Botany

FLORA COSTARICENSIS

William Burger, Editor

Family #80 Lauraceae

William Burger

Henk van der Werff

Family #81 Hernandiaceae

W illiam Burger

January 31, 1990

Publication 1406

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

171 i

-

,

FIELDIANA

Botany

NEW SERIES, NO. 23

FLORA COSTARICENSIS

William Burger, Editor

Family #80 Lauraceae

William Burger Henk van der Werff

Curator Missouri Botanical Garden

Department of Botany St. Louis, Missouri 63166

Field Museum of Natural History

Chicago, Illinois 60605-2496

Family #81 Hernandiaceae

William Burger

Accepted June 1, 1988

Published January 31, 1990

Publication 1406

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

© 1990 Field Museum of Natural History

Library of Congress Catalog Card Number: 78-172358

ISSN 00 15-0746

PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

List Of Illustrations

INTRODUCTION v

ACKNOWLEDGMENTS v

LAURACEAE by William Burger and Henk

van der Werff 1

Diagnostic Key to Genera of Lauraceae ... 3

Artificial Key to Woody Genera and Species

4

Key to Comparative Figures of Costa Ri-

can Lauraceae 13

Aiouea 36

Aniba 38

Beilschtniedia 39

Caryodaphnopsis 43

Cassytha 44

Cinnamomum 45

Endlicheria 45

Licaria 46

Litsea 53

Nectandra 54

Ocotea 67

Persea 101

Phoebe 109

Pleurothyrium 114

Povedadaphne 117

Williamodendron 118

A Species of Uncertain Generic Position

119

List of Accepted Species and Acronyms . 1 20

Index to Exsiccatae 121

HERNANDIACEAE by William Burger ... 129

LITERATURE CITED 135

INDEX 1 36

LAURACEAE

1 . Tripliveined and palmately veined

species 14

2. Large-leaved species 15

3. Species with hollow distal stems 16

4. Montane species with small leaves 17

5. Montane species with stiff coriaceous

leaves 18

6. Montane species with puberulent leaves

19

7. Lower elevation spp. with puberulent

leaves 20

8. Lower elevation spp. with puberulent

leaves 21

9. Lower elevation spp. with puberulent

leaves and two species of Persea 22

10. Species with decurrent lamina bases

(larger) 23

1 1 . Species with decurrent lamina bases

(smaller) 24

12. Montane species, laminae often lustrous

above 25

13. Glabrous leaves often drying dark or

grayish 26

14. Glabrous leaves often drying gray or

greenish 27

15. Species of Beilschmiedia and Persea, the

fruits not subtended by a cup 28

16. Species of Licaria, the fruit cup usually

with a double margin 29

1 7. Nectandra salicifolia and similar species

30

18. Nectandra globosa and similar species

31

19. Williamodendron glaucophyllum 32

20. Ocotea dentata 33

2 1 . Persea silvatica 34

22. Povedadaphne quadriporata 35

HERNANDIACEAE

23. Species of Hernandia, Gyrocarpus, and

Sparattanthelium 1 30

in

Introduction

This is the sixth issue of "Flora Costaricensis."

The first dealt with the Piperaceae (Fieldiana, Bot.

35, 1971). The second included families numbered

42 through 53, Chloranthaceae through Urtica-

ceae (Fieldiana, Bot. 40, 1977). The third issue

covered the Gramineae and was authored by Rich-

ard Pohl (Fieldiana, Bot., new series, No. 4, 1980).

The fourth issue included families numbered 54

through 70, Podostemaceae through Caryophyl-

laceae (Fieldiana, Bot., new series, No. 13, 1983).

The fifth issue covered families 200 and 201, the

Acanthaceae by L. H. Durkee and the Plantagi-

naceae (Fieldiana, Bot., new series, No. 18, 1986).

In the figures of Lauraceae, fruits are illustrated

along the left side over a grid of square centimeters.

Leaves and twigs of each full-page figure are drawn

to the same scale, shown with a horizontal cen-

timeter bar. Stamens are depicted along the right

at varying scales in millimeters. The stamens shown

are part of the outer whorls (series I-II), except in

Licaria. The drawings are based on dried herbar-

ium specimens. The stamens are drawn from boiled

flowers and are diagrammatic; puberulence may

not be accurately represented.

Acknowledgments

We are especially grateful for the financial as-

sistance from the National Science Foundation,

which has aided this program for many years, both

at Field Museum and in Costa Rican fieldwork.

The program was supported most recently by NSF

grant DEB-8103184, through the Biological Re-

search Resources Program. A recent grant from

the National Geographic Society (#3465-86) sup-

ported fieldwork and a review of the Lauraceae in

Costa Rican herbaria which led to the discovery

of a new genus.

The staff and the facilities of the Museo Na-

cional de Costa Rica have been a central resource

and most helpful to our work for more than two

decades. Pablo Sanchez V., in charge of the Nat-

ural History section and the Herbario Nacional,

has given our work on Lauraceae much support.

Jorge Gomez-Laurito provided access to the her-

barium of the Universidad de Costa Rica and has

been very helpful in many other ways. Luis Poveda

led two successful field trips specifically focused

on collecting Lauraceae. Recent collections made

by programs of the Missouri Botanical Garden

have added greatly to our knowledge of the Lau-

raceae, as has the Flora of La Selva project. We

thank the Missouri Botanical Garden for allowing

Henk van der Werff to contribute his time and

effort to our treatment of the Lauraceae.

Finally, we thank the late Timothy Plowman,

Jens Rohwer, Jorge Gomez-Laurito, and two

anonymous reviewers for the many corrections and

improvements they provided for an earlier draft

of this manuscript. However, despite their efforts

and our own, many problems remain in our pre-

sentation and in our understanding of the plants

in these two families.

FLORA COSTARICENSIS

Family #80 Lauraceae

Family #81 Hernandiaceae

LAURACEAE

By William Burger and Henk van der Werff

REFERENCES— C. K. Allen, Studies in the Lau-

raceae, VI. Preliminary survey of the Mexican and

Central American Species. J. Arnold Arbor. 26:

280-434. 1945. L. Bernardi, Lauraceas. 355 pp.

Universidad de los Andes, Facultad de Ciencias

Forestales, Merida, Venezuela. 1962. B. Hammel,

New species and notes on Lauraceae from the Ca-

ribbean Lowlands of Costa Rica. J. Arnold Arbor.

67(1): 123-136. 1986; The vascular flora of La

Selva Biological Station, Costa Rica. Lauraceae.

Selbyana 9: 218-233. 1986. A. J. G. H. Koster-

mans, Lauraceae. Reinwardtia 4: 193-256. 1957;

Bibliographia Lauracearum. 1 ,450 pp. Ministry of

Natl. Research, Bogor, Indonesia. 1964. C. Mez,

Lauraceae Americanae. Jahrb. Konigl. Bot. Gart.

Berlin 5: 1-556. 1889. J. G. Rohwer, Prodromus

einer Monographic der Gattung Ocotea Aubl.

(Lauraceae), sensu lato. Mitt. Inst. Allg. Bot. Ham-

burg 20: 1-278. 1986.

Medium-sized trees, less often tall canopy trees or

shrubs (slender twining parasites with yellowish stems

in Cassythd), bisexual or unisexual (dioecious); the wood

often yellowish, the shoot apex usually with minute lus-

trous appressed-ascending hairs, the hairs always simple

and usually unicellular, stems glabrous or puberulent,

often with aromatic oils in bark and leaves; stipules ab-

sent. Leaves alternate, less often fasciculate or whorled,

rarely consistently opposite (as in Caryodaphnopsis), ev-

ergreen or deciduous, new leaves often produced in flush-

es, always simple, usually petiolate, the petioles often

with adaxial or lateral margins continuous with the lam-

ina margins, sulcate to flat or rounded on the adaxial

surface, rarely clasping the stem; leaf blades always sim-

ple and entire, occasionally undulate (lobed in Sassafras)

but never crenate, serrate or incised, often dark green

and lustrous above in life, never scabrous above, some-

times glaucous beneath, usually stiffly chartaceous to co-

riaceous in texture, the venation pinnate to tripliveined

(rarely palmate), often with veinlike tissue along the edge

of the lamina, glabrous to densely puberulent beneath,

"domatia" of tufted hairs or pits sometimes present in

the axils of proximal veins on the abaxial (lower) surface,

the leaves often turning yellow or orange before falling.

Inflorescences usually solitary and axillary, sometimes

fasciculate or pseudoterminal, on distal branchlets, usu-

ally paniculate with a prominent peduncle and central

rachis, less often racemose or umbellate (very rarely spi-

cate or capitulate), distal flower groups often cymose or

umbellate and subtended by bracteoles, the flowers ses-

sile or more often pedicellate, (the inflorescence enclosed

at first in broad overlapping bracts in a few genera, cf.

Litsea). Flowers bisexual or less often functionally uni-

sexual (rarely with the unisexual flowers lacking pistil-

lode or staminodes), radially symmetrical, minute (1-3

mm) to small (3-12, rarely 20 mm broad), white to yel-

lowish or greenish (rarely pinkish to red), hypogynous

to perigynous with the development of a floral cup or

tube; perianth usually of 6 parts in 2 whorls of 3 tepals

each, the parts equal or subequal (the outer much shorter

than the inner in Caryodaphnopsis and a few species of

Persea, among Costa Rican species), rarely the perianth

of 4 or 9 parts in 2 or 3 whorls, free or united at the base

above the floral cup, glabrous or puberulent on either or

both surfaces; androecium usually of 9 (rarely 1 2) func-

tional stamens in 3(-4) whorls, the outer 2 whorls (series

I-II) of 6 stamens usually similar and appearing as a

single whorl (absent in Licaria and Williamodendrori),

the outer 6 stamens free and with introrse dehiscence

(dehiscence lateral or variable in Pleurothyrium), the sta-

mens sessile or with filaments, anthers narrow and rect-

angular to broad, flat and tepal-like, 4-thecous or 2-the-

cous and dehiscing by 4 or 2 valves (flaps), valves opening

from the base to the apex, the inner 3 stamens (series

III) usually with 2 glands at the base of each filament,

inner stamens free (united in some Licaria sp.) and usu-

ally dehiscing extrorsely, less often laterally or apically,

a whorl of 3 staminodes (series IV) present and with

well-developed apex or minute or absent; pistil simple

and solitary, often narrowed at the base, borne above

BURGER: FLORA COSTARICENSIS

the receptacle or within the floral cup or tube, free and

superior to perigynous (united to the tube and inferior

only in Hypodaphnis of West Africa), the ovary with a

single locule and solitary pendulous anatropous ovule,

the style short or long, the stigma simple to discoid or

capitate (rarely deeply lobed). Fruits borne on a thick-

ened pedicel (as in Beilschmiedia, Caryodaphnopsis, and

Persea) or subtended by a flattened receptacle or borne

in a cupulate receptacle, the perianth parts deciduous or

persisting but not enlarging in fruits, the cup fleshy or

becoming woody, the margin entire or undulate, rarely

with multiple ridges (as in Licaria), often red-colored at

maturity in contrast to the green to blackish fruits; fruits

1 -seeded berries, usually ellipsoid to ovoid or globose,

often flattened at the base and abruptly rounded at the

apex, the style base rarely persisting, exocarp usually

glossy and smooth, often becoming black or purplish,

mesocarp succulent and fleshy; the seed without endo-

sperm, the cotyledons large, flat on the inner faces and

convex on the outer side, longitudinally parallel with the

axis of the pedicel, white or sometimes pink within.

A family of about 2,000 species of trees (only

Cuscuta contains herbaceous parasites), abundant

in the evergreen tropics and subtropics, with a few

species in seasonally very dry and temperate re-

gions. The family's greatest diversity is in South-

east Asia and in South America. The simple, al-

ternate, stiff, entire, aromatic leaves (often dark

green and glossy above in life), the lack of stipules,

6-parted perianth (sometimes irregular in number

in Litsea), nine free stamens (in most), the anthers

that always open by valves (flaps), the simple pistil

with single style and stigma, solitary pendulous

ovule, and fruits often borne in a cup make the

Lauraceae a very distinctive family. The petioles

are often sulcate above; the shoot apex is usually

covered with small (0.1-0.5 mm), slender, ap-

pressed-ascending, lustrous simple hairs; and the

bark and foliage are usually aromatic. Small

"domatia" are present on the lower leaf surfaces

of many species. They are too small to accom-

modate ants; they appear to be associated with

mites (cf. Pemberton & Turner, 1 989).

The family is an important component of rain

forests and cloud forests in Costa Rica. Only about

six species occur above 2600 m elevation, and a

similar number are found in the seasonally very

dry lowlands of Guanacaste. A number of species

are important sources of timber, and the avocado

(Persea americana) is widely cultivated for its nu-

tritious fruits. Cinnamomum camphora is a source

of camphor and C verum is the source of cinna-

mon, but they are only occasionally cultivated in

Central America.

The Lauraceae are one of the most difficult fam-

ilies in the Neotropics as regards the identification

of genera and species (Burger, 1988). Some of the

genera are artificial and linked by intermediate

species, but better ways of organizing the species

are not apparent. We disagree with the submer-

gence of some of these poorly defined genera at

this time (cf. Howard, 1981; Kostermans, 1957).

We believe that major taxonomic changes must

reflect an improved understanding of relationships

and should result in better systems of information

retrieval. In addition, the perspectives gained in

our study of Costa Rican species are insufficient for

making decisions regarding generic circumscrip-

tion. Because many species are large trees with

small flowers, their representation in herbarium

collections is poor, making the delimitation of

species difficult. A number of species groups are

very difficult to interpret, and the treatments pre-

sented here can only be considered tentative; see

the discussions under the species.

Costa Rica differs from neighboring areas in

having had a long tradition of resident botanical

collectors. The collections of Paul Allen, Alberto

Brenes, Gary Hartshorn, Leslie Holdridge, Alfon-

so Jimenez, Luis Poveda, Alexander Skutch, and

Austin Smith have been especially important in

working with this family. Determinations by

Holdridge and Poveda were very helpful during

the early stages of our work. Barry Hammel's col-

lection and study of Lauraceae at La Selva has

been a major contribution. We thank the following

herbaria for access to their collections or loans for

this study: AA, CR, DUKE, F, GH, MO, NY, us, usj.

The following keys can be used to identify flow-

ering collections in which the androecium and

fruiting condition are known (Key 1, Diagnostic

Key to Genera), or in which fruit and some floral

morphology are apparent (Key 2, Artificial Key to

Woody Genera and Species). In addition, there is

a key which should aid in use of the figures (Key

3, Key to Comparative Figures); these are grouped

according to vegetative similarity and altitudinal

range. Identification of individual specimens can

be very difficult. Individual trees of the same species

can differ greatly in some characteristics, and

species with very similar foliage can have signif-

icantly different flowers and fruits. Ultimately, the

most certain method of identifying a specimen is

by careful comparison with properly identified

herbarium collections.

FIELDIANA: BOTANY

Key 1: Diagnostic Key to Genera of Lauraceae

la. Slender twining parasites with yellowish to orange or dull green stems 1-3 mm thick, attached to

small shrubs and herbs by haustoria; leaves reduced to scales; each flower with 9 stamens opening

by 2 valves; fruits enclosed in a perianth tube Cassytha

1 b. Trees and shrubs with woody stems and green leaves, not parasitic; mature fruits not completely

enclosed in a tube 2a

2a. Flowers with only 3 stamens, the stamens free or united and forming a central column around the

style (in Licaria spp.), flowers only 1-3.5 mm long 3a

2b. Flowers with 6, 9, or 12 stamens, or with 6, 9, or 12 staminodes in female flowers, stamens rarely

connivent and united only at the base, flowers 1-15 mm long 4a

3a. Each stamen 2-thecous and opening laterally or distally with 2 valves (each flower with 6

valves), the staminodes exterior to the stamens and not sagittate; fruits borne in a deep cup,

the cup often with a double-margined rim; leaves never obovate in our species, not closely

clustered distally Licaria

3b. Each stamen with 4 thecae and opening distally with 4 minute valves (each flower with 1 2

small valves), with 3 sagittate staminodes interior to the stamens; fruits unknown; leaves

large and obovate, usually closely clustered distally \Villiamodendron

4a. Each stamen opening by 2 valves, anthers 2-thecous 5a

4b. Each stamen opening by 4 valves, anthers 4-thecous (rarely with a few stamens with only 2 valves)

8a

5a. Fruits subtended by a thickened pedicel, a fruit cup or disclike receptacle absent; staminodes

present and conspicuous, stamens often with the connective slightly prolonged; stigma simple;

dried leaves usually with the minor venation forming an elevated reticulum

Beilschmiedia

5b. Fruits subtended and partly enclosed by a fruit cup or disc; staminodes absent or slender,

connective rarely prolonged beyond the thecae; stigma simple or discoid 6a

6a. Flowers functionally unisexual and the trees dioecious; leaves persistently puberulent in the

Costa Rican species placed here Endlicheria

6b. Flowers bisexual, trees bisexual; leaves glabrous or glabrescent (in Costa Rica) 7a

7a. Flowers glabrous in Central American species; staminodes present or absent, outer stamens

with narrow short filaments; ovary ellipsoid to ovoid and borne in an open shallow cup;

margin of the fruit cup entire or with persisting perianth lobes Aiouea

7b. Flowers puberulent; staminodes absent, outer stamens lacking a differentiated filament and

puberulent; ovary very slender ellipsoid to ovoid and included within the narrow floral tube;

margin of the fruit cup entire Aniba

8a. Inflorescences pedunculate umbels, the umbel of flowers at first enclosed in an involucre of broadly

imbricate bracts and resembling a flower bud on a pedicel; stamens 9 or 1 2 Litsea

8b. Inflorescences paniculate to racemose, rarely umbellate and never involucrate; stamens or stam-

inodes 9 (rarely 6) 9a

9a. Stamens with 4 small distal valves which open apically, stamens thick and hairy with a filament

not clearly differentiated; fruits over 6 cm long, subtended by a small disc or cup

Povedadaphne

9b. Stamens opening by 4 lateral (usually vertical) valves, the anthers and filaments usually clearly

differentiated; fruits 6 cm long only in Persea and lacking a basal cup lOa

lOa. Outer stamens adjacent to large glands which are a part of the periphery of the androecium, the

stamens variously bent and usually with lateral dehiscence, outer stamens with the lower pair of

valves dehiscing lateral-extrorse, stamens and glands often tightly congested; fruits a deep cup .

Pleurothyrium

1 Ob. Outer stamens and the periphery of the androecium without conspicuous glands, the glands present

only near the bases of the inner stamens, the outer stamens with all valves introrse, stamens and

glands tightly congested only in Nectandra 1 la

1 la. Outer stamens with the thecae superposed, the upper valves directly above the lower valves, rarely

BURGER: FLORA COSTARICENSIS

with the lower valves somewhat lateral to the upper valves (in an arcuate arrangement) and then

usually with some outer anthers with superposed thecae; filaments shorter than the anthers 1 2a

1 1 b. Outer stamens with the thecae in a single horizontal row or with the lower lateral to the upper in

an arcuate arrangement, anthers usually broader than long 1 5a

12a. Fruits borne on a thickened pedicel (lacking a disclike or cupulate receptacle), tepals equal

or unequal in length, the tepals often persisting at the base of the fruits; stamens with slender

puberulent filaments often exceeding the length of the anthers, staminodes large and sagittate;

surfaces of the dried leaves with a raised reticulum in some spp Persea

1 2b. Fruits borne in cups or on a disclike expansion of the receptacle, tepals deciduous or persisting

on the margin of the cup; the tepals equal or subequal in length; filaments usually equalling

or shorter than the anthers 1 3a

1 3a. Flowers without staminodes or the staminodes small and linear, staminodes lacking a sagittate

or thickened apex and not consistently 3 per flower; leaves rarely tripliveined .... Ocotea

1 3b. Flowers with 3 conspicuous staminodes, the apex of the staminodes cordate-ovate or sagittate;

leaves often tripliveined 1 4a

14a. Trees and shrubs of native vegetation; crushed bark not smelling like camphor or cinnamon

Phoebe

14b. Trees and shrubs of parks and gardens; crushed bark smelling like camphor or cinnamon .

Cinnamomum

1 5a. Leaves alternate and never tripliveined; perianth whorls equal or subequal; stamens often subsessile

and crowded close together; fruits enclosed in a cup or subtended by a disc Nectandra

15b. Leaves opposite and tripliveined; outer perianth whorls smaller than the inner; stamens with

conspicuous filaments; fruits borne on a thickened pedicel Caryodaphnopsis

Key 2: Artificial Key to Woody Genera and Species

(Measurements and leaf color based on dried material; leaf lengths and widths do not include petiole

length.)

la. Leaves densely puberulent beneath with conspicuous hairs (0.3-1 mm long), the hairs spreading

or appressed, the puberulence of the abaxial (lower) leaf surface soft or slightly rough to the touch

2a

1 b. Leaves glabrous to sparsely and minutely puberulent beneath, if densely puberulent the hairs minute

(0.05-0.2 mm) and difficult to see, puberulence of the lower leaf surfaces not discernable to the

touch 29a

2a. Largest leaves rarely more than 1 0 cm long 3a

2b. Largest leaves usually more than 1 3 cm long 7a

3a. Hairs spreading; leaf base not decurrent on the petiole, leaves drying chartaceous; fruit

cups 6-12 mm broad, fruits 15-25 mm long; flowers usually with well-developed

staminodes 4a

3b. Hairs appressed and usually parallel on the lower leaf surfaces, lamina base slightly

decurrent on the petiole, laminae often drying subcoriaceous; fruit cups 10-16 mm

broad, fruits 20-30 mm long; flowers lacking staminodes 5a

4a. Hairs slightly rough to the touch on the lower surface; laminae drying yellowish

to dark brown or black; 1400-3200 m elevation Ocotea pittieri

4b. Hairs soft to the touch and slightly grayish on the lower surface; laminae usually

drying dark brown above; 1400-2300 m elevation Ocotea mollicella

5a. Perianth of 2 different sizes and persisting beneath the globose fruits, a fruit cup not

developed; leaves narrowly elliptic-oblong, to 2.5 (3.5) cm broad; 1000-3300 m ele-

vation Persea spp.

5b. Perianth whorls equal or subequal; fruits subtended by a cup; 1400-3000 m elevation

6a

6a. Leaves usually elliptic with an acute apex, 1 .5-4 cm broad, the lower surfaces remaining

FIELDIANA: BOTANY

densely puberulent; (900-) 1400- 1600 m in the Cordillera de Tilaran and western Mes-

eta Central Ocotea monteverdensis

6b. Leaves usually oblong with an obtuse apex, 1.5-6 cm broad, the lower surface often

becoming glabrescent; 1 700-3000 m in the central highlands and Cordillera de Tala-

manca Ocotea austinii

7a. Margin of the leaf base often broadly revolute to form 1 or 2 pocket-like flaps on the lower

surface just above the petiole, leaves up to 36 cm long, usually long-acuminate at the apex,

with 6-16 pairs of major secondary veins; fruits borne in shallow cups; stamens tepal-like

Nectandra reticulata

7b. Margin of the leaf base flat to narrowly revolute near the petiole beneath but not forming

flaps or auriculae, laminae rarely long acuminate at the apex, with 5-10(-12) pairs of major

secondary veins 8a

8a. Base of the leaf blade decurrent on the petiole and the petiole often poorly defined, margin

of the leaf base often strongly revolute 9a

8b. Base of the leaf blade not clearly decurrent on the petiole 1 3a

9a. Trees of montane formations above 1 400 m elevation; leaf blades elliptic to oblong

(rarely obovate) lOa

9b. Trees of lower elevations (below 1 200 m) in Costa Rica; leaf blades often slightly

obovate 1 la

1 Oa. Puberulence of the lower leaf surface silky and lustrous; leaves subsessile with

poorly differentiated petiole, stiffly coriaceous; only found above 2000 m elevation

Ocotea calophylla

1 Ob. Puberulence of the lower leaf surface not lustrous; leaves petiolate and chartaceous

to coriaceous; go back to 5a

I la. Petioles well defined (the leaf base not long-decurrent), leaves drying stiffly chartaceous

and with 4-8 pairs of major secondary veins; fruits borne in a deep cup 10-16 mm

broad Ocotea hartshorniana

I 1 b. Petioles poorly defined, leaf base decurrent on the petiole, leaves drying subcoriaceous

and with 9-12 pairs of secondary veins 1 2a

1 2a. Fruits in a deep cup with lobed rim, berry ellipsoid or oblong Ocotea dentata

1 2b. Fruits on a shallow saucer-like cup, berry globose Ocotea stenoneura

1 3a. Inflorescences with densely puberulent, peduncles to 1 6 cm long; laminae often oblong and

abruptly rounded at both apex and base; fruits puberulent near the base; flowers becoming

rotate and 1-2 cm broad, often pink; trees of seasonally dry forest formations

Nectandra sinuata

1 3b. Inflorescences with peduncles to 9 cm long or if longer growing in wet forests; fruits glabrous;

flowers rarely more than 1 2 mm broad, never pink 14a

14a. Trees from 1000 m elevation or above 1 5a

14b. Trees from below 1000 m elevation 19a

1 5a. Petioles 1 5-60 mm long; fruits 4-10 cm in diameter and never subtended by or included

in a cup; outer stamens 3-5 mm long; leaves drying chartaceous to subcoriaceous;

inflorescences much branched 1 6a

15b. Petioles 4-25 mm long; fruits 1.5-2.5 cm in diameter, subtended or included in a cup;

outer stamens 1.2-2.5 mm long; leaves usually drying subcoriaceous; inflorescences

often with few or widely distant flowers 1 7a

1 6a. Flowering pedicels 1 0-26 mm long; leaves usually bluntly rounded at the apex,

densely ferruginous-tomentellous beneath; fruits globose . . . Persea schiedeana

1 6b. Flowering pedicels 2-5 mm long; leaves usually acute to short-acuminate or obtuse

at the apex, sparsely puberulent beneath; fruits globose to pyriform

Persea americana

1 7a. Flowers glabrous on the outside; fruits borne in a cup with entire margins; leaves usually

widest at or above the middle; 1 200-2200 m elevation Ocotea valeriana

1 7b. Flowers densely puberulent on the outside; fruit cups often with lobes on the margins;

800-1600 m . . 18a

BURGER: FLORA COSTARICENSIS

1 8a. Laminae usually widest above the middle, often obovate; hairs of lower leaf surface

less than 0.4 mm long Pleurothyrium palmanum

18b. Laminae usually widest at the middle, often suborbicular; hairs of lower leaf surface

more than 0.5 mm long Ocotea gomezii

1 9a. Inflorescences pendulous on long (to 1 5 cm) thin (1-2 mm) peduncles with conspicuous long

(1-2 mm) thin straight hairs; perianth usually glabrous on the outside; fruits borne in a

shallow cup; leaves often narrowly obovate to pandurate, acuminate at the apex and often

slightly rounded at the base Ocotea helicterifolia

19b. Inflorescences pendulous only in fruit, the peduncles not so thin, hairs rarely more than 1

mm long; perianth usually puberulent on the outside; laminae obovate and rounded at the

base only in Nectandra belizensis and Ocotea valerioides 20a

20a. Fruits avocado-like, more than 5 cm long, never subtended by a cup or saucer-like disc;

leaves sparsely puberulent beneath, leaves often broadly elliptic-oblong to ovate, rounded

and short-acuminate at the apex, stiffly chartaceous 2 la

20b. Fruits less than 4 cm long, globose to ellipsoid and always subtended by a cup or saucer-like

disc; leaves sparsely to densely puberulent beneath, drying chartaceous to coriaceous . 22a

2 la. Flowers 5-8 mm long; petioles 1-6 cm long, lower leaf surfaces with slender grayish

hairs 0.1-0.4 mm long, leaves with 5-9 pairs of secondary veins . .Persea americana

21b. Flowers ca. 3 mm long; petioles 1-3 cm long, lower leaf surfaces with slender brownish

hairs 0.2-0.6 mm long, leaves with 4-14 pairs of major secondary veins

Beilschmiedia anay

22a. Leaves usually tapering to a long-acuminate apex, 12-32(-40) cm long with 9-14 pairs of

major secondary veins, often drying dark brown; fruit cups 6-10 mm deep; inflorescences

paniculate with many (over 50) flowers Nectandra kunthiana

22b. Leaves rarely long acuminate (except in TV. belizensis and Endlicheria sp.?), not so long and

usually drying yellowish brown; fruit cups rarely more than 5 mm deep 23a

23a. Leaves gradually narrowed at the base and slightly rounded, narrowly to broadly obovate or

pandurate, with short (4-14 mm) petioles; inflorescences racemose with short lateral branches

and thick ferruginous peduncles 24a

23b. Leaves gradually or abruptly narrowed at the base but not rounded at the base, rarely slightly

obovate, 4-12 cm broad 25a

24a. Leaves 1 3-38 cm x 10-22 cm, with 8-1 2 pairs of major secondary veins, tertiary veins

often subparallel; petioles 4-14 mm long; flowers 8-10 mm broad

Ocotea valerioides

24b. Leaves 9-16 cm x 3.5-6 cm, with 4-8 pairs of secondary veins, tertiary veins not

subparallel; petioles 4-8 mm long; flowers 5-7 mm broad Nectandra belizensis

25a. Petioles to 3 cm long, leaves often drying subcoriaceous; inflorescences racemose or paniculate

and usually with short unbranched lateral branches, peduncles often densely short-puberulent

and ferruginous 26a

25b. Petioles to 2 cm long, leaves usually drying chartaceous; inflorescences paniculate, the primary

branches often with secondary branches, peduncles glabrous to puberulent 27a

26a. Leaves often broadly elliptic, the major veins broadly impressed above and the surface

rounded between the major veins; fruits on a disclike receptacle 6-8 mm broad and

1-2 mm deep; stamens not congested Ocotea babosa

26b. Leaves elliptic-oblong to obovate, the major veins and the upper surface flat; fruit cups

2 cm broad and 1 cm deep; glands and stamens closely congested into an androecial

dome Pleurothyrium golfodulcense

27 a. Leaves broadly elliptic, 6-14 cm broad, rounded at the apex and short-acuminate, the 5-9

pairs of secondary veins not loop-connected near the margin; fruit cups ca. 1.5 cm broad

and with persisting perianth-bases along the margin Ocotea mollifolia

27b. Leaves narrowly oblong (2.5-6 cm broad) and often long-acuminate, with 8-14 pairs of major

secondary veins; fruit cups without persisting perianth on the margins 28a

28a. Secondary veins loop-connected near the margin, leaves chartaceous; fruit cups 12-14 mm

broad, with a simple margin lEndlicheria sp.

