mm broad, ovate-oblong, acute, thin, gla-

brous; petals 7-14 mm long, obovate to subspatulate;

longer stamens 2.5-4.5 mm long, minutely puberulent,

shorter stamens 1-2 mm long, glabrous; pistil 2-5 mm

long, ovary ca. 3 mm in diameter, each locule with 7-

10 ovules, stigma capitate. Fruits 10-12 mm long, ob-

long to ellipsoid, the persisting styles 1-2 mm long, sepals

V2-2/3 of the length of the capsule; seeds 1-1.2 mm long,

ca. 1 mm broad, obovoid and slightly flattened on the

2 sides, with 5 or 6 weakly developed longitudinal ribs,

the transverse ridges weakly developed, orange-brown.

Plants of drier deciduous forest vegetation and

in semideciduous areas, ranging from central Pan-

ama to Peru and in the Galapagos Islands.

Oxalis dombeii is recognized by its short, erect

habit, almost glabrous parts, pinnately trifoliolate

leaves, white punctate cystoliths, and yellow flow-

ers. Although not recorded from our area, it may

become introduced into the seasonally dry Pacific

lowlands.

Oxalis filiformis H.B.K., Nov. gen. sp. 5: 190, t.

460. 1822. O. bradei Knuth, Repert. Spec. Nov.

Regni Veg. 23: 276. 1927. Figure 3.

Short, creeping herbs to 10 cm tall, stems horizontal

or rhizomatous, to 50 cm long, internodes 0-5 cm long,

0.5-1 mm thick, dark brown or reddish, sparsely pu-

berulent, outer layers splitting off in age; stipules 1.5-3

mm long, 0.5-1.7 mm broad, adnate to the petiole, cil-

iate. Leaves alternate or fasciculate, palmately trifolio-

late, petioles 8-25 mm long, slender, sparsely to densely

puberulent with thin ascending whitish hairs, petiolules

ca. 0.5 mm long; leaf blades (leaflets) 2-10(-15) mm

long, 3-15(-20) mm broad, broadly obcordate to ob-

ovate and retuse distally, rounded at the notched apex,

cuneate at the base, both surfaces with thin, white, ap-

pressed hairs 0.5-1 mm long or the upper surface gla-

brous, ciliate along the edge. Inflorescences 1 -flowered,

l-4(-8) cm long and exceeding the leaves, peduncles

(measured to the bracteoles) about twice the length of

the pedicels, filiform and with thin hairs, bracteoles 1-

3 mm long, linear, pedicels 0.5-2.5 cm long. Flowers

yellow, sepals 3-5 mm long, 1-2 mm wide, oblong, ob-

tuse at the apex, usually with thin whitish hairs; petals

8-14 mm long, narrowly obovate; longer stamens 4-5.5

mm long, shorter stamens 3—4 mm long and connate to

V4 of their length; pistils 5-7 mm long, all longistylous,

each locule with 3-6 ovules, style densely puberulent,

stigmas bifid. Fruits 10-15 mm long, 2-3 mm thick,

densely minutely puberulent, calyx V2-2/3 as long as the

capsule, persistent styles 2-5 mm long; seeds 1-1.5 mm

long, broadly ellipsoid and flattened on 2 sides, reddish

brown, with 6 longitudinal ribs and transverse ridges.

Small plants of open sites in highland evergreen

areas from 1 1 00 to 2000 m elevation in Costa Rica

and Panama. Flowering and fruiting material has

been collected from December through May. The

species ranges from Costa Rica southward to Ec-

uador at elevations from 1000 to 3000 m.

Oxalis filiformis is recognized by its small stat-

ure, creeping stems with older exfoliating surfaces,

small obcordate leaflets, single-flowered inflores-

cences, and yellow petals. It appears to be native,

and it prefers open sunny habitats, such as road-

sides.

Oxalis frutescens L., Sp. PI. 435. 1753, subsp. an-

gustifolia (H.B.K.) Lourteig, Phytologia 29: 463-

471, fig. 3. 1975. O. angustifolia H.B.K., Nov.

gen. sp. 5: 193. 1822. O. neaei DC., Prodr. 1:

690. 1 824. O. coccinea Woodson & Schery, Ann.

Missouri Bot. Gard. 28: 431. 1941. Figure 4.

Herbaceous subshrubs or small shrubs to 40 cm tall,

usually with several erect or ascending branches from a

short (2-10 cm), woody base, leafy internodes 0.8-2 mm

thick, with thin, ascending, straight or crooked whitish

hairs to 1 mm long (shorter and curved in age); stipules

obscure. Leaves alternate, subopposite or verticillate,

pinnately trifoliolate, petioles 1-4 cm long, ca. 0.3 mm

thick (dry), puberulent as the stems, rachis 2-5(-10) mm

long, petiolules minute; distal leaf blades (leaflets) 7-

23(-45) mm long, 3-12(-20) mm broad, about twice as

long as the lateral leaflets, narrowly ovate to narrowly

triangular or ovate-oblong, tapering to the apex and

emarginate (or slightly notched), rounded to obtuse (acute)

at the base, the smaller lateral leaflets usually more ob-

12

FIELDIANA: BOTANY

O. spiralis

ssp. vulcanicola

2mm

O. f rutescens

ssp. angustifolia

5mm

FIG. 4. Oxalidaceae: four larger species of Oxalis.

BURGER ET AL.: FLORA COSTARICENSIS

13

long and asymmetric, densely puberulent on both sur-

faces or glabrescent above. Inflorescences somewhat ex-

ceeding the leaves, cymes or umbelliform cymes, (l-)3-

7(-15)-flowered, peduncles to 3.5 cm long, ca. 0.3 mm

thick, puberulent, bracteoles ca. 1 mm long, pedicels 3-

6 mm long, articulate ca. 1 mm above the base. Flowers

yellow, sepals 2.5-5 mm long, ca. 1 mm broad, narrowly

oblong, carpels ciliate on the back, calyx '/2-3/4 the length

obovate, emarginate at the apex, narrowed to the base;

longer stamens ca. 3 mm long, shorter stamens ca. 1.5

mm long, connate to the lower !/3; pistil 3-5 mm long,

minutely pilose, each locule with up to 8 ovules, stigmas

bifid. Fruits 5-8 mm long, ca. 3 mm thick, ovoid to

oblong, carpels ciliate on the back, calyx '/2-% the length

of the capsule; seeds 1.5-2 mm long, ovoid and convex

on two sides, apiculate, with 6 or 8 longitudinal ribs and

10-12 transverse ridges, pale brown.

Herbs and subshrubs of open and shaded sites

in seasonally dry deciduous and partly deciduous

forest formations from near sea level to 800(-

?1 600) m elevation on the Pacific slope of northern

and central Costa Rica. Flowering material has

been collected throughout the year except October

and November. The species ranges from the Mex-

ican-U.S. border through Central America and the

West Indies to Argentina.

Oxalisfrutescens is recognized by the preference

for seasonally dry vegetation, herbaceous or

shrublike habit with a slightly woody base, pin-

nately trifoliolate leaves with the terminal leaflet

usually narrowed to the apex, and the yellow pet-

als. It is found in open savannas as well as partly

shaded sites.

(Kalis latifolia H.B.K., Nov. gen. sp. 237, t. 467.

1822. O. ramonensis Knuth, Notizbl. Bot. Gart.

Berlin-Dahlem 7: 313. 1919. O. chiriquensis

Woodson, Ann. Missouri Bot. Card. 24: 190.

1937. Figure 3.

Herbs, stems not apparent, leaves and inflorescences

from an ovoid or globose bulb to 25 mm long and 20

mm thick, with scales 7-1 5 mm long and 4-6 mm broad

near the base, conspicuously 3-veined and with a long

narrow tip; stipules connate to the petioles, often difficult

to see among the leaf-bases, translucent brownish, au-

riculate at the apex. Leaves closely clustered on the bulb,

erect or ascending, petioles 5-17(-25) cm long, 0.5-1.5

mm thick (dry), glabrous or with scattered thin hairs,

petiolules ca. 1 mm long, thick; leaf blades (leaflets) 10-

25(-45) mm long, 15-45(-75) mm broad, broadly ob-

cordate to triangular in outline, the distal margin round-

ed and emarginate, slightly notched or deeply V-shaped

with divergent lobes, cuneate to obtuse with straight sides

at the base, glabrous or rarely with few hairs and cilia.

Inflorescences exceeding the leaves in length, erect, to

35 cm long, umbelliform cymes with 5-20 flowers, pe-

duncles 1 0-30 cm long, similar to the petioles, bracteoles

1-2 mm long, at the base of the umbel, pedicels 14-28(-

40) mm long, filiform, glabrous. Flowers pink to purplish

or lavender distally, sepals 3-6 mm long, 0.5-1.5 mm

wide, narrowly oblong, thin translucent, glabrous, usu-

ally with 2 brown or reddish calli 0.3-1 mm long near

the tip (the calli sometimes resembling anthers); petals

10-20 mm long, obovate to spatulate; longer stamens 4-

7 mm long, shorter stamens 3-4 mm long and connate

near the base; pistils 4-6(-8) mm long, usually microsty-

lous, each locule with 4-8 ovules, stigmas bifid and pa-

pillose. Fruits 4-9 mm long, oblong and acute, glabrous,

calyx '/4-'/2 the length of the capsule; seeds ca. 1 mm long,

ellipsoid and slightly lenticular, with 8-9 longitudinal

ribs and 9-12 transverse ridges, pale brown.

Plants of moister situations in evergreen and

partly deciduous vegetation from 1000 to

2000(-2700) m elevation. Most of our flowering

collections have been made from May through

August. The species ranges from Mexico and the

Antilles southward along the Andes to Bolivia.

Oxalis latifolia is recognized by the bulbous base

(without stems or internodes), largely glabrous

parts, long peduncles, palmately trifoliolate leaf-

lets with triangular or V-shaped form, and pink-

lavender petals. Denton interpreted O. latifolia

more narrowly in her monograph (see reference at

the beginning of the genus). Central American ma-

terial here considered O. latifolia was placed under

O. galeotti Turcz. in that monograph.

Oxalis microcarpa Benth., PI. hartw. 115. 1839.

Herbs or subshrubs, to 0.7(-1.2) m tall, internodes 2-

1 6 mm long, ca. 1 mm thick, with thin, straight or curved

whitish hairs 0.2-0.5 mm long, stems becoming slightly

woody, 2-3.5 mm thick, gray and glabrescent; stipules

obscure. Leaves alternate to subopposite, pinnately tri-

foliate, petioles 2-4 cm long, filiform (only 0.2-0.3 mm

thick when dry), puberulent, slightly thickened and ar-

ticulate at the base, rachis 4-10 mm long, petiolules ca.

0.7 mm long, thick and puberulent; terminal leaf blades

(leaflets) 10-24(-38) mm long, 6-14(-20) mm wide, ovate-

elliptic to somewhat rhombic or elliptic, tapering to the

rounded apex, obtuse to slightly rounded at the base,

lateral leaflets smaller and slightly asymmetric at the

base, sparsely to densely puberulent with thin appressed-

ascending hairs 0.4-0.9 mm long on both surfaces. In-

florescences solitary and axillary, to 12 cm long in fruit,

bifid cymes with 5-12(-40) flowers, peduncles to 6 cm

long, filiform, the 2 branches to 6 cm long in fruit, brac-

teoles linear, pedicels 1.5-3 mm long, slender, articulate

near the base. Flowers yellow, sepals 2-3 mm long, 0.5-

1 mm broad, narrowly ovate to elliptic, acuminate, gla-

brous; petals 5-7 mm long, obovate to subspatulate;

longer stamens ca. 2.5 mm long, shorter stamens 1.5-2

mm long; pistil ca. 2.5 mm long, ovary glabrous, each

locule with 1 or 2 ovules, stigmas slender. Fruits 2-3

mm long, oblate to subglobose, glabrous, calyx usually

equaling or slightly exceeding the capsule, walls of the

14

FIELDIANA: BOTANY

capsule thin; seeds ca. 1.5 long and 1 mm thick, ellipsoid,

with 6-8 prominent longitudinal ribs and transverse

ridges.

A species of seasonally dry areas of the Pacific

slope in Central America, to 1500 m elevation,

and flowering in the wet season. The species has

not been reported from Costa Rica, but is found

in Mexico, Honduras, and from Panama to Ec-

uador.

Oxalis microcarpa is recognized by the small

stature, pinnately trifoliolate leaflets that are

rounded distally, glabrous yellow flowers, and

short, few-seeded capsules. The unusual distri-

bution may imply that the species is not native to

Central America. This species resembles O. fru-

tescens.

Oxalis rhombifolia Jacquin, Oxalis 22, pi. 2. 1 794.

O. maxonii Standley, J. Wash. Acad. Sci. 17:

311-312. 1927. Figure 4.

Shrubs, erect or climbing over others, to 2(-6) m high,

internodes (l-)3-15(-40) cm long along the main

branches, 1-3 mm thick, puberulent with thin curved

hairs, dark brown; stipules obscure. Leaves alternate or

fasciculate on short shoots, palmately trifoliolate, peti-

oles 1-6 cm long, ca. 0.3 mm thick (dry), puberulent,

thickened and articulate at the base, petiolules ca. 1 mm

long, thick and puberulent; distal leaf blades (leaflets)

14-38(-^5) mm long, 10-20(-25) mm broad, lateral leaf-

lets slightly shorter, rhombic-ovate to ovate-elliptic, ta-

pering to the obtuse or acute apex, acute to obtuse at the

base, glabrescent on both surfaces or ciliate-puberulent

with thin hairs ca. 0.5 mm long. Inflorescences solitary

and axillary, bifid cymes to 9 cm long, 3-5 -flowered,

peduncles to 6 cm long, resembling the filiform petioles,

bracts 3-7 mm long and linear, bracteoles 1.5-2 mm

long, pedicels 2-5 mm long, puberulent, articulate at the

base. Flowers yellow, sepals 4-6 mm long, 1-1 .5(-2) mm

wide, narrowly oblong, puberulent and densely ciliolate

at the tip; petals 10-18 mm long, obovate to subspatu-

late; longer stamens 5-8 mm long, shorter stamens 3-6

mm long, connate for V3 above the base; pistils macro-,

meso-, or microstylous, 6-10 mm long, ovary and styles

puberulent, each locule 3-ovulate, stigmas bifid. Fruits

6-9 mm long, oblong or ovoid, densely puberulent, calyx

'/2 as long as the capsule; seed ca. 2.2 mm long, lenticular,

with 13 or 14 longitudinal ribs.

Plants of open and partially shaded sites in ev-

ergreen forest formations from 1000 to 2300 m

elevation in our area. Flowering material has been

collected from December to August in Central

America. The species ranges from Mexico through

Central America to Colombia and Venezuela.

Oxalis rhombifolia is recognized by its shrublike

or clambering habit, palmately trifoliolate leaves

with somewhat rhombic leaflets, filiform petioles

and peduncles, few-flowered inflorescences, and

yellow petals. This species appears to be rare in

Costa Rica, though common in the Chiriqui high-

lands of Panama.

Oxalis spiralis Ruiz & Pavon ex G. Don, Gen.

hist. 1: 755. 1831. O. vulcanicola J. D. Smith,

Bot. Gaz. 23: 241. 1897. O. spiralis subsp. vul-

canicola (J. D. Smith) Lourteig, Ann. Missouri

Bot. Card. 67: 846. 1980. Figure 4.

Herbs, scandent, decumbent, or erect to 50 cm tall or

1.5 m long, internodes 2-30(-80) mm long, succulent,

with thin yellowish to brown hairs 0.4-0.9 mm long,

often reddish in life; stipules adnate to the petiole, thin

and translucent, reddish, 2-5 mm long, narrow and acute

at the free apex. Leaves alternate, palmately trifoliolate,

petioles l-5(-7) cm long, 0.3-1.2 mm thick (dry), gla-

brous or puberulent, petiolules 0.5-1 mm long; distal

leaf blades (leaflets) 8-25(-38) mm long, 5-18(-25) mm

broad, oblong-obovate to suborbicular-obovate, with a

shallow (1-4 mm) distal notch and rounded distal lobes,

cuneate at the base, glabrous or sparsely puberulent above,

puberulent beneath with thin whitish appressed-ascend-

ing hairs ca. 0.5 mm long, laminae often purplish be-

neath. Inflorescences solitary and axillary or terminal,

exceeding the foliage in length, cymes with 3-17 flowers

or sometimes with only 1 or 2 flowers, peduncles 1.5-

9(-12) cm long, puberulent, bracteoles ca. 1 mm long,

linear, pedicels 5-12(-16) mm long, articulate above the

base. Flowers yellow, sepals 4-8 mm long, 0.7-1.5 mm

broad, narrowly lanceolate, gradually narrowed to the

acute apex, usually glabrous; petals 11-18 mm long, ob-

ovate, veins usually red or dark violet; longer stamens

4-6 mm long, shorter stamens 2-3 mm long and connate

for half their length; pistils micro-, meso-, and macrosty-

lous, 6 mm long, ovary glabrous, each locule with 3 or

4 ovules, stigmas capitate and minutely papillose. Fruits

4-8 mm long, ellipsoid-oblong, glabrous, calyx about the

same length as the capsule; seeds ca. 1.2 mm long, el-

lipsoid, somewhat muricate, without prominent longi-

tudinal ribs or transverse ridges, dark reddish brown.

Plants of evergreen montane forests and sub-

paramo formations from 1400 to 3300 m eleva-

tion. Flowering material has been collected in all

months but June; most flowering collections have

been made between November and March. The

species is found in the Cordillera de Talamanca,

around Volcan Irazu and, in a few collections, as

far west as the Alajuela-Heredia border (Vara

Blanca). The species ranges from southern Mexico

to Peru and is grown as an ornamental.

Oxalis spiralis is recognized by its reddish stems,

sparse (less often dense) puberulence, palmately

trifoliolate leaves with oblong-obovate leaflets

slightly notched distally, yellow petals with red-

dish veins, and seeds without ribs or ridges. Dif-

BURGER ET AL.: FLORA COSTARICENSIS

15

ferent plants may differ greatly in the size of their

leaves, probably as a result of environmental fac-

tors; some have terminal leaflets averaging only

10 mm long, others have terminal leaflets aver-

aging 30 mm. There is also great variation in the

degree of pubescence and growth pattern.

All our material belongs to subspecies vulcani-

cola, which is found in the highlands of Chiapas,

El Salvador, Costa Rica, and adjacent Panama.

This species can be grown in hanging pots; its pen-

dulous reddish stems and yellow flowers make an

attractive display.

(Kalis tetraphylla Cavanilles, Icon, descr. 3: 19-

20, t. 237. 1794. O. deppei Loddiges, Hot. cab.

15, t. 1500. 1828. O. hayi Knuth, Notizbl. Bot.

Gart. Berlin-Dahlem 7: 316. 1919.

Herbs, stems not apparent, leaves and inflorescences

from the apex of an ovoid to globose bulb to 4 cm long

and 3 cm in diameter, bulb covered with scales and leaf-

bases, outer scales 4-18 mm long, 1-12 mm wide, 3-5-

veined, sometimes lost during collection, inner scales

with 5-14 prominent veins, acute; stipules adnate to the

petioles, 12-30 mm long, 4-13 mm wide at the base,

thin or scarious. Leaves palmately 4-foliolate (rarely

3-foliolate in young plants), petioles 5-20(-30) cm long,

1-1.5 mm thick (dry), glabrous or sparsely puberulent,

pctiolulcsca. 1 mm long, puberulent; leaf blades (leaflets)

20-40(-70) mm long, 25-55(-75) mm broad, broadly

obovate to subtriangular (rarely suborbicular or with di-

vergent distal lobes), straight (truncate) to slightly con-

cave distally with rounded corners, cuneate to the base,

sparsely puberulent or glabrescent. Inflorescences to 50

cm tall, umbelliform or bifid cymes, 3-12-flowered, pe-

duncles 10-45 cm long, similar to the petioles, bracts

and bracteoles 2-4 mm long, pedicels to 17 mm long,

articulate at the base. Flowers pink to lavender or violet,

sepals 4-7.5 mm long, 1-2 mm broad, narrowly oblong,

bluntly acute at the apex and usually with 2 dark calli

ca. 0.5 mm long, glabrous or with a few hairs; petals to

3 cm long, usually glabrous; longer stamens 2.5-4 mm

long, shorter stamens 1.5-2.5 mm long, with filaments

connate '/i-'/2; pistils mostly macrostylous, 5-7 mm long,

each locule with 3-6 ovules, stigma bifid and papillose.

Fruits ca. 13 mm long, cylindrical, acute at the apex,

styles ca. 1.5 mm long; seeds ca. 1.2 mm long, ovoid-

lenticular, with 9-14 longitudinal ribs and 12-16 trans-

verse ridges, yellowish brown.

Plants escaped from cultivation in Panama and

growing wild; the species is a native of Mexico and

Guatemala at elevations from 1000 to 3000 m.

Oxalis tetraphylla is recognized by its bulbous

base, long-petiolate, palmately 4-foliolate leaves,

almost triangular rounded leaflets with slightly in-

dented distal margins, and pink to violet flowers.

This species appears to have become naturalized

in the Chiriqui Highlands of Panama, but it has

not yet been found in Costa Rica.

GERANIACEAE

By William Burger

REFERENCE— K. R. Robertson, The genera of

Geraniaceae in the southeastern United States. J.

Arnold Arbor. 53: 182-201. 1972.

Annual, biennial, or perennial herbs, subshrubs (rarely

small trees), stems sometimes succulent, often puberu-

lent with simple or gland-tipped hairs; rhizomes, tubers,

or thickened roots often present; stipules present and

paired at the leaf-base. Leaves alternate or opposite (and

often unequal at the node when opposite), basal or cau-

line, simple and lobed to deeply divided or pinnately or

palmately compound, petioles usually present; leaf blades

lobed or toothed to deeply sinuate (rarely entire), pal-

mately or pinnately veined. Inflorescences terminal or

axillary, generally cymose to umbellate, rarely with only

1 or 2 axillary flowers, pedicels usually subtended by

subulate stipulelike bracts. Flowers bisexual (rarely uni-

sexual), radially symmetrical (bilaterally symmetrical in

Pelargonium), hypogynous, perianth 5-parted, sepals free

or united at the base, imbricate (valvate), often awned

or with a narrow tip, persisting in fruit; petals 5 (rarely

4, 2, or 0), free, imbricate or convolute in bud, often

narrowed at the base and emarginate at the apex, ca-

ducous, usually with 5 small nectariferous glands alter-

nating with the petals; androecium usually with 10 sta-

mens and the outer opposite the petals (rarely with 1 5

stamens), all the stamens fertile or some reduced and

staminodial, filaments slender, united at the base, an-

thers attached at the center of the back (versatile),

2-thecous and dehiscing longitudinally, introrse; pistil 1 ,

of 3-5 (rarely 2 or 6) carpels united by their styles to a

central axis (column) and forming a "beak" on the ovary

and fruit, ovary superior and 3-5 -lobed, 3-5-loculate

with 2 ovules pendulous from axile placentae in each

locule, style present (rarely, absent), 5-parted near the

apex and with 5 slender ligulate stigmas. Fruit a schizo-

carp usually breaking up into 5 mericarps (carpel-bodies)

and separating from the persisting central column with

a part of the style forming an awn on the apex of the

mericarp, dehiscing (often explosively) by the separation

and contraction of the long stylar elements; seed smooth

or minutely reticulate, without endosperm, embryo usu-

ally curved.

A family of five genera and 700 species widely

distributed in temperate, subtropical, and mon-

tane tropical areas. Geranium and Erodium have

native New World species; Pelargonium of the Old

World has several widely cultivated ornamental

species. Erodium cicutarium has been used as a

forage plant, but has become a weed in many areas.

This family is closely related to Vivianiaceae of

South America and Dirachmaceae of Socotra, and

these taxa are sometimes included in a broader

circumscription of the Geraniaceae. The Oxali-

daceae are also closely related.

16

FIELDIANA: BOTANY

Key to the Genera of Geraniaceae Found in Central America

la. Leaves not deeply dissected, usually ovate to reniform and palmately veined; petals slightly unequal

with the upper 2 usually larger than the lower 3, 1 sepal calcarate at the base and with a spur

attached to the pedicel, lacking a disc or extrastaminal glands; cultivated ornamentals

Pelargonium

1 b. Leaves deeply dissected; petals all the same size and shape, sepals never calcarate and without an

adnate spur, a disc or extrastaminal glands present; wild or weedy (rarely cultivated) 2a

2a. Leaves palmately or radially dissected or lobed, usually (3-)5-7-parted or lobed, the lobes usually

dentate; flowers with 1 0 stamens and 1 0 anthers; styles simply coiled in fruit, glabrous on the inner

face; with 2 indigenous species Geranium

2b. Leaves pinnately divided, the pinnate divisions pinnately veined and lobed; flowers usually with

only 5 anthers; style spirally coiled in age and pubescent on the inner side; not yet collected in Costa

Rica . . . Erodium

Kr odium L'Heritier

Annual or perennial herbs, beginning as acaulescent

rosette plants and later with elongate stems, often with

several elongate stems arising from unbranched taproots;

stipules paired at each leaf. Leaves opposite or verticil-

late, basal and cauline leaves similar, pinnately com-

pound or deeply pinnately lobed (rarely simple or pal-

mately lobed), petioles short; leaf blades (leaflets) toothed,

lobed, or divided, sessile or petiolulate. Inflorescences

solitary and axillary, umbellate, the ultimate peduncles

usually with more than 2 flowers. Flowers bisexual (rare-

ly unisexual and the plants dioecious), radially or slightly

bilaterally symmetrical, sepals 5, free, often aristate-

awned or mucronate at the apex; petals 5, glabrous, equal

or subequal, pink to violet or purple, venation often

more darkly colored, sometimes with a dark spot at the

base of the upper 2 (or all) petals; androecium of 5 outer

staminodia or scales and 5 inner fertile stamens; pistil

with a puberulent ovary. Fruit a schizocarp, mostly de-

hiscing downward from the top, mericarps and their awns

becoming completely detached from the central axis,

mericarps tapering toward the base and usually with per-

sisting awns, awns puberulent on the inner (adaxial) sur-

face and the lower part coiling into a spiral when freed

from the central axis; seeds smooth.

A temperate zone genus of 60-80 species, cen-

tered in the area of North Africa and Mediterra-

nean Europe to central Asia, with some species in

Australia and southern South America. A few spe-

cies have become weedy throughout the world.

The elongate pinnately compound leaves with the

leaflets deeply pinnatisect are very distinctive. Ma-

terial of this genus has not been seen from Costa

Rica, but two species have become naturalized in

northern Central America at elevations from 1 500

to 3500 m. These two species, E. cicutarium (L.)

L'Her. ex Aiton and E. moschatum (Burm.f.)

L'Her. ex Aiton, can be distinguished by the fol-

lowing key.

Key to the Species of Erodium in Northern Central America

la. Leaflets deeply pinnatisect; tips of sepals with setiferous hairs, inflorescences lacking viscid hairs

E. cicutarium

Ib. Leaflets serrate or incised; tips of the sepals lacking setiferous hairs; inflorescences with viscid nairs

. E. moschatum

Geranium Linnaeus

REFERENCE— H. E. Moore, Jr., A revision of the

genus Geranium in Mexico and Central America.

Contr. Gray Herb. 146: 1-108. 1943.

Annual or perennial herbs (rarely small shrubs), stems

of 2 kinds: (1) herbaceous and leafy flowering stems with

slightly swollen nodes, and (2) thickened basal or un-

derground stems and rhizomes, often with dichotomous

branching, with thick taproots or adventitious roots;

stipules paired. Leaves alternate, opposite or closely con-

gested, often alternate in the inflorescences, cauline or

from the basal stem, simple but often deeply lobed or

palmately compound, petioles expanded at the base, long

in the basal leaves and shorter in the cauline leaves; leaf

blades mostly circular to pentagonal in outline, pal-

mately to radiately deeply cleft or lobed, the lobes usually

lobulate or incised. Inflorescences terminal or axillary,

BURGER ET AL.: FLORA COSTARICENSIS

17

often cymose and umbelliform, ultimate peduncles 1- or

2-flowered and with 2 or 4 stipulelike bracteoles, usually

long-pedunculate or long-pedicellate. Flowers bisexual

and radially symmetrical, small (5 mm) to large (40 mm),

sepals 5, free, imbricate, often with a narrow tip or awn

at the apex; petals 5, free, imbricate, puberulent at the

base and often emarginate at the apex, white to pink or

reddish purple, a disc and glands present; stamens 10 in

2 whorls, often of 2 lengths, filaments slender and con-

nate near the base, anthers versatile; pistil with a 5-lobed

ovary, often puberulent, with 2 ovules in each locule but

only 1 developing, the 5 styles united to the central col-

umn, stigmas 5. Fruit a schizocarp with a very unusual

mode of dehiscence, the basal carpels splitting into 5 (or

fewer) mericarps (carpel-bodies) and carried upward (of-

ten explosively) by the bending of the revolute styles,

which remain attached to the apex of the central column

(carpophore), styles and central column form the beak

on the pistil and developing fruit, mericarps rounded at

the base and with a persisting awn (part of the style)

distally; seeds usually ejected from the mericarps, surface

reticulate or smooth, with little or no endosperm.

A genus of 250-300 species widely distributed

in temperate, montane tropical, and polar regions

of the world. Major areas of diversification are the

mountains of Europe, the Mediterranean region,

and the major cordilleras of South America and

western North America. The genus is much in

need of a modern revision, both as regards the

delimitation of species and their relationships. Only

two species are recognized here, and these are like-

ly to become submerged within larger more broad-

ly defined species when the Neotropical taxa are

studied more carefully.

Geranium is a very distinctive genus with its

herbaceous habit, simple but palmately deeply

lobed leaves with incised margins, few-flowered

inflorescences, 5 -parted hypogynous flowers with

usually ten stamens, and beaked fruit with unusual

mode of dehiscence.

Key to the Species of Geranium in Costa Rica

la. Leaves essentially glabrous on the upper and lower surfaces, but often minutely ciliate along the

edges; short plants with stems covered with persisting and overlapping stipules, the distal internodes

not usually visible; sepals glabrous or ciliate along the edge G. costaricense

1 b. Leaves sparsely to densely puberulent on their surfaces; erect or decumbent plants with long, slender,

conspicuous distal internodes; sepals puberulent and often with gland-tipped hairs

G. guatemalensis

Geranium costaricense H. E. Moore, Gentes. Herb.

8: 253. 1951, based on G. cucullatum H.B.K.

var. multifidum Suessenguth, Bot. Jahrb. Syst.

72: 276. 1942 (non G. cucullatum L., nee G.

multifidum Sweet). Figure 5.

Perennial herbs, stems short (to 10 cm tall or 30 cm

long) and partly horizontal, 5-10 mm thick (including

the persisting stipules), arising from a vertical slightly

woody rootstock; stipules (6-)8-15 mm long, lanceolate

to subulate, thin, brownish and covering the stem. Leaves

closely clustered at the apex of the stem, petioles (l-)3-

10 cm long, 0.5-0.7 mm thick, expanded at the base and

united with the stipules, minutely (0.1-0.3 mm) puber-

ulent with whitish retrorse hairs, the hairs dense at the

apex of the petiole; leaf blades 1 2-20 mm long, 1 5-25

mm broad, rounded in outline, deeply divided, the pri-

mary divisions forming 3 or 5 parts that are also deeply

lobulate or divided, the secondary divisions usually with

3 prominent lobules and each leaf with as many as 25

distal lobules, acute and often reddish at the tips of the

lobules, glabrous and the major veins impressed above,

glabrous or ciliate along the distal margins, glabrous be-

neath. Inflorescences apparently solitary and axillary from

among the crowded leaves, 1 -flowered, peduncles (0.5-)

3-5 cm long (to the narrowly lanceolate paired bracte-

oles), bracteoles ca. 8 mm long, pedicels 1 .5-5.5 cm long,

sparsely and minutely puberulent, more densely retrorse

whitish puberulent beneath the perianth. Flowers ca. 10

mm long, to 24 mm broad, sepals 6-8 mm long, 2-3

mm broad at the base, narrowly ovate, with a short

narrow tip, green and glabrous but ciliate along the trans-

lucent margin and just beneath the tip; petals 9-15 mm

long, 5 mm wide, obovate, rounded or truncated distally,

white to pink with dark red or purple longitudinal lines;

stamens ca. 6 mm long; pistil 7-8 mm long. Fruits (12-)

14-17 mm long, united styles (beak) 0.8-1.2 mm thick,

free stigmas 1-2 mm long, mericarps (carpel bodies) 2-

2.5 mm long, sparsely puberulent.

Plants of evergreen higher montane forest for-

mations from 2600 to 3400 m elevation. Flowers

and fruits have been collected from January to

August. This species is only known from the Cor-

dillera de Talamanca; it has been collected from

the area near El Empalme to the area near Chirripo

Grande.

Geranium costaricense is recognized by its deep-

ly divided and many-lobed leaves, short stipule-

covered stems, and relative lack of pubescence.

This species may prove to be a subspecific element

of an Andean species when the genus is properly

revised. The smaller leaves with more narrow lobes

18

FIELDIANA: BOTANY

Geranium costancense

FIG. 5. Geraniaceae: two species of Geranium.

BURGER ET AL.: FLORA COSTARICENSIS

19

help distinguish this species from Geranium gua-

temalense.

Geranium guatemalense K nut h, /// Engler, Pflanz-

enr. 4(129): 200. 1912. G. repens H. E. Moore,

Contr. Gray Herb. 146: 78. 1943, based on G.

pulchrumC. V. Morton, Phytologia 1: 147. 1935,

non G. pulchrum N. E. Brown, 1895. Figure 5.

Perennial herbs, stems 1 0-80(-1 50) cm long, erect when

young, ascending to procumbent in age, lower stems ca.

4 mm thick, distal stems 0.5-1 mm thick, often rooting

at the nodes, internodes to 1 5 cm long, sparsely to dense-

ly puberulent with thin whitish (sometimes gland-tipped)

hairs ca. 0.5 mm long, roots fibrous to woody; stipules

5-10 mm long, subulate or lanceolate, with a slender tip

1-2 mm long, glabrous and ciliate to puberulent over

the surfaces. Basal leaves few to many, not long persist-

ing, with petioles 5-25 cm long and laminae deeply di-

vided into 3-7 nearly equal narrowly rhombic lobes;

cauline leaves with petioles (l-)2-8(-l 4) cm long, strigil-

lose to hirsutulous, the hairs more dense beneath the leaf

blades; cauline leaf blades (l-)2-5.5(-8) cm long, 2-6

(-1 1) cm broad, deltoid to pentagonal or hastate in out-

line, mostly with 3 or 5 major lobes (parts), distal lobes

often rhombic and conspicuously toothed or cleft above

the middle, lateral lobes broader than the distal but not

as long, usually drying darker above than below, upper

surface with slender ascending hairs paralleling the major

veins, lower surface densely hirsutulous on the veins and

surfaces beneath, the hairs thin, slightly curved or straight

and 0.4-0.8 mm long. Inflorescences solitary and axil-

lary, each peduncle usually 2-flowered, (l-)3-5(-10) cm

long, 0.3-0.5 mm thick (dry), bracteoles 2.5-4 mm long,

subulate, pedicels (3-)5-20(-35) mm long, with short

pilose and longer (0.7 mm) gland-tipped hairs more dense

beneath the flowers. Flowers 6-15 mm long and 8-20

mm broad, cam pan u late to rotate, sepals (4.5-)6-7(-8)

mm long (including a slender awn-tip 0.5-1 .5 mm long),

1.5-3 mm broad, narrowly oblong, usually with glan-

dular hairs ca. 0.7 mm long on the veins and margins;

petals (6-)8-14 mm long (3-)4-7 mm wide, narrowly

obovate, lilac to pale lavender, apex entire to deeply

notched; stamens with filaments shorter than the sepals;

stigmas ca. 2 mm long. Fruits 1 8-20 mm long, beak 0.8-

1 .5 mm thick, hispidulous or with thin gland- tipped hairs

0.5 mm long, mericarps 2.5-4 mm long and 1.5-2 mm

thick; seeds 2-3 mm long, dark brown and reticulate.