FIELDIANA: BOTANY

28b. Secondary veins not loop-connected near the margin, leaves subcoriaceous; fruit cups 16-

22 mm broad, with a double margin Licaria multinervis

29a. Leaves tripliveined with 2 major lateral veins arising from near the base and extending beyond

the middle of the lamina to the distal half of the lamina, and with additional secondary veins

arising from the distal half of the lamina 30a

29b. Leaves pinnately veined, rarely with the basal secondary veins strongly ascending, and with several

pairs of secondary veins usually arising in the proximal half of the lamina 34a

30a. Trees of parks and gardens, rare in our area; bark with the odor of camphor or cinnamon;

leaves often opposite Cinnamomum

30b. Trees of native vegetation; bark lacking the odor of cinnamon or camphor when crushed .

3 la

3 la. Leaves consistently opposite, the lateral veins reaching the apex of the lamina; domatia

absent Caryodaphnopsis

31b. Leaves alternate, rarely subopposite, the lateral basal veins often reaching only the middle

of the lamina; pit domatia and tufted domatia often present in the vein axils 32a

32a. Leaves drying stiffly chartaceous and usually dark in color; inflorescences racemose and few-

flowered, the flowers 7 mm long and 10 mm broad; 1600-2700 m elevation

Ocotea holdridgeiana

32b. Leaves drying yellowish brown to olive green; flowers less than 5 mm long and 6 mm broad

33a

33a. Leaves usually drying subcoriaceous and often yellowish brown, secondary veins not loop-

connected distally; inflorescences paniculate to racemose; 0-2000 m elevation

Phoebe spp.

33b. Leaves drying chartaceous and usually dark olive green, secondary veins loop-connected

distally; inflorescences paniculate; 0-500 m, moist forests of the Pacific lowlands

Aiouea obscura

34a. Distal leafy stems hollow and often harboring small ants; leaves narrowly oblong to elliptic-oblong

or narrowly obovate, 1 0-50 cm long, glabrous beneath; small trees in evergreen forest understory

35a

34b. Distal leafy stems solid, the center with wood or pith, not consistently hollow; small to large trees

in deciduous or evergreen forests 40a

35a. Leaves usually drying chartaceous, and often with a slender acuminate tip 1-3 cm long; fruit

cups 1-3 mm deep; 0-1 100 m elevation 36a

35b. Leaves usually drying subcoriaceous and grayish green to orange brown in color, with long

acuminate tips only in O. dendrodaphne 37a

36a. Leaves often drying very dark, usually obovate, 12-32(-55) cm long; fruit cups 6-10

mm broad Ocotea atirrensis

36b. Leaves drying grayish to dark brown, usually elliptic-oblong, 1 2-27 cm long; fruit cups

5-8 mm broad Ocotea wedeliana

37a. Distal stems strongly angled with 3-5 prominent longitudinal ridges; leaves narrowly obovate,

14— 40(-55) cm long, with 7-12 pairs of secondary veins; fruit cups ca. 8 mm broad and 1-

2 mm deep and often with persisting perianth on the rim; 0-1 100 m

Ocotea nicaraguensis

37b. Distal stems without prominent longitudinal ridges and not strongly angled in cross section;

leaves with 5-1 1 pairs or major secondary veins 38a

38a. Leaves drying grayish green, usually elliptic-oblong, 14-36 cm long and 5-14 cm broad; fruit

cups ca. 1 5 mm broad and 3-7 mm deep with a single or double margin; flowers 4-6 mm

long; common, 0-1 500 m Ocotea dendrodaphne

38b. Leaves drying grayish to brown, usually narrowly oblong, 10-30 cm long and 3-8 cm broad;

flowers 2-3 mm long; rarely collected 39a

39a. Fruit cups 6-12 mm broad and 1-2 mm deep, without a flaring ridge beneath the distal rim;

600-2300 m Ocotea paulii

39b. Fruit cups 20-25 mm broad and 5 mm deep, with a prominent ridge around the periphery

just below the distal rim; 0-1400 m elevation Licaria brenesii

BURGER: FLORA COSTARICENSIS

40a. Leaf blades 20-50 cm long, usually becoming more than 30 cm long (not including the petiole-

length), 10-23 cm broad; 0-500 m in evergreen forests 4 la

40b. Leaf blades less than 30 cm in length, rarely exceeding 1 3 cm in width 43a

4 la. Petioles 2-6 cm long; leaves broadly elliptic to elliptic-oblong, drying grayish green and

subcoriaceous, with 6-10 pairs of major secondary veins; Caribbean slope

Phoebe chavarriana

41b. Petioles ca. 1 cm long; leaves obovate, drying chartaceous and brownish, with 9-14 pairs of

major secondary veins; Golfo Dulce area 42a

42a. Leaves broadly obovate, 15-23 cm broad, lamina attenuate at the petiole, the secondary

veins not loop-connected near the margin Ocotea rivularis

42b. Laminae narrowly oblanceolate, 10-15 cm broad, lamina slightly rounded at the petiole,

secondary veins often loop-connected near the margin . . Pleurothyrium hexaglandulosum

43a. Fruits borne on a thickened pedicel and not subtended by a cup or expanded disc; dried leaves

with the minor venation nearly always raised on the upper and/or lower surfaces and often forming

a reticulum of well-defined isodiametric areolae; petioles often more than 25 mm long (in Persed)

44a

43b. Fruits borne in a deep or shallow cup or subtended by a disclike receptacle expanded beyond the

thickened pedicel; minor venation inconspicuous or slightly raised on the dried leaf surfaces but

rarely forming a reticulum of well-defined areolae (compare dichotomy 74a); petioles exceeding

25 mm in length only in unusually large leaves or in Williamodendron 45a

44a. Fruits becoming ellipsoid; perianth deciduous and not persisting at the base of the fruits;

rarely collected trees, except for a common species with smaller (10x3 cm) leaves at 800-

2000 m elevation Beilschmiedia spp.

44b. Fruits becoming globose to ovoid, pyriform or reniform; the perianth bases often persisting

at the base of the fruits; both common and rare species, but those with smaller leaves above

2000 m elevation Persea spp.

45a. Fruits usually borne in small umbellate groups of 3 at the apex of a short (1 cm) peduncle, the

pedicels slightly expanded beneath the fruits to form a disc 2-4 mm broad; leaves small (to 1 2 x

3 cm) and stiff; rare unisexual trees of the Pacific slope, 1500-2000(-3000) m

Litsea glaucescens

45b. Fruits never borne in umbellate groups on such short peduncles; peduncles more than 2 cm long,

fruiting receptacle forming a cup or disc more than 4 mm broad 46a

46a. Distal branches strongly alate with narrow longitudinal wings 2-3 mm high, the stems 3-5-angled

in cross section; laminae narrowly oblong to narrowly obovate, drying coriaceous; Pacific slope of

Costa Rica, 0-700 m Ocotea aurantiodora

46b. Distal branches not alate, occasionally with longitudinal ridges and angular in cross section but

not winged 47a

47a. Leaf base decurrent on the petiole, the leaf base and petiole poorly differentiated, the leaf margin

often revolute near the base, the leaves petiolate, subsessile or sessile 48a

47b. Leaf base not usually decurrent on the petiole, the leaf blades acute to obtuse or rounded at the

base and usually clearly differentiated from the petiole, the leaves petiolate, never sessile or sub-

sessile 60a

48a. Leaf base broadly revolute (auriculate) and forming 2 broad flaps beneath just above the

petiole, petiole poorly differentiated and the leaf subsessile, broadly obovate and often co-

riaceous; 700-2300 m Ocotea endresiana

48b. Leaf base flat or revolute beneath but not forming broad flaps beneath, usually petiolate . .

49a

49a. Leaves usually with slender appressed ascending hairs on lower surfaces, hairs often parallel

with the secondary veins; trees often with prop roots 50a

49b. Leaves glabrous or with thin irregular hairs on the lower surfaces in early stages; trees without

prop roots 52a

50a. Leaf blades with the base usually long-decurrent, the narrowed portion of the leaf base

FIELDIANA: BOTANY

to 5 cm long, leaf narrowly elliptic-obovate to elliptic, 2-5 cm broad, with 5-9 pairs

of secondary veins; 600-1400 m elevation (if from higher elevations go to 5 la)

Ocotea skutchii

50b. Leaf blades not long decurrent at the base, broadly obovate, 4.5-1 1 cm broad, with 7-

1 2 pairs of major secondary veins 5 la

5 la. Leaves drying dull above and the minor venation not raised on the upper surface; fruits

ovoid and 3-4 cm long; Caribbean lowlands Ocotea caracasana

51b. Leaves usually drying lustrous above and with the minor venation slightly raised on

the upper surface; fruits globose to oblong, 2-3 cm long; 1 500-2500 m on the Pacific

slope and in Chiriqui Ocotea glaucosericea

52a. Leaves often rounded at the apex or bluntly obtuse (rarely acute to short-acuminate in smaller

leaves), often obovate and rarely oblanceolate, usually drying coriaceous to subcoriaceous

and often yellowish or grayish brown; fruiting receptacles cupulate and often with persisting

perianth bases along the rim of the cup, fruits ellipsoid, 1-2 cm long 53a

52b. Leaves bluntly acute to acuminate at the apex, sometimes bluntly obtuse but never rounded,

often oblanceolate to narrowly elliptic-obovate or oblong; fruiting receptacles flat and disclike

or cupulate but without persisting perianth at the rim 54a

53a. Stamens opening by 4 valves; leaves (2-)5-9(-12) cm broad and often obovate, drying

dark reddish brown to pale yellowish brown; fruit cups with or without persisting

perianth parts along the rim; 0-2000 m Ocotea insularis

53b. Stamens opening by 2 valves; leaves 1.5-6(-8.5) cm broad and often oblong-obovate,

drying dark brown to yellowish brown; fruit cups with persisting perianth; 1 100-2500

m Aiouea costaricensis

54a. Mature fruits usually more than 15 mm long; leaves often drying subcoriaceous 55a

54b. Mature fruits usually less than 15 mm in length, globose to oblong, often borne on flat

receptacles; leaves usually drying stiffly chartaceous 56b

55a. Fruits becoming ellipsoid, 1-2 cm in diameter, in cupulate receptacles 8-14 mm broad;

domatia rarely present; 1200-2500 m elevation Ocotea whitei

55b. Fruits becoming globose-pyriform, to over 5 cm in diameter, the receptacle only slightly

expanded below the fruits; leaves with pit domatia; 500-1 100 m

Povedadaphne quadriporata

56a. Leaves narrowly oblanceolate to elliptic-obovate, gradually narrowed at the base, leaves often

drying dark (gray to blackish); fruit cups gradually thickened to the flat or concave apex and

4-6 mm broad distally (but not forming a flat disc); 0-1000 m elevation 57a

56b. Leaves usually elliptic to oblong-obovate and rarely oblanceolate, gradually to abruptly

narrowed at the base, the leaves drying grayish to brownish (58a) or dark (60a) 58a

57a. Fruits globose, 5-8 mm in diameter (dried); 0-500 m, Caribbean lowlands in Costa

Rica Ocotea bijuga

57b. Fruits oblong, 9-18 mm long and 7-9 mm in diameter; (0-)400-1000 m

Ocotea oblonga

58a. Fruits globose or subglobose, 1-2 cm in diameter and borne on a flat disc or shallow cup 6-

1 2 mm broad; 0-1 500 m elevation 59a

58b. Fruits and receptacle otherwise 60a

59a. Fruits subglobose, 10-16 mm in diameter and usually with persisting style base at the

apex, borne on a thick double-rimmed flat disclike receptacle 6-10 mm broad; leaves

usually drying grayish green and the stems black Ocotea jloribun da

59b. Fruits globose, 1 2-20 mm in diameter, without persisting style base, borne on a shallow

cup 9-12 mm broad; leaves drying yellowish brown to grayish, stems drying gray or

brown Nectandra cissiflora

60a. (from 47b and 58b) Leaves drying dark (almost black) and thin-chartaceous; fruiting receptacle

gradually expanded (obconic) to the 1 cm broad apex, concave or slightly cupulate, fruits becoming

2-3 cm long and 1-2 cm thick 6 la

BURGER: FLORA COSTARICENSIS

60b. Leaves drying grayish to dark brown, stiffly chartaceous to coriaceous if drying very dark . . 62a

6 la. Leaves elliptic to elliptic-oblong, 7-16(-20) cm long, glabrous, with 4-7 pairs of major

secondary veins; flowers 1.5-3 mm long; 0-1700 m elevation Ocotea tenera

61b. Leaves ovate to broadly elliptic, 6-12(-15) cm long, sparsely puberulent, with 3-5 pairs of

major secondary veins; flowers 3-5 mm long; 900-2000 m elevation Ocotea brenesii

62a. Fruit cups with a double margin distally, an elevated ridge or flange present in addition to the

distal edge of the cup and encircling the cup close to the edge, the distal edge usually entire and

the subtending ridge or margin often undulate 63a

62b. Fruit cups without a conspicuous double margin distally or the fruits subtended by a flat disc and

a cup absent 69a

63a. Cups well developed and 5-10 mm deep, rounded to conical in form; species of Licaria . .

64a

63b. Cups shallow or saucer-like, rarely more than 5 mm deep 68a

64a. Leaves becoming narrowly elliptic-oblong to lanceolate, usually 3—4 times longer than

broad 65a

64b. Leaves elliptic to elliptic-oblong, usually only 2-3 times longer than broad 66a

65a. Leaves with 8-14 pairs of major secondary veins and minutely puberulent be-

neath; 500-1000 m elevation L. multinervis

65b. Leaves with 5-10 pairs of major secondary veins, glabrous beneath; 150-2300

m elevation L. excelsa

66a. Fruit cups 8-12 mm broad; leaves 10-20 cm long, 3-7 cm broad, often lustrous above;

Caribbean slope, 0-700 m L. sarapiquensis

66b. Fruit cups 1 1-28 mm broad at the top 67a

67a. Trees of the Caribbean slopes and central highlands, 0-1400 m elevation; leaves 5-16

cm long, 2-6 cm broad L. triandra

67b. Trees of southwestern (Pacific) Costa Rica, 0-1000 m elevation; leaves 9-18 cm long,

3-7 cm broad L. cufodontisii

68a. Leaves obtuse to bluntly acute at the apex; fruit cups shallow and with ellipsoid fruits; very

dry deciduous to partly deciduous forests of the Pacific slope Ocotea veraguensis

68b. Leaves acute to short-acuminate at the apex; fruit cups very shallow and often disclike, fruits

subglobose; evergreen and partly deciduous forests Ocotea floribunda

69a. Leaves usually less than 9 cm long and 3 cm wide, narrowly elliptic to narrowly elliptic-oblong,

usually drying dark brownish with the minor venation slightly raised above 70a

69b. Leaves usually becoming much more than 9 cm long and 3 cm wide, if small the leaves usually

drying pale greenish gray or yellowish 74a

70a. Trees of lower-elevation (0-1600 m) wet evergreen forest formations; fruits 10-18 mm long,

globose to ellipsoid 71a

70b. Trees of montane forest formations above 1600 m elevation 72a

7 la. Leaves with the upper surfaces drying lustrous and with the tertiary venation conspic-

uously raised; fruits ellipsoid in development; 600-1600 m Nectandra salicina

7 1 b. Leaves with the upper surfaces dull or lustrous when dry, tertiary venation only slightly

raised; fruits globose in all stages of development; below 500 m

Nectandra salicifolia

72a. Basal secondary veins often strongly ascending and almost tripli veined, leaves often drying

very dark and lustrous above, pit domatia usually present in vein axils beneath; flowers to

7 mm long and 10 mm broad Ocotea holdridgeiana

72b. Basal secondaries not strongly ascending, the leaves never tripliveined, pit domatia absent;

flowers 2-4 mm long and 2-5 mm broad 73a

73a. Stems minutely puberulent in early stages; leaves 2-8 cm long and up to 2.4 cm broad, acute

to short acuminate, venation flat above; Cordillera de Talamanca . . Aiouea talamancensis

73b. Stems glabrous in early stages; leaves 3-12 cm long and up to 4 cm broad, often long-

acuminate, venation slightly raised above when dry; Chiriqui highlands . Ocotea viridiflora

74a. Minor venation raised (elevated) on the lower leaf surface when dry and forming small areolae

10 FIELDIANA: BOTANY

0.3-1 mm broad, the areolae often with subequal sides, the laminae often drying yellowish green

or grayish 75a

74b. Minor venation not raised and forming a distinct reticulum on the lower (abaxial) leaf surfaces

when dried, areolae very irregular if present 79a

75a. Leaves gradually narrowed to the base, usually narrowly obovate and drying grayish green;

fruits globose with a short persisting style at the apex and subtended by a small, flat, disclike

receptacle Ocotea floribunda

75b. Leaves abruptly narrowed to the base (obtuse to acute); fruits never with a persisting style

base, mostly ellipsoid and borne in cupulate receptacles 8-14 mm broad and 2-5 mm deep

76a

76a. Leaves large (14-30 x 7-18 cm) and obovate, rounded to bluntly obtuse at the apex, drying

yellowish brown and subcoriaceous; fruit cups often with irregular margins; wet forests, 500-

1000 m Ocotea sp. aff. O. laetevirens

76b. Leaves not becoming so large and never broadly obovate, not more than 10 cm broad, usually

acuminate at the apex 77a

77a. Leaves with 4-9 pairs of major secondary veins, petioles 7-15 mm long and 2-3.5 mm thick;

inflorescences densely and very minutely puberulent; 0-1000 m Aniba venezuelana

77b. Leaves with 3-6(-7) pairs of major secondary veins, petioles 5-30 mm long, 1-2.5 mm thick;

inflorescences glabrous or very minutely and sparsely puberulent 78a

78a. Leaves drying stiffly chartaceous and often grayish; fruit cups 2-3 mm deep; 0-2100 m

elevation Ocotea meziana

78b. Leaves drying subcoriaceous or stiffly chartaceous and often yellowish green; fruit cups 3-5

mm deep; 1 200-2000 m Ocotea laetevirens

79a. Trees not known from below 1 400 m elevation; petioles, stems, and inflorescence axes often reddish

brown and glabrescent, the leaves often drying reddish brown to yellowish, often lustrous and

coriaceous 80a

79b. Trees not known from above 1 400 m elevation (except in O. leucoxylon and N. membranacea and

those not drying reddish or yellowish) 8 la

80a. Upper leaf surfaces usually drying lustrous and with the tertiary venation raised; fruits globose

and becoming more than 2 cm in diameter Nectandra cufodontisii

80b. Upper leaf surfaces drying dull and the tertiary venation not raised; fruits ellipsoid, ca. 1 cm

in diameter Phoebe hammeliana

8 la. Leaves conspicuously glaucous beneath, drying chartaceous, to 29 cm long and 12 cm broad;

evergreen forests below 600 m elevation 82a

8 Ib. Leaves not conspicuously glaucous beneath 83a

82a. Petioles 0.5-1.5 cm long, leaves usually elliptic with 3-6 pairs of major secondary veins;

flowers 2.5 mm long Nectandra hypoleuca

82b. Petioles 2-7 cm long, leaves usually obovate with 9-14 pairs of major secondary veins,

clustered at the ends of twigs; flowers ca. 1.5 mm long . . \Villiamodendron glaucophyllum

83a. Fruits borne in deep (3-6 mm) well-developed cups usually more than 10 mm broad at the top;

fruits usually ellipsoid to ovoid; stamens with 2 or 4 valves; leaves and inflorescences essentially

glabrous; trees of the wet Caribbean slope 84a

83b. Fruits borne in shallow (1-3 mm) cups rarely more than 8 mm broad or on shallow saucer-like

receptacles to 12 mm broad; fruits globose to ovoid; stamens with 4 valves 86a

84a. Leaves drying grayish or yellowish green, usually caudate-acuminate at the apex with a narrow

(3 mm) tip to 2 cm long; laminae usually oblong and less than 16 cm long, chartaceous;

flowers usually drying black Ocotea cernua

84b. Leaves not drying grayish and rarely caudate-acuminate (gradually acuminate when with a

long tip) 85a

85a. Leaves drying subcoriaceous to stiffly chartaceous and lustrous above with the tertiary ve-

nation obscure, 1 0-20 cm long and elliptic-oblong; fruit cups sometimes with a double margin,

fruits ca. 1 cm in diameter Licaria sarapiquensis

BURGER: FLORA COSTARICENSIS 1 1

85b. Leaves drying chartaceous and dull above, the tertiary venation usually visible on the upper

surfaces, 10-30 cm long and elliptic; fruits ca. 2 cm in diameter Aiouea sp.?

86a. Fruit cups drying dark and with conspicuous paler colored lenticel-like warts, fruits 8-10 mm in

diameter and subglobose; young stems strongly ridged or 3— 4-angled in cross section; leaves usually

elliptic-oblong and drying grayish, usually subcoriaceous and margin revolute throughout; flowers

unisexual, anthers with superposed thecae Ocotea leucoxylon

86b. Fruit cups not drying dark with conspicuous lighter colored warts or lenticels, fruits often borne

on shallow saucer-like receptacles to 1 2mm broad, fruits globose or (less often) oblong-ellipsoid

and less than 1 5 mm long; young stems not strongly angled or ribbed; outer anthers with the thecae

in a single horizontal plane or arc. (The following are all species of Nectandra and very difficult

to identify in the absence of flowers or with atypical foliage.) 87a

87a. Largest leaves rarely more than 1 7 cm long 88a

87b. Largest leaves usually more than 20 cm long 9 la

88a. Leaves minutely puberulent beneath, never drying lustrous above; flowers 6-14 mm broad

89a

88b. Leaves glabrous beneath (except in early stages); flowers 5-8 mm broad 90a

89a. Leaves slightly decurrent at the base but not revolute, narrowly elliptic to narrowly

elliptic-oblong, to 20 cm long, usually drying dark brown; flowers ca. 8 mm broad; 0-

1 400 m Nectandra globosa

89b. Leaves slightly decurrent on the stem and often revolute at the base, ovate to narrowly

elliptic, to 1 5 cm long, usually drying very pale brown; flowers ca. 10 mm broad; Pacific

slope, 0-1400 m Nectandra ramonensis

90a. Leaves usually drying brownish and often lustrous above, usually with the minor venation

raised beneath when dry, secondary veins usually loop-connected distally

Nectandra salicifolia

90b. Leaves usually drying grayish and rarely lustrous above, the minor venation obscure (in the

central highlands and Caribbean slope) beneath or slightly elevated and clearly visible (on

the Pacific slope where leaves are often very narrow), secondaries not loop-connected ....

Nectandra turbacensis

9 la. Leaves with the major secondary veins often loop-connected near the margin, usually drying dark

brownish and lustrous above with the tertiary venation slightly raised, glabrous beneath, laminae

ovate-elliptic to elliptic-oblong, 13-24 cm long and 5-11 cm broad in Costa Rica; Caribbean

lowlands A^. latifolia

91b. Leaves with the major secondary veins not (or only very weakly) loop-connected near the margin,

tertiary venation prominent above usually only in N. martinicensis; absent from the Caribbean

lowlands (except for N. membranaced) in Costa Rica 92a

92a. Leaves usually drying subcoriaceous, slightly decurrent on the petiole and the margin often revolute

above the petiole, minutely puberulent beneath, often ovate-lanceolate in shape 93a

92b. Leaves usually drying stiffly chartaceous, some leaves may be slightly decurrent on the petiole but

rarely with the margin revolute near the base, usually with 4-8 pairs of major secondary veins,

puberulent or sometimes glabrous beneath in N. martinicensis; flowers 5-8 mm broad .... 94a

93a. Leaves with 4-8(-10) pairs of major secondary veins arising throughout the length of the

midvein, often drying yellowish green or yellowish brown; flowers 6—12 mm broad; common

in deciduous and partly deciduous forest, 0-700 m N. globosa

93b. Leaves with 3-5 pairs of major secondary veins arising from the proximal '/2 (%) of the

midvein and arcuate-ascending, often drying dark brown; evergreen or partly deciduous forest

formations, 0-1 700 m N. membranacea

94a. Leaves usually drying dull or slightly lustrous above and greenish or brown, laminae often elliptic-

oblong, 12-28 cm long and 4-8.5 cm broad; twigs glabrous or sparsely puberulent

N. martinicensis

94b. Leaves usually drying dark and very lustrous on the upper surfaces, often lanceolate or very narrowly

elliptic, 10-30 cm long and 3-7 cm broad; twigs densely puberulent M nitida

12 FIELDIANA: BOTANY

Key 3: Key to Comparative Figures of Costa Rican Lauraceae

la. Leaves tripliveined or with the basal pair of secondary veins strongly ascending Fig. 1

Ib. Leaves rarely tripliveined 2a

2a. Leaves usually exceeding 30 cm in length (not including the petiole length) Fig. 2

2b. Leaves rarely exceeding 30 cm in length 3a

3a. Distal stems hollow; leaves glabrous and often narrowly oblong or narrowly obovate Fig. 3

3b. Distal stems solid; leaves variously shaped 4a

4a. Leaves usually less than 1 0 cm long, elliptic to narrowly oblong; from above 1 000 m elevation .

Fig. 4

4b. Leaves usually becoming more than 1 0 cm long or lowland species 5a

5a. Leaves very stiffly coriaceous; montane species from above 1 500 m elevation Fig. 5

5b. Leaves not stiffly coriaceous and from montane habitats 6a

6a. Leaves densely puberulent beneath, the trichomes easily seen and the vesture apparent to touch

7a

6b. Leaves glabrous beneath or the puberulence very small or difficult to see, lower surface smooth to

touch lOa

7a. Montane species, (1000-)! 500-2500 m elevation Fig. 6

7b. Species of lower habitats, 0-1 500 m elevation 8a

8a. Pubescence usually becoming dark brown beneath Fig. 7

8b. Pubescence rarely becoming dark brown beneath 9a

9a. Pubescence dense on the lower surface Fig. 8

9b. Pubescence sparse to moderately dense beneath Fig. 9

lOa. Leaves usually distinctly decurrent on the petiole and often cuneate at the base 1 la

lOb. Leaves not usually decurrent or only slightly decurrent at the base 12a

1 la. Leaves generally broad, often obovate Fig. 10

1 Ib. Leaves generally narrow, often elliptic (see also fig. 20) Fig. 11

1 2a. Montane species from 1 500 to 3000 m, with leaves often lustrous above, mostly glabrous

Fig. 12

1 2b. Lowland or montane species, if montane the leaves not lustrous on the upper surface when dry

13a

1 3a. Leaves drying grayish green, yellowish green, or very dark (almost black) on the upper surface .

14a

1 3b. Leaves usually drying pale grayish, yellowish or dark 1 5a

14a. Leaves drying very dark Fig. 13

14b. Leaves drying grayish or yellowish Figs. 13-14

1 5a. Fruits not subtended by cups or discs Fig. 15

1 5b. Fruits subtended by a cup or flattened disc 1 6a

16a. Fruit cups usually with a double-rimmed edge; stamens 3 and often connate (Licarid) . . Fig. 16

1 6b. Fruit cups with a single edge 1 7a

1 7a. Nectandra salicifolia and related/similar species Fig. 17

1 7b. Nectandra globosa and similar/other species Fig. 18

BURGER: FLORA COSTARICENSIS 13

P. cinnamomifolia

sensu lato

FIG. 1 . Lauraceae: Tripliveined and palmately veined species.

14

FIELDIANA: BOTANY

FIG. 2. Lauraceae: Large-leaved species.

BURGER: FLORA COSTARICENSIS

15

O. nicaraguensis

I 20cm

FIG. 3. Lauraceae: Species with hollow distal stems.

16

FIELDIANA: BOTANY

Ocotea monteverdensis

FIG. 4. Lauraceae: Montane species with small leaves.

BURGER: FLORA COSTARICENSIS

17

FIG. 5. Lauraceae: Montane species with thick coriaceous leaves.

18

FIELDIANA: BOTANY

FIG. 6. Lauraceae: Montane species with densely puberulent leaves.

BURGER: FLORA COSTARICENSIS

19

FIG. 7. Lauraceae: Lower-elevation species with densely puberulent leaves.

20

FIELDIANA: BOTANY

FIG. 8. Lauraceae: Lower-elevation species with densely puberulent leaves.

BURGER: FLORA COSTARICENSIS

21

r/7 Pleu. golfodulcense

FIG. 9. Lauraceae: Lower-elevation species with sparsely puberulent leaves and two species of Persea.

22

FIELDIANA: BOTANY

FIG. 10. Lauraceae: Species with decurrent lamina bases (larger-leaved species).

BURGER: FLORA COSTARICENSIS

23

FIG. 1 1 . Lauraceae: Species with decurrent lamina bases (smaller-leaved species).

24

FIELDIANA: BOTANY

FIG. 1 2. Lauraceae: Montane species with the laminae often lustrous above.

BURGER: FLORA COSTARICENSIS

25

FIG. 13. Lauraceae: Species with glabrous leaves often drying dark or grayish.

26

FIELDIANA: BOTANY

FIG. 14. Lauraceae: Species with glabrous leaves often drying gray or greenish.

BURGER: FLORA COSTARICENSIS

27

FIG. 15. Lauraceae: Species of Beilschmiedia and Persea, the fruits not subtended by a cup.

28

FIELDIANA: BOTANY

FIG. 16. Lauraceae: Species of Licaria, the fruit cup usually with a double margin.

BURGER: FLORA COSTARICENSIS

29

FIG. 17. Lauraceae: Nectandra salicifolia and similar species.

30

FIELDIANA: BOTANY

FIG. 18. Lauraceae: Nectandra globosa and similar species.

BURGER: FLORA COSTARICENSIS

31

3cm

FIG. 19. Lauraceae: Williamodendron glaucophyllum. A, branchlet of the type collection; B, two flowers.

32

FIELDIANA: BOTANY

1mm

FIG. 20. Lauraceae: Ocotea dentata. A, branchlet of the type; B, base of two leaves; C, lower leaf surface; D, part

of the inflorescence; E, flower viewed from the side; F, flower from above; G, stamens and pistil; H, young fruits; I,

flower of Ocotea insularis for comparison.

BURGER: FLORA COSTARICENSIS

33

FIG. 21. Lauraceae: Persea silvatica. A, branchlet of the type; B, flower with 4 tepals removed showing reflexed

outer stamens and inner stamens with glands; C, pistil; D, inner and outer stamens; E-F, lateral and frontal view of

a staminode; G, base of inflorescence with bracts.

34

FIELDIANA: BOTANY

3 mm

FIG. 22. Lauraceae: Povedadaphne quadriporata. A, branchlet of the type; B, floral diagram; C, two stamens; D,

flower from above; E, flower in cross section; F, domatia on lower leaf surface.

BURGER: FLORA COSTARICENSIS

35

Aiouea Aublet

REFERENCE— S. Renner, Aiouea. Flora Neotro-

pica31: 85-124. 1982.

Small to medium-sized trees (rarely shrubs), bisexual.

Leaves alternate, usually well-spaced along the stems,

leaf blades glabrous above, glabrous or rarely puberulent

beneath, pinnately veined or rarely tripliveined, domatia

present in only 2 species in South America. Inflores-

cences axillary to distal leaves or clustered near the tips

of branchlets, paniculate, 10-15 cm long, each pedicel

subtended by a small bract and with 2 bracteoles. Flow-

ers small (1-4 mm) and bisexual, usually obconic to obo-

void, glabrous or puberulent, floral tube short and pu-

berulent within, perianth of 6 equal parts in 2 whorls;

fertile stamens 9 in 3 whorls in ours (rarely with only 6

or 3 fertile stamens), the outer 6 stamens with 2 large

introrse thecae in each anther, the inner 3 stamens with

2 thecae or with 4 thecae and the upper smaller, inner

3 stamens biglandular, staminodia (series IV) stipiti-

form, clavate, triangular or absent; pistil glabrous, style

slender, stigma simple or capitate (often depending on

stage of development). Fruits borne in a shallow cupulate

receptacle, rim of the cup entire, undulate or with per-

sisting perianth parts, at first fleshy but becoming woody;

berry ovoid to ellipsoid.

Aiouea is a Neotropical genus of about 25 species

ranging from Mexico and the West Indies south-

ward to 30°S latitude. The three whorls of usually

nine fertile 2-thecous stamens, the well-developed

staminodes in most species, and the cupulate fruit-

ing receptacle distinguish this genus. However, it

appears that three Costa Rican species, A. costar-

icensis, A. obscura, and A. talamancensis, may be

independent 2-thecous derivatives of species of

Ocotea; see the discussions under these species.

Key to Costa Rican Species of Aiouea

la. Venation tripliveined, lamina drying thin-chartaceous, dark gray or olive green; lowland forests of

the Golfo Dulce area A. obscura

Ib. Venation pinnate; trees not known to inhabit the Pacific lowlands 2a

2a. Leaves obtuse to rounded at the apex, 4-14(-19) cm long, subcoriaceous and yellowish or pale

grayish when dry; staminodia usually absent; rim of fruiting receptacle with persistent perianth;

(600-)1 100-2300 m alt A. costaricensis

2b. Leaves acute to acuminate at the apex, usually chartaceous and dark when dried; rim of fruiting

cupule entire or unknown 3a

3a. Leaf blades 2-8 cm long and 0.8-2.4 cm broad; staminodes usually absent; known only from the

Cordillera de Talamanca at 1600-2300 m alt A. talamancensis

3b. Leaf blades 1 6-22 cm long and 7-9 cm broad; slender staminodes usually present; known only from

the Caribbean escarpment at 400-700 m alt Aiouea sp.?

Aiouea costaricensis (Mez) Kosterm., Meded. Bot.

Mus. Herb. Rijks. Univ. Utrecht 46: 73. 1938

(and Recueil Trav. Bot. Neerl. 35: 73. 1938).

Bellota costaricensis Mez, Jahrb. Konigl. Bot.

Gait. Berlin 5: 27. 1889. Boldus costaricensis

(Mez) Kuntze, Rev. gen. pi. 2: 569. 1889. Figure

10.

Trees 5-15(-20) m tall, trunk to 50 cm d.b.h., branch-

lets 2-6 mm thick, at first densely brownish sericeous

with thin straight ascending hairs 0. 1-0.4 mm long, lon-

gitudinally ridged but becoming terete and glabrous.

Leaves alternate in a spiral, petioles 2-10 mm long, usu-

ally flat above, with lateral margins continuous with the

lamina margins; leaf blades 4-14(-19) cm long, 1.5-

6(-8.5) cm broad, obovate, elliptic-obovate, oblong or

spatulate, usually bluntly obtuse to rounded at the apex,

tapering gradually to the cuneate base and decurrent on

the petiole, margin entire and often revolute (especially

near the base), drying subcoriaceous and yellowish brown

to gray, smooth and glabrous above, becoming glabrous

beneath, with 5-7 major secondary veins on each side,

tertiary venation raised on the lower surface, domatia

absent. Inflorescences 10-20 cm long, peduncle 4-10 cm

long, paniculate with open widely spaced branches to 6

cm long, glabrous or very minutely and remotely pu-

berulent, pedicels ca. 5 mm long and with a small bract

or scar at the base. Flowers 2.5-4 mm long, 2-3 mm

broad, yellowish and glabrous, perianth parts 1-2 mm

long and 1 .5 mm broad, glabrous on both surfaces; outer

stamens with short (0.3 mm) filaments, outer anthers

0.5-0.8 mm long, usually longer than broad, with only

2 thecae, inner stamens 2-thecous and ca. 1 .4 mm long,

with hairs at the base of the anthers, staminodes absent;

pistil 1.5-2 mm long, style 0.6-1 mm long, stigma simple

or slightly discoid. Fruits borne on a cupulate receptacle

5-6 mm long and 6-8 mm broad, with persisting peri-

anth lobes (ca. 1 mm long and 2 mm broad) on the rim,

obconic and very shallow (1-3 mm deep) but enclosing

the fruits in early stages and only the base at maturity,

becoming red; berry ellipsoid, 1-1.5 cm long and 0.8-

1 . 1 cm thick, green.

36

FIELDIANA: BOTANY

Trees of wet evergreen forest formations be-

tween 1100 and 2500 m elevation; a single col-

lection has been made at 600 m near Cariblanco

in the valley of the Rio Sarapiqui. Flowers have

been collected in March-September; fruiting ma-

terial has been collected in March-May, July, Oc-

tober, and December. The species ranges along the

Caribbean side of the Cordillera Central (Zarcero,

Palmira, Poas, Barva [Barba], Carpintera, Cachi),

the southeastern edge of the Meseta Central (Tab-

lazo, Tarbaca), and the Cordillera de Talamanca,

probably into Panama.

Aiouea costaricensis is recognized by its 2-the-

cous anthers, the generally small, stiff, obovate

leaves with bluntly obtuse or rounded apices, and

cuneate bases decurrent on the petiole. The leaves

usually dry grayish or yellowish brown and with

the tertiary venation slightly raised beneath when

dry (but not forming a conspicuous reticulum).

The small, shallow fruit cups with persisting peri-

anth lobes, the glabrous flowers, and restricted

middle-elevation montane habitat are additional

characteristics. This species has not been found at

Monteverde in the Sierra de Tilaran, but there is

an almost identical population ofOcotea insularis

(O. tonduzii in a more narrow sense) at Monte-

verde with 4-thecous anthers. This near-identity

makes it appear that A. costaricensis may be noth-

ing more than a 2-thecous derivative of some high-

land populations of O. insularis (in a wide sense).

A recent collection from Rio Colon in easternmost

Costa Rica (Davidse et al. 25529, CR, MO) is placed

here because it is consistently 2-thecous. It may

be related to the population of O. insularis which

was the basis of the name Aiouea lundelliana and

in which some anthers are 2-thecous. See the dis-

cussions under O. insularis.

Aiouea obscura van der Werff, Ann. Missouri Bot.

Card. 75: 402. 1988. Figure 1.

Trees ca. 10m tall, leafy branchlets 1-2 mm thick,

glabrous, dark brown, terete. Leaves alternate and dis-

tant, petioles 10-18 mm long, 0.9-1.4 mm thick, terete

but flat or slightly sulcate above, glabrous; leaf blades

(8-)l 1-17 cm long, 2.5-5.3 cm broad, narrowly elliptic

to narrowly elliptic-oblong, tapering gradually to an acute

or acuminate apex, tip 5-10 mm long, acute at the base

and slightly decurrent on the petiole, drying thin-char-

taceous or membranaceous, dark gray or olive green when

dry and dull above, the midvein prominent above, gla-

brous and slightly lustrous beneath, tripliveined with a

prominent pair of basal secondary veins arising 8-15

mm above the petiole and ascending beyond the middle

of the lamina, with 2-3 additional secondaries on each

side in the distal half of the lamina, the secondaries

usually connected to form a submarginal vein 1.5-5 mm

from the margin, minor venation usually flat above and

prominent beneath, inflated domatia with minute pore-

like openings sometimes present in the axils of the major

veins beneath. Inflorescences pseudoterminal in the axils

of deciduous bracts, solitary, open-branched panicles with

relatively few (ca. 50) flowers, 7-15 cm long, peduncle

1-6 cm long and 0.7 mm thick, glabrous and drying dark,

pedicels 5-10 mm long, thin. Flowers bisexual, 2-2.5

mm long, 3 mm broad, glabrous on the outside, tepals

equal, 1-1.5 mm long and ca. 1.2 mm broad; outer sta-

mens 1.2-1.4 mm long with prominent minutely pu-

berulent filaments, outer anthers ca. 0.7 mm long and

0.5 mm broad, with the connective slightly prolonged

beyond the thecae, glands of the inner stamens ca. 0.4

mm in diameter, staminodes absent; pistil ca. 1.6 mm

long, style 0.5 mm long, stigma simple. Fruits and fruit-

ing cupules unknown.

Trees of the southern Pacific slope southeast of

Palmar Norte on steep slopes of evergreen rain

forest at about 50 m elevation. At present the

species is known from only a single flowering col-

lection made in May 1986 by Hammel, Grayum,

and de Nevers (15197, CR, F, MO, the holotype).

Aiouea obscura is distinguished by its thin, nar-

row, glabrous leaves with strongly ascending basal

secondary veins, and the small flowers with nine

2-thecous stamens. This is an unusually glabrous

species; the shoot apex is almost glabrous except

for minutely ciliate margins of the young leaves.

The relationships of this species are rather uncer-

tain.

Aiouea talamancensis W. Burger, sp. nov. Fig-

ure 4.

Arbor 4-1 0 m alta, ramulis foliiferis 0.6-2 mm crassis.

Folia alterna, petiolis 3-7 mm longis, 1 mm latis, laminis

(1.5-)2.5-8 cm longis, 0.8-2.4 cm latis, anguste ellipticis,

ellipticis vel elliptico-oblongis, apice acuto vel breviter

acuminato, subtus glabrescentibus, nervis secondariis 3-

6 paribus. Inflorescentiae paniculatae, 2-6 cm longae.

Flores 2-2.5 mm longi, 2-3 mm lati, extus glabri vel

minute puberuli, tepalis aequalibus; stamina ser. 1-1 1

filamentis brevibus, antheris 0.5-0.7 mm longis, stam-

inodiis nullis; gynoecium ca. 1 .8 mm longum, stylo 0.6-

0.8 mm longo.

Small trees, 4-10 m tall, leafy branchlets 0.7-2 mm

thick, sparsely and minutely (0.1-0.2 mm) puberulent

but quickly becoming glabrous, with ca. 3 prominent

longitudinal ridges but becoming terete and grayish.

Leaves alternate in a spiral, petioles 3-7 mm long, ca. 1

mm thick, glabrous or very minutely puberulent, with 2

lateral ridges continuous with the lamina margins, flat

or broadly sulcate above; leaf blades ( 1 .5-)2.5-8 cm long,

0.8-2.4 cm broad, narrowly elliptic to elliptic or elliptic-

oblong, tapering to an acute or short-acuminate apex,

tapering gradually to the acute base and decurrent on the

BURGER: FLORA COSTARICENSIS

37

petiole, drying stiffly chartaceous, glabrous above and

grayish or dark, dull or lustrous above, glabrescent be-

neath, with 3-6 major secondary veins on each side,

tertiary- venation slightly raised beneath. Inflorescences

2-6 cm long in distal leaf axils, 2-6 cm long, paniculate

but few-branched and few-flowered, peduncle to 3 cm

long, glabrous or minutely puberulent, pedicels 2-4 mm

long. Flowers yellowish, 2-2.5 mm long, 2-3 mm broad,

glabrous or minutely puberulent on the outside, perianth

parts equal in length; outer stamens with short (0.5 mm)

filaments and anthers 0.5-0.7 mm long, inner stamens

ca. 1 .4 mm long, staminodes absent; pistil ca. 1 .8 mm

long, the narrow style 0.6-0.8 mm long, stigma subcap-

itate. Fruits borne in a shallow obconical cup about 5

mm long and 5-7 mm broad; berry ca. 6 mm long and

5 mm in diameter, ovoid (fruit characters based on Oroz-

co 12/10/45 seen at CR).

TYPE— Costa Rica, Prov. San Jose, Pan-Amer-

ican Highway between San Isidro del General and

Division, elevation 1900 m, 4 March 1966, An-

tonio Molina R., William C. Burger & Bruce Wal-

lenta 7W59(holotype, F 1749014; negative, 6 1 1 25;

isotypes, CR, NY).

Trees of montane cloud forest formations from

1600 to 2300 m elevation in the Cordillera de

Talamanca. Flowering collections have been made

in February, September, and November. This

species is known only from Costa Rica; it has been

collected from near Patarra (southwest of San Jose),

northwest of San Isidro del General, and near Cer-

ro Kamuk. Collections in addition to the type are

Chacon & Herrera 1547, CR, F; Davidse & Herrera

29178, CR, MO; and L. Orozco 12/10/45, collected

9 March 1985 and seen at CR).

Aiouea talamancensis is distinguished by its very

small elliptic glabrescent leaves, small few-flow-

ered inflorescences, the 2-thecous anthers, and the

montane habitat. The small narrow leaves, taper-

ing gradually at both ends and decurrent on the

petiole, strongly resemble the leaves of some spec-

imens of Ocotea whitei, and it may be that this

species is a 2-thecous derivative of that species of

Ocotea or one of its close allies. Aiouea parvissima

(Lundell) Renner has similarly small leaves but

much longer petioles, subcoriaceous leaf texture,

clavate staminodes and grows in lowland forests

of Peten, Guatemala.

Aiouea sp.?

Small trees ca. 5 m tall, leafy branchlets 2-3 mm thick,

glabrous. Leaves alternate in a spiral, petioles 1 6-20 mm

long, 1.5-2 mm thick, narrowly sulcate above, glabrous;

leaf blades 16-22 cm long, 7-14 cm broad, elliptic-ob-

long to slightly obovate, acuminate at the apex, obtuse

at the base, drying stiffly chartaceous, glabrous above

and with minute appressed hairs below, with 4-6 major

secondary veins on each side, tertiary veins slightly el-

evated beneath when dry. Inflorescences axillary to distal

leaves or leafless nodes, 12-14 cm long, paniculate with

widely spaced branches and ca. 30 flowers, peduncle 2-

4 cm long, 1-1.5 mm thick, glabrous, pedicels 1-4 mm

long, slender. Flowers greenish, ca. 2.5 mm long and 3

mm broad, glabrous on the outside, perianth parts ca.

1.5 mm long, 1.2 mm broad; outer stamens ca. 1.4 mm

long with anthers 0.8 mm long, connective expanded at

the apex and apiculate, staminodes ca. 1 mm long and

slender; pistil ca. 1.8 mm long with a short (0.5 mm)

style, stigma simple. Fruits borne in a saucer-like cup

1.5-2 cm broad and 2-4 mm deep, abruptly expanded

above the thickened (5-7 mm) pedicel, rose-brown in

color; berry 32 mm long and 20 mm in diameter, ellip-

soid-oblong.

Small trees of the very wet forests of the Carib-

bean escarpment in tropical forest-premontane

forest transition zone. At present, this taxon is

known only from a single collection by I. A. Cha-

con and G. Herrera (1722, CR, F, MO), from Es-

tacion Carillo de 700 a 450 m de la Fila al Cano

del R. Sucio, 12 Nov. 1983, near the juncture of

the provinces of Cartago, Limon, and San Jose.

This collection is provisionally placed in Aiouea

because of its nine 2-thecous stamens. Unusual for

Aiouea is the fact that the thecae had not opened

in any of the four dissected flowers, suggesting that

the flowers are unisexual. The erect tepals and the

well-developed staminodia are arguments against

placing this taxon in Endlicheria; a genus char-

acterized by unisexual flowers with nine 2-thecous

stamens. It is possible that the flowers are abnor-

mal; all dissected flowers contained a small insect

larva. Vegetatively, this taxon is unlike any species

we have seen, and we do not doubt that it is un-

described. Given the fact that it is only known

from one collection, it seems prudent to wait with

a formal description until more collections are

available and a generic determination can be made

without doubt.

Aniba Aublet

REFERENCE— K. Kubitzki, Aniba. Flora Neotro-

pica31: 1-84. 1982.

Small to medium-sized trees (rarely shrubs or very

large trees). Leaves alternate, well-spaced along the stems

or in distal clusters; leaf blades usually elliptic to oblong

or lanceolate (rarely cordate), entire, glabrous above, gla-

brous or puberulent below, pinnately veined. Inflores-

cences in the axils of leaves or caducous bracts, solitary,

paniculate with relatively short lateral branches, bracts

38

FIELDIANA: BOTANY

caducous, pedicels well developed. Flowers usually small,

bisexual, with a conspicuous (often urceolate) floral tube

which enlarges in fruits, perianth of 6 parts in 2 whorls,

equal or the outer smaller than the inner; stamens 9 in

3 whorls, 2-thecous, the outer 6 with introrse or apical-

introrse dehiscence, filaments as wide and thick as the

anthers (less often slender), inner 3 stamens (rarely stam-

inodial) extrorse or extrorse-lateral and each with 2 large

sessile glands, staminodes (series IV) small and stipiti-

form or absent; pistil slender, ovary ellipsoid or ovoid,

glabrous or puberulent, included in the floral tube, style

slender, stigma minute. Fruits borne in a deeply cupulate

or hemispheric receptacle, rim simple or rarely double-

margined, often with wartlike spots or lenticel-like pits

in the outer surface.

A Neotropical genus of 41 species in six species

groupings, ranging from Costa Rica to Bolivia and

Brazil. The genus is distinguished by the nine 2-

thecous anthers, poorly developed or absent stam-

inodes, slender pistils, well-developed floral tube,

and deep fruiting cup. The often thick (poorly dif-

ferentiated) filaments, floral tube, slender pistils,

and occasionally double-rimmed cups indicate a

close relationship with Licaria; also, their foliage

is similar. This genus was unknown in Central

America until 1982.

Aniba venezuelana Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 63. 1889. Figure 13.

Trees 10-20 m tall, leafy branchlets 2-6 mm thick,

very minutely (0.05-0.1 mm) puberulent with brownish

hairs in early stages, becoming terete and pale brown,

with large central pith. Leaves alternate (occasionally

subopposite), well spaced along the stems, petioles 7-15

mm long, 2-3.5 mm thick, orange brown, with 2 adaxial

ridges usually forming a narrow sulcus above; leaf blades

1 1-27 cm long, 5-10 cm broad, elliptic to elliptic-oblong

or elliptic-obovate, short-acuminate at the apex with a

tip 0.5-2 cm long (occasionally caudate-acuminate), acute

to obtuse at the base, margin entire and slightly revolute

(especially at the base), drying very stiffly chartaceous to

subcoriaceous, dull grayish or yellowish green and the

midvein flat or slightly raised above, yellowish green

beneath and glabrous or sparsely puberulent along the

midvein, with 4-9 major secondary veins on each side,

the central secondaries arising at angles of 50°-70°, small-

er veins slightly raised beneath and forming a poorly

defined reticulum with areolae 0.5-0.9 mm in width.