Herbs of evergreen higher montane forest for-

mations from (1100-)1700 to 3300 m elevation

in Costa Rica. Flowers and fruit have been col-

lected in all months of the year except March. The

species ranges from Guatemala to Panama.

Geranium guatemalense is recognized by its pu-

berulent flowers and foliage, deeply lobed leaves

with cleft distal lobes, and spreading stems with

slender internodes. The specimens placed under

this name include a number of different species

according to Moore's monograph, but the flowers

are remarkably alike and the distinctions of length

of the stylar beak in fruit seem insignificant. It

seems best to place this material under a single

name until the genus can be carefully revised.

However, there seem to be ecologically correlated

morphological distinctions among our material.

Specimens from higher elevations (above 2800

m) have smaller, more deeply divided leaves and

the leaf-divisions are more deeply incised or lobed.

Specimens from below 2200 m often have broader

leaves that are less deeply incised and are thinner

in texture. These differences can be quite striking

in some specimens, but it does not appear that

they are correlated with any consistent differences

in flowers or fruit. These differences appear to be

clinal and may be worthy of subspecific recogni-

tion. (Compare the differences between G. gua-

temalense and G. repens as outlined by MacBryde

in the Flora of Panama, Ann. Missouri Bot. Gard.

54: 202. 1967.) Also included here is material as-

cribed to G. mexicanum H.B.K. by Standley in

his Flora of Costa Rica (1937-1938, p. 560).

Pelargonium L'Heritier

Annual or perennial herbs to subshrubs or small shrubs,

stems usually thick, usually puberulent, often viscid and

aromatic; stipules various, often prominent. Leaves op-

posite or alternate, simple (rarely compound) and peti-

olate; leaf blades entire and lobed to crenate or serrate,

or palmately to pinnately lobed or incised, often fleshy,

venation palmate or pinnate. Inflorescences terminal,

axillary or leaf-opposed, 2-many-flowered pseudo-um-

bels (flowers rarely solitary). Flowers bilaterally sym-

metrical, sepals 5, free, imbricate, or valvate, unequal

and 1 forming a short spur or nectariferous tube which

is decurrent on the pedicel and adnate to it; petals 5(0-4),

free, imbricate, unequal, usually narrowed to the base,

a disc or extrastaminal glands absent; stamens 10 but

usually with only 5-7 bearing anthers and the others

sterile, filaments linear, united only at the base; pistil 1 ,

ovary 5-lobed and 5-loculed, each locule with 2 pen-

dulous ovules, styles forming a beak around a persistent

central column, puberulent within, stigmas 5, usually

filiform. Fruit a schizocarp, breaking into 5 basal 1 -seed-

ed mericarps, the mericarps acute at the base and round-

ed at the apex, thrown upward by the spirally coiled style,

which is often puberulent or feathery distally, the central

axis persisting; seeds oblong to ovoid, keeled and with

2 grooves, smooth or foveolate, endosperm absent.

A genus of over 200 species, nearly all from

South Africa, with a few from tropical Africa and

Australia. Many horticultural hybrids are culti-

vated as garden and potted plants.

20

FIELDIANA: BOTANY

Pelargonium x hortorum L. H. Bailey, Stand. Cycl.

Hort. 2531. 1916.

Succulent-stemmed herbs or subshrubs, to 1.5 m tall;

stipules broad and expanded at the leaf-base, to 20 mm

long and 15 mm broad. Leaves alternate or opposite,

petioles 2-20 cm long, puberulent; leaf blades 4-12 cm

long, 6-15 cm broad, ovate to rounded or reniform,

crenate-dentate or scalloped around the edge, cordate at

the base, often with zones of dark and light green on the

leaf surface, puberulent on both surfaces with slender

hairs. Flowers with nearly equal petals, double-flowered

and other forms sometimes grown, red to pink, salmon,

or white.

A popular ornamental of complex hybrid origin

used both for garden flower beds and as a pot plant;

represented by many varieties. The brightly col-

ored flowers, usually variegated foliage, and suc-

culent aromatic vegetative parts make this a very

distinctive plant. It is called geranio in Spanish

and the "garden geranium" in English. According

to Standley (Flora of Costa Rica, 1937-1938, p.

560), Pelargonium graveolens (Thunb.) L'Her. is

also grown in our area; it has deeply lobed and

crisped leaves.

lobes), inserted on the floral tube, equal or the lower 2

larger, imbricate in bud; petals 5 (or the lower 3 rarely

absent), free, imbricate, strongly narrowed to the base

(clawed), the upper 2 adaxial petals usually differing from

the 3 lower (abaxial) petals, the upper 2 often hairy at

the base, with entire to serrate or laciniate distal margins;

stamens 8 in 2 unequal whorls of 4, filaments free, slender

or winged at the base, anthers small, 2-thecous, opening

laterally by longitudinal slits, pollen tricolporate or di-

colporate; pistil 1 (3-carpellate), glabrous, ovary 3-lobed

and 3-locular, each locule with 1 pendulous ovule from

the apex of the locule, style slender, 3-parted distally,

stigmas simple. Fruits indehiscent, separating from the

central axis into 1 -seeded drupaceous or nutlike men-

carps (with a single samaroid mericarp in Magallana);

seed with a large straight embryo and 2 thick cotyledons,

endosperm absent.

A New World family of three genera. Tropaeo-

lum contains 86 species according to Sparre, rang-

ing from Mexico to Chile and southern Argentina.

The two other genera have three species and are

found only in southernmost South America; they

are very similar to Tropaeolum. Magallana pro-

duces winged samara-like carpels, and Trophaeas-

trum has erect peduncles and an almost regular

calyx with a very small spur.

TROPAEOLACEAE

By William Burger

REFERENCE— B. Sparre, Tropaeolaceae, in Flora

of Panama, Ann. Missouri Bot. Card. 62: 15-20,

1975.

Annual or perennial herbs, mostly climbers with long

twining petioles, stems often glabrous and semisuccu-

lent; stipules present and small or absent. Leaves alter-

nate (sometimes opposite near the base of the plant),

simple and peltate or subpeltate, the leaf blades pal-

mately (radiately) veined, often palmately lobed or an-

gled (to orbicular or reniform), margin entire. Inflores-

cences of solitary axillary long-pedunculate (pedicellate)

flowers, rarely umbellate or fasciculate, ebracteate in

nearly all species, bird- or insect-pollinated. Flowers bi-

sexual, bilaterally symmetrical (almost radially sym-

metrical in Trophaeastrum), showy and yellow to orange,

red, or dark purple, hypogynous to somewhat perigynous

(with a floral tube or hypanthium), with a backward-

projecting spur which has been interpreted as ( 1 ) an ex-

tension of the hypanthium or (2) part of the upper se-

pal(s) (the spur is little developed in Trophaeastrum),

sepals 5, free or united near the base (and with 5 calyx

Tropaeolum Linnaeus

Characters of the family (see above). The genus is

easily recognized because of its usually herbaceous twin-

ing habit with long twisted petioles, peltate or subpeltate

laminae, and solitary showy flowers with a conspicuous

backward oriented nectariferous spur.

Tropaeolum tuberosum R. & P. (anu) is an im-

portant tuber crop in the high Andes. Tropaeolum

majus, T. minus L., and T. peltophorum Bentham

and their hybrids are important ornamental plants

(the "garden nasturtiums") grown throughout the

world. Tropaeolum peregrinum L. (the "canary-

bird flower") is also an important garden orna-

mental.

The following treatment follows Sparre, but it

is possible that the native species recognized below

are in reality only different elements of a single

polymorphic species. More and better collections

of the rarer taxa (T. moritziana and T. pendulum)

are needed to assess variability within populations

and the validity of our present species concepts.

Key to the Species of Tropaeolum

la. Flowers 4-7 cm long, petals about the same size, much exceeding the sepals; petioles attached near

the center of the lamina; plants grown for ornament and rarely escaping T. majus

BURGER ET AL.: FLORA COSTARICENSIS

21

Ib. Flowers 2.5-3.5 cm long, petals unequal and only slightly exceeding the sepals; petioles attached

about halfway between the margin and the center of the lamina; wild climbers 2a

2a. Leaves with 5 major veins, the 3 distal major veins not bifid (except near the margin), the leaves

usually with 9 veins radiating from the petiole attachment; lower petals ciliate at the base of the

lamina (lateral sepals ca. 10 mm long) T. moritzianum

2b. Leaves usually with 3 major veins, the 2 lateral veins usually strongly bifid, the leaves usually with

7 or 8 veins radiating from the petiole attachment, usually with 3 major distal veins and 4 or 5

smaller veins; lower petals not ciliate at the base of the laminae 3a

3a. Distal edge of the petals deeply serrate to laciniate, rarely extending beyond the sepals, lateral sepals

ca. 6 mm long and 5 mm broad, outer parts of the flower usually orange to red; common plants . .

T. emarginatum

3b. Distal edge of the petals entire to serrate, usually extending slightly beyond the sepals, lateral sepals

ca. 10 mm long and 7 mm broad, outer parts of the flower usually yellowish; rarely collected plants

T. pendulum

Tropaeolum emarginatum Turcz., Bull. Soc. Imp.

Naturalistes Moscou 31: 425. 1858. T. guate-

malense Suess., Repert. Spec. Nov. Regni Veg.

51: 205. 1942. T. moritzianum sensu auctores,

non Klotsch.

Twining herbaceous climbers to 3 m high (to 8 m

long?) in shrubs and small trees, internodes to 20 cm

long, 0.5-3 mm thick (dry), minutely (0.3 mm) whitish

puberulent at the nodes and glabrous elsewhere; stipules

2-3 mm long and linear or absent. Leaves peltate, petiole

attached 3-12 mm from the proximal margin, petioles

4-8 cm long, 0.4-1 mm thick (dry), glabrous, often twist-

ed in the lower half; leaf blades ( 1 . 5-)3-8 cm long, ( 1 .8-)3-

9 cm broad, bluntly triangular (with rounded corners) to

ovate-triangular in outline, usually with a straight base

and 3 or 5 distal and lateral lobes, sinuses shallow (2-5

mm) and usually broadly rounded, apex obtusely round-

ed, sometimes with a small (0.5-2 mm) tip, drying mem-

branaceous to thin chartaceous, glabrous above and be-

low, often glaucous beneath, venation radiating from the

petiole attachment and with 3 major distal veins, the 2

lateral major veins usually parallel with the base of the

lamina and bifid in the proximal half. Inflorescence of

solitary axillary flowers, peduncles (= pedicels) 3-12 cm

long, filiform, pendant. Flowers 2.5-3.5 cm long, orange

to reddish orange on the outer parts, spur 12-24 mm

long, 1.5-3 mm in diameter, lateral sepals ca. 6 mm long

and 5 mm broad, corolla ca. 8 mm long and 4—7 mm

broad distally, fimbriate distally, yellow or marked with

purple; filaments ca. 7 mm long, anthers ca. 0.7 mm

long. Fruit a deeply lobed schizocarp 5-8 mm long, the

mcricarps to 8 mm long (measured parallel to the central

axis) and 1 2 mm broad, rectangular, fleshy.

Plants of open secondary sites in montane ev-

ergreen forest formations, from 1300 to 2800 m

elevation on the wet Caribbean slope in Costa Rica.

Probably flowering throughout the year. The spe-

cies ranges from Chiapas, Mexico, to Chiriqui,

Panama, and Colombia.

Tropaeolum emarginatum is recognized by its

slender, twining, almost glabrous stems, long

twisted petioles, peltate leaves with three major

veins, long-pendant showy flowers, and colorful

rearward-projecting nectariferous spur. The leaf

blades are usually broader than long with a straight

or convex base and with five broad distal and lat-

eral lobes. This species is very similar to T. moritzi-

anum, and the differences in the flowers are very

difficult to see in dried specimens. Also, the dis-

tinction between five major veins and three major

veins breaks down in some collections of T. emar-

ginatum where the lateral veins are strongly bifid

near the base (see the discussion under T. morit-

zianum).

Tropaeolum majus L., Sp. PL 345. 1753.

Small erect herbs or climbing annual vines, stems of-

ten to 5 mm thick and fleshy, internodes 1-15 cm long,

usually glabrous; stipules absent or obscure. Leaves pel-

tate with the petiole attached near the center of the lam-

ina, petioles 8-20 cm long, 0.5-2 mm thick, usually gla-

brous; leaf blades (3-)5-10 cm long, (3-)5-10 cm broad,

suborbicular with 5-7 rounded (and weakly defined) lobes,

often with 3 broadly rounded distal lobes and the sinuses

only 1-3 mm deep, drying thin chartaceous, glabrous

above, minutely (0. 1-0.2 mm) papillate puberulent be-

neath, venation with 5-7(-9) major veins radiating out-

ward from the peltate base. Inflorescence of solitary ax-

illary flowers, peduncles to 25 cm long, often equaling

the petioles, glabrous. Flowers 4-7 cm long, the spur 25-

35 mm long, sepals 13-18 mm long, 4-8 mm broad,

sepals and spur often yellowish green; petals subequal,

upper petals 3-4 cm long with a lamina 1 5-20 mm long

and equally broad and with a narrow clawed base 1 2-

15 mm long, bright orange to yellow or dark purple.

Fruits to 10 mm long, fleshy.

A species probably of hybrid origin and not

known in the wild, having originated in Peru. It

is now grown as an ornamental throughout the

22

FIELDIANA: BOTANY

world. It has been called capuchina, espuela del

gelan, mastuerzo, and "garden nasturtium." The

plants are sometimes used for medicinal purposes.

Tropaeolum moritzianum Klotzsch, Allg. Garten-

zeitung6: 241. 1838.

Herbaceous climbers to 10 m long, internodes to 25

cm long, 0.5-2.5 mm thick (dry), glabrous; stipules ab-

sent. Leaves peltate with the petiole attached about half-

way between the center and the basal margin, petioles

5-15 cm long, 0.3-1 mm thick, glabrous; leaf blades 3-

9 cm long, 2.5-8.5 cm broad, broadly ovate-triangular

to suborbicular, usually with a straight or rounded (con-

vex) base and 5 or 7 distal lobes, the lobes often weakly

denned with shallow (1-3 mm) sinuses, tips of the lobes

mucronate to emarginate, the leaf blades drying mem-

branaceous and usually glaucous beneath, glabrous above

and below, with usually 9 veins radiating from the petiole

attachment, with 5 major veins, the 3 major distal veins

unbranched and the 2 lateral veins often bifid. Inflores-

cences of solitary axillary flowers, peduncles 7-20 cm

long, ca. 0.5 mm thick, pendant, glabrous. Flowers ca.

3 cm long, glabrous, spur 20-25 mm long, straight or

slightly curved upward, sepals (calyx-lobes) 10-12 mm

long, 4-5 mm broad, broadly elliptic, reddish; upper

petals 8-9 mm long, 4-5 mm broad distally, lower petals

10-12 mm long, with a narrow ciliate base and distal

lamina ca. 7 mm long and 4-5.5 mm broad, the petals

yellow to orange or reddish (with darker purple veins)

and with a deeply serrate-ciliate distal margin. Fruits

becoming 10 mm long, triangular, and deeply ribbed

before separation of the mericarps.

Plants often clambering over shrubs in open for-

est edges in evergreen forest formations, from 1000

to 2000 m elevation. Flowering collections are few

and range from September to January in our area.

The species is said to range from Guatemala to

Colombia and Venezuela (but see below).

Tropaeolum moritzianum is recognized by the

climbing habit with long twisted petioles, weakly

lobed peltate leaves, laminae with five major veins,

lack of pubescence, and solitary pendant reddish

flowers with long nectariferous spurs. This species

seems to be very similar to both T. emarginatum

and T. pendulum, differing in leaf venation and

subtle characters of the flowers. It is possible that

all three are different forms of a single polymor-

phic species, for which T. moritzianum would be

the oldest name. It is highly likely that T. war-

scewiczii Buchenau (Bot. Jahrb. Syst. 26: 582.

1899), based on Warscewicz 2-15 from "Costa

Rica et Veragua" (not seen), is a synonym of this

species.

Tropaeolum pendulum Klotzsch, Allg. Gartenzeit-

ung 18: 377. 1850.