Inflorescences axillary to distal leaves or near-terminal

from leafless nodes, 7-18 cm long with distant lateral

branches and relatively few (30-40) flowers, peduncles

5-8 cm long, very minutely (0.05 mm) papillate-puber-

ulent, reddish brown, pedicels ca. 2 mm long. Flowers

2-3 mm long (including the obconic floral tube), 2-2.6

mm broad, greenish, minutely (0.05 mm) papillate-pu-

berulent on the outside, floral cup 0.5-1 mm long and

glabrous within, perianth parts subequal; outer stamens

ca. 1 mm long and 0.7 mm broad, puberulent, the fila-

ment not differentiated from the anther, thecae dehiscing

near the apex with small flaps, inner stamens poorly

developed in our material, staminodes absent; pistil 1.8

mm long, puberulent, ovary ca. 0.5 mm thick, style ca.

0.7 mm long, stigma minute. Fruits enclosed in the lower

part by a reddish brown cup ca. 2 cm long; berry ellip-

soid, ca. 3.5 cm long and 2 cm in diameter.

Trees of evergreen rain forest formations in the

Caribbean lowlands of Costa Rica, to as high as

2300 m elevation in Venezuela. This species has

only been collected near Siquirres and Puerto Vie-

jo de Sarapiqui from 100 to 300 m elevation (but

see below). Flowers were collected in June (Gom-

ez-Laurito 79, CR, usj) and July (Hammel &

Trainer 13111, DUKE); fruits were collected in Au-

gust (Hammel 13366, DUKE). The species, as here

interpreted, is found in Costa Rica, the Choco of

Colombia, and in Venezuela.

Aniba venezuelana is recognized by the stiff, ob-

long, essentially glabrous leaves on thick sulcate

petioles and the small puberulent flowers with well-

developed floral tube and nine stamens, each with

only two thecae. Costa Rican material is quite

similar to Cuatrecasas 21541 from Choco and also

resembles a photo of Fendler 2394 from Vene-

zuela; the only other collection is the lectotype

Funck & Schlim 569 from Venezuela. Kubitzki

identified our Costa Rican material as Aniba,

probably new species related to A. intermedia

(Meissn.) Mez. In Costa Rican material the anthers

appear to open upwardly, and while the filaments

seem thicker, and the form of the stamen some-

what different, we prefer to place this material

under A. venezuelana at the present time. As Ku-

bitzki mentions in his monograph, a large number

of species in this genus are known only from a few

collections and, despite his excellent monograph,

many specific concepts may have to be revised as

more material is collected. Two sterile collections

from the Caribbean slope are very probably this

species: Gomez- Laurito 9900 (CR) and Poveda 1021

(CR).

Beilschmiedia Nees

REFERENCE— A. J. G. H. Kostermans, A mono-

graph of the genera: Anaueria, Beilschmiedia

(American species) and Aniba. Recueil Trav. Bot.

Neerl. 35: 834-928. 1938.

Trees or shrubs. Leaves alternate or subopposite (rare-

ly opposite); leaf blades chartaceous to coriaceous, gla-

brous to puberulent, often glaucous beneath, venation

pinnate, the minor venation usually elevated (raised) on

the dried leaf surfaces above and/or below and often

BURGER: FLORA COSTARICENSIS

39

forming a raised reticulum with small areolae, domatia

absent. Inflorescences paniculate and usually few-

branched and with relatively few flowers, solitary and

axillary to distal leaves or clustered at the tips of branch-

es. Flowers bisexual and small, narrowly campanulate

in our species, perianth parts 6 in 2 whorls, equal or the

outer somewhat shorter than the inner, deciduous (some-

times coming off as a whorl united at the base); stamens

9, 2-valved and free (said to be 4-valved in B. sulcata),

the 6 outer with filaments or subsessile, outer anthers

usually ovate and flattened, usually with the connective

slightly prolonged distally beyond the thecae, introrse,

the 3 inner stamens with well-developed filaments and

2 basal glands, with narrow anthers dehiscing laterally

or extrorse, staminodes 3 and well-developed, ovate to

triangular and narrowed to an acute or apiculate tip,

cordate to truncate and sessile or stipitate; pistil with

ovoid or subglobose ovary, glabrous (except in B. rigida),

gradually narrowed into a thick style, stigma simple and

often oblique. Fruits not subtended by an enlarged re-

ceptacle, pedicel thickened but not conspicuously ex-

panded beneath the fruits; berry usually ellipsoid and

rounded at the apex, 2-15 cm long, outer fleshy layer

usually thin.

The genus Beilschmiedia ranges from Africa and

southern Asia to Australia, New Zealand, and the

New World tropics. The Costa Rican species dis-

play a series of concordant character states, sug-

gesting that the genus may be a natural and mono-

phyletic group. The well-developed staminodes,

2-thecous anthers often with an extended apical

connective, and simple or oblique stigma char-

acterize the flowers. The usually ellipsoid fruits

subtended only by a stout pedicel, and a general

tendency for the minor venation to be elevated on

the surfaces of the dried leaves, further distinguish

Beilschmiedia. Nevertheless, fruits, flowers, and

vegetative characteristics seem to vary greatly from

tree to tree, and make species delimitation diffi-

cult. The species concepts used here should be

considered no more than tentative; see the dis-

cussions under the species.

Key to Species of Beilschmiedia

la. Leaves relatively thin in texture (usually chartaceous), 10-25 cm long, broadly ovate to obovate,

usually abruptly rounded and short-acuminate at the apex; known only from below 500 m elevation

(in Costa Rica) 2a

Ib. Leaves usually stiffly chartaceous to coriaceous, 5-16(-20) cm long, narrowly to broadly elliptic,

ovate or suborbicular, rarely short-acuminate when abruptly rounded at the apex; known only from

above 500 m elevation 3a

2a. Lower leaf surface with short straight erect hairs to 0.6 mm long, with 4-14 pairs of secondary

veins; fruits becoming more than 6 cm long B. anay

2b. Lower leaf surface glabrous or with minute (0.2 mm) appressed hairs, with 3-7 pairs of secondary

veins; fruits not exceeding 5 cm in length B. sulcata

3a. Leaves usually broadly ovate to suborbicular, usually coriaceous, reticulum of fine venation forming

areolae 0.1-0.3 mm broad; evergreen montane forest formations from (1 100-) 1800-2800 m ele-

vation B. ovalis

3b. Leaves usually elliptic to oblong, often stiffly chartaceous but not usually coriaceous, reticulum of

fine venation forming areolae 0.5-1.5 mm broad on the lower surface (or the areolae not well

developed); evergreen or partly deciduous forest from 600-1800 m elevation (in Costa Rica) ....

B. pendula

Beilschmiedia anay (S. F. Blake) Kosterm., Re-

cueil Trav. Bot. Neerl. 35: 847. 1938. Hufelan-

dia anay S. F. Blake, J. Wash. Acad. Sci. 9: 459.

1919. Figure 7.

Trees to 22 m tall, bark dark brown to reddish brown,

inner bark orange, leafy branchlets 2-5 mm thick, at first

densely puberulent with slender brownish hairs 0.1-0.5

mm long. Leaves alternate and often crowded distally,

petioles 10-22(-35) mm long, 1.5-3 mm thick, densely

brownish puberulent, flat or slightly sulcate above; leaf

blades (9-) 1 3-25 cm long. 5.5-9(-l 2) cm broad, broadly

elliptic to ovate, gradually or abruptly narrowed and

rounded to a short acuminate (ca. 1 cm) apex, obtuse to

acute at the base, drying stiffly chartaceous, upper surface

minutely puberulent above the veins, but glabrous and

with the minor venation slightly raised between the veins,

lower surface conspicuously soft puberulent with slender

brownish hairs 0.2-0.6 mm long, with 4-10(-14) major

secondary veins on each side, tertiary venation often

subparallel, smaller veins forming a slightly raised retic-

ulum with areolae 0.2-0.5 mm broad, sometimes glau-

cous beneath. Inflorescences axillary to distal leaves,

paniculate but few-branched, 6-15 cm long, primary

peduncle to 9 cm long, ca. 1 mm thick and minutely

40

FIELDIANA: BOTANY

brownish, puberulent, pedicels 2-5 mm long. Flowers

ca. 3 mm long and 3 mm broad, narrowly campanulate,

sparsely puberulent on the outside, perianth parts 2-2.5

mm long, 1-1.5 mm broad; outer stamens subsessile on

very short (ca. 0.3 mm) puberulent filaments, anthers

ca. 1 mm long, with or without a distally prolonged (0. 1-

0.4 mm) connective, inner stamens ca. 2 mm long and

biglandular, staminodes 1-1.5 mm long, triangular to

ovate and apiculate at the apex, truncate and sessile to

stipitate and cordate, puberulent at the base; pistil ca. 2

mm long, ovary gradually narrowed to the style and

equal in length, stigma simple and oblique. Fruits not

seen, said to be 6-15 cm long and 2.5-6 cm thick, ellip-

soid-pyriform, skin thin and the seed very large, becom-

ing black.

Rarely collected trees of wet evergreen forest

formations from near sea level to 500 m elevation

on both the Pacific and Caribbean slopes in Gua-

temala. Hammel (1986) reports that the species

flowers in July and fruits in February at La Selva.

Blake (1919) cited reports of flowering in May; he

also listed Guatemalan specimens flowering in De-

cember-January and fruiting in August-Septem-

ber. At present the species is only known from

Guatemala and Costa Rica.

Beilschmiedia anay is recognized by its large

avocado-like fruits, small flowers with nine 2-the-

cous anthers, and the broad thin-textured leaves

with soft brownish hairs beneath. The leaves, often

rounded and short-acuminate at the apex, resem-

ble those of Ocotea mollifolia. Beilschmiedia al-

loiophylla (Rusby) Kostermans from Colombia

may prove to be a synonym of this species.

Beilschmiedia ovalis (S. F. Blake) C. K. Allen, J.

Arnold Arbor. 26: 418. 1945. Hufelandia ovalis

S. F. Blake, J. Wash. Acad. Sci. 9: 461. 1919.

Perseaaustin-smithiiSiandley, Publ. Field Mus.

Nat. Hist., Bot. Ser. 18: 1552. 1938. B. austin-

smithii (Standley) C. K. Allen, loc. cit. 418.1 945.

Figure 5.

Small to medium-sized trees, 7-20 m tall, leafy

branchlets 3-6 mm thick, minutely (0.2 mm) brownish-

puberulent at first, becoming glabrescent and dark in

color. Leaves alternate and usually clustered at the tips

of branchlets, petioles 6-20 mm long, 2-3 mm thick,

with 2 adaxial ridges forming a shallow sulcus or flat

above; leaf blades 5-14(-18) cm long, 3-8(-10) cm broad,

broadly ovate to broadly elliptic or suborbicular, obtuse

to rounded at the apex (rarely short-acuminate), obtuse

to truncate or rounded at the base, drying subcoriaceous

and often yellowish or reddish brown, glabrous or mi-

nutely puberulent above the veins, flat or with the minor

venation slightly raised to form an obscure reticulum,

lower surface often whitish-glaucous, with slender hairs

0.3-0.5 mm long near the veins or with minute (0. 1 mm)

appressed ascending hairs over the lower surface, with

(3-)4-6(-8) major secondary veins on each side, the

smaller venation often raised beneath and forming a

reticulum of minute (0.1-0.3 mm) areolae. Inflores-

cences axillary to distal leaves, 4-20 cm long, paniculate

with short lateral branches and relatively few flowers,

peduncle 2-9 cm long, 1-2.3 mm thick, minutely pu-

berulent, pedicels 1-2 mm long. Flowers ca. 3.5 mm

long and 3.5 mm broad, puberulent, perianth parts ca.

1.8 mm long and 1.3 mm broad; outer stamens 1.4-2

mm long, anthers oblong and 1-1.4 mm long, often with

the connective slightly developed beyond the two thecae,

inner stamens biglandular, staminodes subsessile, 0.8-

1.2 mm long, ovate with an apiculate apex; pistil ca. 2

mm long, ovary gradually narrowed into the style, stigma

simple. Fruits borne on a slightly thickened (ca. 3 mm)

pedicel, the receptacle not expanded; berry ovoid or short-

ellipsoid, ca. 3 cm long and 2 cm in diameter, becoming

black.

Infrequently collected trees of montane ever-

green wet forest formations along the northern edge

of the Cordillera Central (from Palmira in Alajuela

to Cerro de las Vueltas in San Jose), and in the

southern part of the Cordillera de Talamanca and

adjacent Chiriqui, at altitudes of ( 1 1 00-) 1 800-2800

m. Flowers have been collected in March-May;

fruits have been collected in September and March.

This species appears to range from Costa Rica into

the northern Andes.

Beilschmiedia ovalis is recognized by the stiff,

almost glabrous, rounded leaves with fine reticu-

lation on the lower surfaces, small puberulent

flowers with nine 2-thecous anthers, fruits sub-

tended by a narrow stalk, and higher-elevation

habitat. This species resembles some specimens

of Persea vesticula, which is found at similar ele-

vations. Submersion of Beilschmiedia ovalis under

B. sulcata, as suggested by Kostermans (1938),

seems ill advised. A Ruiz and Pavon collection at

F has larger somewhat obovate leaves with short-

acuminate apices, closely resembling the original

illustration of Laurus sulcata Ruiz & Pavon, but

atypical of B. ovalis. In addition, this Ruiz and

Pavon collection has thin-textured leaves very un-

like the leaves of B. ovalis.

An unusual collection of Beilschmiedia, Gomez-

Laurito 9800 (CR, F), is tentatively placed here. It

has narrower elliptic leaves with more veins (eight

per side) and much longer (6.5 x 2.8 cm) narrowly

ellipsoid fruits. This collection was made between

1 300 and 1 500 m at Cerros de La Palma de San

Ramon in late January. Another unusual collec-

tion is Zamora 1215 (CR) from Muneco de Car-

tago, with fruiting in March and with larger (to 23

x 16 cm) subopposite leaves. This last collection

resembles Hartshorn 2 166 (CR, F), and the two may

represent larger-leaved trees from lower elevation.

BURGER: FLORA COSTARICENSIS

41

Beilschmiedia pendula (Sw.) Hemsley, Biol. centr.

amer., Bot. 3: 70. 1882. Laurus pendula Sw.,

Prodr. 65. 1783. Hufelandia costaricensis Mez

& Pittier, Bull. Herb. Boissier, ser. 2, 3: 228.

1903. B. costaricensis (Mez & Pittier) C. K. Al-

len, J. Arnold Arbor. 26: 415. 1945. B. brenesii

C. K. Allen, loc. cit. 415. 1945. Cryptocarya

kostermansiana C. K. Allen, loc. cit. 423. 1945.

Figure 13.

Trees 6-30(-40) m tall, bark smooth in younger trees

and developing square patches in age, inner bark reddish,

leafy branchlets 1.5-3.5 mm thick, minutely yellowish

brown, appressed, puberulent at first but quickly becom-

ing glabrous. Leaves alternate (subopposite or opposite

in B. brenesii, see below), petioles 6-15 mm long (to 28

mm in Panama), 1-2 mm thick, with 2 adaxial ridges

forming a shallow sulcus above; leaf blades 5-16(-20)

cm long, 3-6(-9) cm broad, elliptic to oblong-elliptic or

occasionally elliptic-obovate, short-acuminate to acute

or bluntly obtuse at the apex, acute to obtuse at the base

and slightly decurrent on the petiole, margin entire or

undulate, the laminae drying stiffly chartaceous (coria-

ceous), glabrous above or with minute hairs above the

major veins, usually with the minor venation raised above

but not forming a well-defined reticulum, lower surface

glabrous or with minute (0.2 mm) straight slender ap-

pressed-ascending hairs, with 4-7 major secondary veins

on each side, smaller venation raised beneath and often

forming an irregular reticulum of areolae 0.5-1.5 mm

broad. Inflorescences axillary to distal leaves, 2-8(-15)

cm long, paniculate or racemose with short lateral

branches, peduncle 1 .5-3 cm long and 1 mm in diameter,

sparsely and minutely puberulent, pedicels 1.5-4 mm

long. Flowers 2.3-3.5 mm long, 2-3 mm broad, sparsely

and minutely puberulent on the outside, perianth parts

ca. 1.7 mm long and 1.3 mm broad, sometimes dehiscing

as a whorl united at the base; outer stamens with short

filaments or subsessile, anthers 0.8-1 mm long, ca. 0.6

mm broad, oblong or narrowed at the apex and often

with the connective prolonged distally beyond the the-

cae, inner stamens 1.5-2.8 mm long, staminodes 0.8-1

mm long and ca. 0.7 mm broad, triangular to ovate and

often apiculate at the apex, cordate to truncate and sessile

or subsessile, puberulent at the base; pistil 1.5-2.1 mm

long, style 0.4-0.9 mm long, stigma simple. Fruits borne

on slightly thickened (to 5 mm) pedicels but the recep-

tacle not expanded and the perianth deciduous; berry

ellipsoid, 2-5.5 cm long, 1-3 cm in diameter, green be-

coming dark purple or black, pericarp thin (ca. 1.5 mm).

Trees of evergreen or partly deciduous forest

formations from 600 to 2000 m elevation in Costa

Rica, but close to sea level in central Panama and

elsewhere. Flowering collections have been made

in December-May, and fruits have been collected

in February-September in Costa Rica. The species

occurs in the Cordillera de Tilaran, the north-

western part of the Meseta Central (San Ramon

to Zarcero) and the Cordillera de Talamanca. The

species ranges from Mexico to Panama, and in the

West Indies from Cuba to northern Venezuela.

Beilschmiedia pendula is recognized by the often

smaller elliptic, essentially glabrous, leaves with

tertiary veins raised on one or both surfaces, the

nine 2-thecous anthers, subsessile staminodes, and

ellipsoid fruits lacking an expanded receptacle. This

species may resemble small-leaved specimens of

Ocotea meziana, Nectandra cufodontisii , and

Aiouea costaricensis, which share some of the same

montane habitats but differ in leaf venation. This

species displays considerable variation, both in the

West Indies and in our area. The Costa Rican

collections differ from Caribbean material in hav-

ing more elliptic and slightly more puberulent

leaves. Our highland plants (above 1000 m) may

be worthy of subspecific recognition (but see be-

low). Common names recorded in Costa Rica are:

Chanco bianco, Come negro, Tiguissaro, and Vo-

lador.

The present circumscription of Beilschmiedia

pendula in Costa Rica includes a variety of ma-

terial, and it is probable that more than one species

is included here. Beilschmiedia brenesii may be a

separate species; it often has opposite or clustered

leaves with short petioles and thickened nodes.

Unfortunately, there is so much variation in B.

pendula that it is difficult to know if the charac-

teristics of B. brenesii represent a different species

or a local variety. Likewise, some of the collections

from the General Valley below 1 000 m elevation

resemble collections from Barro Colorado Island

in Panama, and may represent another entity.

These plants deserve careful study in the field.

Beilschmiedia sulcata (Ruiz & Pavon) Kosterm.,

Recueil Trav. Bot. Neerl. 35: 850. 1938. Laurus

sulcata Ruiz & Pavon, Laurografia t. 11. ca.

1830, and Fl. peruv. prodr. 4: pi. 356; text pub-

lished in Anal. Inst. Bot. Cavanilles 13: 21. 1954.

Figure 15.

Trees or shrubs, to 30 m tall, leafy branchlets 1.5-4.5

mm thick, minutely brownish, appressed, puberulent.

Leaves alternate, petioles 6-18 mm long, 1.5-2.5 mm

thick, minutely (0. 1 mm) appressed, puberulent with 2

adaxial margins usually forming a narrow (0.5 mm) and

deep (0.7 mm) sulcus above distally; leaf blades 8-18

cm long, 3.5-9(-l 2) cm broad, broadly elliptic to elliptic-

obovate or elliptic-oblong, abruptly narrowed to an ob-

tuse or rounded and short-acuminate apex, obtuse and

very slightly decurrent at the base, margin entire or un-

dulate, drying stiffly chartaceous, glabrous above but with

minute hairs usually present on the slightly raised mid-

vein, the minor venation slightly raised above, lower

42

FIELDIANA: BOTANY

surface glabrous or with minute (0.2 mm) thin appressed

parallel hairs, with 4-7 major secondary veins on each

side, smaller veins raised beneath and forming a loose

reticulum with areolae 0.2-0.7 mm broad. Inflorescences

shorter than the leaves, solitary in distal leaf axils, pan-

iculate, minutely appressed puberulent. Flowers not seen

(depicted as having 4-thecous anthers by Ruiz and Pa-

von). Fruits borne on slightly thickened (2-6 mm) ped-

icels; berry 3.5-4.5 cm long, ca. 2 cm in diameter, be-

coming purple at maturity.

Rarely collected trees of evergreen rain forest

formations in the Caribbean lowlands of Costa

Rica (La Selva and near Limon) and ranging into

middle elevation ( 1 000 to 2000 m) evergreen for-

ests on the eastern slopes of the Andes in Peru.

Hammel reports (1986) flowering at La Selva in

January-February, with fruits maturing in Sep-

tember. The present circumscription of this name

is very uncertain (see below).

Beilschmiedia sulcata is recognized by its rela-

tively thin broad nearly glabrous leaves usually

short-acuminate at the apex, raised minor vena-

tion on both leaf surfaces, nine 2-thecous anthers,

and ellipsoid fruits on slightly thickened pedicels.

The following collections are tentatively placed

here: Hammel 7/705, 7/766 (DUKE), Hartshorn

1519 (CR, F), Little 20060 (us), and Pittier 16140

(us). They are quite similar to both the original

illustration and to a Ruiz and Pavon collection

from Muna, Peru, bearing the original name at

Field Museum. Hammel (1986) placed his collec-

tions under the name B. mexicana (Mez) Koster-

mans, but that species will probably prove to be

a synonym of B. pendula. Both Kostermans and

Bernardi included B. ovalis as a synonym under

B. sulcata, but this appears to be incorrect (see the

discussion under that species). The original illus-

tration shows the anthers as 4-thecous and a recent

collection from Ecuador is also 4-thecous. Unfor-

tunately, flowering material has not been seen from

Central America. Lack of collections and the vari-

ability found in other species of Beilschmiedia make

the present circumscription of this species very

uncertain.

Caryodaphnopsis Airy-Shaw

REFERENCES— A. J. G. H. Kostermans, A mono-

graph of Caryodaphnopsis. Reinwardtia 9: 123-

137. 1974. H. van der Werff & H. G. Richter,

Caryodaphnopsis Airy-Shaw (Lauraceae), a genus

new to the Neotropics. Syst. Bot. 10(2): 166-173.

1985.

Trees, branchlets terete or quadrangular. Leaves op-

posite or subopposite, petiolate, elliptic to ovate-elliptic,

glabrous to very sparsely puberulent, often tripliveined.

Inflorescences axillary or pseudoterminal, paniculate with

opposite branches, bracts and bracteoles minute and ca-

ducous, pedicels slender. Flowers bisexual, floral tube

short, perianth of 6 parts in 2 whorls of 3, strongly un-

equal, the outer whorl much shorter than the inner; sta-

mens 9 in 3 series, anthers 4-thecous (2-thecous in the

South American C. inaequalis) outer stamens introrse

with the thecae in a horizontal row or arc, inner stamens

with extrorse dehiscence and biglandular, 3 staminodes

relatively large and stipitate, cordate-sagittate; pistil gla-

brous, style long and slender, stigma minute. Fruits sub-

tended by an obconical or slender pedicel, perianth de-

ciduous, a disc or cupule not developed; berry small and

globose (in ours) to large and pyriform, green to yellowish

green, glabrous, seed large.

A genus of about 14 species ranging from south-

ern China and Indochina to the Philippines and

Borneo in the Old World, and from Costa Rica to

Ecuador and Peru in the upper Amazon Basin.

This genus is distinguished by the opposite leaves,

usually with three prominent veins from the base,

and the flowers with strongly unequal tepals. Two

of the Neotropical species were originally placed

in Persea, but Persea does not have opposite leaves.

Also, the flowers of Caryodaphnopsis differ from

those of Persea in having shorter stamens and

broader anthers, with the lower thecae usually lat-

eral to the upper. Recent collections of Caryoda-

phnopsis show that the Neotropical species fall

into two groups. One group includes species with

pinnately veined (or subtripliveined) leaves and

avocado-like fruits; the other group includes species

with strongly tripliveined leaves and small, glo-

bose fruits. Our species belongs in this latter group.

Caryodaphnopsis burger! Zamora & Poveda, Ann.

Missouri Bot. Gard. 75: 1160. 1988. Figure 1.

Trees to 30 m tall and 80 cm d.b.h., leafy branchlets

1.3-6 mm thick, glabrous or very minutely (0.1 mm)

appressed, puberulent in early stages, becoming dark in

color; dichotomous branching sometimes present (from

adjacent axillary branches at a node with an aborted

apex). Leaves opposite or subopposite, petioles 7-12(-18)

mm long, 1 .3-2.4 mm thick, glabrous and dark, rounded

or narrowly sulcate above; leaf blades 5- 1 2(- 1 6) cm long,

1.5-5(-8) cm broad, elliptic to elliptic-oblong or oblong,

short-acuminate at the apex, the narrowed tip 5-12 mm

long, acute to obtuse at the base, margin entire and with

a thickened texture when dry, drying stiffly chartaceous,

glabrous and grayish green to pinkish brown above, gla-

brescent and whitish green beneath (minutely appressed,

puberulent on the veins beneath in young foliage), with

3 major primary veins extending from near the base to

the apex (tripliveined), secondary veins difficult to see

BURGER: FLORA COSTARICENSIS

43

and weakly subparallel between the primary veins. Flow-

er buds ca. 5 mm long, flowers rotate and ca. 10 mm

broad at anthesis, outer perianth whorl only 1 mm long,

inner whorl ca. 5 mm long; outer stamens ca. 5 mm long

with long slender filaments, outer anthers ca. 1.5 mm

long, the lower thecae lateral to the upper, inner stamens

5 mm long, with glands attached to their filaments above

their base, staminodes ca. 2 mm long, with triangular

apex and hairy stipe ca. 1 mm long; pistil ca. 4 mm long,

sparsely puberulent, ovary ellipsoid, style slender and

2.5 mm long, stigma bilobed. Fruits borne in short (2-

4 cm) infructescences with only 1 or 2 fruits, pedicels

ca. 5 mm long and 1 mm thick, receptacle not expanded;

berry ca. 15mm long and 1 3 mm in diameter, subglobose

or globose-oblong, pale olive green and with a smooth

surface when dry.

Trees of evergreen forest formations of the Pa-

cific slope of central and southern Costa Rica from

near sea level to 500 m elevation. Flowers and

fruits were collected in March. The species is en-

demic to Costa Rica.

Caryodaphnopsis burgeri is unique among our

native species of Lauraceae with its opposite or

subopposite tripliveined leaves. The small round-

ed fruits lacking a cup or disc, the flowers with

very short outer tepals, and the stamens with long

filaments and with lower thecae lateral to the upper

are also unusual. This species was first discovered

by Luis Poveda in January 1984, and represented

the first record for the genus in Central America.

The type is Zamora et al. 1208 (CR, the holotype,

F).

Cassytha Linnaeus

Slender stemmed herbaceous plants, the stems with

small haustoria attaching and obtaining nourishment

from host plants, yellowish to orange or greenish. Leaves

reduced to minute sessile scales, alternate in a spiral.

Inflorescences solitary to several in the axils of scale

leaves, spicate to racemose or capitate; flowers sessile or

on short pedicels, subtended by a small bract and 2 brac-

teoles. Flowers bisexual and small, perianth of 2 whorls

with 3 parts each, the outer whorl smaller and resembling

the bracts, perianth persisting, floral tube shallow but

developing to enclose the fruits; fertile stamens 9 in 3

series (whorls), 2-thecous, the outer 6 (series I-II) with

introrse dehiscence, the inner 3 with extrorse dehiscence

and each biglandular, staminodia (series IV) present and

sessile or stalked; pistil simple, stigma small and discoid.

Fruits completely included in the enlarged and succulent

floral tube (calyx tube or hypanthium) with a small open-

ing at the apex with erect persisting perianth parts; testa

membranous to coriaceous, cotyledons thick and becom-

ing united (giving the impression of a carnose endo-

sperm).

A genus of about 20 species with all but one

native to the Old World tropics; the largest num-

ber occur in Australia, while one species is pan-

tropical. The plants are profoundly different from

all other Lauraceae in habit and method of nutri-

tion, but the flowers are typical of the family, re-

sembling the flowers of Cryptocarya in particular.

These plants bear a striking resemblance to another

twining herbaceous parasite: Cuscuta of the Con-

volvulaceae. The slender yellow orange stems

climbing over low herbaceous vegetation are very

similar in the two genera but the flowers and fruits

are very different.

Cassytha filiformis L., Sp. PI. 35. 1753.

Herbaceous twining parasites to 1 or 2 m long, peren-

nial, stems 0.3-3 mm thick (dry), minutely puberulent

with thin straight hairs ca. 0.2 mm long or glabrous,

yellowish to orange or olive green, cupulate or discoid

haustoria ca. 1 mm broad and 0.5 mm high present on

the stems. Leaves scalelike, 1-2 mm long, 1 mm broad

at the base. Inflorescences solitary or paired, 1-5 cm

long, spicate, puberulent or glabrous, the flowers sessile

and subtended by 3 tepal-like bracts (1 bract and 2 brac-

teoles). Flowers ca. 2 mm long, greenish or white, outer

perianth parts ca. 0.7 mm long, inner perianth 1.3-1.7

mm long and ca. 1.5 mm broad, thin and pellucid-punc-

tate; outer stamens 0.9-1 . 1 mm long with short (0.3 mm)

broad filaments, anther narrowly ovate to triangular and

narrowed apically, inner stamens 1.2-1.4 mm long with

narrowly triangular anthers and distally prolonged nar-

row connective, glands sessile, staminodia minute or not

readily visible; pistil ovoid, ca. 1.3 mm long, ovary ca.

0.6 mm thick and gradually narrowed apically (the style

not clearly differentiated), stigma slightly discoid. Fruits

included in the expanded fleshy floral tube 5-6 mm long

and 4-6 mm in diameter, urceolate but becoming glo-

bose in late stages, perianth persisting, greenish.

Twining parasitic plants in herbaceous vegeta-

tion and low shrubs in evergreen or partly decid-

uous formations from near sea level to 1000 m

elevation. Flowering and fruiting collections have

been made in all months of the year except De-

cember in southern Mexico and Central America.

This species is now pantropical; it ranges from

southernmost Florida (U.S.A.) and southern Mex-

ico to the West Indies and South America.

Cassytha filiformis is recognized by its slender

yellow to greenish twining stems with small haus-

toria that parasitize herbaceous and small woody

plants. The 6-parted flowers, nine stamens with 2-

thecous anthers opening by flaps, and 1 -seeded

berries enclosed in a fleshy floral tube distinguish

this species from the very similar species of Cus-

cuta (Convolvulaceae). No other genera of Lau-

raceae are either herbaceous or parasitic. This

species was first collected in Costa Rica in January

44

FIELDIANA: BOTANY

1987 along Laguna Gandoca in easternmost Li-

mon Province (Grayum et al. 8064, CR, MO).

Cassytha paradoxa Proctor, published in Mos-

cosoa 2:20, 1983, from Honduras, Surinam, and

Brazil, with solitary flowers, may prove to be a

depauperate variant of C. filiformis.

Cinnamomum Schaeffer, nomen conservandum

Shrubs or trees, usually with aromatic bark and fo-

liage. Leaves alternate or opposite, petiolate, usually co-

riaceous and tripliveined, less often pinnately veined.

Inflorescences axillary or pseudoterminal, solitary or fas-

ciculate, paniculate. Flowers bisexual (rarely unisexual

and the female larger than the male), floral tube funnel-

form and enlarging in fruits, perianth 6-parted in 2 whorls,

the parts equal or subequal; fertile stamens 9 (rarely 6),

usually 4-thecous with the thecae superposed in an arc,

with slender filaments usually equalling the anthers in

length, 6 outer stamens dehiscing introrse and without

glands, 3 inner stamens with stipitate glands and de-

hiscing extrorse, staminodes stipitate and ovate to sag-

ittate; pistil with sessile ovary and slender style, stigma

discoid to peltate. Fruits usually ellipsoid, fruiting re-

ceptacle a cup with an entire margin or with persisting

tepals or perianth-bases.

Cinnamomum is a genus with more than 200

species in Southeast Asia, Malaysia, Australia, and

the Pacific Islands. The genus includes a number

of important economic and ornamental trees. The

bark of C. verum J. Presl (syn. C. zeylanicum

Blume) provides the cinnamon of commerce, while

C. cassia (Nees) Nees & Eberm. ex Blume is often

used as a substitute. Cinnamomum camphora (L.)

J. Presl has been the source of camphor. These

trees resemble some of our native species of Phoebe,

but they are not common in Central America where

they are occasionally planted in parks and gardens.

Specific descriptions are not provided but the fol-

lowing key should serve to differentiate the species

likely to be found in cultivation.

Key to Commonly Cultivated Species of Cinnamomum

1 a. Apical and lateral buds at first enclosed in large broad-based imbricate scales, leaves usually alternate,

ovate-elliptic, to 1 4 cm long, often with pinnate venation; bark with the odor of camphor; trees to

25 m tall C. camphora

1 b. Apical and lateral buds not enclosed by large scales; leaves conspicuously tripliveined and usually

opposite; bark with odor of cinnamon; trees to 1 2 m tall 2a

2a. Leaves oblong to lanceolate and long acuminate to caudate-acuminate, to 1 4 cm long; panicles as

long as the leaves C. cassia

2b. Leaves ovate to narrowly ovate-oblong, obtuse to acute, to 18 cm long; panicles longer than the

leaves . . C. verum

Endlicheria Nees, nomen conservandum

Trees or shrubs, dioecious. Leaves alternate or rarely

verticellate, laminae usually puberulent beneath, vena-

tion usually pinnate (rarely tripliveined or palmate). In-

florescences axillary or subterminal, few- to many-flow-

ered, the female inflorescence usually smaller than the

male and with shorter pedicels, bracts and bracteoles

deciduous or persisting. Flowers small and unisexual,

with a small floral tube, perianth 6-parted in 2 whorls

and the whorls equal or subequal; male flowers with 9

fertile stamens in 3 series (the outer 2 series often ap-

pearing as a single whorl, rarely the inner whorl sterile

and with 6 fertile stamens), all stamens 2-thecous (the

inner series 4-thecous in E. anomala), the outer 2 series

with introrse dehiscence, sessile or with filaments, the

inner 3 stamens with extrorse dehiscence and biglandular

at the base, staminodes absent, ovary slender (stipiti-

form) and nonfunctional; female flowers usually slightly

smaller and with a broader floral tube than the male,

stamens similar to those of the male but smaller and

sterile, ovary included in the floral tube, style thick and

short or long, stigma discoid to 3-lobed. Fruit cups

shallow and concave to deeply cupulate, perianth lobes

deciduous or persisting, the rim often thin, simple; berry

ovoid to ellipsoid, glabrous or rarely puberulent.

A Neotropical genus of about 40 species, nearly

all South American. The unisexual flowers on

dioecious plants, nine 2-thecous stamens, lack of

well-developed staminodes and cupulate fruiting

receptacles distinguish this genus. Endlicheria for-

mosa A. C. Smith has just been collected near

Palmar Norte (Grayum et al. 9153 CR, MO), too

late to be included in the keys and descriptions.

The glabrous elliptic leaves are 12-35 x 4-10 cm

with petioles 1 5-40 mm long, and the tertiary ve-

nation is raised on both surfaces. The thinly

branched inflorescences are 10 cm long with mi-

nute (2x2 mm) flowers having erect tepals ca.

0.5 mm long; the ellipsoid fruits become 3 cm long.

BURGER: FLORA COSTARICENSIS

45

The following species, known only from sterile and

fruiting collections, is placed here provisionally.

Endlicheria sp.? Figure 9.

Trees to ca. 25 m tall and 70 cm d.b.h., leafy branchlets

1.2-3.5 mm thick, at first densely tomentulose with

brownish or reddish brown ascending hairs 0.3-1 mm

long. Leaves alternate and mostly in a single plane, pet-

ioles 3-5 mm long, 1-2.2 mm thick, densely brownish

puberulent, with a narrow sulcus above; leaf blades

(10-) 12-23 cm long, 2.5-6 cm broad, narrowly elliptic-

oblong or lanceolate, acuminate to long-acuminate at the

apex, the tip 8-25 mm long, obtuse to acute at the base,

margin entire, drying chartaceous and dark brownish or

dark grayish brown, glabrous above and with the major

veins impressed, conspicuously brownish puberulent be-

neath with straight slender ascending hairs 0.3-0.8 mm

long over the veins and undersurface (but not obscuring

the surface), with 10-14 major secondary veins on each

side, the secondaries strongly loop-connected about 2-

8 mm from the edge of the lamina. Inflorescences and

flowers unknown. Fruits borne in a deeply (5 mm) cu-

pulate receptacle 12-14 mm broad, ca. 8 mm long and

abruptly expanded above the slightly thickened (3-4 mm)

pedicel, becoming red; berry ovoid but flattened at the

base, 1 7-20 mm long, ca. 1 3 mm in diameter, basal scar

5-7 mm broad, becoming purplish black.

Known only from above the Rio Reventazon

below the CATIE agricultural research institute

near Turrialba, at about 600 m elevation on the

Caribbean slope of central Costa Rica. Mature fruits

were collected by Luis Poveda and Gerardo Sal-

sedo (3795, CR, F) in January. Two sterile collec-

tions from the same area are Holdridge 6590 (CR,

NY) and Poveda 3498 (CR).

Endlicheria sp.? is unique among our species of

Lauraceae because of its thin, narrowly oblong,

acuminate leaves, puberulent beneath, with usu-

ally more than 1 0 pairs of secondary veins strongly

loop-connected near the margin and almost form-

ing a submarginal or "collecting" vein 2-8 mm

from the leaf edge. Further distinctions are the cup

with broad base and thin simple margins, and the

ovoid fruits flattened at the bottom and with a

broad scar at the base. In fact, the dried fruits

resemble an acorn (Quercus). While the generic

placement of this species remains uncertain, the

leaves and fruiting receptacles are reminiscent of

a few South American species of Endlicheria; com-

pare E. multiflora (Miq.) Mez, E. sprucei (Meissn.)

Mez, and E. verticillata Mez. However, it is also

possible that this species might belong in Pleu-

rothyrium.

Licaria Aublet

REFERENCE— Holger Kurz, Fortpflanzungsbiol-

ogie einiger Gattungen neotropischer Lauraceen

und Revision der Gattung Licaria (Lauraceae).

Ph.D. Thesis, Universitat Hamburg, 251 pp. 1982.

Shrubs or trees, bisexual (but see below), glabrous or

puberulent with simple transparent or brownish hairs,

stems terete (in ours). Leaves alternate in a spiral (in

ours) or rarely opposite, simple and often elliptic to lan-

ceolate or oblong in shape, usually acuminate at the apex,

acute to obtuse at the base, margins entire, drying char-

taceous to coriaceous, upper surface glabrous, under-

surface glabrous to hirsutulous, pinnately veined (in ours),

domatia absent. Inflorescences paniculate with few to

many flowers (rarely capitate), solitary in the axils of

distal leaves or caducous scales. Flowers very small (in

ours) or up to 8 mm long, bisexual (perhaps unisexual

in an undescribed Costa Rican species), obovoid to glo-

bose with the tepals opening only near the top, glabrous

or puberulent, outer perianth parts (tepals) often larger

and overlapping the inner, floral tube short to long and

surrounding the ovary; androecium of 3 inner fertile sta-

mens (series III) which may be free or variously connate

and forming a thick column around the slender style,

with 0-6 small or flattened staminodes in the outer series

but usually very difficult to see in ours, filaments usually

not clearly differentiated in the fertile stamens, each fer-

tile stamen with 2 thecae opening longitudinally or api-

cally and extrorse (introrse in subgenus Canella), apex

of the stamen (anther) usually broadly rounded, form of

the stamens appears to vary considerably within some

species, a fourth inner whorl of minute staminodes may

be present in a few species, glands may be present at the

abaxial base of the fertile stamens and free (6) or var-

iously united (0-5); pistil slender, ovary glabrous (in ours)

or sericeous, style slender, stigma simple and small. Fruits

at first enclosed in the enlarged floral tube, fruiting re-

ceptacle forming a deep rounded (often hemispheric) cup

enclosing the lower Vj or '/2 of the fruits, the distal margin

of the cup with 2(-3) usually distinct margins (occasion-

ally poorly developed as in L. sarapiquensis), the outer

rim associated with the perianth bases and the inner rim

developed from the androecial whorls, outer surface often

marked by lenticel-like warts and becoming reddish or

bluish; berry ellipsoid to narrowly ovoid (rarely globose),

smooth, cotyledons plano-convex, filling most of the seed

and enclosing the minute plumule and radicle.

Licaria is a Neotropical genus of about 37 species

in three subgenera (according to Kurz), ranging

from Florida (U.S.A.) and Mexico through Central

America and the West Indies to Bolivia and Brazil.

The unusual androecium of only three stamens

with two thecae each, and the deeply cupulate

fruiting receptacle with two usually distinct mar-

gins on its rim make this a very well-characterized

genus. The stamens are often united to form a

staminal column in the center of the flower, sur-

rounding the style. This column can be difficult to

interpret. In some species the androecium opens

by six very small apical valves that may be hard

to see. In these instances the flower may appear

to be abnormal or teratological. In addition, the

fruit cups may fail to show the double margin,

46

FIELDIANA: BOTANY

especially in L. sarapiquensis. Foliage, also, seems

to vary greatly within species, and it may be very

difficult to be certain of an identification in the

absence of mature flowers.

Acrodiclidium Nees, Chanekia Lundell, and

Misanteca Schlecht. & Cham, are generic syn-

onyms that have been used in Central America

and Mexico.

We have followed Kurz's monograph, but differ

in accepting Licaria cufodontisii as a valid species

and not as a part of the more broadly defined L.

misantlae. Hammel has discovered two new en-

tities at La Selva since Kurz's thesis was written:

L. sarapiquensis and a single problematic collec-

tion, here called Licaria sp. A. The trees of this

genus appear to be uncommon, and collections of

our species are few in number. The paucity of

material, variability of foliage, and the difficulty

of dissecting the minute flowers contribute to mak-

ing Licaria a poorly understood genus. This treat-

ment should be considered provisional; we believe

that there may be more species in Costa Rica than

are recognized here. For a key based on fruits and

leaves see dichotomy 64a in Key 2 at the beginning

of the family.

Key to Species of Licaria

la. Anthers opening by large (0.5 mm) longitudinal extrorse valves, stamens or staminal column often

becoming slightly exserted beyond the perianth; 0-1400 m elevation 2a

Ib. Anthers opening by smaller (0.1-0.4 mm) rounded apical valves, stamens not exserted at anthesis

3a

2a. Leaves glabrous beneath, with 3-8 pairs of secondary veins L. triandra

2b. Leaves usually minutely puberulent beneath, with 1 0-20 pairs of secondary veins

L. multinervis

3a. Stamens clearly narrowed at the base and with erect globose or obovoid glands at their base, anther

valves oblong and 0.2-0.4 mm long; flowers ca. 2 mm long; 0-700 m elevation 4a

3b. Stamens thick at the base, glands broad and flat or absent, anther valves 0. 1-0.2 mm long, rounded;

flowers 2-3 mm long 5a

4a. Anthers apical and extrorse; pistil with a slender ovary; laminae lustrous above and often oblong

L. sarapiquensis

4b. Anthers apical and introrse; pistillode slender and without an ovary; laminae dull and elliptic

L. sp. A

5a. Leaves usually 2-3 times longer than broad, often oblong and caudate-acuminate; 0—400 m elevation

in southeastern Costa Rica L. cufodontisii

5b. Leaves usually 3-4 times longer than broad, often narrowly oblong, rarely caudate-acuminate 6a

6a. Leaves subcoriaceous; distal stems usually solid; often large trees of montane forest formations,

600-2800 m elevation L. excelsa

6b. Leaves drying chartaceous; small trees of evergreen forest formations 7a

7a. Distal stems hollow, leaves glabrous beneath; 500-1400 m on the Caribbean slope . . . L. brenesii

7b. Distal stems solid, leaves minutely puberulent beneath; 10-500 m, Golfo Dulce area

L. pergamentacea

Licaria brenesii W. Burger, sp. nov. Figure 3.

Arbor ca. 3 m alta, ramulis foliiferis 1.5-4 mm crassis,

glabris, fistulosis. Folia alterna, petiolis 5-1 2 mm longis,

2-3 mm latis, laminis 16-30 cm longis et 4-8 cm latis,

anguste oblongis vel anguste oblongo-ellipticis, apice

saepe acuminato, subtus glabris vel leviter puberulis,

nervis secondariis 6-1 1 paribus. Inflorescentiae pani-

culatae, 7-16 cm longae, pedunculis 3-8 cm longis. Flo-

res 2-3 mm longi et 1.5-2.5 mm lati, extusglabri; stam-

ina 3, crassa, conniventia, 0.5-1 mm longa, staminodiis

nullis vel parvulis; gynoecium 1 .2-1 .6 mm longum, ova-

no globoso, ca. 0.8 mm crasso. Fructus ignotus; cupula

2-2.5 cm lata, ca. 5 mm profunda.

Small or medium-sized trees, 3-7 m tall, leafy branch-

lets 1.5-4 mm thick, with prominent longitudinal ribs

or obscurely ribbed, glabrous, becoming grayish and ter-

ete, the young stems usually hollow. Leaves alternate or

occasionally opposite or whorled, petioles 5-12 mm long,

2-3 mm thick, dark in color, with 2 lateral ridges but

sulcate only near the base; leaf blades 1 6-30 cm long,

4-8 cm broad, very narrowly oblong to narrowly oblong-

elliptic, tapering gradually or abruptly to a long-acu-

BURGER: FLORA COSTARICENSIS

47

minate apex (rarely acute), acute to obtuse or slightly

rounded at the base, margin entire, drying chartaceous,

glabrous above with the midvein slightly elevated, gla-

brous beneath or very slightly puberulent on the mid-

vein, with 6-1 1 major secondary veins on each side,

tertiary' venation slightly raised beneath. Inflorescences

solitary in the axils of distal leaves or occasionally borne

in the axils of caducous scales on terminal shoots and

forming compound inflorescences, 7-16 cm long, pani-

culate, few-branched and few-flowered, primary pedun-

cle 3-8 cm long and ca. 1 mm thick, glabrous. Flowers

2-3 mm long and 1.5-2.5 mm broad, yellowish, urceo-

late in shape with a narrow ( 1 mm) opening at anthesis,

drying dark and glabrous on the exterior, outer perianth

parts broader than the inner; stamens 0.5-1 mm long,

ca. 0.8 mm broad, strongly or weakly connivent, filament

broad and thick (not differentiated), anthers rounded

distally and the small (0. 1 5 mm) valves opening distally

or slightly extrorse near the apex of the stamen, stami-

nodes and glands poorly developed or absent; pistil 1 .2-

1 .6 mm long, ovary globose and ca. 0.8 mm in diameter,

style slender, stigma occasionally subcapitate. Fruits

borne in a cupulate receptacle 2-2.5 cm broad with a

conspicuously wide-flaring (3 mm) outer margin, ca. 5

mm deep (but probably flattened in pressing), bright rose

red; fruits unknown.

TYPE— Costa Rica, Alajuela Province, Cataratas

de San Ramon, 26 February 1931, A. M. Brenes

13523 (holotype, F 857810; negative, 61121; is-

otype, CR).

Understory trees of wet evergreen forest for-

mations of the Caribbean slope, between 600 and

1400 m elevation. Flowers have been collected in

February and a mature fruiting receptacle was col-

lected in September. This species has also been

collected from the upper Rio Penas Blancas valley

(below the Monteverde reserve, Burger et al.

10718), and below Los Angeles de San Ramon

(Burger & Antonio 11170) in Alajuela Province.

Licaria brenesii is recognized by its long narrow,

essentially glabrous, thin-textured short-petiolate

leaves, hollow stems, small flowers with partly

united stamens and six minute valves opening near

the top, and the fruiting receptacle with flared dou-

ble-margin. The slender treelet habit, long narrow

leaves, and hollow stems are very similar to those

of Ocotea paulii and its relatives. An unusual col-

lection, Khan et al. 1983, with stiff many- veined

leaves, hollow stems, and anthers with small distal

pores, may key out here but is presently placed

with L. multinervis.

Licaria cufodontisii Kosterm., Recueil Trav. Bot.

Neerl. 34: 602. 1937. Figure 16.

Shrubs or trees (2-)5-20 m tall, leafy branchlets 0.7-

5 mm thick, essentially glabrous, grayish and terete,

sometimes with small (0.5 mm) grayish lenticels. Leaves

alternate, petioles 5-12 mm long, 0.8-1.6 mm thick,

usually sulcate above with 2 adaxial ridges, glabrous,

usually drying dark; leaf blades (8-) 10- 18 cm long, 3-7

cm broad, elliptic-oblong to oblong or ovate-oblong, ta-

pering abruptly to the acuminate or caudate-acuminate

apex with a narrow tip 1-3 cm long, acute to obtuse at

the base, margin entire or slightly undulate, drying stiffly

chartaceous and usually dull grayish green or grayish

brown, glabrous above and below, midvein prominent

above, with 4-7 major secondary veins on each side.

Inflorescences solitary and axillary or pseudoterminal,

to 8 cm long, paniculate with few branches and few

flowers, peduncle 1-3 cm long, 0.5-1 mm thick, glabrous

or sparsely puberulent with minute (0.1-0.2 mm) ap-

pressed hairs, flowers usually in umbellate or cymose

clusters at the ends of slender hairs, flowers usually in

umbellate or cymose clusters at the ends of slender lateral

branches, pedicels 2-5 mm long. Flowers 1.5-3 mm long,

2-2.8 mm broad, yellowish and glabrous on the outside,

the outer tepals to 2 mm broad, imbricate, inner tepals

smaller; stamens connivent, 0.7-1 .3 mm long, 0.6-1 mm

broad, oblong-cylindrical (not narrowed at the base) and

thick, with short ascending hairs, the small (0.2 mm)

thecae on the upper distal surface of the stamen, broad

rounded glands present at the exterior (abaxial) base of

the stamens, staminodes usually absent; pistil slender,

1-1.6 mm long, ovary ca. 0.7 mm long and 0.5 mm

thick, stigma simple. Fruits borne in a cupule 10-15 mm

long, 15-28 mm broad and 6-10 mm deep, conical to

broadly cupulate, with or without lenticels on the surface,

outer margin undulate, inner margin entire, becoming

bright red; berry ellipsoid, 2-3 cm long, 1-2.2 cm thick,

black or black with a bluish tint.

Trees of evergreen forest formations around

Golfo Dulce and in the General Valley between

sea level and 900 m elevation. Flowering material

has been collected in April, June, September, and

December, while fruiting material has been col-

lected in February-March, September, and De-

cember. This taxon is endemic to the Pacific Slope,

from the Reserva Biologica Carara southward to

western Panama. However, it has also been inter-

preted as a subspecific element of L. misantlae

(Brandegee) Kostermans, and that species is said

to range from central and southern Mexico to Co-

lombia (see below).

Licaria cufodontisii is recognized by the very

small flowers with thick connivent stamens and

apical dehiscence, fruiting cups with double mar-

gins, and often oblong leaves with caudate-acu-

minate apices. Kurz (1983) placed this species un-

der L. misantlae and discussed the distinctive traits

of L. cufodontisii, and several other elements with-

in L. misantlae. There is no doubt, based on sta-

men morphology, that these species are closely

related, but we prefer to keep them separate. In

addition to a disjunct geographical distribution,

Costa Rican material has much larger leaves with

48

FIELDIANA: BOTANY

duller surfaces that tend to dry grayish. Leaves

and fruits of this species may be confused with

Ocotea veraguensis.

A collection received after this text (above) had

been completed (Haberet al. 4396, CR, F, MO; from

the Cordillera de Tilaran) looks very much like

typical material of Licaria misantlae (sensu stric-

to), and indicates that this species is a part of the

Costa Rican flora. The leaves are small (ca. 6 cm

long and 2 cm broad) and the flowers fit Kurz's

description for L. misantlae. In addition, recent

collections from the Carara reserve in central Pun-

tarenas Province (Grayum et al. 7605, CR, F, MO;

Hammel et al. 1428, CR, F, MO) might be inter-

preted to be intermediate between L. misantlae

and L. cufodontisii. A similar interpretation can

be made of an earlier Brenes collection (12262, F,

GH) from the Osa Peninsula. Thus it is quite prob-

able that Kurz's interpretation is correct and L.

cufodontisii must be synonymized under L. mis-

antlae. In this event, the thicker larger leaves and

restricted range of L. cufodontisii would support

its recognition as a subspecies of L. misantlae.

Licaria excelsa Kosterm., Recueil Trav. Bot. Neerl.

34: 595. 1937. Acrodiclidium excelsum (Kos-

term.) Lundell, Amer. Midi. Naturalist 19: 428.

1938. Misanteca excelsa (Kosterm.) Lundell,

Wrightia 1: 147. 1946. Licaria alata Miranda,

Ceiba4: 128. 1954. Figure 16.