Annual herbaceous climbers to 5(-8) m long, inter-

nodes 3-18 cm long, 0.5-2.5 mm thick (dry), sparsely

puberulent at the nodes and glabrous elsewhere; stipules

small (0.5 mm) and caducous. Leaves peltate with the

petioles attached halfway between the center and margin

(4-14 mm from the edge), petioles 4-10 cm long, often

much twisted in the lower half, sparsely puberulent with

thin whitish hairs near the base or glabrous; leaf blades

3-7 cm long, 4-9 cm broad, pentagonal to very broadly

ovate-triangular (suborbicular), usually 5- or 3-lobed

distally, the lobes broadly rounded and separated by very

shallow (0-3 mm) sinuses, often short-mucronate with

tips 0.3-1.5 mm long, base straight to slightly concave

or rounded, lamina drying membranaceous and green-

ish, glabrous, major veins 3(-5) with the lateral veins

usually bifid. Inflorescences of solitary axillary flowers,

peduncles 2.5-5(-7) cm long, pendant, and filiform (dry),

glabrous. Flowers 2.5-3 cm long, yellowish, spur 12-16

mm long, 0.7-1.2 mm thick distally, greenish, lower se-

pals ca. 12 mm long and 3-4 mm broad, upper sepals

ca. 10 mm long and 6-8 mm broad; upper (larger) petals

8-12 mm long and exceeding the calyx-lobes, with a

narrow base ca. 5 mm long and expanded lamina to 8

mm broad, entire to slightly serrate on the distal petal

margins. Fruits 5-6 mm long, longitudinally ribbed (be-

fore separation of mericarps).

Rarely collected climbing plants of montane ev-

ergreen forest formations, from 1600 to 2300 m

elevation on the Pacific slope and along the central

highlands in Costa Rica. Flowering material has

been collected in July-September, December, and

January. The species ranges from Costa Rica to

Colombia and Ecuador.

Tropaeolum pendulum is distinguished by its

yellow flowers with greenish spurs, peltate leaves

with rounded, often mucronate lobes, long calyx-

lobes, and petals with entire to serrate distal mar-

gins. This species resembles the more common T.

emarginatum and T. moritzianum, but those spe-

cies tend to have deep orange flowers with fim-

briate petal margins that barely exceed the calyx-

lobes. The flowers of T. pendulum tend to dry

thinner, more whitish, and more translucent than

the other two species. However, there is the pos-

sibility that all three are part of a single polymor-

phic complex (see the discussion under T. moritzi-

anum). The original description of T. pendulum

was based on plants grown in Europe from seeds

originating in Costa Rica.

LINACEAE

By William Burger

REFERENCE— K. R. Robertson, The Linaceae in

the Southeastern United States. J. Arnold Arbor.

52: 649-665. 1971.

BURGER ET AL.: FLORA COSTARICENSIS

23

Herbs, rarely subshrubs or shrubs, stipules present and

small (sometimes glandlike or interpetiolar) or absent.

Leaves alternate or opposite (whorled), simple, sessile or

petiolate, leaf blades entire, often narrow. Inflorescences

cymose or paniculate, few-flowered to racemose or spi-

cate, axillary or terminal. Flowers bisexual, radially sym-

metrical, hypogynous, 5-parted (4-parted in Radiola),

sepals 5 (4), free or united near the base, imbricate, per-

sisting; petals 5 (4), free, contorted or convolute in bud,

narrowed to the base or clawed, fugacious, a disc absent;

stamens the same number as the petals, alternate with

the petals (opposite in Anisadenia), sometimes alternat-

ing with small staminodes, filaments united to form a

short tube or cup, anthers basifixed and 2-thecous, de-

hiscing by longitudinal slits, introrse; pistil 1 , composed

of (2-)3-5 united carpels, ovary superior, (2-)3-5-locular

or unilocular near the apex, with 2 axile pendulous ovules

in each locule, styles (2-)3-5, free or united below, stig-

mas capitate. Fruit a septicidal capsule, usually 3-5-

valved (of 2 indehiscent 1 -seeded mericarps in Anisa-

denia); seed with a usually straight embryo and with flat

cotyledons, endosperm oily, little, or none.

This family is now delimited more narrowly

than in the past, and only one genus is represented

in Central America. Cronquist (198 1) included six

genera within the family and estimated 220 spe-

cies. The largest genus by far is Linum, with about

200 species in temperate and subtropical areas of

the world. Woody genera formerly considered part

of the family have now been separated as Hugonia-

ceae (Old World tropics), Ixonanthaceae (five pan-

tropical genera), and Humiriaceae (see the follow-

ing family).

Linum Linnaeus

Annual or perennial herbs or small shrubs woody at

the base, usually glabrous; stipules absent or reduced to

Key to the Species of Linum

1 a. Petals yellow, ca. 6 mm long, sepals 2-3 mm long; rarely collected wild plants of higher (ca. 2000

m) elevations L. guatemalense

Ib. Petals blue or white, 10-15 mm long, sepals 5-7 mm long; plants cultivated for ornament, fiber, or

oil seeds . . L. usitatissimum

small paired glandlike structures. Leaves alternate, less

often opposite or in whorls of 4, simple, sessile or pet-

iolate, leaf blades elliptic or ovate to lanceolate or linear,

entire, pinnately veined. Inflorescences terminal or ax-

illary, racemose to corymbose, less often cymose to fas-

ciculate or in spicate heads, bracts present and with 2

dark glands at the basal margins. Flowers bisexual and

radially symmetrical, blue to white or yellow (red), hy-

pogynous, sepals 5, free or united at the base, lanceolate

to ovate, usually with long narrow apices, often with

gland-tipped teeth, with 1-5 veins and persisting in fruit;

petals 5, free or slightly united at the base, obovate and

narrowed at the base, with 5 small nectar-glands opposite

the petals and adnate to the staminal tube; stamens 5,

alternating with the petals, united at the base to form a

short tube or cup, sometimes with small filamentlike

staminodia on the edge of the tube, filaments linear,

anthers dehiscing introrsely by longitudinal slits; pistil

superior, ovary 5-locular or 10-locular by the formation

of false septa, styles 5, stigmas 5, capitate to clavate.

Fruits 5-valved or apparently 10-valved, 5-locular with

2 seeds in each locule or 10-locular with 1 seed in each

locule, usually dehiscing along the false septa or locules;

seed with a straight embryo and mucilaginous testa.

A genus of about 200 species in temperate and

subtropical regions, especially the Northern

Hemisphere. About 40 species are found in North

America, and 36 in the Mediterranean. Three spe-

cies are native to Guatemala, of which one ranges

southward into Honduras and Nicaragua and one

into Costa Rica. These slender, few-branched herbs

can be confused with Schwenkia of the Solanaceae

or Evolvulus of the Convolvulacea, but those gen-

era have corolla tubes.

Linum guatemalense Benth., Bot. voy. Sulphur 67.

1844.

Herbs 50-110 cm long, annual or perennial, erect,

branched at the base or distally (within the inflorescence),

internodes 1-3 mm thick, terete, glabrous or sparsely

and minutely (0.2 mm) puberulent, with slightly elevated

and slender longitudinal ridges. Leaves alternate and as-

cending (occasionally fasciculate on the short shoots),

petioles absent; leaf blades 14-30 mm long, 4-8 mm

broad, lanceolate to linear-lanceolate, acute at the apex,

contracted or slightly rounded at the base, drying mem-

branaceous and grayish brown, with 3-6 obscure sec-

ondary veins on each side, glabrous above and below.

Inflorescences terminal, 7-30 cm long, 2.5-1 5 cm broad,

racemose to corymbiform panicles, with slender alter-

nate lateral branches subtended by linear-lanceolate leaf-

like bracts 6-20 mm long, 0.7-4 mm broad, glabrous.

Flowers glabrous, sepals 2-3 mm long, 1.5-2 mm broad,

ovate with a sharp acuminate or acute apex, margin

occasionally with a few minute (0.2 mm) teeth; corolla

yellow; petals ca. 6 mm long. Fruits ca. 3 mm long and

3 mm broad, ovate with a conical apex, appearing to

24

FIELDIANA: BOTANY

split into 10 parts; seeds 1.3-1.5 mm long, 0.8-1 mm

broad, lenticular and lustrous brown.

Linum guatemalense was collected in 1 989 by

G. Hen-era and W. Gamboa (3606 CR, MO) at 1 900

m elevation in the Sabanas de Murur-Biruk, north

of Buenos Aires, southern Puntarenas Province.

This is the first record of the species south of Gua-

temala. The unbranched erect stems with broad

distal inflorescence, alternate sessile lanceolate

leaves, lack of pubescence, yellow flowers with five

separate sepals and petals, and 1 0-locular capsules

are characteristic.

Linum usitatissimum L., Sp. PI. 277. 1753.

Annual herbs, 20-80(-120) cm tall, single-stemmed

or branched only at the base or below the inflorescence,

stems slender, internodes 0.5-1.5 mm thick, glabrous;

stipules absent. Leaves alternate and sessile; leaf blades

6-20(-25) mm long, (0.5-)l-4 mm broad, linear to lan-

ceolate, glabrous, with 3 major veins from the base. In-

florescences distal corymbs or panicles, pedicels 8-14

mm long, slender and elongating in fruit. Flowers to 1.5

cm long, ca. 1.2 cm broad, sepals 5.5-7.5 mm long, ca.

2 mm broad, acuminate at the apex, inner sepals with

scarious or denticulate margins but without glandular

dentations; petals blue or white, 10-15 mm long; stigma

clavate. Fruits 7-10 mm long, ca. 8 mm in diameter,

pale yellowish brown, capsule slightly longer than the

persisting sepals, usually 1 0-locular with 5 thicker walls

and 5 thinner walls (false septa).

Linum usitatissimum is recognized by its slen-

der erect stems, narrow sessile leaves, usually bright

blue 5 -parted flowers, and capsular fruit with usu-

ally 1 0 locules. These plants are not known to be

naturalized in Costa Rica, though they are occa-

sionally cultivated. This species has been in cul-

tivation for more than 5,000 years in Europe and

western Asia. Today, special varieties are grown

for their oil seeds or their excellent fiber. Linseed

oil is used for paints, varnishes, and coatings as

well as an edible oil. Flax fiber is flexible, strong,

and durable and absorbs water, making it very

useful for towels and table coverings. The bright

blue flowers have made this species an ornamental

as well. This species is called linaza, linaza lino,

and lino in Spanish, and flax or linseed in English.

HUMIRIACEAE

By William Burger and Nelson Zamora

REFERENCES—!. Cuatrecasas, A taxonomic re-

vision of the Humiriaceae. Contr. U.S. Natl. Herb.

35: 25-214. 1961. A. H. Gentry, Humiriaceae, in

Flora of Panama, Ann. Missouri Bot. Gard. 62:

35^4, 1975.

Evergreen trees or shrubs, wood hard and often red-

dish; stipules absent or minute, paired, deciduous. Leaves

alternate, often distichous, simple petiolate or rarely ses-

sile; leaf blades often coriaceous, entire to slightly serrate,

pinnately veined, often glandular punctate near the mar-

gin on the undersurface. Inflorescences axillary or pseu-

doterminal (rarely terminal), paniculate, often corym-

biform with trichotomous, dichotomous, or alternate

branching, bracts and bracteoles small, persistent or de-

ciduous, pedicels short. Flowers bisexual, radially sym-

metrical, hypogynous, calyx 5-lobed, united to form a

cup or short tube, glabrous to puberulent, equal or the

outer 2 lobes smaller, imbricate in bud; petals 5, free,

oblong-lanceolate to oblong or linear, white to yellowish

(red), contorted or imbricate along the edges in bud (but

appearing to be valvate); stamens 10-30 in 1-2 series or

many and multiseriate, filaments united in the lower part

to form a short staminal tube, free filaments usually

alternating in length, sterile filaments (antherless stam-

inodia) often present, anthers dorsifixed or basifixed,

2-thecous and bilocular with longitudinal dehiscence, or

2- or 4-thecous and unilocular and dehiscing by detach-

ment (the 4 thecae separate on the connective with 2

proximal and 2 distal), connective usually thick and pro-

longed beyond the thecae, an intrastaminal disc sur-

rounding the ovary and cupular to tubular and dentate

to laciniate or of 10-20 free scales; pistil 1, composed

of (4-)5(-7) united carpels, ovary sessile, ovoid to ellip-

soid, with (4-)5(-7) locules and 1 or 2 ovules from axile

placentae in each locule, style 1 , stigmas (4-)5(-7) and

often capitate. Fruits drupaceous with a hard, usually

plurilocular stone, small ( 1 cm) to large ( 1 2 cm), exocarp

pulpy to fibrous and coriaceous in texture, endocarp

(stone) hard and woody, the surface smooth to deeply

rugose or costate, (4-)5(-7)-locular but with only l-2(-

5) seeds, valves of the endocarp visible as grooves on

the surface of the hard stone; seeds oblong, embryo

straight or slightly curved, endosperm fleshy and oily.

A family of eight genera and about 50 species,

ranging from Nicaragua to southern Brazil and

with a single species in western Africa. The family

includes tall trees of lowland rain forests as well

as smaller trees of deciduous forests and savannas.

Because our species are tall trees of lowland rain

forests, some have been little collected and are

poorly known. For this reason, many of the floral

details described below are based on the excellent

monograph by Jose Cuatrecasas (see reference

above).

The family is recognized by the stiff" simple al-

ternate leaves, these glabrous and lustrous with

entire to undulate, serrulate, or crenulate margins,

the 5-parted flowers with short calyx cup, narrow

petals, ten or more stamens with filaments united

at the base, cupular or scaly disc, 4-7-locular ova-

ry, and fruit with an unusually sculptured hard

BURGER ET AL.: FLORA COSTARICENSIS

25

stone. The petals are imbricate only at their edges

and may appear valvate. The anthers are quite

unusual, with the thecae usually borne on the low-

er part of a thick and distally expanded connective.

The wood is very hard and heavy.

Key to the Genera of II umiriaceae

la. Flowers with 50-180 stamens, anthers with 2 bilocular thecae; locules 2-ovulate; endocarp with 5

separate Ungulate valves; leaves bluntly obtuse to rounded at the apex, margins usually entire . . .

Vantanea

Ib. Flowers with 10-20 stamens, anthers with 2 unilocular thecae; locules 1-ovulate; leaves acuminate

or abruptly short-acuminate at the apex, margins undulate to sinuate, bluntly serrulate, or crenulate

2a

2a. Stamens 10, anthers with lateral thecae; endocarp with inconspicuous furrows, without dorsal fo-

ramina (apertures), alternating ribs thin; leaf margins mostly undulate to sinuate Sacoglottis

2b. Stamens 20, anthers with basal thecae; endocarp with 5 foramina at the apex, with 5 short opercular

valves; leaf margins mostly bluntly serrulate to crenulate Humiriastrum

Humiriastrum (Urban) Cuatrecasas

Evergreen trees, stipules small and deciduous or lack-

ing. Leaves petiolate to subsessile, subcoriaceous to co-

riaceous, entire to denticulate. Inflorescences axillary or

pseudoterminal, paniculate and usually with dichoto-

mous or trichotomous (cymose) branching, bracts de-

ciduous or persistent. Flowers with calyx united to form

a cup, calyx-lobes broadly rounded and imbricate; petals

free, oblong to linear, stiff; stamens 20 and of 2 alternate

lengths, glabrous, filaments united at the base, anthers

ovate-lanceolate to oblong, basifixed, thecae 2 and uni-

locular, ellipsoid or rounded, borne at the base of the

expanded connective, connective well developed beyond

the thecae, thick and acute at the apex, disc a dentate

ring around the ovary or of free scales; pistil with carpels

(locules) opposite the sepals, ovary 5-locular, 1 pendu-

lous ovule in each locule, style short, stigma smooth and

capitate. Fruit a small to medium-sized (1-5 cm) drupe,

ellipsoid to subglobose, smooth, exocarp fleshy to hard-

carnose, endocarp woody, usually lacking resinous cav-

ities, with 5 foramina (openings) around the apex and 5

oblong germination valves (opercula) on the upper half;

seeds oblong, usually 1 or 2 per fruit.

A genus of 1 2 species, mostly in South America,

but with one species reaching Costa Rica and Pan-

ama.

Humiriastrum diguense (Cuatr.) Cuatrecasas,

Contr. U.S. Natl. Herb. 35:141.1961. Sacoglot-

tis diguense Cuatr., Trop. Woods 96: 38. 1950.

H. diguense subsp. costaricense Cuatr., Contr.

U.S. Natl Herb. 35: 142. 1961. Figure 6.

Small to large trees, 25-40 m tall, with trunks 50-70

cm d.b.h., heartwood reddish brown, leafy branchlets

0.7-4 mm thick, internodes 1-6 cm long, glabrescent,

becoming grayish; stipules obscure or absent. Leaves al-

ternate, subsessile with petioles only 1-2 mm long; leaf

blades (4-)5-8.5(-9.5) cm long, 2-4.5(-5.5) cm broad,

elliptic to obovate-elliptic, acuminate (acute) at the apex,

cuneate (obtuse) to subtruncate at the base, margin blunt-

ly serrulate or crenulate with 2-5 teeth per centimeter,

laminae drying stiffly chartaceous to subcoriaceous and

dark brown, glabrous above, with sparse, thin, incon-

spicuous hairs ca. 0.2 mm long beneath, with 12-25

obscure secondary veins on each side, central second-

aries arising at angles of 50-70°, the secondaries difficult

to differentiate from the parallel tertiary veins. Inflores-

cences paniculate to corymbiform, usually with trichot-

omous branching, (5-)8-15 cm long, peduncles 1-2 cm

long, 1-2 mm thick, minutely (0.2 mm) puberulent, ra-

chis with 2 longitudinal ridges, pedicels 0.5-1 mm long.

Flowers ca. 3 mm long, greenish white, calyx ca. 0.5-0.7

mm long, calyx-lobes ca. 0.5-1.1 mm broad, glabrous

on the surfaces (in ours) and minutely ciliolate along the

margin; petals ca. 2.3 mm long and 1 mm broad, oblong,

sparsely puberulent with thin whitish ascending hairs;

stamens 20, glabrous, filaments connate at the base, un-

equal, the larger 1.2 mm long and the shorter 0.9 mm

long, anthers 0.7-0.8 mm long, ovate-lanceolate, disc

made up of small (0.2 mm) scales; ovary ca. 1 mm long

and puberulent, stigma 5-lobed. Fruits 20-26 mm long,

11-16 mm in diameter, ellipsoid to obovoid with bluntly

rounded or obtuse apex and truncated base, exocarp dry-

ing hard, smooth, and glabrous, becoming black at ma-

turity, endocarp hard woody, with 5 rounded apertures

near the apex.