Trees 5-30 m tall, leafy branchlets 3-6 mm thick,

usually glabrous or minutely appressed, puberulent in

early stages, with numerous grayish lenticels 0.5-2 mm

long. Leaves alternate, petioles 8-24 mm long, 2-3.5 mm

thick, glabrous or minutely appressed puberulent, often

deeply sulcate above, drying dark in color; leaf blades

7-20(-28) cm long, 3-7(-9) cm broad (Kurz, 1983, re-

ports leaves to 40 cm long and 1 5 cm broad on petioles

to 35 mm long), narrowly oblong to oblong-lanceolate,

narrowly elliptic-oblong or elliptic-lanceolate, tapering

gradually to a bluntly acute or short-acuminate apex,

obtuse at the base, margin entire and slightly revolute

(especially at the base), drying subcoriaceous to coria-

ceous, glabrous and slightly lustrous on both surfaces

(sparsely puberulent beneath in early stages), midvein

broad (3 mm) and slightly impressed above, with 5-10

major secondary veins on each side. Inflorescences ax-

illary from distal leaves or apparently several from axils

of caducous bracts, (5-)10-18(-22) cm long, primary pe-

duncle 2-3 mm thick, distal parts with slender grayish

or yellowish-brown hairs ca. 0.2 mm long, pedicels 1.5-

4 mm long, densely puberulent. Flowers 2.5-3.5 mm

long, 1.5-2.5 mm broad, obovoid, often puberulent be-

neath but glabrous distally, outer perianth parts broadly

imbricate (almost valvate), 0.8 mm long and 1.5 mm

broad, inner perianth parts 1 mm broad; stamens united

to form a thick column 0.7-1.2 mm high, each stamen

ca. 1 mm broad and with an equally broad thick filament,

the small circular valves dehiscing upwards and outward

(at an angle of 45° abaxially), glands minute and free (6)

or united (3), staminodes absent, floral tube slightly pu-

berulent within; pistil ca. 1.8 mm long, ovary 0.6-1 mm

thick, style ca. 0.5 mm long, stigma simple. Fruits borne

in a cupulate or hemispheric receptacle, 1.3-2 cm long,

1.8-2.5 cm broad at the apex and 10-16 mm deep, the

rim entire with inner and outer margins 1-2 mm distant

and weakly developed, pinkish but drying dark with many

conspicuous pale lenticels 0. 5-2.5 mm long, fruiting ped-

icels to 1 5 mm long, 5 mm thick; berry ovoid-ellipsoid,

to 3.5 cm long and 2.3 cm in diameter.

Trees of evergreen forests of the Pacific slope of

southern Costa Rica and the adjacent Chiriqui

highlands, from (600) 1 100 to 2300 m elevation.

Flowers have been collected in June, and fruits

have been collected in January, March, and May

in Costa Rica and Panama. The species ranges

from southern Mexico to Panama.

Licaria excelsa is recognized by the very stiff

narrow leaves with dark deeply sulcate petioles,

the small flowers with a staminal column in which

the small circular thecae open at an angle upwards

and outwards, and the large fruiting cups with

poorly defined double margin. The laminae are

usually more than three times longer than broad.

While the flowers appear to be distinctive, vari-

ability in leaf form and fruit development may

make it very difficult to distinguish this species

from L. multinervis. This species is poorly known;

we have only five collections from Costa Rica. The

following collections from the Caribbean slope with

stiff narrowly oblong leaves and hollow stems with

longitudinal ridges are placed here provisionally:

Hammel & Grayum 14111 (CR, MO), Poveda et al.

3637 (CR).

Licaria multinervis H. Kurz, Mitt. Inst. Allg. Bot.

Hamburg 23: in press. Figure 16.

Trees to 30 m tall, with trunks to 1 m d.b.h., leafy

branchlets 1.3-6 mm thick, minutely (0.2 mm) puber-

ulent with thin brownish or yellowish hairs, becoming

glabrous and terete. Leaves alternate, petioles 4-12(-20)

mm long, 1.2-2.5 mm thick, puberulent or glabrous,

often drying dark; leaf blades 7-14(-22) cm long, 2-

4(-5.5) cm broad, very narrowly oblong to narrowly el-

liptic-oblong or lanceolate, gradually tapering to an acute

or short-acuminate apex, acute at the base, margin entire

and often somewhat revolute, drying subcoriaceous, and

lustrous above, glabrous above, lower surface with a cov-

ering of thin straight appressed ascending hairs 0.1-0.3

mm long and often difficult to see, midvein slightly im-

pressed above, with (9-) 1 2-20 (often obscure) secondary

veins on each side, tertiary venation obscure above and

BURGER: FLORA COSTARICENSIS

49

below. Inflorescences terminal or axillary or from leafless

distal nodes, usually on the basal parts of new shoots,

paniculate, ca. 5 cm long and with 50-100 flowers, pe-

duncle ca. 1 mm thick and with many yellowish brown

hairs, pedicels 1-2 mm long. Flowers ca. 2 mm long and

1 .3 mm broad, obovoid to funnelform, the outer 3 tepals

broadly ovate, ca. 0.8 mm long and 0.6 mm broad, pu-

berulent, inner tepals much narrower, (0.4 mm) and thin-

ner, erect at anthesis, floral tube sericeous within; sta-

mens free or connate only at the short (0.4 mm) broad

filaments, slender hairs present at the base, each anther

ca. 0.6 mm long and 0.5 mm broad, ovoid with 2 large

vertical thecae and extrorse dehiscence, slightly exserted,

glands 2-6 but minute, staminodes absent; pistil ca. 1.4

mm long, ovary ca. 0.6 mm long and 0.3 mm thick, style

thick, stigma slightly discoid. Fruits borne in a cupulate

or hemispheric receptacle 10-14 mm long, 16-25 mm

broad and 8 mm deep, abruptly expanded above the

short (6 mm) thick (5 mm) pedicel, outer surface with

pale lenticel-like spots 0.3-1 mm broad, rim of the cup

3 mm thick and with 2 margins but the margins not

expanded and difficult to see when dry, cup minutely

puberulent within; berry ovoid, ca. 1 5 mm long and 1 2

mm thick.

Trees of evergreen wet forest formations on the

Caribbean slope and in the General Valley, be-

tween 500 and 1000 m elevation. Immature in-

florescences were collected in September, flowers

have been collected in May (Leon 1104, the des-

ignated type), and fruits were collected in February

and October. The species is only known from four

collections in central Costa Rica, near Tuis in the

province of Cartago and near Volcan in southern-

most San Jose Province.

Licaria multinervis is distinguished by its long

narrow stiff leaves with dense but minute puber-

ulence on the undersurfaces, and the thick cupu-

late fruiting receptacles with two weakly denned

margins. Vegetatively, this species is very similar

to L. excelsa, but that species has glabrous leaves

and very different stamens. The almost free sta-

mens with large longitudinally dehiscing extrorse

anthers are more like those of L. triandra and may

be slightly exserted at anthesis. This species is an

important timber tree often called quina. We be-

lieve this species is a good one, but new collections

from the Caribbean slope in Costa Rica have com-

pounded the difficulties in denning the Costa Ri-

can species of Licaria, and we suspect there are

other new species. Variability of leaves and fruit-

ing material within each species makes it very dif-

ficult to recognize and separate different species in

the absence of flowers. The proper circumscription

of all these species requires more and better col-

lections.

Licaria pergamentacea W. Burger, sp. nov. Figure

16.

Arbor ca. 20 m alta, ramulis foliiferis 2-3 mm crassis,

puberulis. Folia alterna, petiolis 8-14 mm longis, ca. 2

mm crassis, dense puberulis; laminis 24-26 cm longis,

5-6 cm latis, anguste oblongis, apice acuminate, papyr-

aceis, supra glabris, infra costa minute puberula, nervis

secondariis 8-10 paribus. Inflorescentiae paniculatae, 8-

14 cm longae, pedunculis puberulis. Flores ca. 2 mm

longi et 1 .6 mm lati, extus puberuli, stamina 3, filamentis

ca. 0.2 mm longis, antheris ca. 0.4 mm longis et 0.6 mm

latis, staminodiis complanatis; gynoecium ca. 1.1 mm

longum, ovario angusto. Fructus absens in typo.

Trees 12-20 m tall, leafy branchlets 2-3 mm thick,

densely puberulent with brownish hairs 0. 1-0.3 mm long,

twigs solid. Leaves alternate, petioles 8-14 mm long, 2-

3 mm thick, densely puberulent; leaf blades 22-26 cm

long, 5-7.5 cm broad, narrowly oblong or narrowly el-

liptic-oblong, broadest at or near the middle, acuminate

at the apex, acute to obtuse and slightly rounded at the

base, drying chartaceous and brown, dull and glabrous

above, tertiary venation slightly raised above (dry), gla-

brous beneath but with minute appressed hairs along the

midvein, with 8-14 major secondary veins beneath, cen-

tral secondaries arising at angles of 35°-50°, tertiary ve-

nation becoming raised beneath and forming a reticulum

with areolae ca. 1 mm broad but not well defined. In-

florescences 8-14 cm long (but perhaps not fully mature),

paniculate, peduncles ca. 3 cm long and 1 mm thick,

densely and minutely puberulent with brownish hairs,

lateral branches 2-3 cm long, pedicels 0.5-1 mm long.

Flowers small and yellow, ca. 2 mm long and 1.6 mm

broad, sparsely and minutely puberulent on the outside,

inner tepals slightly smaller than the outer; functional

stamens 3, loosely connivent with short (0.2 mm) broad

filaments and broadly rounded anthers 0.4-0.8 mm long

and 0.6 mm broad, valves opening apically or slightly

extrorse, small flattened or glandlike structures present

on the exterior (abaxial) side of the stamens; pistil ca.

1.1 mm long, ovary slender, style slender and 0.5 mm

long, stigma simple. Fruits borne in a deep hemispheric

double-margined cup 15-18 mm long and 16-20 mm

broad; fruits only slightly longer than the cup and with

broadly rounded apex (but not fully mature).

TYPE— Costa Rica. Puntarenas Province, hills

above Palmar Norte de Osa, 22 Feb. 1951, Paul

H. Allen 5950 (holotype, F, 1439749; isotype, EAP).

Trees of lowland rain forest formations of the

Golfo Dulce area and Osa Peninsula in south-

western Costa Rica, from near sea level to 500 m

elevation. Flowers were collected in February (Al-

len 5950; Burger et al. 12366, CR, F, MO, NY), and

an immature fruiting receptacle was collected in

May (Gomez- Laurito & Bermudez 2711, usj). The

species is known only from the three collections

cited above.

50

FIELDIANA: BOTANY

Licaria pergamentacea is recognized by the thin

narrowly oblong leaves on short puberulent peti-

oles, the small flower with only three functional

stamens dehiscing apically by small pores, and

deep fruiting cup with double margin. The flow-

ering and fruiting collections match each other very

well, except that they differ somewhat in the num-

ber of secondary veins. The fruiting collection was

made near Quebrada Zavala, Sierpe, near the Osa

Peninsula. The thinner, pergamentaceous leaves

are unusual in Licaria. This species was referred

to as Ocotea pergamentacea Standl. & L. O. Wms.

(an unpublished name) by Paul Allen (1956, p.

276).

Licaria sarapiquensis Hammel, J. Arnold Arbor.

67: 124. 1986. Figure 16.

Slender trees 5-20 m tall, trunks 5-20(-30) cm d.b.h.,

leafy branchlets 1.3-3 mm thick, glabrous, dark grayish,

terete and smooth. Leaves alternate, petioles 4-12 mm

long, 1-2 mm thick, usually sulcate above, glabrous and

drying very dark in color; leaf blades 1 0-20 cm long, 3-

7 cm broad, elliptic-oblong to oblong, narrowly elliptic

or rarely lanceolate, tapering gradually or abruptly to a

long (8-30 mm) acuminate apex, acute to obtuse at the

base, margin entire and slightly revolute (dry), drying

stiffly chartaceous and lustrous above (dark green in life),

glabrous on both surfaces, with 3-5(-6) major secondary

veins on each side, loop-connected near the margin,

smaller veins forming a slightly raised reticulum on the

lower surface (dry) with areolae 1-3 mm broad. Inflo-

rescences axillary or extra-axillary, solitary and very

slender, paniculate or racemose, 2.5-5 cm long, few-

flowered, the flowers on lateral branches solitary, few or

umbellate, primary peduncle ca. 0.5 mm thick (dry),

glabrous, secondary peduncles to 2 cm long, pedicels 1-

6 mm long. Flowers 1 .5-2.5 mm long, ca. 1 .5 mm broad,

yellowish green and inconspicuous, campanulate, gla-

brous, perianth lobes ca. 1 .2 mm long, the outer broadly

imbricate; stamens 0.8-1.2 mm long, with filaments be-

coming 1 mm long and short (0.4 mm) oblate anthers

0.6-0.7 mm broad, the 2 thecae opening upward and

outward (abaxially), each stamen with 2 stipitate glands

and the stipes equalling the glands in length, the 6 outer

stamens (series I & II) represented by ligulate stamino-

dia, inner staminodia (series IV) absent; pistil 1.5-1.8

mm long, ovary equalling the length of the style, stigma

slightly discoid. Fruits borne on a conical receptacle 6-

12 mm long, 8-16 mm broad at the top and ca. 6 mm

deep, margin entire but with a clearly differentiated in-

ternal ridge, fruiting pedicels 8-16 mm long and 1-2 mm

thick, cups red at maturity; berry narrowly ovoid or

ovoid-ellipsoid, 15-20 mm long and 7-10 mm in di-

ameter, becoming bluish black.

Small trees of the very wet forests of the Carib-

bean slope and lowlands from near sea level to

about 800 m elevation, often found on ridges and

steep slopes within the forest. Flowers have been

collected in April-May, while fruits have been ob-

served in August-November and collected in No-

vember, February, and March. This species is en-

demic to Costa Rica and only known from the

following sites: La Selva (Hammel 8663, DUKE,

the type; 10532, 12235, DUKE, F; Hartshorn 1588,

F), slopes below Volcan Barva (Hammel 6923),

the Rio Reventazon below Cairo (Standley & Val-

eria 48784, us), and near Turrialba (Poveda et al.

3489, CR, F) in the provinces of Heredia and Car-

tago. Provisionally placed here is Gomez et al.

21 143 (CR, MO) with unusual much-branched in-

florescence and long-pedicellate (10 mm) flowers.

Licaria sarapiquensis is recognized by the lus-

trous leaves much darker above than below (in

life), the lack of pubescence (except at the shoot

apex), the slender little inflorescences, the incon-

spicuous flowers with only three stamens, and the

conical fruit cup with double-margined rim.

Crushed leaves and bark smell like Sassafras or

sarsaparilla. Quizarrd torita has been reported as

a common name. Dried material of this species

resembles Ocotea cernua, O. floribunda, and O.

veraguensis, and sterile material may be difficult

to separate. Kurz (1983) cited a collection of this

species (Hartshorn 5249, us) as Licaria guate-

malensis Mez, but that is a species of northern

Central America.

Licaria triandra (Sw.) Kosterm., Recueil Trav. Bot.

Neerl. 34: 588. 1937. Laurus triandra Sw., Prodr.

65. 1788. Misanteca triandra (Sw.) Mez, Jahrb.

Konigl. Bot. Gart. Berlin 5: 103. 1889. Acrod-

iclidium triandrum (Sw.) Lundell, Contr. Univ.

Michigan Herb. 7: 12. 1942. Misanteca pittieri

Mez, Bull. Herb. Boissier, ser. 2, 3: 230. 1903.

L. pittieri (Mez) C. K. Allen, J. Arnold Arbor.

26: 427. 1945. Misanteca costaricensis I. M.

Johnston, Contr. Gray Herb. 70: 70. 1924. Fig-

ure 16.

Trees 10-15(-20) m tall, leafy branchlets

mm thick, essentially glabrous and drying dark brown

or grayish. Leaves alternate, petioles 4-1 5(-20) mm long,

1-2 mm thick, glabrous, flat or slightly sulcate above

with 2 adaxial or lateral margins; leaf blades 5-16 cm

long, 2-5(-7.5) cm broad, elliptic to elliptic-oblong or

narrowly ovate, tapering gradually to abruptly to a short

acuminate apex, the tip ca. 1 cm long, obtuse to acute

at the base, margins entire or undulate, drying subcor-

iaceous and slightly lustrous above, often grayish brown,

BURGER: FLORA COSTARICENSIS

51

glabrous above and below (minutely puberulent beneath

in early stages), midvein slightly elevated above, with

3-8 major secondary veins on each side. Inflorescences

solitary and axillary to distal leaves or terminal, pani-

culate, small (6 cm), and few flowers (ca. 20) to racemose-

paniculate with a long (16 cm) central rachis and alter-

nating lateral panicles with up to ca. 200 flowers, mi-

nutely puberulent, pedicels 1-4 mm long and glabrous

or minutely puberulent. Flowers 1.8-2.8 mm long, ca.

1.5 mm broad with the perianth parts erect at anthesis,

yellowish, tepals glabrous (rarely puberulent) on the out-

side; stamens united in the lower half to form a short

column surrounding the slender pistil, androecium 1.1-

1.7 mm long and 1 mm in diameter, thecae free distally

and becoming elevated above the perianth, dehiscing on

the exterior (abaxial) side of the column, valves ca. 0.5

mm long, broad glands present at the abaxial base of the

staminal column, the glands free and 2 per stamen or

united and 1 per stamen, staminodes absent, hairs often

present at the narrowed base of the stamens; pistil 1.5

mm long, ovary narrowly ellipsoid, style ca. 0.7 mm

long, stigma simple. Fruits borne in a cupulate or hem-

ispheric receptacle, (6-)8-12(-15) mm long, 1 1-18(-25)

mm broad, 5-9 mm deep, outer margin often conspic-

uous and entire or undulate, 1-3 mm broad, inner (distal)

margin entire and 1-2 mm high, becoming red and often

with small (1 mm) conspicuous lenticels on the outer

surface; berry 1.5-2.8 cm long, 10-18 mm thick, ovoid

to ovoid-ellipsoid, the lower '/3 or '/2 immersed within

the cup, becoming dark purplish or black.

Trees of evergreen wet forest formations from

near sea level to 1400 m elevation on the Carib-

bean side of Costa Rica and in the central high-

lands. Flowering material has been collected in

April-May in Costa Rica, and fruits in January,

April-July, and October. This species ranges from

Florida (U.S.A.) and the West Indies through

Mexico and Central America to Bolivia.

Licaria triandra is recognized by its stiff, essen-

tially glabrous, and usually elliptic-oblong leaves,

the very small flowers with three stamens united

to form a tubular column around the slender style,

the extrorse and exserted thecae, and the conspic-

uously double-rimmed fruiting cup. Foliage of this

species varies greatly and can make identification

difficult. Hammel's lowland collections from La

Selva have laminae averaging 8 cm long and 3 cm

broad, while the highland collections are consid-

erably larger. In addition, the fruiting receptacles

of the highland material is consistently larger than

in lowland and West Indian collections. The high-

land collections are well characterized by the de-

scription and type of L. pittieri and, at first, it

seemed best to recognize the highland material as

a distinct species. However, Kurz (1982) inter-

preted L. triandra quite broadly, and the same

pattern of larger-lea ved highland collections, con-

trasting with smaller-leaved lowland collections,

is also found in Guatemala. Thus, we follow Kurz

in submerging L. pittieri, though it may be worthy

of subspecific rank. Kurz also included the follow-

ing species as synonyms of L. triandra: L. cervan-

tesii (H.B.K.) Kosterm., L. limbosa (Ruiz & Pa-

von) Kosterm., L. redinata Lundell, and L.

tikalana (Lundell) Lundell. It seems that L. cori-

acea (Lundell) Kosterm. also belongs here, and not

under L. misantlae as suggested by Kurz.

Licaria sp. A.

A slender shrub 3 m tall, stem 3 cm d.b.h., leafy

branchlets 0.6-2.5 mm thick, glabrous and grayish. Leaves

alternate, petioles 5-1 5 mm, 0.6-1.3 mm thick, glabrous,

sulcate above; leaf blades (4-)7-14 cm long, (1.3-)2-4

cm broad, narrowly elliptic to narrowly elliptic-oblong,

tapering gradually to the acuminate apex, the narrow tip

0.6-2 cm long, acute at the base, margin undulate, drying

chartaceous and grayish green, glabrous above and be-

low, midvein slightly elevated above, with 5-9 major

secondary veins, tertiary venation slightly elevated be-

neath. Inflorescences solitary and axillary, small (2 cm)

and few-flowered, racemose but with a distal umbel of

flowers, peduncle only 0.3 mm thick (dry), glabrous, ped-

icels to 3 mm long. Flowers apparently unisexual and

the plants dioecious, female flowers not seen; male flow-

ers ca. 2 mm long and 1.5 mm broad, obovoid and

glabrous, perianth parts ca. 0.8 mm long; fertile stamens

connivent, 0.7-1 mm tall, with a short thick filament

and short (0.5 mm) broad (0.5 mm) anther, thecae apical

and the valves with slightly introrse dehiscence, large

glands and hairs present at the base of the stamens, stam-

inodes not apparent; pistillode very slender, ca. 0.5 mm

long. Fruits (based on Zamora 399) borne in a red cup

1 2 mm long and 20 mm broad with a prominent ridge

around the distal margin; fruits ca. 20 mm long and 12

mm in diameter.

A small shrub in gallery forest at the edge of the

Rio Puerto Viejo near its confluence with the Rio

Sarapiqui at about 100 m elevation (Hammel

10670, DUKE, F). It was collected with flowers in

early December, at the La Selva research site in

the Caribbean lowland rain forest formation. The

very slender pistil and the apical thecae that open

inwardly (adaxially) are unusual in the genus. The

small glabrous narrow and often long-acuminate

leaves with undulate margins and lack of a strong

aroma when crushed are further distinctions. It is

possible that larger individuals of this species will

prove to have bisexual flowers. (This is the first

report of unisexual flowers in Licaria, and addi-

tional collections to verify this condition are great-

ly desired.) A second collection tentatively placed

here (Zamora 399, CR) is a 10m tree flowering in

late November at Horquetas-Sarapiqui, at 400 m

elevation.

52

FIELDIANA: BOTANY

Litsea Lamarck

Shrubs or small trees (in Central America), dioecious.

Leaves alternate or rarely subopposite, laminae small

and of 2 types in American species, elliptic and up to 14

cm long or ovate and orbicular to 8 cm long, usually

coriaceous, pinnately veined (in ours) or tripliveined.

Inflorescences solitary or fasciculate, in leaf axils or on

short leafless shoots in a racemose arrangement, sessile

or pedunculate, umbellate or capitate, at first enclosed

in an involucre of 4-6 broadly imbricate bracts. Flowers

pedicellate, unisexual, the floral tube short or absent, the

perianth 6-parted (with 2 whorls of 3) or irregular, peri-

anth parts (tepals) equal or subequal; male flowers usu-

ally with 9 or 1 2 stamens in 3 or 4 series (whorls), fila-

ments well differentiated, anthers 4-thecous and introrse

(the inner series sometimes with lateral dehiscence), the

innermost whorl of stamens usually with stipitate glands

borne on the base of the slender filaments, a pistillode

present or absent; female flowers with 9 or 1 2 staminodes

in 3 or 4 series, the outer 2 series usually lacking glands

and the inner series usually with stipitate glands, pistil

with rounded ovary, stigma often discoid and lobed or

subcapitate. Fruits borne on a slightly thickened pedicel

or subtended by a small receptacular disc or cupule; berry

usually globose and fleshy.

The genus is reported to contain around 400

species, and ranges from southern and eastern Asia

(as far north as Korea and Japan) through Malay-

sia to Australia and New Zealand; in the New

World it ranges from the southeastern United States

and Mexico to Costa Rica. Only one species has

been reported from Costa Rica and that species

has not been collected there for over 50 years. The

small trees with stiff aromatic little leaves, the glo-

bose inflorescence buds with their imbricate in-

volucre and borne on a short peduncle, and the

functionally unisexual flowers with 9 or 12 sta-

mens/staminodes make these plants easy to rec-

ognize among our species of Lauraceae.

Litsea glaucescens H.B.K., Nov. gen. sp. 2: 133.

1817. Tetranthera glaucescens Spreng., Syst. veg.

2: 267. 1825. L. neesiana Hemsl., Biol. centr.

amer., Bot. 3: 76. 1882. L. guatemalensis Mez,

Jahrb. Konigl. Bot. Gart. Berlin 5: 479. 1889.

L. flavescens Bartlett, Proc. Amer. Acad. Arts

44: 599. 1909. L. acuminatissima Lundell,

Contr. Univ. Michigan Herb. 4: 3. 1940. L.

glaucescens \^r. flavescens (Bartlett) C. K. Allen,

J. Arnold. Arbor. 26: 413. 1945. Figure 4.

Shrubs or small trees 1.5-6 m tall, dioecious (unisex-

ual), leafy branchlets 1.3-3 mm thick, glabrous or with

slender grayish ascending hairs, reddish brown to black,

often with small dark lenticels, terete. Leaves alternate

and well spaced along the stems, petioles (3-)5-16(-20)

mm long, ca. 1 mm thick, glabrous or puberulent, slightly

sulcate above; leaf blades 3-8(-12) cm long, l-3(-3.8)

cm broad, narrowly elliptic to lanceolate or ovate-ellip-

tic, tapering gradually to the acute or acuminate apex,

obtuse or acute at the base, margin entire and often

defined with a veinlike edge, drying very stiffly charta-

ceous to subcoriaceous, pale grayish to dark brown above,

glabrous and slightly lustrous above with the midvein

flat or slightly impressed, paler in color or glaucous be-

neath and glabrous or sparsely puberulent, with 4-8 ma-

jor secondary veins on each side, the minor venation flat

beneath but sometimes forming a slightly areolate sur-

face on the glaucous underside. Inflorescences solitary

in distal leaf axils or several on leafless short-shoots (and

racemosely arranged), to 2.5 cm long, umbellate, pe-

duncles 6-12(-15) mm long, ca. 0.5 mm thick, glabrous

or puberulent, broadly imbricate bracts borne at the apex

of the peduncle and forming an ovoid-globose bud 3-6

mm in diameter which encloses the immature umbel,

bracts caducous, reddish brown, each umbel with 3-6

flowers, pedicels l-3(-5) mm long. Male flowers ca. 4

mm long and 6 mm broad, with 6 narrow tepals ca. 3

mm long, outer stamens with anthers 1.3-1.7 mm long

and ca. 0.6 mm broad, with slender filaments ca. 1.5

mm long; pistillode ca. 1 .4 mm long, with ovoid ovary

and slender style. Female flowers ca. 4 mm long and 4

mm broad, perianth with 6 or 4 tepals, staminodes usu-

ally 9 and ca. 1.2 mm long, flat; pistil ca. 2.3 mm long,

ovary subglobose and 1.3 mm in diameter, style often

crooked, stigma discoid and dark in color. Fruits subgl-

obose, narrowed at the apex and base, 8-12 mm in di-

ameter, fleshy; fruiting peduncles 3-5(-7) mm long and

1-2 mm thick, receptacle slightly expanded (2-4 mm

broad) below the fruits and flat or saucer-like.

Small trees of evergreen but seasonally dry mon-

tane forest formations, between 1 500 and 2000 m

elevation on the Pacific slope of the western por-

tion of the Cordillera de Talamanca (Candelaria,

Sta. Maria de Dota, and Tarrazu) in Costa Rica.

This species reaches 3000 m elevation in Guate-

mala, and flowers in November-April in Central

America. Mature fruits have been collected in Au-

gust in Costa Rica. This species ranges from east-

ern Mexico through Guatemala and Honduras to

Costa Rica.