Large trees of lowland rainforest formations from

near sea level to 250(-400) m elevation in Costa

Rica. Flowers have been collected in January-Feb-

ruary and July-November. Fruits have been col-

lected in January-April. This species is found on

both the Caribbean slope (near Tirimbina, Here-

dia) and the southern Pacific slope (General Valley

to Osa Peninsula) in Costa Rica; it ranges south-

ward through Panama to western Colombia.

Humiriastrum diguense is recognized by its small

26

FIELDIANA: BOTANY

Humiriastrum diguense

Sacoglottis trichogyna

5 mm

10cm

Vantanea barbourii Sacoglottis holdridgei

FIG. 6. Humiriaceae: four Costa Rican species.

BURGER ET AL.: FLORA COSTARICENSIS

27

alternate subsessile leaves with bluntly serrulate

laminae with many weakly denned secondary

veins, small 5-parted flowers with 20 stamens, and

unusual drupes with hard outer surface and woody

endocarp with five distal apertures. The large size

of these trees explains why they are poorly rep-

resented in herbaria. They are important timber

trees and their hard wood has been used for heavy

construction. In Costa Rica the names chiricano,

lorito, laurelito, and mspero have been used for

this species. All our material belongs to subspecies

costaricense, which is distinguished from other el-

ements of the species by its glabrous calyx-surface,

narrow leaves, less puberulent petals, and more

puberulent stems. These are not very significant

characteristics, but they do seem to be consistent.

Paul Allen mistakenly placed this species under

the name Sacoglottis excelsa Ducke in his book,

The Rain Forests of Golfo Dulce (p. 317. 1956).

Sacoglottis Martius

Evergreen trees; stipules minute or absent. Leaves al-

ternate, petiolate; leaf blades subcoriaceous to coria-

ceous, margin entire to crenate. Inflorescences axillary

or subterminal, paniculate with dichotomous or tri-

chotomous branching, bracts persisting or deciduous.

Flowers with united calyx, calyx-lobes 5, broadly round-

ed and imbricate; petals 5, free, imbricate (or appearing

valvate), stiff; stamens 10, unequal, with the 5 opposite

the sepals longer than those alternate with the sepals,

glabrous, filaments flattened and united near the base,

anthers ovoid to ovoid-oblong, dorsifixed near the base,

thecae 2 and unilocular, ellipsoid, attached to the lower

side of the expanded connective, dehiscing by detach-

ment, connective prolonged, acute, disc forming a thin

cup with a dentate margin, surrounding the base of the

ovary; pistil with 5 united carpels opposite the sepals,

ovary 5-locular, with 1 pendulous ovule in each locule,

style equaling or exceeding the length of the stamens,

stigma capitate and lobed. Fruits medium to large drupes,

ellipsoid to subglobose, exocarp carnose and smooth,

drying coriaceous, endocarp woody and bullate, without

apical openings, usually with 10 obscure longitudinal

furrows, resinous cavities present within the woody tis-

sue; seeds oblong, usually 1 or 2 per fruit.

A genus of about ten species, most South Amer-

ican, but with one species in western Africa and

the two listed below reaching Costa Rica. One of

our species marks the northern limits of the genus

in Central America and the other is endemic to

Cocos Island. The relatively few stamens, small

inflorescences, slightly undulate leaf-margins, loop-

connected secondary veins, and woody endocarps

without apertures distinguish this genus from our

other representatives of the family.

Key to the Species of Saccoglottis

la. Laminae elliptic-oblong, 2.5-5 cm broad, acute to cuneate at the base, major secondary veins arising

at angles of 50-70°; isthmian Costa Rica S. trichogyna

Ib. Laminae oblong, 4-8 cm broad, rounded to truncate at the base, major secondary veins arising at

angles of 70-90°; Cocos Island S. holdridgei

Sacoglottis holdridgei Cuatrecasas, Ciencia (Mex-

ico) 27: 171. 1972. Figure 6.

Medium to large trees, to 25 m tall and with trunks

to 1 m d.b.h., leafy branchlets with internodes 1-3.5 cm

long, 1.5-5 mm thick, glabrous, subterete; stipules 0.5-

1 mm long, caducous. Leaves alternate and distichous,

petioles 4-7 mm long, 2-4 mm thick, with adaxial ridges

continuous with the lamina margins, glabrous; leaf blades

7-13(-15) cm long, 4.2-6.5(-8) cm broad, oblong-ovate

to ovate-elliptic, abruptly short-acuminate at the apex,

rounded to truncate at the base, margin entire or slightly

sinuate-crenate, minutely glandular punctate at the mar-

gin, drying dark grayish brown to almost black above,

paler beneath, glabrous above and below, with 8-1 1 ma-

jor secondary veins on each side, the central secondaries

arising at angles of 70-90°, weakly loop-connected near

the margin. Inflorescences axillary, 0.5-3 cm long, cy-

mose-paniculate, subsessile or sessile (and appearing as

several inflorescences in the leaf axil), peduncle 0-4 mm

long, branches dichotomous, 1-4 mm long, sparsely and

minutely papillate-puberulent, bracts ca. 2.5 mm long,

1 mm broad, deciduous, pedicels ca. 0.5 mm long. Flow-

ers green to greenish yellow, calyx 1.7-2 mm high, calyx-

lobes ca. 2 mm broad, broadly rounded and suborbicu-

lar, glabrous but minutely ciliolate along the edge; petals

ca. 5.5 mm long and 2 mm broad, narrowly oblong;

stamens 10, glabrous, the longer filaments 4 mm long

and sepal-opposed, the shorter filaments 3 mm long and

petal-opposed, anthers ca. 1 mm long, ovate, thecae or-

ange, attached at the base of the connective, annular disc

cupulate, ca. 0.8 mm high, with denticulate margin; ova-

ry ca. 1.5 mm long, ovoid, style ca. 2.5 mm long, co-

lumnar. Fruits 32-40 mm long, 21-32 mm in diameter,

oblong-ellipsoid, sepals persisting at the base, exocarp

2-4 mm thick, endocarp woody, irregularly 5-septate,

resin vesicles with lustrous interior surfaces; seeds ca.

12 mm long, 4 mm in diameter, oblong.

A dominant large tree on Cocos Island. Flowers

have been collected in January, March, April, and

28

FIELDIANA: BOTANY

October. Fruits were collected in January, April,

August, and October. This species is said to in-

clude the tallest trees on Cocos Island; it is en-

demic there.

Sacoglottis holdridgei is recognized by its stiff

oblong to ovate leaves abruptly rounded at the

base and with crenulate-undulate margin, lack of

pubescence, small inflorescences, hard endocarp

with internal resinous chambers, and isolated hab-

itat. Sterile specimens of this species were origi-

nally placed under S. ovicarpa Cuatr. in the mono-

graph (pp. 168-169), but additional material has

shown the Cocos Island population to merit the

status of an endemic species. Sacoglottis amazon-

ica Mart, is closely related, but has larger fruit and

thinner elliptic leaves. Sacoglottis ovicarpa also

has larger fruit (5x4 cm), and the leaves are

similar to those of S. holdridgei but not as abruptly

truncated at the base.

Sacoglottis trichogyna Cuatrecasas, Ciencia (Mex-

ico) 27: 171. 1972. Figure 6.

Trees to 35 m tall, trunks 40-90 cm d.b.h., lower

branches often drooping, leafy branchlets 0.8-3.7 mm

thick, glabrous (the terminal bud minutely puberulent),

tuberculate-lenticellate, becoming terete; stipules minute

(ca. 0.5 mm) and caducous. Leaves alternate, petioles

(4-)5-l 1 mm long, 0.9-1.7 mm thick, with adaxial mar-

gins continuous with the lamina-margins, glabrous; leaf

blades (4-)5-12(-15) cm long, (1.5-)2-5(-7) cm broad,

elliptic-oblong to narrowly elliptic, gradually narrowed

to the acuminate apex, the tip 0.5-2 cm long, acute to

obtuse at the base, margin subentire to crenate-undulate

with a few reddish glands along the edge beneath, leaf

blades drying stiffly chartaceous to subcoriaceous and

dark grayish brown above, paler beneath, glabrous above

and below, major secondary veins (5-) 7-1 1 on each side,

central secondaries arising at angles of 50-70°, arcuate

and loop-connected near the margin. Inflorescences ax-

illary, paniculate, 1-3 cm long, peduncle and rachis to

1 cm long, minutely puberulent, lateral branches 1-3

cm, bracts 0.7-1 mm long, triangular, deciduous, pedi-

cels 1 mm long. Flowers in bud to 4.5 mm long and 2

mm in diameter, calyx ca. 1 mm long, calyx-lobes broad-

ly rounded, 1.5 mm broad, minutely ciliate along the

margin, usually eglandular, greenish; petals 4-6 mm long,

1-1.6 mm broad, oblong-linear, yellow; stamens 10, gla-

brous, united near the base, longer stamens 3.5-4.3 mm

long and 0.5-0.6 mm broad at the base, shorter stamens

3-3.2 mm long and 0.7-0.8 mm broad at the base, an-

thers 1-1.3 mm long, connective thick and acute, thecae

ca. 0.5 mm long, disc closely surrounding the ovary-

base, ca. 0.8 mm high, glabrous and minutely denticu-

late; ovary 1.5 mm long, ovoid, minutely puberulent,

style ca. 4 mm long, stigma 5-lobed. Fruits 3-5 cm long,

2-2.8 cm in diameter, oblong-ellipsoid to oblong-ob-

ovate, glabrous, exocarp 1-2 mm thick, endocarp with

5 longitudinal ridges and pitted warty surface, woody

interior with spherical resin cavities 1-4 mm in diam-

eter; seeds 1 cm long.

Trees of lowland rainforest formations, from near

sea level to 500 m elevation. Flowers have been

collected in February, June, and August; fruits were

collected in June, August, and December. The spe-

cies ranges from the Departments of Rio San Juan

and Zelaya in Nicaragua, the northern Caribbean

lowlands, and southern Pacific slopes of Costa Rica

through Panama.

Sacoglottis trichogyna is recognized by its stiff

alternate, slightly crenulate-undulate, and acu-

minate leaves with weakly loop-connected sec-

ondary veins, small flowers in small axillary in-

florescences, ten stamens of two lengths, thecae

near the base of an expanded connective, and el-

lipsoid drupe with unusual woody endocarp. The

reddish glands (punctate to linear) terminating

small veins in the proximal sinuses beneath the

leaf margin can help in the determination of sterile

collections. The young foliage has a conspicuous

pinkish red coloring, and leaf size can vary greatly.

This species is called campano, danto plomillo,

lorito, manteca, rosita, and titor in Costa Rica.

Material of this species was assigned to 5. ama-

zonica Mart, in the monograph of the family.

Vantanea Aublet

Tall trees, wood hard; stipules absent or obscure. Leaves

alternate, petiolate or sessile; leaf blades subcoriaceous

to coriaceous, usually entire. Inflorescences axillary or

terminal, paniculate or corymbiform, with alternate or

dichotomous branches, bracts deciduous. Flowers white

or whitish (reddish in K guianensis), calyx cupular, calyx

5-lobed distally; petals 5, free, oblong to linear-oblong,

stiff, contorted in bud; stamens (15-18-)50-120(-180),

filaments united at the base, linear, glabrous, anthers

ovate-lanceolate, basifixed, thecae 2 and bilocular, at-

tached near the base of the anther, dehiscing by longi-

tudinal slits, connective extended beyond the thecae and

acuminate to bluntly obtuse, disc cupular and closely

surrounding the base of the ovary, dentate to fimbriate

at the margin; pistil with 5 carpels opposite the sepals,

ovary glabrous or puberulent, 5-locular, each locule with

2 pendulous ovules, the lower ovule with a longer funicle,

style simple and straight, stigma thickened and 5-lobed.

Fruit a medium to large (2-8 cm) drupe, pericarp smooth

(tuberculate in V. tuberculata), exocarp carnose and co-

riaceous when dry, thick or thin, endocarp woody, with-

out resiniferous cavities, opening (when the seeds ger-

minate) by linear or oblong valves or operculae pushed

out by the emerging radicle; seeds l(-3) in each fruit.

A genus of 14 species in lowland (below 1000

m) tropical rainforest formations. The genus rang-

BURGER ET AL.: FLORA COSTARICENSIS

29

es from southeastern Nicaragua to southern Brazil.

It is distinguished by its small flowers with cupular

calyx, five free (almost valvate), long, narrow pet-

als, many stamens of uniform length, 5-locular

ovary with two pendulous ovules in each locule,

and the hard (when dry) drupe with unusual woody

stone. A species with only 15-18 stamens has re-

cently been described from central Panama (see

Ann. Missouri Bot. Gard. 75: 1 148-1 150. 1988).

The large number of stamens is probably a derived

feature and not primitive as many authors have

suggested. The unusual expansion of the connec-

tive with the thecae at the base is similar to that

in other genera of the family.

Vantanea bar hour ii Standley, Trop. Woods 75: 5.

1943. Figure 6.

Large trees up to 50 m tall, trunks to over 1 m d.b.h.,

leafy branchlets 2-5 mm thick, glabrous; stipules ap-

parently absent (or reduced to small tuberculae). Leaves

with petioles 4-10 mm long, 1-2 mm broad, with lateral

margins continuous with the lamina-margins, thickened

at the base; leaf blades 5-14 cm long, 2.4-7 cm wide,

broadly elliptic to oblong-elliptic, bluntly obtuse to

rounded and emarginate at the apex, obtuse to cuneate

at the base and decurrent on the petiole, margins entire

or slightly undulate, the leaves drying dark brownish

above and slightly paler beneath, subcoriaceous, slightly

lustrous above, glabrous above and below, with 7-10

major secondary veins on each side, the central second-

aries arising at angles of 45-65°, not clearly loop-con-

nected near the margin. Inflorescences axillary or sub-

terminal, cymose and paniculate to corymbiform, 2-8

cm long, often subtended by caducous small leaves, pe-

duncles 1-2 mm thick, branches of the inflorescences

dichotomous, very minutely (0.2 mm) puberulent, ped-

icels 1-2 mm long. Flowers whitish, ca. 12 mm long,

calyx 1-1.5 mm high, with rounded ciliolate lobes ca.

0.5 mm broad; petals 7-9 mm long, 1.5-2.5 mm wide,

oblong and narrowed toward the apex, glabrous within

and appressed retrorse puberulent on the outside except

for the thin glabrous margins; stamens ca. 50-60, fila-

ments 5-7 mm long, glabrous, anthers ca. 0.8 mm long,

ovate-lanceolate, thecae 0.4 mm long, disc 1 mm high,

thick, with short denticulate margin; ovary 1.5-2 mm

high, tomentulose-hirsutulous, style ca. 4 mm long, slen-

der. Fruits 2.8-3 cm long, 1.8-2 cm in diameter, ovoid-

oblong to ellipsoid, narrowed to the apex, abruptly

rounded at the base, smooth, endocarp hard woody, ca.

2.5 cm long and 1.6 cm in diameter, with 5 oblong

U-shaped grooves (valves).

Tall trees of lowland rainforest formations on

the northern Caribbean and southern Pacific slopes

of Costa Rica, from near sea level to 800 m ele-

vation. Flowers are probably produced in Novem-

ber-June; fruits have been collected in June and

September. The species ranges from Costa Rica to

Panama.

Vantanea barbourii is recognized by its tall stat-

ure, stiff glabrous alternate leaves with entire or

slightly undulate margins, small flowers with many

stamens, unusual anthers, and fruit with woody

endocarps that have five longitudinal U-shaped

grooves. The short calyx-cup with broadly round-

ed lobes, narrow petals puberulent along the back,

and the apically blunt to emarginate leaves are also

distinctive. Paul Allen noted that these trees tend

to lose all their leaves for a brief period at the end

of the heavy rains in November or December (The

Rain Forests of Golfo Dulce, p. 351. 1956). The

species has been called ira chiricana, chiricano,

chiricano triste, and nispero. The very hard and

durable wood has been used for bridge timbers.

Only the following Costa Rican collections have

been seen: Allen 6546 and 6681, Barbour 1018,

Dayton & Barbour 3129 (the type), and Hartshorn

2139. Vantanea occidentalis Cuatr. of Colombia

is very similar to V. barbourii and may prove to

be a synonym.

ERYTHROXYLACEAE Kunth

By Timothy Plowman

REFERENCES— N. L. Britton, Erythroxylaceae, N.

Amer. Flora 25: 59-66. 1907. W. G. D'Arcy & N.

Schanen, Erythroxylaceae, in Flora of Panama,

Ann. Missouri Bot. Gard. 62: 21-33. 1975. T.

Plowman, Erythroxylaceae, in R. A. Howard, Flo-

ra of the Lesser Antilles, 543-551. 1988. O. E.

Schulz, Erythroxylaceae, in A. Engler, Pflanzenr.

4(134): 1-176. 1907. P. C. Standley & J. A. Stey-

ermark, Erythroxylaceae, in Flora of Guatemala,

Fieldiana: Bot. 24(5): 390-393. 1946.