Litsea glaucescens is recognized by its small stat-

ure, small stiff usually glabrous leaves, umbellate

inflorescences at first enclosed in broadly imbri-

cate bracts at the apex of a short peduncle, and

the unisexual flowers with both male and female

parts on dioecious trees. The leaves have been

used like bay (Laurus) leaves for flavoring food,

especially soup and meat. It has been called len-

tisco in Costa Rica.

The Costa Rican specimens are distinguished

from the more northern collections by their prom-

inent marginal vein, yellowish venation, and the

nonglaucous lower leaf surfaces. Caroline Allen

(1945) recognized the Costa Rican material as va-

BURGER: FLORA COSTARICENSIS

53

riety flavescens. The collections seen from Costa

Rica are: Oersted Lauraceae no. 10 (us), Standley

42525 (F, us), Tonduz 7796 (us) and 11638, also

distributed as J. D. Smith's number 7352 (CR, F,

GH, NY, us). We have seen no collections more

recent than that made by Paul Standley in 1925,

and it may be that the endemic Costa Rican va-

riety of this species is now extinct.

Nectandra Rolander ex Rottboel

REFERENCE—!. G. Rohwer, Prodromus einer

Monographic der Gattung Ocotea Aubl. (Laura-

ceae), sensu lato. Mitt. Inst. Allg. Bot. Hamburg

20: 1-278. 1986.

Trees or shrubs, monoecious or rarely dioecious. Leaves

alternate or rarely subopposite or opposite, petiolate, the

laminae entire and pinnately veined, chartaceous to sub-

coriaceous, puberulent to glabrous, with or without dom-

atia. Inflorescences usually solitary in axils of distal leaves,

rarely pseudoterminal, usually paniculate, pedicels pres-

ent. Flowers bisexual or rarely functionally unisexual,

generally small (<1 cm broad), white to yellowish or

greenish, perianth tube well developed or absent but

accrescent in fruits, perianth 6-parted in 2 whorls of 3

tepals each, the whorls equal or subequal, usually re-

flexed or rotate (spreading) at anthesis, tepals usually

thick and often papillate-puberulent on the inner surface;

fertile stamens 9 in 3 series, the 6 stamens of the outer

whorls (often appearing as a single whorl) similar in size

and shape and with introrse dehiscence, filaments short

or absent (very rarely longer than the anthers), the an-

thers usually thick and papillate, reniform to ovate or

laminate, the connective sometimes developed beyond

the thecae or the anthers sometimes emarginate, thecae

in a horizontal row or in an arc with the lower thecae

lateral to the upper, opening by flaps attached at the top,

the 3 stamens of the inner series (series III) usually with

filaments and quadrate or rectangular anthers and each

with 2 glands at the base, the upper 2 thecae usually

dehiscing laterally or lateral-extrorse and the 2 lower

usually extrorse, the inner whorl of staminodes (series

IV) small and stipelike or absent; pistil with subglobose

to ovoid or ellipsoid ovary, glabrous (puberulent in N.

reticulata s.l.), style usually shorter than the ovary, stig-

ma usually discoid to peltate or capitate. Fruits borne

in a cupulate receptacle with simple margin, perianth

parts deciduous (rarely persisting on the margin), pedicel

accrescent in fruits; berry a single-seeded berry, ellipsoid,

subglobose, to oblong or obovoid.

A New World genus of 100-1 50 species ranging

from Florida (U.S.A.) to Argentina and well rep-

resented in Mesoamerica and the West Indies, but

with the great majority of species in South Amer-

ica. This genus has been submerged under Ocotea

by Kostermans (1957) and Howard (1981). Such

a taxonomic change creates many nomenclatural

problems and clarifies little. Until we understand

all the genera of Lauraceae much better, it seems

best to continue using Nectandra in its traditional

sense. Species in Costa Rica that have staminal

characteristics intermediate between Nectandra

and Ocotea are generally placed under Ocotea in

this treatment. There may also be a Costa Rican

species of Nectandra related to a few species of

Ocotea formerly placed in Phoebe; see the discus-

sion under Nectandra belizensis.

Some of the most difficult problems of species

delimitation in the Costa Rican Lauraceae are

found in Nectandra. Local differentiation within

wide-ranging species or species groups and overall

similarity of different lineages make species cir-

cumscription very tentative or even arbitrary in

some groups of Nectandra. The following account

attempts to recognize locally distinct populations

with as little nomenclatural change as possible.

Some of our "local species" are undoubtedly sub-

species of more widely ranging species; but the

circumscription of these widely ranging entities

requires much more study and better sampling

over their entire range before we can erect mean-

ingful subspecies. Jens Rohwer is currently study-

ing this genus, and many names and specific con-

cepts are likely to be changed as a result of this

new work.

Key to Species of Nectandra

la. Outer stamens with the connective conspicuously to slightly expanded beyond the thecae, the

connective minutely papillate or minutely puberulent, the stamens often flat and tepal-like . . 2a

1 b. Outer stamens with the connective not conspicuously prolonged distally 8a

2a. Outer stamens with anthers more than 1.5 mm long, flat and tepal-like, sessile or with a

narrow base; leaves and flowers densely puberulent 3a

2b. Outer stamens with anthers less than 1.5 mm long, not tepal-like 4a

3a. Outer stamens 2-3 mm long, thecae often superposed as in Ocotea, flowers 10-20 mm

broad, peduncle much exceeding the length of the flowering rachis; growing in semi-

deciduous forests . . N. sinuata

54

FIELDIANA: BOTANY

3b. Outer stamens 1.5-1.8 mm long, thecae in an arc, flowers 8-12 mm broad, peduncles

usually equal in length to the flowering rachis of the panicle; growing in semi-deciduous

and evergreen forests N. reticulata

4a. Outer stamens with the connective prolonged conspicuously beyond the thecae, the anther

usually curved or obtuse at the apex 5a

4b. Outer stamens with the connective slightly prolonged beyond the thecae, apex of the anther

usually flat or undulate; flowers 4-8 mm broad 7a

5a. Leaves often drying pale grayish or yellowish green, with (4-)7-l 1 pairs of major

secondary veins, leaf base often slightly revolute; flowers 6-12 mm broad; in deciduous

and partly deciduous forests N. globosa

5b. Leaves drying very pale brown to dark brown, with 3-7 pairs of major secondary veins

6a

6a. Anthers 1-1.6 mm long, flowers to 12(-15) mm broad; leaves to 15 cm long, ovate-

elliptic to elliptic-oblong, usually drying very pale brown; Pacific slope, 600-2400 m

elevation TV. ramonensis

6b. Anthers 0.7-0.8 mm long, flowers ca. 8 mm broad; leaves to 20 cm long, usually

narrowly elliptic and drying dark brown; evergreen forests, 0-1400 m

N. globosa (in part)

7a. Leaves essentially glabrous and drying grayish, ovate to oblong or lanceolate; anthers ca. 0.7

mm long; evergreen and semi-deciduous forests N. turbacensis

7b. Leaves puberulent beneath and drying brownish, obovate to elliptic-oblong; anthers ca. 0.9

mm long; evergreen Caribbean lowlands N. belizensis

8a. Leaves conspicuously puberulent beneath with hairs more than 0.3 mm long, lower surface soft

to the touch 9a

8b. Leaves glabrous or minutely puberulent beneath, pubescence not evident to the touch 1 Oa

9a. Leaves 12-40 cm long and obtuse at the base, oblong and drying dark brown, with 9-14

pairs of major secondary veins; flowers functionally unisexual N. kunthiana

9b. Leaves 9-24 cm long and decurrent at the base, broadly elliptic to elliptic-obovate and drying

pale brown, with 5-9 pairs of major secondary veins; flowers bisexual N. cissiflora

lOa. Anthers 0.8-0.9 mm long and usually borne on well-developed (0.4 mm) filaments, flowers 6-8

mm broad; fruits ca. 20 mm in diameter; leaves often drying lustrous and with the tertiary venation

prominent on the upper surface; 1 400-2400 m elevation N. cufodontisii

lOb. Anthers usually less than 0.8 mm long, often subsessile, flower rarely becoming 8 mm broad

(smaller); fruits rarely exceeding 15 mm in diameter; trees rarely found above 1500 m 1 la

1 la. Leaves silvery white beneath (in life) and conspicuously glaucous when dried, the leaf blades 10-

28 cm long but with only 3-6 pairs of major secondary veins, long-acuminate at the apex; fruits

ovoid-ellipsoid, to 25 mm long; rare trees of lowland evergreen forests ./V. hypoleuca

lib. Leaves never silvery white beneath and not conspicuously glaucous beneath when dried; fruits

rarely 20 mm long 1 2a

1 2a. Leaves drying subcoriaceous, decurrent on the petiole and with the base often revolute, oblong-

elliptic to lanceolate, with 3-5 pairs of major secondary veins arising from the proximal half of

the midvein; flowers 3-5 mm broad; fruit cups to 12 mm broad; 0-1700 m elevation

N. membranacea

1 2b. Leaves usually drying stiffly chartaceous, not decurrent or revolute at the base, major secondary

veins arising throughout the length of the midvein; fruit cups ca. 6 mm broad 1 3a

13a. Leaves with the major secondary veins slightly loop-connected near the margin in the distal half

of the lamina, tertiary venation usually slightly elevated on both surfaces when dry 14a

13b. Leaves with the major secondary veins not distally loop-connected near the margin 17a

1 4a. Leaves 5-1 1 cm broad (in Costa Rica), major secondary veins usually weakly loop-connected;

trees only known from the Caribbean lowlands in our area (but compare 16b, below) . . 15a

14b. Leaves 5-15 cm long and 1-6 cm broad, major secondary veins usually directly loop-connected

near the margin distally 1 6a

15a. Petioles 10-25 mm long, leaves 8-17 cm long; twigs glabrous; tepals ca. 3 mm long

N. longipetiolata

BURGER: FLORA COSTARICENSIS 55

15b. Petioles 5-12 mm long, leaves 13-24 cm long; twigs minutely puberulent; tepals ca. 2

mm long N. latifolia

16a. Leaves narrowly elliptic or ovate; anthers to 0.7 mm broad; fruits usually globose in devel-

opment, to 16 mm long; flowers appressed puberulent on the outside; 0-600(-900) m in

Costa Rica, widely distributed N. salicifolia

16b. Leaves narrowly elliptic to lanceolate; anthers to 1 mm broad; fruits usually ellipsoid in

development, to 20 mm long (dry); flowers sparsely puberulent on the outside; 600-1600 m

elevation in central and western Costa Rica N. salicina

1 7a. Flower buds densely white puberulent; stems sparsely puberulent; leaves usually drying dull above

and with the tertiary veins slightly elevated, narrowly ovate-elliptic to elliptic-oblong, with 6-10

pairs of major secondary veins N. martinicensis

1 7b. Flower buds yellowish puberulent; young stems densely puberulent; leaves usually drying dark,

lustrous and flat on the upper surface with the tertiary venation little elevated, usually narrowly

elliptic to lanceolate, with 4-7(-9) pairs of major secondary veins N. nitida

Nectandra austinii C. K. Allen, J. Arnold Arbor.

26: 374. 1945.

This species was typified by A. Smith P 2226.

Two additional collections were cited: A. Smith

240 and P 2114. We have only seen Smith's 240

and that appears to be Ocotea valeriana. Rohwer

(pers. comm.) believes that this species is part of

the Ocotea helicterifolia complex, of which O. val-

eriana is a part. Whether that complex of species

is better treated as a complex of smaller species

or a single very variable species with many sub-

specific elements cannot be determined at this time.

See the discussion under O. helicterifolia.

Nectandra belizensis (Lundell) C. K. Allen, J. Ar-

nold Arbor. 26: 400. 1945. Phoebe belizensis

Lundell, Contr. Univ. Mich. Herb. 6: 20. 1941.

N. schippiiC. K. Allen, loc. cit. 373. 1945. Fig-

ure 8.

Trees ca. 10-16 m tall and ca. 20 cm d.b.h., leafy

branchlets 2.5-4.5 mm thick, densely tomentulose with

brownish or ferruginous hairs ca. 1 mm long, the longer

hairs deciduous to expose very short (0.2 mm) grayish

hairs, the stems becoming grayish and glabrescent. Leaves

alternate and subverticellate, usually clustered at the ends

of branchlets, petioles 4-8 mm long, 1.5-2 mm thick,

densely tomentulose; leaf blades 9-16 cm long, 3.5-6

cm broad, elliptic-oblong to elliptic obovate, acute to

acuminate at the apex (rarely obtuse), obtuse to cuneate

at the base and slightly rounded at the petiole, margins

entire or undulate, drying stiffly chartaceous and brown-

ish, the upper surface with small dense hairs above the

impressed primary and secondary veins but grayish and

glabrous between the veins, darker brown and conspic-

uously puberulent beneath with slender hairs ca. 0.5 mm

long, with 4-8 major secondary veins on each side, ter-

tiary veins prominent beneath. Inflorescences solitary in

distal leaf axils, from leafless nodes, or on a terminal

shoot with poorly developed leaves and made up of sev-

eral panicles, each panicle (3-)6-12 cm long, few-flow-

ered and racemose in outline, peduncle 2-6 cm long,

0.5-1.5 mm thick, densely velutinous, pedicels 4-8 mm

long and ca. 0.3 mm thick (dry). Flowers ca. 3 mm long

and 5-7 mm broad, white, tepals ca. 4.5 mm long and

3 mm broad, sericeous on the exterior and minutely

papillate-puberulent within; outer stamens with fila-

ments only 0.1-0.2 mm long, outer anthers ca. 0.9 mm

long and 1 . 1 mm broad (in ours), broadly obovate with

the connective slightly prolonged and papillate, thecae

in an arc, staminodes very short (0.3 mm); pistil ca. 1.2

mm long with a short (0.3 mm) style and slightly discoid

stigma. Fruits and fruiting receptacle unknown.

Trees of lowland rain forest formations along

the Caribbean slope and coastal plain, from 20 to

400 m elevation. We have a single flowering col-

lection (Utley & Utley 4046, DUKE, F) from Costa

Rica, made in February at about 300 m elevation

along the road between Rio Naranjo and Canalete

(road to Upala) in northern Alajuela Province. The

only other collections are the types from Belize,

with immature inflorescences in December and

mature flowers in March.

Nectandra belizensis is recognized by its dense

puberulence, narrow and often obovate short-pet-

iolate leaves, and long-pedunculate racemose pan-

icles. The leaves clustered near the ends of branch-

lets and the laminae slightly rounded at the petioles,

usually with acuminate apices, also help to identify

the species. The broad stamens with the connec-

tive papillate and slightly expanded distally resem-

ble those found in some elements of the Ocotea

helicterifolia complex. These and other similarities

with that complex probably reflect the real rela-

tionships of this species. Costa Rican material has

leaves more narrowly obovate in form than either

the type of N. belizensis (Gentle 3304, F) or the

type of N. schippii (Schipp 856, GH, NY).

56

FIELDIANA: BOTANY

Nectandra cissiflora Nees, Syst. laur. 296. 1836.

N. paulii C. K. Allen, J. Arnold Arbor. 26: 400.

1945. Figure 11.

Trees to 30 m tall, trunks ca. 30 cm d.b.h., leafy

branchlets 2.5-6 mm thick, densely puberulent in early

stages with slender ascending hairs 0.1-0.3 mm long,

with longitudinal ridges from beneath the leaf bases but

becoming terete. Leaves alternate and somewhat clus-

tered distally, petioles 1 1-28 mm long, 1 .3-3.5 mm thick,

velutinous-tomentulose, the adaxial ridges forming a sul-

cus above; leaf blades 9-21 (-24) cm long, 4-8(-9) cm

broad, broadly elliptic to elliptic-oblong, oblanceolate or

elliptic-obovate, abruptly narrowed or rounded to the

bluntly obtuse to short-acuminate apex, tapering grad-

ually to the acute or obtuse base and strongly decurrent

on the petiole, margin entire and revolute (especially near

the base), drying stiffly chartaceous to subcoriaceous and

grayish green to pale brown, densely puberulent on the

midvein above with slender appressed or spreading hairs

ca. 0.2 mm long, becoming (sub)glabrous in age, the

lower surface minutely puberulent with slender incon-

spicuous hairs, with 6-9(-l 1) major secondary veins on

each side, the secondary veins slightly impressed above

and prominent beneath. Inflorescences axillary to leaves

(occasionally pseudoterminal), 12-25 cm long, many

flowered panicles with branches in the distal half, pe-

duncle to 11 cm long, 1.8-3 mm thick, densely and

minutely velutinous, pedicels 1-5 mm long (to 12 mm

in fruits). Flowers 4-6 mm broad, white, densely pu-

berulent on the outside, tepals 1.3-2 mm long, 0.9-1.5

mm broad, broadly rotate at anthesis, densely papillate-

puberulent within; outer stamens subsessile with anther

0.4-0.6 mm long and 0.6-0.9 mm broad, thecae in a

single row or arcuate, connective papillate distally, inner

stamens with anthers broader than long and with con-

spicuous glands, staminodes small (0.5 mm) and some-

what broadened and flattened distally; pistil ca. 1.3 mm

long, ovary globose and 0.9-1.1 mm in diameter, style

very short (0.2-0.3 mm), stigma simple and flat. Fruits

borne in a flat, saucer-like or shallow cupulate receptacle,

abruptly expanded above the thickened (3 mm) pedicel,

10-12 mm broad and 1-3 mm deep, green; berry globose

or subglobose, 1 .3-2 cm in diameter, dark green becom-

ing black.

Rarely collected trees of wet evergreen or partly

deciduous forest formations, between 50 and 1 200

m elevation on both the Caribbean and Pacific

slope in Costa Rica. Flowering is in January-March

with fruit set in April-June. In Costa Rica the

species is only known from La Selva (Hammel

1 1168, DUKE, 1 1665, DUKE, F), the General Valley

near San Isidro (Skutch 2605, us, the type of N.

paulii) and Las Alturas de Coto Brus (Burger et

al. 12187, CR). The species ranges from Mexico to

Peru and Brazil.

Nectandra cissiflora is recognized by the larger,

minutely puberulent, stiffleaves with usually more

than six pairs of secondary veins, decurrent leaf

base, small subsessile anthers with thecae in a hor-

izontal row, ovary with very short style, and glo-

bose fruits subtended by a very shallow cup. This

species is similar to Ocotea sp. aflf. caracasana and

Ocotea glaucosericea in general appearance (C. K.

Allen used fruits of the latter species in describing

N. paulii). Even in forests that have been carefully

studied, these trees are rarely collected. Hammel

(1986) states that they grow on ridge tops.

Nectandra cufodontisii (O. C. Schmidt) C. K. Al-

len, J. Arnold Arbor. 26: 393. 1945. Ocotea cu-

fodontisii O. C. Schmidt, Arch. Bot. (Forli) 1 1 :

50. 1935. O. seibertii C. K. Allen, loc. cit. 336.

1945. Figure 12.

Trees 8-35 m tall, often with a well-developed spread-

ing crown, leafy branchlets 2-5 mm thick, at first with

minute (0.3 mm) slender appressed-ascending hairs but

quickly glabrescent. Leaves alternate, petioles 8-2 1 mm

long, 1.2-2.3 mm thick, flat or slightly sulcate above

with lateral margins continuous with the lamina margins,

very minutely puberulent; leaf blades (5-)6-15(-21) cm

long, (2-)3-7(-9) cm broad, narrowly elliptic to ovate-

elliptic, acute to short-acuminate at the apex, obtuse to

rounded at the base and decurrent on the petiole, margins

entire and with a veinlike revolute edge, drying subcor-

iaceous and the midvein yellow orange above, smooth

and lustrous (rarely dull) above, essentially glabrous be-

neath but sometimes with longer whitish hairs in the

basal vein-axils (domatia), with 4-7 major secondary

veins on each side, central secondaries arising at angles

of 20°-40°, tertiary veins slightly raised on the upper

surface. Inflorescences axillary to distal leaves, 5-15(-25)

cm long, peduncle to 10 cm long, reddish in life and

drying dark, lateral branches short (1-3 cm) or long and

the panicle narrow or broad, glabrous or minutely ap-

pressed puberulent, pedicels 1.5-4.5 mm long. Flowers

ca. 3 mm long and 6-8 mm broad, sweet-scented and

cream-white to yellowish green, floral tube short and

funnelform, tepals ca. 2.5 mm long and 1.5 mm broad,

papillate-puberulent within; androecium 2-2.5 mm broad,

outer stamens broadly ovate, 0.8-0.9 mm long on fila-

ments ca. 0.4 mm long, thecae in an arcuate configura-

tion or somewhat superposed, inner stamens biglandu-

lar, small staminodes often present; pistil 1.7-2.1 mm

long, ovary ellipsoid to ovoid, style 0.8-1 mm long, stig-

ma simple. Fruits borne in a flat or cupulate receptacle

8-1 1 mm broad, 0-3 mm deep, abruptly expanded above

the thickened pedicel, a very short (1-2 mm) broad stipe

often developed beneath the fruits, cup red at maturity;

berry (13-)20-40 mm long and 10-20 mm in diameter

when dry (said to become 5 cm long and globose), green

becoming blue black or purple.

Distinctive trees of montane cloud forest for-

mations, from 1400 to 2600 m elevation. Flow-

ering in May-August; fruits have been collected

in January-September. The species ranges along

the Caribbean side of the Meseta Central, from

Palmira and Poas eastward through the Cordillera

BURGER: FLORA COSTARICENSIS

57

de Talamanca to the Chiriqui highlands in Pan-

ama, and it has recently been found in the Cor-

dillera Central de Nicaragua.

Nectandra cufodontisii is distinguished by the

dried leaves usually lustrous and with the minor

venation slightly raised on the upper surface to

produce a characteristic texture, and the montane

habitat. The inflorescences are bright orange in life

and the leaves often dry an orange brown color.

The stamens are usually more like those charac-

teristic of Nectandra, with the thecae forming an

arcuate pattern, but superposed (Ocotea-\ike) the-

cae can be found in many collections. The short

flat cup beneath the large round fruits is another

unusual feature of this species. The outer part of

the fruiting receptacle (cup) may break off, pro-

ducing a smaller (7 mm) flat base as in the type

collection (Cufodontis 315, F). Common names are

quizarrd and yema de huevo.

Nectandra davidsoniana C. K. Allen, J. Arnold

Arbor. 26: 369. 1945.

This small-leaved species of the cloud forests in

the Chiriqui highlands in Panama resembles N.

salicina but has Ocotea-like stamens. The correct

disposition of this species is uncertain at this point;

it appears to be endemic in western Panama. See

the discussion under Ocotea viridiflora.

Nectandra globosa (Aubl.) Mez, Jahrb. Konigl. Bot.

Gart. Berlin 5:415.1889. Laurus globosa Aubl.,

Hist. pi. Guiane 1: 364. 1775. See also Lourteig

in Phytologia 63: 153-154. 1987. Figure 18.

Trees 4-1 5(-25) m tall, leafy branchlets 2-7 mm thick,

at first grayish or yellowish puberulent with a covering

of very minute (0.05-0. 1 mm) thin appressed hairs that

may be difficult to see, becoming terete. Leaves alternate,

petioles 6-18 mm long, 1-2 mm thick, slightly flattened

or sulcate above; leaf blades (7-)l l-24(-30) cm long,

(2-)3-8(-10) cm broad, ovate lanceolate to elliptic-lan-

ceolate, narrowly elliptic or narrowly elliptic-oblong, ta-

pering gradually to the long-acuminate (acute) apex, acute

to obtuse at the base, margin entire and often revolute

just above the petiole, drying stiffly chartaceous to sub-

coriaceous and often yellowish or grayish, dull, flat and

glabrous above with the tertiary venation obscure, usu-

ally with a fine tomentum of minute (0. 1 mm) hairs

beneath, with (4-) 7-1 1 arcuate ascending major second-

ary veins on each side and very prominent beneath, ter-

tiary veins subparallel between the secondaries and usu-

ally obscure, domatia of tufted hairs often present in the

vein axils beneath. Inflorescences axillary to distal leaves

or pseudoterminal, 8-15(-20) cm long, broadly pani-

culate, peduncle ca. 1.5 mm thick and minutely puber-

ulent, usually equal to the length of the distal flowering

rachis, pedicels 1-7 mm long, minutely puberulent.

Flowers white or cream colored, puberulent, 5-7 mm

long, 6-9(-12) mm broad, tepals 3-5 mm long, densely

papillate within, inner tepals often narrower and more

obovate than the outer; outer stamens sessile or subses-

sile, outer anthers 0.5-1.4 mm long and 0.8-1.4 mm

broad, the 4 thecae in a horizontal row at the base of

the distally expanded thick and minutely papillate con-

nective, distal part of the connective usually more than

half the length of the anther, staminodes present or ab-

sent, to 1 mm long and acute at the apex; pistil 1.3-1.7

mm long, globose or turbinate, the style very short (0.4

mm) or absent, stigma simple or slightly discoid. Fruits

borne on a short (1-4 mm) flat or slightly cupulate re-

ceptacle 0.5-3 mm deep, 5-8 mm broad at the apex,

margin entire; berry becoming globose and ca. 1 cm in

diameter (dry), perhaps ovoid-ellipsoid in some popu-

lations (fruits are rarely collected), becoming black.

Trees of deciduous and partly deciduous forest

formations, from near sea level to 700 m elevation

along the Pacific slope of Costa Rica. Flowering

material has been collected in December-March.

Fruits have been collected in February-April (and

in June at Turrialba where it has been planted).

This species ranges from Mexico to Bolivia and

Brazil.

Nectandra globosa is recognized by its large stiff

leaves usually broadest below the middle and with

the margin often curled under just above the pet-

iole, the minute puberulence on many parts, the

large flowers (for Nectandra) with tepal-like sta-

mens, and the restriction to forests with a dry sea-

son of several months. (The distal development

of the anther connective is an unusual feature

among Costa Rican species but typical for many

Nectandra species of other areas.) There are some

specimens in Costa Rica that may be interpreted

as intermediate between N. globosa and N. ra-

monensis of higher elevations. Both of these species

have the anther connective expanded distally.

Present evidence seems to indicate that the two

species are well separated ecologically, with only

occasional and problematic intermediates. Roh-

wer has recently annotated material (at F & MO)

as Nectandra caucana (Meissn.) Mez, and this may

become the accepted name for the material placed

here.

In the earlier part of this study Nectandra gla-

brescens Bentham was thought to differ from N.

globosa by the thinner dark brown leaves with

obscure minor venation (when dried), and a pref-

erence for moister forest habitats (cf. dichotomy

6b in the key to Nectandra species; and fig. 18,

second from the top). It now appears that these

differences are not significant. Also, the syntypes

58

FIELDIANA: BOTANY

from Mexico and Nicaragua may not represent the

same species (Rohwer, pers. comm.).

Nectandra hypoleuca Hammel, J. Arnold Arbor.

67: 126. 1986. Figure 18.