Glabrous shrubs or small trees, bisexual (hermaph-

roditic) or rarely dioecious; stipules on twigs persistent

or caducous. Leaves alternate (opposite in Aneulophus),

simple, stipulate, petiolate, entire, pinnately veined. In-

florescences fasciculate at nodes, with 1-many flowers,

sometimes short-pedunculate, subtended by small scar-

ious bracteoles, flowers pedicellate. Flowers radially

symmetrical, bisexual or unisexual, often heterostylous;

calyx persistent, the 5 valvate sepals united below; petals

free, 5, alternate with sepals, imbricate in bud, caducous,

usually appendaged on the inner (adaxial) surface and

narrowed at the base; stamens 10, biseriate, united at the

base by the filaments and usually forming a short tube,

anthers 2-locular, longitudinally dehiscent; pistil soli-

tary, ovary superior, 3-locular (2-locular in Nectarope-

talum), usually only one locule ovuliferous; ovules sol-

itary (2 in Nectaropetalum), axile, pendulous, styles 3 (2

in Nectaropetalum), free or connate from the base, stig-

30

FIELDIANA: BOTANY

mas capitellate, rarely subsessile. Fruits small, drupa-

ceous, and 1 -seeded (in Aneulophus rarely capsular and

2- or 3-seeded); embryo straight, with or without en-

dosperm.

Four genera with about 230 species; three of the

genera, with few species, are restricted to tropical

Africa and are incompletely known. Erythroxylum

is by far the largest genus and occurs in tropical

parts of Australia, Asia, Africa, and the Americas.

Erythroxylaceae are recognized by the lack of

indumentum, the intrapetiolar stipules, the usu-

ally alternate, simple, entire leaves, the fascicled,

axillary flowers, the persistent 5-lobed calyx, the

appendaged petals, and the small, drupaceous fruit.

Erythroxylum P. Browne

Glabrous shrubs or small trees, evergreen or decidu-

ous, rarely dioecious or subdioecious, the twigs com-

pressed at apex, often bearing persistent, distichous, im-

bricated cataphylls and stipules; stipules intrapetiolar,

united and appearing as a single organ, dorsally bicostate,

often 2- or 3-setulose at apex, persistent or caducous,

often leaving an obliquely transverse scar. Leaves alter-

nate, often distichous, petiolate, simple, entire, with in-

volute vernation that sometimes imprints 2 parallel lines

and/or a distinct central panel on lower surface, pin-

nately veined. Inflorescences of fasciculate or solitary

axillary flowers, rarely short-pedunculate, the flowers

arising from small, scarious, often persistent bracteoles,

pedicellate. Flowers radially symmetrical, small, heter-

ostylous; calyx of 5 valvate sepals, united below, persis-

tent; petals 5, free, imbricate in bud, appendaged on the

adaxial surface with a 2-lobed ligule, caducous; stamens

10 in 2 whorls of 5, the outermost alternate with the

petals, the filaments united at base and forming a short

tube surrounding the ovary, persistent, anthers 2-locular,

longitudinally dehiscent; ovary 3-locular but with only

one locule ovuliferous, the ovule solitary in the fertile

locule, pendulous from an axile placenta, anatropous,

epitropous, styles 3, free or partly connate from base,

stigmas capitellate. Fruit a small, fleshy, reddish, one-

seeded drupe, the endocarp 1 - or 3-locular but with only

one fertile locule; seed with or without endosperm, the

embryo straight.

About 230 species, of which at least 180 are

found in the Neotropics; the centers of diversity

for the genus are found in Venezuela, eastern Bra-

zil, and Madagascar; six species, including one cul-

tivated, occur in Costa Rica. Costa Rica seems to

be one of the less diverse areas of Central America

in species of Erythroxylum, being surpassed even

by Nicaragua. Most if not all of the species have

originated elsewhere (South America, Antilles) with

subsequent dispersal to Costa Rica.

In Costa Rica, Erythroxylum species fall into

two main groups: those with relatively large, api-

cally acute or acuminate leaves and conspicuously

striate-nerved stipules (E. macrophyllum, E. citri-

folium, E. fimbriatum), and those with small to

medium-sized, apically obtuse or rounded leaves

and smooth, nonstriate stipules (remaining spe-

cies). Identification within either of these groups

may be difficult, especially if complete material is

not available. I therefore provide multiple char-

acters in the key to help distinguish flowering,

fruiting, and/or vegetative material. The impor-

tant stipule characters should be observed in newly

formed (subapical), undamaged stipules or cata-

phylls. The character of relative positions of calyx

and staminal tube is easily observed in flowers

whose petals have fallen, or in young fruit.

Key to the Species of Erythroxylum in Costa Rica

la. Stipules longitudinally striately veined, caducous or persisting; mature endocarps terete 2a

Ib. Stipules smooth, not striately veined, persistent (sometimes withering but not caducous); mature

endocarps variously ribbed or sulcate, not terete 4a

2a. Stipules apically acute or tapered to a long point; bracteoles 2-5.5 mm long; calyx-lobes broadly

ovate, oblong or obovate and abruptly acuminate at the apex E. macrophyllum

2b. Stipules apically obtuse or rounded and 2- or 3-setulose; bracteoles 0.5-2 mm long; calyx lobes

triangular to triangular-ovate or ovate-lanceolate, acute or acuminate at the apex 3a

3a. Stipules persistent, 1.5-2 mm long (not including seti), the 2 lateral apical seti conspicuous,

often fimbriate and strongly recurving, persisting; staminal tube equaling or longer than the

calyx; endocarps acute to obtuse at the apex E. fimbriatum

3b. Stipules usually caducous, 4-1 2 mm long, the apical seti slender, weakly filamentous, evanescent;

staminal tube shorter than the calyx; endocarp obtuse at apex E. citrifolium

4a. Flowers manifestly unisexual, produced near apex of current season's shoots, often on short shoots;

mature drupe 3-locular with 1 fertile and 2 empty locules; petiole longer than '/5 the length of leaf

blatle E. rotundifolium

BURGER ET AL.: FLORA COSTARICENSIS

31

4b. Flowers bisexual, produced on year-old twigs of past season; mature drupe unilocular; petiole shorter

than '/5 the length of the leaf blade 5a

5a. Staminal tube less than 50% as long as the calyx; plants completely deciduous; flowers produced

on last season's twigs before the leaves, never in leaf-bearing axils; leaf blades never bilineate

nor with distinct central panel on lower surface; a native species E. havanense

5b. Staminal tube 50-1 00% as long as the calyx; plants evergreen or incompletely deciduous; flowers

produced in axils of year-old twigs (in Costa Rica), often with leaves present; leaf blades usually

bilineate and/or with distinct central panel on lower surface; cultivated species

E. novogranatensis

Erythroxylum citrifolium St. Hil., Fl. bras. mend.

94. 1829. E. citrifolium St. Hil. var. minus O.

Schulz, 1/1 Engl., Pflanzenr. 4(134): 37. 1907.

D'Arcy & Schanen, Ann. Missouri Bot. Gard.

62: 29. 1975.

Evergreen shrubs or small trees to 10 m tall, the

branchlets smooth, the bark dark reddish or grayish

brown, developing a tan suberous outer bark, the len-

ticels if present minutely punctate; cataphylls scattered

at base of shoots, 5-7, early caducous, similar to foliar

stipules; foliar stipules caducous, sometimes tardily so,

(2-)4-8(-12) mm long, oblong-lanceolate, membrana-

ceous, densely longitudinally striate-nerved, obtuse at

apex with three slender, short, often evanescent setae 1-

2 mm long. Leaves with petiole 4-7 mm long, the leaf

blades 8-14 cm long, 2-7 cm wide, oblong, oblong-el-

liptic or lanceolate, long-acuminate at apex, short-acu-

minate to rounded at base, coriaceous or subcoriaceous,

the upper surface drying dark grayish green or brown,

the secondary nerves often obscure, the lower surface

drying somewhat ferruginous, not bilineate and a central

panel lacking or faint, the lateral nerves prominulous,

more or less shiny on both surfaces. Inflorescence pro-

duced in the axils of leaves or cataphylls on mature twigs

of current season, many-flowered; bracteoles 0.5-2 mm

long, lightly striate-nerved, strongly keeled, at apex acute

or obtuse, 1-setulose; pedicels 3-5 mm long, strongly 5-

ribbed. Flowers bisexual; calyx ca. 1.5 mm long, trian-

gular to ovate-lanceolate, often spreading after anthesis;

Staminal tube ca. lh as long as calyx. Drupe 7-10 mm

long, 4-5 mm in diameter, red, the endocarp oblongoid,

obtuse at apex, terete.

Evergreen forest formations of the Pacific slope,

700-1 200 m elevation, the General Valley (Skutch

4351 F, Skutch & Barrantes 5057 F), and near Las

Alturas in Puntarenas Province ( Wilbur et al. 22658

F). It appears to be uncommon in Costa Rica.

Flowering specimens have been collected between

June and August. This species is widely distrib-

uted, often in secondary forest, from Nicaragua to

Bolivia; the Guianas; and southeastern Brazil.

Erythroxylum citrifolium is recognized by its ca-

ducous, striate-nerved, apically obtuse, 3-setulose

stipules, by relatively large (8-14 cm long), oblong,

apically acuminate leaves, and by dense axillary

clusters of small (ca. 1.5 mm long) flowers.

Erythroxylum fimbriatum Peyr., in Mart., Fl. bras.

12(1): 162. 1878.

Evergreen shrubs or treelets, 2-6(-10) m tall, the

branchlets initially with smooth bark without discrete

lenticels, becoming superficially suberose, light tan; cat-

aphylls scattered at base of new shoots, 4 or 5, similar

to foliar stipules; foliar stipules persistent, 1.5-2 mm

long (excluding setae), broadly ovate, subcoriaceous, dis-

tinctly longitudinally striate-nerved with 4-6 nerves per

side, obtuse or truncate at apex, 3-setulose, the 2 lateral

setae persistent, conspicuous, (l-)3-5 mm long, ribbon-

like, extending from subalate keels, often fimbriate, man-

ifestly recurved, the medial seta filamentous, evanescent.

Leaves persistent, scattered on twigs, the petiole 3-5 mm

long; leaf blades 5-10(-14) cm long, 2.5-5.5 cm wide,

elliptic to elliptic-oblong, short to long-acuminate at

apex, rarely acute, the tip itself obtuse or acute, broadly

to narrowly cuneate at base, chartaceous, the upper sur-

face drying dark grayish green, the adaxial midrib some-

times impressed-sulcate with a slender medial ridge, the

lateral nerves obscure or sometimes impressed-sulcate,

the lower surface drying ochreous or ferruginous, without

parallel lines, the lateral nerves sometimes conspicuous.

Inflorescences in axils of leaves or cataphylls on current

or year-old shoots, with l-3(-8) flowers per node; brac-

teoles ca. 1 mm long, striate-nerved, acute and 1-setulose

at apex; pedicels 4-6 mm long, pentagonal in cross sec-

tion, the fruiting pedicels 5-7 mm long. Flowers bisexual;

calyx 1-2 mm long, the lobes narrowly triangular to

lanceolate, narrowly acute or acuminate at apex; sta-

minal tube equaling or slightly longer than the calyx.

Drupes 10-12 mm long, 4-5 mm in diameter, red, the

endocarp oblongoid, acute or obtuse at apex, terete at

maturity.

In Central America known only from Costa Rica,

Provincia de Heredia, Finca La Selva, in lowland

moist primary forest, 45-100 m elevation (Hart-

shorn 1243 and 1413 F, Opler 1740 F, Kress 76-

526 DUKE). In Costa Rica, Erythroxylum fimbria-

tum flowers in March-May and fruits in April-

June. In South America, the species is sporadically

distributed throughout the greater Amazon basin,

from Colombia to French Guiana, south to Peru

and western Brazil.

Erythroxylum fimbriatum is easily recognized

by the well-developed (3-5 mm long), persistent,

fimbriate, and recurved lateral setae on the striate-

32

FIELDIANA: BOTANY

nerved stipules. The Costa Rican and some Pe-

ruvian material of this species differs modestly

from the typical, lowland Amazonian form in hav-

ing smaller leaves and the stipular setae much

shorter and not so markedly fimbriate and re-

curved. Moreover, the lower surface of the leaves

in the Amazonian collections shows conspicuous

secondary nerves diverging at near right angles

from the midrib.

Erythroxylum havanense Jacq., Enum. Syst. PI.

21. 1760. Select. Stirp. Amer. Hist. t. 87, fig. 2.

1763. D'Arcy & Schanan, Ann. Missouri Bot.

Gard. 62: 32. 1975. E. overturn Cav., Diss. 404,

t. 233. 1789. E. obtusum DC., Prodr. 1: 574.

1824. E. havanense Jacq. var. continentis O.

Schulz, in Engl., Pflanzenr. 4(134): 92. 1907. E.

chiapense Lundell, Wrightia 4: 175. 1971. Fig-

ure 7.

Deciduous shrubs or treelets 1-5 m tall, the branchlets

1.5-2.0 mm in diameter, weakly differentiated into long

and short shoots, smooth, light grayish brown to dark

reddish brown, dotted with whitish punctate or elongate

lenticels, becoming longitudinally fissured; cataphylls

persistent, produced at base of long shoots or congested

on short shoots, similar to foliar stipules, turning dark

brown with age; foliar stipules persistent, 2.0-3.5 mm

long, triangular-ovate, not striate-nerved, acute and brief-

ly 2-setulose at apex, drying reddish brown. Leaves de-

ciduous, sometimes tardily so, scattered on long shoots

or 1-3 produced at tips of short shoots, the petiole 2.5-

5 mm long; leaf blades (3-)3.5-8 cm long, (1.2-)1.5-3(-

4) cm wide, obovate or elliptic, rounded, obtuse or slight-

ly retuse at apex, acute at base, chartaceous, strongly

bicolorous, the upper surface drying medium green, dull,

the lower surface drying very pale green or glaucous to

whitish, without parallel lines nor with a distinct central

panel, the lateral nerves 12-14, inconspicuous. Inflores-

cence produced in axils of leaves or cataphylls on leafless,

year-old twigs just prior to leaf flush, with 1-8 flowers

per node; bracteoles 1-1.5 mm long, ovate, acuminate

at apex; pedicels slender, 4-9 mm long. Flowers bisexual;

calyx-lobes triangular-ovate, reflexing after an thesis; sta-

minal tube much shorter than calyx (in Costa Rica).

Drupes 5-7 mm long, 4-5 mm in diameter, the endocarp

ellipsoid to oblongoid, rounded or obtuse at apex, un-

equally and shallowly 4-sulcate at maturity.

Lowland Guanacaste and adjacent Puntarenas,

seasonally dry, deciduous forest and adjacent sa-

vannas, often along water courses, sea level to 250

m elevation. This species flowers in March-June

and fruits in May-June. It ranges over Cuba, the

Lesser Antilles, and Mexico to Panama and the

northern coast of South America.

Erythroxylum havanense is recognized by its de-

ciduous habit, with flowers appearing before the

leaves, by the relatively small, apically obtuse or

rounded, strongly bicolorous leaves that are

elineate beneath, by the staminal tube much shorter

than the calyx lobes, and by the short, obtuse,

sulcate endocarps. The illustrations of E. hava-

nense (fig. 7) are based on the following collections:

A, E, F, drawn from Frankie 380c from Costa

Rica; B, C, drawn from Sandino 2888 from Nic-

aragua; D, drawn from Plowman 3597 from Co-

lombia; G, H, I, drawn from Berry 3502 from

Venezuela. All the specimens are represented at F.

In the Flora of Panama (Ann. Missouri Bot.

Gard. 62: 32. 1975), D'Arcy and Schanen erro-

neously placed several species in synonymy under

E. havanense. Erythroxylum cumanenseKunlh and

E. hondense Kunth are distinct species that occur

in northern South America, and E. mexicanum

Kunth is found in dry areas of Mexico, El Salva-

dor, and Nicaragua. Erythroxylum pringlei Rose

is here considered a synonym ofE. rotundifolium

Lunan sens. lat. (see below).

Erythroxylum macrophyllum Cav., Diss. 401, t.

227. 1789. E. lucidum H.B.K., Nov. Gen. sp.

5: 138. 1822. E. costaricense J. D. Smith, Bot.

Gaz. 23: 240. 1897. E. ellipticum Ramirez, An-

ales Inst. Med.-Nac. Mexico 3: 36. 1897. E.

tabascense Britton, N. Amer. Fl. 25: 66. 1907.

Standl. & Steyerm., Fieldiana: Bot. 24(5): 393.

1946. E. lucidum H.B.K. var. costaricense (J.

D. Smith) O. Schulz, in Engl., Pflanzenr. 4(1 34):

25. 1907. D'Arcy & Schanen, Ann. Missouri

Bot. Gard. 62: 28. 1975. E. skutchii Standl.,

Publ. Field Mus. Nat. Hist., Bot. Ser. 22: 344.

1 940. E. multiflorum Lundell, Amer. Midi. Nat-

uralist 29: 474. 1943. D'Arcy & Schanen, Ann.

Missouri Bot. Gard. 62: 24, t. 1. 1975.

Shrubs 2-3 tall or small trees to 10 m tall; branchlets

2-4 mm in diameter, smooth, becoming tan or light

grayish brown, with obliquely transverse scars of fallen

stipules and cataphylls, without distinct lenticels; cata-

phylls produced at base of shoots, caducous, 10-60 mm

long, similar to but sometimes larger than foliar stipules;

foliar stipules 4-35 mm long, longitudinally striate-

nerved, at apex acutely tapered to a point, sometimes

with 2-3 filamentous setae, caducous, leaving obliquely

transverse scars on twigs. Leaves with petiole 3-10 mm

long; leaf blades (5-)7-30 cm long, (2-)3-15 cm wide,

oblong to elliptic-oblong, long- to short-acuminate at

apex, the tip itself sharply acute, acuminate to acute,

obtuse or rounded at base, the upper surface often drying

rather shiny and leaden gray, the lateral nerves incon-

spicuous, rarely impressed-sulcate, the lower surface dry-

ing dull and ferruginous, usually without parallel lines

BURGER ET AL.: FLORA COSTARICENSIS

33

E

Fio. 7. Erythroxylaceae: Erythroxylum havanense. A, flowering branch; B, fruiting branch; C, leaf showing ve-

nation; D, stipule; E, short-styled flower; F, petal; G, drupe with attached calyx; H, cross section of endocarp; I,

embryo.

34

FIELDIANA: BOTANY

or distinct central panel, the lateral nerves inconspicuous

or rarely prominulous. Inflorescences produced in the

axils of leaves or cataphylls on new shoots, often ap-

pearing "knobby" at the nodes, the flowers usually nu-

merous; bracteoles 2.0-5.5 mm long, persistent, 1 -keeled,

striate-nerved, apically acute, 1-setulose; pedicels 3-10

mm long, 5-ribbed. Flowers bisexual; calyx 2.5-5.0 mm

long, the lobes broadly ovate, obovate or oblong, often

subfoliose, lightly striate-nerved, abruptly acuminate at

apex, the margins often overlapping or touching; sta-

minal tube about half as long as the calyx. Drupes 8-1 1

mm long, 4-5 mm in diameter, the endocarp oblongoid

or ovoid-oblongoid, obtuse or acutish at apex, terete at

maturity.

In Costa Rica, widespread on both Atlantic and

Pacific slopes (Guanacaste, Puntarenas, Alajuela,

San Jose, Heredia, Limon) in evergreen moist to

wet forests, seasonally dry or cloud forests, sea

level to 1600 m elevation. Rowers in January-

September; fruits in February-November. The

species ranges from southern Mexico to Bolivia

and Brazil.

Erythroxylum macrophyllum is recognized by

the relatively large (4-30 mm long), apically acute,

striate-nerved, caducous stipules that leave

obliquely transverse scars on the stem, by the rath-

er thick, apically acuminate leaves, by the persis-

tent, striate-nerved bracteoles, and by the broadly

ovate, abruptly acuminate calyx-lobes.

Erythroxylum macrophyllum is a highly poly-

morphic, complex, and wide-ranging species, oc-

curring from southern Mexico to Bolivia. Many

names have been proposed to accommodate the

numerous minor and major variants in both Cen-

tral and South America. I am treating the species

here in the widest sense. In Central America, sub-

specific taxa are difficult to characterize with any

degree of constancy. However, in Costa Rica two

general forms occur, corresponding to E. costari-

cense J. D. Smith, with leaves generally less than

1 4 cm long, and E. skutchii Standley, with leaves

more than 1 5 cm long. The former, small-leaved

form is far more widespread, occurring from sea

level to 1 600 m elevation in most of Costa Rica.

Erythroxylum skutchii was described from a

specimen (Skutch 4847 F) collected in the General

Valley. The leaves are distinguished adaxially by

impressed-sulcate nerves and abaxially by prom-

inulous, reticulate venation. Two other collections

from Limon Province (Gomez et al. 20463 F) and

the Osa Peninsula (Gomez 19681 F) more or less

match the type of E. skutchii. Several other col-

lections from the lowlands of Heredia and Limon

Provinces have exceptionally large leaves but

without the impressed-sulcate nerves of E. skut-

chii(Chacon 716, Hartshorn 1198, Kennedy 3790A,

Stevens 23797, 23964, Wilbur 34408, all at F).

These collections differ from the common high-

land forms primarily in the larger leaf size and

greater number of flowers per node. A number of

intermediate specimens connect the extremes of

variation. Until this group can be thoroughly re-

vised, I decline to designate formally subspecific

taxa in Costa Rica.

Erythroxylum multiflorum Lundell from Pan-

ama is also included in synonymy under E. mac-

rophyllum. This distinctive plant with large leaves

and cataphylls is obviously related to both E.

skutchii and two distinctive South American va-

rieties: E. macrophyllum var. macrocnemium

(Mart.) Plowman and var. ecuadorense Plowman.

Bawa and Opler (Evolution 29: 167-179. 1975)

erroneously reported that " Erythroxylum lucidum

var. costaricense" from Guanacaste is dioecious.

This error resulted from a misidentification of E.

rotundifolium Lunan.

Erythroxylum novogranatense (Morris) Hieron.,

Bot. Jahrb. Syst. 20, Beibl. 49: 353. 1895. O.

Schulz, i/i Engl., Pflanzenr. 4(134): 85, t. 18.

1907. E. coca Lam. var. novogranatense Morris,

Bull. Misc. Inform. 5, t. 2. 1889.

Evergreen shrubs or small trees to 6 m tall, the branch-

lets ca. 2 mm in diameter, without short shoots, flexuous,

often appearing zigzag, reddish brown, the lenticels

punctate or absent, if present rarely breaking the surface;

cataphylls lacking or few, if present resembling foliar

stipules; foliar stipules 2.5-3.5 mm long, narrowly ovate

to triangular, acute to obtuse at apex, minutely 2-setu-

lose, membranous, not striate-nerved, soon withering

and disintegrating. Leaves persistent, scattered on twigs,

the petiole 2-5 mm long; leaf blades 2.5-7.5 cm long,

1.2-3.6 cm wide, elliptic to obovate or oblong, at apex

obtuse, rounded or retuse, at base acute to attenuate,

nrm-membranaceous, the upper surface bright yellowish

green when fresh, the adaxial midrib flat not ridged, the

lower surface very pale green, usually bilineate and/or

with a distinct central panel, the laminas drying pale

green or yellowish and dull on both surfaces, the lateral

nerves 10-15, inconspicuous. Inflorescence produced in

axils of the previous season's twigs, with or without leaves

present, l-3(-10) flowers per node; bracteoles 1-1.5 mm

long, cymbiform-deltoid, acuminate at apex, pedicels 3-

7(-12) mm long. Flowers bisexual; calyx 1.5-2.5 mm

long, divided V2-% its length, the lobes narrowly to broad-

ly ovate; staminal tube half as long as to equaling the

calyx. Drupes 8-13 mm long, 4-7 mm in diameter, the

endocarp ovoid to ellipsoid, rounded to obtuse at apex,

unequally 4-sulcate.

Ornamental shrubs cultivated at I.I.C.A., Turri-

alba, ca. 635 m, flowering in June-August (Brown

CR-221 F, Rossbach 3570 GH, Leon 591 us). Na-

BURGER ET AL.: FLORA COSTARICENSIS

35

live to Colombia and cultivated throughout the

tropics and in Central America as an ornamental

or medicinal plant, or as a minor commercial

source of the alkaloid cocaine.

Erythroxylum novogranatense is recognized by

the relatively delicate, bright yellowish green (when

fresh), apically obtuse leaves that are usually bilin-

eate beneath, by the thin stipules that soon dis-

integrate, and by the sulcate endocarps. This spe-

cies has long been confused with E. coca Lam.,

which is the commercially important species from

which both coca leaves and cocaine are principally

derived (Plowman, J. Linn. Soc., Hot. 84: 329-

353. 1982. Bohm et al., Syst. Bot. 7: 121-133.

1982). Because it is more difficult to grow, E. coca

is rarely seen outside of the tropical Andean mon-

tana where it is native. No unambiguous collec-

tions of E. coca are known from Central America.

The common name of E. novogranatense is "coca."

Erythroxylum rotundifolium Lunan, Hort. Jamaic.

2:116.1814. Standl. & Steyerm., Fieldiana: Bot.

24(5): 393. 1946. E. obovatum Macfad., Fl. Ja-

maica 143. 1837, non Griseb.. E. pallidum Rose,

Contr. U.S. Natl. Herb. 8: 314. 1905. Standl. &

Steyerm., Fieldiana: Bot. 24(5): 392. 1946. E.

pringlei Rose, Contr. U.S. Natl. Herb. 8: 314.

1905. E. compactum Rose, Contr. U.S. Natl.

Herb. 8: 313. 1905. E. suave O. Schulz, in Ur-

ban, Symb. Antill. 5: 197. 1907. E. suave O.

Schulz var. compactum (Rose) O. Schulz, in

Engl., Pflanzenr. 4(134): 68, t. 15. 1907. E. ses-

silijlorum O. Schulz, in Engl., Pflanzenr. 4(134):

69. 1907. E.fiscalense Standl., Publ. Field Mus.

Nat. Hist, Bot. Ser. 22(1): 33. 1940. Standl. &

Steyerm., Fieldiana: Bot. 24(5): 391. 1946. E.

tikalense Lundell, Wrightia 4: 177. 1971.

Semideciduous, dioecious shrubs or small trees to 8

m tall, highly variable in habit with branches long and

slender, or short and compact with well-developed short

shoots, the branchlets dark brown to dark gray, abun-

dantly supplied with lenticels; cataphylls persistent, scat-

tered at base of long shoots or crowded on short shoots,

similar to foliar stipules; foliar stipules persistent, 1.5-

2.5 mm long, at apex obtuse and briefly 2-setulose, fim-

briate at margin. Leaves persistent or partly deciduous,

the petioles 3-6 mm long, very slender, often drying

orange or ferruginous; leaf blades 8-30 mm long, 7-25

mm wide, obovate, elliptic, or rounded, rounded or shal-

lowly retuse at apex, acute at base, bicolorous, drying

medium green on upper surface, very pale green on lower

surface, without parallel lines or central panel on lower

surface, the lateral nerves 4-6, inconspicuous. Inflores-

cence produced in the axils of leaves or cataphylls near

apex of current season's shoots, the flowers 1 (or rarely

2 or 3) per node; bracteoles ca. 0.8-1.0 mm long, ovate,

fimbriate; pedicels 0.5-4 mm long, 5-angled, thickened

at the apex into the calyx. Flowers unisexual; calyx 1.2-

1.5 mm long, the lobes triangular to triangular-ovate,

acute at apex; pistillate flowers bearing a staminal tube

with very short filaments and rudimentary anthers, the

ovary with subsessile, flattened stigmas, 0.5 mm long;

stamina te flowers with 10 stamens in two unequal series,

the staminal tube about half as long as the calyx, and a

pistil rudiment. Drupes 5-8 mm long, 3-5 mm in di-

ameter, the endocarp ellipsoid or curved, rounded at

apex, ± trigonal in cross section, subequally 3-locular

with 1 fertile and 2 empty locules.

In Costa Rica, known only from Guanacaste and

northwestern San Jose provinces, in dry forest,

often along water courses, 50-900 m elevation.

Flowering specimens have been collected in late

May and in August-September, fruits in Septem-

ber. This species ranges from the Bahamas, Great-

er Antilles, and Mexico to Costa Rica.

Erythroxylum rotundifolium is recognized by the

small, rounded leaves with long, slender, orangish

petioles and by the small, unisexual, solitary, sub-

terminal flowers and trilocular drupes. Erythrox-

ylum rotundifolium is treated here in the widest

sense as a highly polymorphic species with either

slender and open or compact and dense branching

habit, and the leaves variable in shape and size of

lamina and relative length of petiole. After further

study, some of the Central American segregate

populations such as E. pallidum Rose and E. com-

pactum Rose may need to be recognized at least

at the varietal level. Much Central American ma-

terial of this species has been misidentified as E.

brevipes DC., a species from Puerto Rico and His-

paniola closely related to E. havanense.

ZYGOPHYLLACEAE

By William Burger

REFERENCE— D. M. Porter, The genera of the

Zygophyllaceae in the southeastern United States.

J. Arnold Arbor. 53: 531-552. 1972.

Annual or perennial herbs, subshrubs, shrubs, or trees,

often strongly scented and with sticky resin, growth usu-

ally sympodial, branches often with swollen or articu-

lated nodes, with simple hairs; stipules paired at each

leaf base, free, persisting or rarely deciduous, sometimes

modified as spines. Leaves opposite or less often alter-

nate, often distichous, usually evenly pinnate, occasion-

ally simple to bifoliolate or 3-7-parted, petiolate; leaf

blades (leaflets) petiolulate or subsessile, inequilateral,

usually entire, stiff to fleshy in texture, often strongly

36

FIELDIANA: BOTANY

resinous. Inflorescences racemose or fasciculate (rarely

cymose), usually with the flowers solitary on axillary

peduncles/pedicels. Flowers bisexual (rarely unisexual

and dioecious), radially symmetrical or rarely somewhat

bilaterally symmetrical, hypogynous, perianth (4-)5(-6)

parted, sepals free or united at the base, imbricate or

valvate in bud; petals imbricate or convolute in bud

(rarely valvate), narrowed at the base, a disc usually

present and often with extra- or intrastaminal glands;

androecium usually with twice as many stamens as the

petals (rarely 3 x or 1 x as many), free, the outer stamens

opposite the petals, filaments with or without append-

ages at the base, anthers 2-thecous, versatile or basifixed,

dehiscing introrsely with longitudinal slits; pistil solitary,

of (2-)4-5(-6) united carpels, ovary sessile or short-stip-

itate (with a gynophore), 2-6-locular, with axile placen-

tation and l-several(-many) ovules in each locule, ovules

pendulous or ascending, style slender and terminal, stig-

ma solitary and lobed or 2-6-parted. Fruit a 2-6-lobed

I ocu 1 icidal or septicidal capsule, or a schizocarp breaking

up into few to several mericarps or nutlets (rarely ber-

rylike or drupaceous), the mericarps often thickened and

spinose to tuberculate on the outer (abaxial) surface; seed

with a straight or curved embryo, cotyledons linear or

oblong, endosperm hard and oily or absent, an aril pres-

ent or absent.

A family of about 25-30 genera and 250 species,

best represented in arid, saline, or seasonally dry

tropical and subtropical environments. The family

consists of somewhat diverse elements, several of

which are often segregated as separate families (such

as Balanites of Africa-Eurasia and Nitraria of the

Old World subtropical deserts). This family has

been classified with both the Geraniales and the

Sapindales; the Rutaceae of the Sapindales may

be the most closely related family.

Key to the Genera of Zygophyllaceae

la. Trees with very hard heavy wood; petals blue or purplish Guaiacum

Ib. Herbs or subshrubs, usually prostrate on the ground; petals white to yellow or orange 2a

2a. Fruiting mericarps armed with spines; each leaf with more than 1 0 leaflets (in our species)

Tribulus

2b. Fruiting mericarps without spines; each leaf with 8 or fewer leaflets (in our species) Kallstroemia

Guaiacum Linnaeus

Small to medium-sized trees, wood very hard, heavy

and resinous, branches often with thickened nodes; stip-

ules small. Leaves opposite, parapinnate, petiolate, leaf-

lets in 2-6 opposite pairs. Inflorescences of fasciculate

flowers at distal nodes, each flower borne on a slender

unbranched peduncle (= pedicel) from the axil of a distal

node. Flowers 4- or 5 -parted, sepals slightly united at

the base, petals 4 or 5, blue or purple, strongly narrowed

at the base (clawed); stamens 8 or 10, filaments slender,

anthers cordate or sagittate at the base; pistil with a short

stipe (gynophore), ovary 2-5-lobed and 2-5-locular,

ovules 8-10 in each locule. Fruits slightly fleshy when

ripe, but drying smooth and coriaceous, with 2-5 prom-

inent longitudinal lobes or rounded ridges (rarely wings);

seeds ovoid to ellipsoid.

A genus of about six species, ranging from

southern Florida, USA, and the West Indies to

northern South America. This is the genus of lig-

num vitae, the hardest of commercial timbers, of-

ten used for construction in salt water. The med-

ical resin guaiacum is obtained from these plants

by heating. Unfortunately, the usefulness of these

trees has resulted in their decimation in many ar-

eas. Guaiacum sanctum grows wild in northwest-

ern Costa Rica, while G. officinale L., with larger

more rounded leaflets, occurs in the West Indies.

These species are sometimes planted as ornamen-

tals for their showy blue flowers.

Guaiacum sanctum L., Sp. PI. 382. 1753. Figure 8.

Small trees up to ca. 10m tall, leafy branchlets 0.8-

3 mm thick, minutely puberulent with thin ascending

hairs ca. 0.2 mm long, becoming pale gray and glabrous;

stipules 2-4 mm long, triangular and acute, appressed

puberulent distally, thick and persisting. Leaves oppo-

site, 3-9 cm long, pinnately compound with (2-)3-5(-6)

pairs of leaflets, petiole 3-8 mm long, petiole and rachis

to 5 cm long, rachis 0.3-0.8 mm thick, minutely puber-

ulent or glabrescent, deeply sulcate above; leaf blades

(leaflets) 12-28(-32) mm long, 4-14(-16) mm broad,

asymmetric and narrowly oblong to oblong-obovate, the

middle leaflets the longest on each leaf, bluntly obtuse

to acute at the apex, subsessile and unequal on the thick

petiolule, with the proximal side usually rounded at the

base and the distal side more straight, leaflets drying

stiffly chartaceous, glabrous above and below or sparsely

sericeous near the base, venation palmate with 1-3 major

veins and more weakly defined lateral veins, all these

veins strongly ascending and loop-connected distally. In-

florescences fascicles of (l-)4-8 flowers at distal nodes,

peduncles (= pedicels) 12-20 mm long, slender (0.3-0.5

mm thick when dry), sparsely and minutely puberulent.

Flowers ca. 15 mm long and 20 mm broad, sepals 4-6

mm long and ca. 3 mm broad, broadly imbricate in bud;

BURGER ET AL.: FLORA COSTARICENSIS

37

Tribulus cistoides

Kallstroema maxima

Kallstroemia pubescens Guaiacum sanctum

FIG. 8. Zygophyllaceae: four Central American species.

38

FIELDIANA: BOTANY

petals 8-12 mm long and 6-8 mm broad, broadly ob-

ovate and clawed at the base, bright blue; stamens ca. 6

mm long, anthers 1.5-2 mm long, becoming curved;

pistil ca. 10 mm long, with a short (2 mm) stipe, and

slender style 2-3 mm long, ovary obovoid and drying

dark. Fruits 14-16 mm long and 12-18 mm broad, ob-

ovoid and prominently 2-5-lobed or ridged, the longi-

tudinal lobes 7-9 mm thick and rounded, fleshy or moist

at maturity but drying to a yellowish and lustrous hard

surface; seeds ellipsoid, ca. 1 cm long, brown to black

and with a red aril.

Trees of the seasonally very dry and deciduous

forest formations in northwestern Costa Rica. The

species is found from 1 0-200 m elevation in Costa

Rica, and up to 700 m in Nicaragua. Flowering

material has been collected in March; fruit has

been collected in July. The species ranges from

southern Florida and eastern Mexico through the

West Indies and Central America to northern South

America.

Guaiacum sanctum is recognized by its opposite

compound leaves with 2-5 pairs of small subses-

sile asymmetric leaflets, the bright sky-blue flow-

ers, and the restriction to lowland deciduous forest

formations. The leaflets resemble those of some

Leguminosae, but the opposite leaves and lobed

capsular fruit are quite different. The hard and

heavy heartwood is said to become bluish on ex-

posure to air (Janzen & Liesner, Brenesia 18: 90.

1980). For a discussion of uses of this species and

the closely related G. officinale see the Flora of

Guatemala (in Fieldiana: Bot. 24(4): 395. 1946,

reprinted 1985).

Kallstroemia Scopoli

REFERENCE— D. M. Porter, The genus Kallstroe-

mia (Zygophyllaceae). Contr. Gray Herb. 198: 41-

153. 1969.

Annual or less often perennial herbs, stems herbaceous

or slightly woody, prostrate or decumbent to ascending,

often spreading outward from a central taproot, terete

and slightly succulent, becoming longitudinally striate

when dry; stipules paired at each leaf-base, free. Leaves

opposite and parapinnate, petiolate, with 2-10 pairs of

opposite subsessile leaflets, the distal leaflets larger and

often somewhat falcate and asymmetric, the basal (prox-

imal) leaflets often unequal in size; leaf blades (leaflets)

entire, pubescent to glabrescent, subsessile on short slen-

der petiolules. Inflorescences represented by usually sol-

itary axillary or pseudoaxillary flowers, each peduncle

(= pedicel) with a single flower. Flowers bisexual and

radially symmetrical, sepals 5(-6), free, pubescent, usu-

ally persisting in fruit; petals 5(-6), free, rotate, white to

bright yellow or orange, convolute in bud, broadly ob-

ovate and rounded distally, narrowed to the clawed base,

fugaceous and quickly withering but often persisting; sta-

mens 10(-12), the outer 5(-6) opposite the petals and

somewhat larger than the inner 5(-6), these latter often

subtended at the base by small lobed glands, filaments

filiform (rarely winged near the base), inserted on the

disc, anthers ovoid to oblong-linear; a fleshy annular disc

present, obscurely 10(-12)-lobed; pistil 1, ovary broadly

sessile, 10(-12)-ribbed and 10(-12)-locular, globose to

conical, glabrous to pubescent, 1 pendulous ovule in each

locule, style simple and cylindrical to conical, persisting

in fruit, stigma capitate to clavate, with 10(-12) ridges

or lobes. Fruits broadly ovoid to conical, with 10(-12)

longitudinal ridges or lobes and the persisting style (beak),

glabrous or puberulent at maturity, breaking septicidally

into 10(-12) or fewer mericarps and those separating

from the persisting central axis, each mericarp with 1

locule and 1 seed, the mericarps wedge-shaped (trian-

gular in cross section) with a rounded tuberculate or

rugose abaxial surface; seeds oblong to ovoid, testa mem-

branaceous, endosperm absent.

An American genus of 1 7 species, ranging from

the southern United States, Mexico, and the West

Indies through Central America to central Argen-

tina. The genus has become naturalized in western

Africa and India. Nearly all the species are found

in arid and seasonally very dry environments, or

in early stages of open secondary succession. The

mericarps are quite unusual in form, and a mer-

icarp may leave an oblong opening in the side of

the fruit if adjacent locules fail to develop. In ad-

dition, the mericarps can be mistaken for seeds.

The two species of Kallstroemia found in Costa

Rica are quite similar in overall appearance; they

appear to be native to Central America. These

plants may be mistaken for Portulaca oleracea L.

(verdolaga or purslane), of similar habit but with

alternate or clustered leaves.

Key to the Species of Kallstroemia

la. Ovary and fruit glabrous or strigose only at the base; sepals lanceolate in fruit, often curved over

the fruit; stipules narrowly lanceolate; leaflets as many as 8 per leaf . K. maxima

Ib. Ovary and fruit usually with appressed thin pilose hairs; sepals linear-lanceolate with involute

margins and usually spreading laterally beneath the fruit; stipules usually linear; leaflets as many as

6 per leaf K- Pubescens

BURGER ET AL.: FLORA COSTARICENSIS

39

Kallstroemia maxima (L.) Hook. & Arnott, Bot.

Beechey Voy. 282. 1838. Tribulus maximus L.,

Sp. PI. 386. 1753. Figures.

Annual herbs or herbaceous vines, stems prostrate to

decumbent, to 1 m long, leafy internodes 1-3 mm thick,

with thin whitish ascending hairs of varying (0.3-1 mm)

length; stipules 3-4 mm long, lanceolate. Leaves 1.2-6

cm long and 1-3.5 cm broad, obovate or oblong in gen-

eral outline, paripinnate with (2-)3-4 pairs of leaflets,

terminal leaflets the largest, a slender tip ca. 1 mm long

often present at the end of the rachis; leaf blades (leaflets)

2-1 6(-l 8) mm long, 1 .5-7(-l 1) mm broad, distal leaflets

strongly asymmetric, oblong-obovate to obovate-falcate,

obtuse at the apex, unequal at the base with the proximal

(outer) side rounded and the distal (inner) side straight

or concave, surfaces with slender appressed hairs ca. 1

mm long, with 1 midvein and several ascending sec-

ondary veins. Inflorescences of solitary axillary flowers,

peduncles (= pedicels) (7-)12-20(-28) mm long, gla-

brescent. Flowers with sepals 4-5 mm long, ca. 2 mm

wide, narrowly ovate, with stiff slender hairs to 1.5 mm

long, persisting, petals 4-6 mm long, to 6 mm wide,

obovate, pale yellow to orange; ovary ca. 1 mm in di-

ameter, style cylindrical, to 2 mm long, stigma capitate.

Fruits broadly ovoid to broadly conical, 6-9 mm long,

4-7 mm broad near the base, pale yellowish white, gla-

brous or strigose at the base, the beak 3-4 mm long and

darker in color, mericarps 3-4 mm high, 1-1.5 mm wide,

abaxially tuberculate and transversely ridged, with pitted

sides.

Plants of open recently cleared ground and early

secondary growth in seasonally dry and deciduous

formations and in evergreen areas; ranging from

sea level to 500(-1400) m elevation. The species

probably flowers throughout the year in Central

America, but has been collected most often in May-

August and in November. The species ranges from

the southeastern United States through Mexico,

Central America, and the West Indies to northern

South America.

Kallstroemia maxima is recognized by the low-

growing habit on open ground, the opposite evenly

pinnate leaves, the 3 or 4 pairs of leaflets with the

distal leaflets largest, and the unusual fruit (see

discussion under the genus).

Kallstroemia pubescens (G. Don) Dandy, in Keay,

KewBull. 10: 138. 1955. Tribulus pubescens G.

Don, Gen. Hist. 1 : 769. 1 83 1 . K. caribaea Rydb.,

in Vail and Rydb., N. Amer. Fl. 25: 111. 1910.

Figure 8.

Annual herbs, stems prostrate to decumbent, up to 1

m long, leafy internodes 0.5-2 mm thick, with thin as-

cending hairs 0.5-1 mm long; stipules 3—4 mm long,

linear, with thin straight hairs. Leaves opposite, to 4.5

cm long and 2.5 cm broad, often rectangular to obovate

in outline, paripinnately compound with 2-3 pairs of

subsessile leaflets, the terminal leaflets the largest, peti-

olules ca. 0.5 mm long; leaf blades (leaflets) 5-14(-20)

mm long, 3-8(-10) mm broad, proximal (lower) leaflets

broadly oblong or ovate-oblong, distal leaflets asym-

metric and oblong-falcate or obovate-falcate, rounded

at the apex or with an apiculate tip, unequal at the base

with the proximal (outer) part rounded and the distal

(inner) side straight or convex, with straight slender stiff

appressed ascending hairs ca. 0.5-1 mm long on the

surfaces and along the edges, midvein with 1-3 diverging

secondary veins on each side. Inflorescences represented

by solitary, axillary flowers, peduncles (= pedicels) 5-15

mm long. Flowers with sepals 3.5-5 mm long, 0.5-1.5

mm broad, lanceolate, with thin hairs ca. 1 .2 mm long;

petals ca. 6 mm long and 5 mm broad, obovate, white

to yellow, withering but persistent; pistil 4-6 mm long,

ovary broadly ovoid to conical, densely appressed-pu-

berulent with fine white hairs, style 3—4 mm long, gla-

brous distally. Fruits 5-6 mm in diameter near the base,

with 10 longitudinal ridges, the ridges with tubercles or

transverse striations, with dense short hairs, mericarps

3-4 mm long and 1 mm thick, the subtending persisting

sepals linear-lanceolate with involute margins.

Plants of open sunny early successional sites in

seasonally dry deciduous formations and in ev-

ergreen areas, from sea level to 200 m elevation

in Central America. Flowers and fruits have been

collected in March-October in Central America.

The species ranges from central Mexico through

Central America and the West Indies to Colombia,

Venezuela, Ecuador, and Peru. This species has

become naturalized in West Africa and India.

Kallstroemia pubescens is recognized by the low

habit on open ground, the paripinnate leaves with

only four or six leaflets, the yellow flowers, the

pubescent ovary and fruit, and the unusual mer-

icarps. All our Costa Rican collections come from

lowland Guanacaste.

Tribulus Linnaeus

Annual herbs (occasionally perennial, sometimes

woody at the base, rarely small shrubs), plants diffusely

branched and often spreading radially from a central

taproot, prostrate to decumbent or ascending, stems her-

baceous to slightly woody, often somewhat succulent,

terete and longitudinally striate on drying, densely pu-

berulent to glabrescent; stipules paired at the leaf-bases,

leaflike. Leaves opposite with one of each pair usually

shorter than the other, occasionally alternate when an

opposing leaf fails to develop, evenly pinnate (paripin-

nately compound) with 3-10 pairs of opposite leaflets

(rarely 1 leaflet of a pair aborting), petiolules very short

or the leaflets subsessile, rachis often extending beyond

the terminal pair of leaflets; leaf blades (leaflets) oblong

to ovate-oblong or elliptic, often somewhat oblique and

asymmetric at the base, the terminal pair usually point-

40

FIELDIANA: BOTANY

ing forward, puberulent with thin straight whitish hairs,

with a single midvein. Inflorescences represented by sol-

itary axillary flowers (usually from the axil of the smaller

leaf of the node), borne on long peduncles (= pedicels).

Flowers bisexual, radially symmetrical, sepals 5, free,

puberulent, caducous or sometimes persisting; petals 5,

free, bright yellow (white), spreading to rotate, imbricate

in bud, narrowed at the base, deciduous; stamens 10

(rarely 5), the 5 outer longer and united to the base of

the opposing petals, the 5 inner (antesepalous) stamens

with nectariferous glands at the base, anthers cordate to

oblong, interstaminal glands free or connate beneath the

base of the ovary; pistil sessile, globose to ovoid, 5-lobed

and 5-locular, densely puberulent, with 3-5 lobules in

vertical rows on axile placentae in each locule, style cy-

lindrical, stigma 5-angular or 5-lobed, pyramidal to ob-

long. Fruits usually breaking up into 5 (or fewer) hard

mericarps but not leaving a central axis, mericarps broadly

triangular, each mericarp divided internally by trans-

verse septa to produce 2-5 1 -seeded locules, the meri-

carps with hard spines or wings (or tuberculate) on their

outer (abaxial) surface; seeds oblong-ovoid, testa mem-

branaceous, endosperm absent, embryo straight.

A genus of about 25 species of seasonally dry

habitats and desert communities. Originally re-

stricted to the Old World; a few species have be-

come widespread and are now found throughout

the tropics and warm-temperate areas of the world.

The genus is recognized by the bright yellow flow-

ers with free parts, the somewhat unequal usually

opposite leaves with (3-)5-10 pairs of small (4-

1 5 mm) opposite subsessile leaflets, and the fruit

breaking up into hard spiny mericarps. Two spe-

cies (T. cistoides L. and T. terrestris L.) have been

found in Central America, but neither has been

collected in Costa Rica. The following key distin-

guishes these two distinctive species.

Key to the Species of Tribulus Collected in Central America

la. Rowers borne on peduncles usually longer than the leaves, sepals 5-12 mm long; intrastaminal

glands united and forming a lobed cup at the base of the ovary; stigma oblong, on a short prominent

style T. cistoides

Ib. .Flowers borne on peduncles equaling or shorter than the leaves, sepals 3-6 mm long; intrastaminal

glands free; stigma hemispheric, subsessile T. terrestris

Literature Cited

CRONQUIST, A. 1981. An Integrated System of Clas-

sification of Flowering Plants. Columbia University

Press, New York, 1262 pp.

SIMPSON, B. B. 1982. Krameria flowers: Orientation

and elaiophore morphology. Taxon, 31: 517-528.

STANDLEY, P. C. 1937-1938. Flora of Costa Rica. Publ.

Field Mus. Nat. Hist., Bot. Ser., 18: 1-1571.

BURGER ET AL.: FLORA COSTARICENSIS

41

Index

The index includes all accepted names (in Roman type), synonyms (in italics), and vernacular names

(Spanish names in italics). The page numbers of illustrations are in boldface. Hyphenated words and

multiple words are alphabetized by letter.

Acedera 8, 1 1

Acederilla 1 1

Aneulophus 3 1

ami 21

Averrhoa 4

bilimbi 4

carambola 5

Balanites 37

Biophytum 5

dendroides 6, 7

falcifolium 6, 7

panamense 6

campano 29

canary-bird flower 2 1

capuchina 23

carambola 5

Centris bees 1

chiricano 28, 30

chiricano trieste 30

coca 36

cocaine 36

danto plomillo 29

Dirachmaceae 16

Erodium 17

cicutorium 17

moschatum 17

espuela del gelan 23

Erythroxylaceae 30

Erythroxylum 31

brevipes 36

chiapense 33

citrifolium 32

citrifolium var. minus 32

coca 36

coca var. novogranatense 35

compactum 36

costaricense 33, 35

cumanense 33

ellipticum 33

fimbriatum 32

fiscalense 36

havanense 33, 34

havanense var. continents 33

hondense 33

lucidulum var. costaricense 33

lucidum 33

macrophyllum 33

macrophyllum var. ecuadorense

35

macrophyllum var. macrocne-

mium 35

mexicanum 33

multiflorum 33, 35

novogranatense 35

obovatum 36

obtusum 33

ovatum 33

pallidum 36

pringlei 33, 36

rotundifolium 36

sessiliflorum 36

skutchii 33, 35

suave 36

swave var. compactum 36

tabascense 33

tikalense 36

flax 25

forage 16

garden geranium 21

garden nasturtium 2 1

Geraniaceae 16

geranio 21

Geranium 17

costaricense 18, 19

cucullatum 18

cucullatum var. multifidum 1 8

guatemalense 19, 20

mexicanum 20

multifidum 18

pulchrum 20

repens 20

Guaiacum 37

officinale 37

sanctum 37, 38

Hugoniaceae 24

Humiriaceae 25

Humiriastrum 25

diguense 26, 27

diguense var. costaricense 26

Hypseocharis 4

/ra chiricana 30

Ixonanthaceae 24

Kallstroemia 39

caribaea 40

maxima 38, 40

pubescens 38, 40

Krameria 1

cuspidata 2

ixine 2

revoluta 2

Krameriaceae 1

laurelito 28

Linaceae 23

linaza 25

linaza lino 25

///jo 25

linseed 25

linseed oil 25

Linum 24

guatemalense 24

usitatissimum 25

lorito 28, 29

Magallana 2 1

manteca 29

mastuerzo 23

medicinal plants 1, 23

mimbro 5

nasturtium 23

Nectaropetalum 30

nispero 28, 30

Nitraria 37

Oxalidaceae 2

Oxalis 6

angustifolia 12

articulata 9

barrelieri 9, 13

bradei 12

chiriquensis 14

coccinea 12

corniculata 10, 1 1

darienensis 12

debilis 10, 1 1

dendroides 6

deppi 16

dombei 12

filiformis 10, 12

frutescens subsp. angustifolia 12,

13

galeotii 14

hayi 166

latifolia 10, 14

martiana 1 1

maxonii 15

microcarpa 14

42

FIELDIANA: BOTANY

neaei 12 diguense 26 moritzianum 23

ramonensis 14 excelsa 28 peltophorum 21

rhombifolia 13, 15 holdridgei 27, 28 pendulum 23

spiralis 1 5 ovicarpa 29 peregrinum 2 1

spiralis subsp. vulcanicola 13, 1 5, trichogyna 27, 29 tuberosum 2 1

16 warscewiczii 23

tetraphylla 16 Trophaeastrum 2 1

vulcanicola 1 5 tiriguro 5

Tribulus 40 Vantanea 29

Pelargonium 20 cistoides 38, 41 barbourii 27, 30

graveolens 21 maxima 40 occidentalis 30

x hortorum 2 1 pubescens 40 tuberculata 29

Portulaca oleracea 39 terrestris 4 1 vinagrillo 8

Tropaeolaceae 21 Vivianiaceae 16

Tropaeolum 21

rosita 29 emarginatum 22

guatemalense 22 Zygophyllaceae 36

majus 22

Sacoglottis 28 minus 2 1

amazonica 29

BURGER ET AL.: FLORA COSTARICENSIS 43

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