Trees 5-15 m tall, trunks 10-35 cm d.b.h. with dark

brown bark, leafy branchlets 2-5 mm thick, very mi-

nutely (0. 1 mm) papillate-puberulent. Leaves alternate,

petioles 5-15 mm long, sulcate above, dark brown; leaf

blades 10-28 cm long, 4-1 1 cm broad, elliptic to broadly

elliptic, elliptic-oblong or narrowly elliptic, tapering

gradually to the long-acuminate apex, the tip often 2 cm

long, tapering gradually to the acute base, drying stiffly

chartaceous and dull grayish above, glabrous above and

below but with minute appressed hairs along the midvein

and tufts of hairs (domatia) in some vein axils beneath,

conspicuously glaucous beneath when dry, with 3-6 ma-

jor secondary veins on each side, tertiary veins obscure

and occasionally subparallel. Inflorescences axillary to

distal leaves (sometimes pseudoterminal with distal leaves

failing to develop and forming larger compound inflo-

rescences), 4-18 cm long, peduncles 3-15 mm long,

branches of the primary rachis becoming shorter distally,

somewhat distant (5-1 5 mm) and forming an open pan-

icle, ultimate flower clusters with 3-10 flowers, pedicels

ca. 2 mm long, minutely puberulent. Flowers 2-2.5 mm

long and 4-6 mm broad, perianth white or greenish white,

tepals 2-2.5 mm long, minutely papillate-puberulent

within; outer stamens subsessile, outer anthers 0.3-0.4

mm long and 0.6-0.7 mm broad, distinctly broader than

long and with the thecae in 1 plane with the outer opening

laterally, inner stamens with narrower thecae, stami-

nodesca. 0.4 mm long and clavate; pistil 1-1.5 mm long,

ovary 0.7 mm broad, style narrow, stigma simple or

slightly lobed. Fruits borne in a shallow or conical cup

5-8 mm long, 7-12 mm broad and 2-4 mm deep, en-

closing only the base of the fruits, pedicel and cup red;

berry 16-25 mm long, 12-1 6 mm thick, ovoid to broadly

ellipsoid, becoming purple.

Trees of the wet evergreen forests of the Carib-

bean lowlands and perhaps in the Golfo Dulce

area. Flowering collections have been made in

May-August and October-November; fruits have

been collected in September-October. At present,

this species is known from the La Selva site where

it is found in old secondary woods and alluvial

forest along the Rio Puerto Viejo at about 100 m

elevation in the province of Heredia, and it may

occur on the Osa Peninsula (see below).

Nectandra hypoleuca is recognized in life by its

gray green leaves which are strikingly silvery white

or glaucous on the undersurfaces, according to

Hammel (1986). The leaves almost glabrous, with

occasional tufted domatia, grayish and smooth

above, tapering gradually to both apex and base

(but not decurrent), long-acuminate tip, and the

small flowers on somewhat distant branches of an

open panicle help distinguish dried specimens. An

unusual collection (Knapp & Mallet 2208, CR, MO)

from Parque Nacional Corcovado has foliage ex-

actly like that of N. hypoleuca but the small (0.7

mm) anthers have only two valves! Two sterile

collections^. Gentry 48524, 48567, MO) from this

same locality probably also represent this popu-

lation; all three are placed here tentatively.

Nectandra kunthiana (Nees) Kosterm., Meded.

Bot. Mus. Utrecht 25: 19. 1936. Acrodiclidium

kunthianumNees, Syst. laur. 269. 1936. Ocotea

kunthiana (Nees) Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 29 1 . 1 889. Ocotea cooperi C. K. Allen,

J. Arnold Arbor. 26: 335. 1945. Rhodostemon-

odaphne kunthianum (Nees) Rohwer, Mitt. Inst.

Allg. Bot. Hamburg 20: 84. 1986. Figure 7.

Trees to 25 m tall, dioecious, leafy branchlets 2.5-7

mm thick, sparsely to densely puberulent with dark gray

or brown hairs to 0.3 mm long. Leaves alternate, petioles

8-37 mm long, 2-4 mm thick, somewhat sulcate above,

minutely puberulent; leaf blades 12-32(-40) cm long, 6-

12(-16) cm broad, elliptic to elliptic-oblong, tapering

gradually to a long (1-4 cm) acuminate apex, obtuse at

the base, margin entire or somewhat undulate, drying

subcoriaceous, becoming glabrous and lustrous above,

puberulent beneath with stiff slender hairs 0.1-0.5 mm

long, with 9-14 major veins on each side, the central

secondaries arising at angle of 40°-60°, major veins im-

pressed above, tertiary veins often subparallel and slight-

ly raised above. Inflorescences 1 1-22 cm long, axillary

or pseudoterminal, thyrsiform panicles with the flowers

somewhat clustered on the lateral branches, peduncles

4-9 cm long, ca. 2 mm thick, densely and minutely pu-

berulent. Flowers unisexual but with organs of both sexes,

urceolate and with a well-developed floral tube; male

perianth campanulate, 4-5 mm broad, outer anthers

subsessile, 0.6 mm long and 0.6-0.9 mm broad, thecae

in a line or arc, staminodia absent, pistillode linear; fe-

male flowers more urceolate, ca. 4 mm long and 3 mm

broad, anthers smaller than in the male and with incon-

spicuous valves, pistil 2.5 mm long, slender, style short,

stigma discoid. Fruits borne in a cup ca. 16 mm long

and 16 mm broad, campanulate and 6-10 mm deep,

fruit stalk 6-20 mm long, margin of cup entire or slightly

lobed, becoming orange red; berry 2.5-4 cm long, ca. 1 .5

cm thick, oblong-ellipsoid, the lower third enclosed with-

in the cup, glabrous, becoming black.

Infrequently collected trees of wet evergreen for-

est formations, between 50 and 900 m elevation

on the Caribbean slope and foothills, and in the

General Valley in Costa Rica. Flowering material

has been collected in January-March, and fruits

have been collected in July-August. The species

ranges from Costa Rica and Panama to the Guian-

as and Ecuador, and may extend to Bolivia.

BURGER: FLORA COSTARICENSIS

59

Nectandra kunthiana has distinctive foliage that

dries dark in color, laminae that are often quite

large and with a narrow acuminate tip, numerous

secondary veins, and often with the subparallel

veins raised on the leaves' lustrous upper surface.

The densely puberulent inflorescences, function-

ally unisexual flowers, and the deep cups are fur-

ther distinctions. A local name is quizarrd negra.

Rohwer and Kubitzki (1985) erected the genus

Rhodostemonodaphnelo replace Synandrodaphne

Meissn., and they distinguish this genus from Nec-

tandra by its dioecious habit, inner surface of the

tepals hairy rather than papillate, and thecae along

the upper edge of the anther. While transfer to a

segregate genus may be justified, we continue the

traditional placement of this species in Nectandra

for the sake of convenience.

Nectandra latifolia (H.B.K.) Mez, Jahrb. Konigl.

Bot. Gart. Berlin 5: 454. 1889. Ocotea latifolia

H.B.K.,Nov.Gen.Sp.2: 165. IS 11. N. purpurea

auct. Figure 17.

Small or medium-sized trees 5-17 m tall, 15-20(-40)

cm d.b.h., leafy branchlets 1.5-4.5 mm thick, minutely

(0.1-0.2 mm) appressed puberulent with thin ascending

brownish hairs, becoming (sub)glabrous and dark gray-

ish brown. Leaves alternate, petioles 5-12 mm long, 1.5-

2 mm thick, with 2 adaxial ridges continuous with the

lamina margin and usually forming a slight sulcus above,

minutely appressed, puberulent; leaf blades 13-24 cm

long, 5-11 cm broad, elliptic-oblong to narrowly ovate-

elliptic or broadly oblong, short to longer acuminate or

caudate-acuminate at the apex, tip 0.5-2 cm long, acute

to obtuse (rarely rounded and subtruncate) at the base,

shortly decurrent on the petiole, margin entire or slightly

undulate (dry), drying stiffly chartaceous, the upper sur-

face glabrous, lustrous, and with the minor venation

often slightly raised, lower surface glabrous and with the

minor venation slightly raised, with (4-)5-8 major sec-

ondary veins on each side, the distal secondaries often

loop-connected near the margins, pit domatia with a few

hairs often present in the proximal and central vein axils

on the lower surface. Inflorescences axillary to distal

leaves or pseudoterminal, 6-1 7 cm long, paniculate with

spreading lateral branches, peduncle 1-6 cm long, red-

dish, lateral branchlets grayish puberulent, pedicels 1-2

mm long. Flowers ca. 3 mm long, 4-5 mm broad, white

at anthesis and becoming orange, tepals (1-) 1.3-2 mm

long, densely papillate-puberulent within; outer anthers

0.4-0.6 mm long and 0.5-0.9 mm broad, on short (0.2

mm) filaments, lower thecae lateral to the upper or oc-

casionally superposed, inner stamens ca. 1 mm long with

filaments 0.6 mm long, staminodes slender with a narrow

or thickened apex, 0.4-0.7 mm long; pistil 1-1.6 mm

long, ovary ca. 0.7 mm in diameter, style ca. 0.4 mm

long, stigma discoid. Fruits not seen from Costa Rica

and the following based on Barro Colorado Island (Pan-

ama) collections: fruiting receptacle short (3-7 mm) and

conical, only 2-3 mm deep and 6-10 mm broad at the

apex, becoming red; berry globose-ellipsoid, 14-18 mm

long and 10-14 mm in diameter when dry.

Trees of evergreen lowland rain forest and partly

deciduous forests, but only known from La Selva

at 50 to 1 50 m elevation at the edge of the Carib-

bean coastal plain in Costa Rica. Flowering in May-

June in Costa Rica; flowering in December-July

in Central Panama with two peaks: December-

March and May-July. This species is said to range

from Nicaragua to Colombia, Ecuador, Peru, and

Bolivia (but see below).

Nectandra latifolia is recognized by the rela-

tively larger (in ours), almost glabrous leaves, lus-

trous above and with the minor venation usually

slightly elevated on both surfaces (dry), secondary

veins loop-connected distally, the sparsely puber-

ulent domatia, the many-flowered panicles with

small flowers, very small stamens, and the slender

staminodes. Costa Rican material is quite different

from the smaller-leaved material from Barro Col-

orado Island (which resembles N. salicifolia in some

ways), and it may be an error to base our fruiting

description on the Panamanian material. Speci-

mens from La Selva are intermediate in leaf size

between the Barro Colorado Island collections, and

the much larger-leaved TV. lundellii C. K. Allen

from the Caribbean side of Honduras, Guatemala,

and Belize. Also, there may be trees intermediate

between our lowland representatives of this species

and TV. cufodontisii in the General Valley and Las

Alturas de Coto Brus (cf. Burger et al. 12183, CR,

F, determined as N. cufodontisii).

Costa Rican collections have been called Nec-

tandra purpurea (Ruiz & Pavon) Mez based on

Laurus purpurea Ruiz & Pavon, in Laurografia

Peruviana t. 7. A Ruiz and Pavon collection at

Field Museum, with foliage much like that in the

original illustration, has stamens with a more Oco-

tea-like form than in material from Panama and

Costa Rica. Thus, N. latifolia, both in the sense

of the collections from Panama and those from La

Selva, appears to be more closely related to N.

salicifolia than to TV. purpurea. This species was

mistakenly called N. purpurascens in the Flora of

Barro Colorado Island (Croat, 1978).

Nectandra longipetiolata van der Werff, sp. nov.

Figure 13.

Arbor ad 10 m alta. Folia altema, chartacea, 9-17 x

5-8 cm, glabra, elliptica vel ovato-elliptica, basi obtusa,

apice acuta vel breviter acuminata; costa nervique (4-

8) super immersi, subtus elevati; reticulatio elevata. Pe-

60

FIELDIANA: BOTANY

tioli glabri, 10-25 mm longi. Inflorescentiae ad 4 cm

longae, parviflorae. Flores extus minute puberuli vel

papillati. intus dense papillati; tepala ca. 3.5 mm longa,

patentia vel reflexa per anthesim; stamina 9, omnia 4-

locellata, 6 exterioribus filamentis brevissimis, locellis

introrsis; 3 interioribus locellis lateralibus vel laterali-

extrorsis; staminodia 3; ovarium ovoideum, glabrum.

Fructus ellipsoideus, ad 2 x 1.4 cm; cupula parva, pa-

telliformis.

Small trees 4-10 m tall, leafy branchlets 1.2-3 mm

thick, glabrous and reddish brown. Leaves alternate, pet-

ioles (8-) 10-25 mm long, 0.9-1.7 mm thick, glabrous;

leaf blades (7.5-)9-l 7 cm long, (4-)5-8 cm broad, ovate-

elliptic to elliptic or slightly oblong, tapering to a short

and bluntly acuminate apex, obtuse at the base, drying

stiffly chartaceous, slightly lustrous on the upper surface

with the secondary and tertiary venation distinctly raised

when dry and grayish green to olive green, drying paler

grayish green or yellowish beneath, with 4-8 major sec-

ondary veins on each side, central secondaries arising at

angles of 40°-60°, minor venation prominent beneath

but not forming a raised reticulum, pit domatia with a

few hairs sometimes present in the axils of the basal

secondaries beneath. Inflorescences ca. 4 cm long (only

one seen), a racemose panicle with few (ca. 25) flowers,

axillary to a distal leaf, glabrous. Flowers ca. 6 mm broad

at anthesis, perianth parts ca. 3.5 mm long and 2 mm

broad, sparsely puberulent on the exterior and minutely

papillate puberulent on the interior (adaxial) surfaces;

outer stamens ca. 0.9 mm long with anthers 0.7 mm long

and 0.8 mm broad, the thecae in a horizontal plane or

very low arc, staminodes prominent, ca. 0.8 mm long

with a short stipe and broad (0.6 m) apex; pistil ca. 1.4

mm long, ovary 1 mm in diameter, style short (0.4 mm)

and poorly denned, stigma slightly thickened. Fruits borne

on a short (2-6 mm) cup 6-7 mm broad at the apex,

conical or abruptly expanded, puberulent within, rose

red; berry broadly ellipsoid or slightly obovoid, 1 5-20

mm long and 1 1-14 mm in diameter, green.

TYPE— Costa Rica, Prov. Limon, Hitoy Cerere

Reserve, 31 July 1985, M. Grayum & B. Hammel

5769 (holotype, MO; isotype, CR).

Understory trees of ridges in evergreen rain for-

est of the Caribbean lowlands near Manzanillo de

Talamanca and the Hitoy Cerere Reserve, between

50 and 200 m elevation. Known from only three

collections (Grayum et al. 4381, 4396, 5769, all

CR, MO), from the Caribbean lowlands of south-

eastern Costa Rica.

Nectandra longipetiolata is distinguished by its

completely glabrous leaves drying grayish to yel-

lowish green, distinctive venation on the dried leaf

surfaces, anther form, and small fruiting cups. At

first glance TV. longipetiolata does not seem to be

a Nectandra, because of the long petioles (for a

Nectandra), the weakly extended secondary veins

which do not come close to the lamina margin,

and the raised minor venation on both leaf sur-

faces. However, flowers and fruits leave no doubt

that this species belongs to the N. salicifolia species

group, characterized by relatively well-developed

staminodia, inner stamens with lateral or lateral-

extrorse dehiscence and the small, saucer-like cu-

pule. Within the N. salicifolia group, N. longipe-

tiolata stands apart by its long petioles, glabrous

twigs and leaves, almost glabrous flowers, upper

surface of the leaves not drying dark, the broad

leaves with an obtuse base, and the small inflo-

rescences.

Nectandra martinicensis Mez, Mitt. Bot. Vereins

Kreis Freiburg 47^18: 421. 1888, Jahrb. Konigl.

Bot. Gart. Berlin 5: 459. 1889. N. woodsoniana

C. K. Allen, J. Arnold Arbor. 26: 394. 1945.

Figure 17.

Small to medium-sized trees 5-15(-25) m tall, leafy

internodes 0.5-4 cm long, 1.5-4.5 mm thick, minutely

grayish tomentulose but quickly becoming (sub)glabrous,

grayish or brown and minutely longitudinally striate.

Leaves alternate, petioles 6-1 3 mm long, 1-2 mm thick,

with 2 adaxial ridges and sulcate or flat above; leaf blades

12-28 cm long, 4-8.5 cm wide, elliptic-oblong to nar-

rowly elliptic or narrowly ovate-elliptic, tapering grad-

ually to an acuminate apex, the tip 0.5-1 .5 cm long, acute

to obtuse at the base, drying chartaceous and dull grayish

green or darker and slightly lustrous above, glabrous or

with slender appressed hairs 0.2-0.4 mm long on the

upper surface, glabrous or with minute slender hairs be-

neath, with 6-10 major secondary veins on each side,

pit domatia or tufted hairs often present in the axils of

proximal veins beneath, the minor venation often slight-

ly raised above (dry). Inflorescences solitary in distal leaf

axils or pseudoterminal, 8-15(-20) cm long, paniculate

with open distal branching and the peduncle about half

the length (3-8 cm) of the rachis, peduncle 1.5-2 mm

thick, reddish brown or yellowish, minutely puberulent,

pedicels 0.5-4 mm long. Flowers 3-4 mm long and 5-8

mm broad, white and showy at anthesis, tepals ca. 2.5

mm long, puberulent on the outside and densely papil-

late-puberulent on the inner surfaces; androecium ca. 2.5

mm in diameter, outer stamens subsessile or with short

(0.4 mm) filaments, outer anthers 0.4-0.6 mm long and

0.7-0.9 mm broad, subreniform to rectangular, the lower

thecae lateral to the upper or all in a single plane, anthers

emarginate or not, inner stamens ca. 1 mm long with

filaments 0.4-0.5 mm long, staminodes slender or with

a broadened apex, ca. 0.5 mm long; pistil 1.2-1.7 mm

long, style 0.3-0.5 mm long, stigma discoid. Fruits borne

on a flat disclike receptacle 2-4 mm long and 6 mm

broad; berry ellipsoid, 15-18 mm long and 10 mm in

diameter.

Trees of evergreen forests or partly deciduous

forests of the Pacific slope in Costa Rica, from

near sea level in Panama to 1 300 m in the central

highlands of Costa Rica. Costa Rican collections

are primarily from the eastern part of the Meseta

Central and the western part of the General Valley

BURGER: FLORA COSTARICENSIS

61

in the province of San Jose. Flowering collections

have been made in March-July; fruits have been

collected in October-November and January. The

species range is poorly understood because of the

difficulty of separating closely related species.

Nectandra martinicensis is recognized by the

larger sparsely puberulent chartaceous laminae with

as many as 1 0 pairs of major secondary veins, the

larger open many-flowered inflorescences with

whitish puberulent buds, and short outer stamens

with thecae often in a single row. Specimens of

this species may be difficult to separate from un-

usual individuals of the N. salicifolia-N. latifolia

species group. Moreover, there may be interme-

diates with material placed in N. nitida, which has

narrow leaves that are usually very lustrous and

often dry dark on the upper surface. Despite the

name, this species does not grow in Martinique.

Howard (198 1) discusses the problems associated

with the name of this species and, because he does

not accept Nectandra as a genus, he places this

taxon under the name Ocotea tabascensis (Lun-

dell) Howard, since Mez had already described an

Ocotea martinicensis. We follow the interpretation

of Bernardi (1967, pp. 69-72).

Nectandra membranacea (Sw.) Griseb., Fl. Brit.

W. I. 282. 1860. Laurus membranacea Sw.,

Prodr. 65. 1788. N. skutchii C. K. Allen, J. Ar-

nold Arbor. 26: 396. 1945. N. standleyi C. K.

Allen, loc. cit. 1945. Figure 17.

Shrubs and small to medium-sized trees, 4-18 m tall,

leafy branchlets 1.5-7 mm thick, at first minutely (0.1

mm) appressed puberulent, becoming (sub)glabrous and

dark in color, terete. Leaves alternate, petioles 5-20 mm

long, flat or slightly sulcate above, densely and very mi-

nutely puberulent in early stages, lateral margins contin-

uous with the decurrent lamina margins; leaf blades 8-

24(-30) cm long, 4-7(-13) cm broad, oblong-elliptic to

lanceolate, ovate-lanceolate, narrowly ovate or oblong,

broadest at or below the middle, usually tapering grad-

ually to a short or long-acuminate apex, acute to obtuse

at the base, margin entire and decurrent on the petiole,

margin usually revolute near the base of the lamina,

drying subcoriaceous and dull grayish or brown above,

the major veins flat or impressed above and the tertiary

venation obscure, very minutely (0. 1 mm) puberulent or

appearing glabrous, with 3-5 major secondary veins on

each side and arising at angles of 20°-40°, usually with

the most distal secondaries arising from the middle of

the lamina, tertiary veins subparallel and perpendicular

to the midvein (often perpendicular to the secondaries

in larger leaves), domatia absent. Inflorescences pseu-

doterminal or axillary to distal leaves, 4-14(-20) cm

long, paniculate, peduncles to 8 cm long, minutely ap-

pressed puberulent with yellowish or brownish hairs,

pedicels 1.5-4 mm long. Flowers 3.5-5.5 mm broadi

yellowish green in life, tepals 1 .5-2.5 mm long, ca. 1 mm

wide, densely papillate-puberulent within; outer anthers

0.3-0.5 mm long and 0.6-0.8 mm broad, on short fila-

ments, staminodes (0.2-)0.4-0.7 mm long, slender and

with a slightly triangular tip; pistil 1.1-1.6 mm long,

ovary 1 mm thick, glabrous, style 0.7-1 mm long, stigma

simple. Fruit cup (8-) 10-1 2 mm broad, very shallow (1-

3 mm), mostly conical in form, ca. 10 mm long, appar-

ently remaining green; berry 1 2-1 5 mm long, globose to

ovoid (rarely ellipsoid), green becoming black.

Shrubs or trees often encountered along forest

edges and in more open sites or secondary woods

in wet evergreen forest formations, from 50 to

1700 m elevation along the Caribbean slopes of

Costa Rica and in the highland forests; not known

from the Pacific slope below 600 m elevation.

Flowering collections have been made in May-

December with an apparent peak in July-August.

Fruits have been collected in December-April, but

appear to mature mostly in March-April. The

species ranges from Mexico and the West Indies

to Peru and Brazil.

Nectandra membranacea is recognized by the

narrow leaves with relatively few major secondary

veins arising from the proximal half of the leaf

blade and strongly arcuate-ascending. The decur-

rent leaf base and revolute margin (near the base),

minute puberulence, small globose-ovoid fruits,

and restriction to moister evergreen forests further

characterize this species. Nectandra gentlei Lun-

dell with narrower, usually lanceolate, leaves is

probably no more than a variety of this species.

A number of collections from the Caribbean slope

with leaves drying orange brown and venation not

so arcuate-ascending are tentatively placed here

(Gomez- Laurito 11262, 11272, both at CR, usj;

Grayum et al. 8060, CR, MO).

Nectandra nitida Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 461. 1889. N. perdubia Lundell, Lloy-

dia4:47. 1941. Figure 17.

Trees (rarely shrubs) (3-)5-20 m tall, 4-30(-90) cm

d.b.h., leafy branchlets 1.7-5 mm thick, densely puber-

ulent with minute (0. 1-0.4 mm) yellowish brown or gray-

ish appressed-ascending hairs, longitudinally ridged but

becoming terete, glabrescent and grayish. Leaves alter-

nate, petioles 4-10 mm long, 1-2 mm thick, densely

brownish puberulent, sulcate or flat above; leaf blades

10-23(-30) cm long, 3-6(-7) cm broad, lanceolate to

narrowly elliptic, elliptic-oblong, narrowly oblong or

narrowly ovate-elliptic, tapering gradually to an acu-

minate tip (rarely acute or caudate-acuminate), the tip

to 3 cm long, obtuse to acute at the base, the margin

entire, drying stiffly chartaceous to subcoriaceous, usu-

ally dark and lustrous above, usually sparsely appressed-

62

FIELDIANA: BOTANY

puberulent above and becoming glabrescent, sparsely ap-

pressed-puberulent beneath with thin ascending hairs

(rarely glabrous), with (3-)4-7(-9) major secondary veins

on each side, the central secondaries arising at angles of

25°-45° and strongly ascending (rarely loop-connected

distally), with pits and tufted hairs in the axils of the

major veins beneath (not always present). Inflorescences

axillary to distal leaves or pseudoterminal, sometimes

appearing to be several per axil when the peduncle is

very short, to 1 5 cm long, paniculate and widely branch-

ing to racemose in form, peduncles 3-50 mm long, pu-

berulent, pedicels 1-4 mm long. Flowers 5-7 mm broad,

white, tepals 2-2.3 mm long, ca. 1.3 mm broad, puber-

ulent on the outside and densely papillate within; outer

stamens 0.6-0.8 mm long with filaments 0.2-0.3 mm

long, outer anthers ca. 0.3 mm long and 0.5 mm broad,

rectangular to reniform, the thecae in 1 plane or arcuate,

connective flat or emarginate distally, inner stamens with

quadrangular anthers and large glands, staminodes mi-

nute or small (0.6 mm), often with a sagittate apex; pistil

1-1.3 mm long, ovary rounded and ca. 0.7 mm thick,

style ca. 0.3 mm long, stigma discoid. Fruits borne on

a short (2-4 mm) conical or slightly cupulate receptacle

4-6 mm broad at the apex and only 1-2 mm deep, mar-

gin entire, becoming red; berry (7-)8-12 mm long and

(6-)7-10 mm in diameter (dry), globose or subglobose,

becoming dark green.

Small or medium-sized trees often found at for-

est edges and in secondary formations, from sea

level to 1 300 m elevation in evergreen and partly

deciduous formations. The primary flowering pe-

riod for this species in Mexico and Guatemala is

March-June, and fruiting range is September-

February. The species ranges from middle Mexico

through Belize and Guatemala to Costa Rica and

adjacent Panama (but see below).

Nectandra nitida is recognized by its narrow

leaves usually gradually narrowed at both apex

and base, the glossy upper leaf surfaces that dry

dark, the densely puberulent young stems, the small

broad anthers on short filaments, and the small

globose fruits on short-conical or shallow cupulate

receptacles. This species has been little collected

in Costa Rica, and it is difficult to separate from

N. martinicensis and TV. salicifolia in our area; in

fact, there may be intergradation between N. nitida

and TV. martinicensis in our highlands, and inter-

gradation with N. salicifolia in our lowlands. While

difficult to recognize and separate in Costa Rica,

this species seems more easily identified and much

more common in Guatemala and southern Mex-

ico.

Nectandra ramonensis Standley, Publ. Field Mus.

Nat. Hist. Hot. Ser. 18: 453. 1937. Figure 18.

Trees 5-1 5 m tall, leafy branchlets (0-)5-35 mm long,

1-4 mm thick, densely appressed puberulent with short

(0.2 mm) grayish or pale brown hairs, older twigs gla-

brescent, a reddish brown inner bark sometimes becom-

ing exposed by flaking off of outer bark. Leaves alternate

or subopposite, petioles 4-12(-17) mm long, 1-1.8 mm

thick, minutely puberulent, with lateral margins only

near the lamina base, terete near the base and rarely

sulcate above; leaf blades (4-)7-15 cm long, 2.5-6(-7)

cm broad, elliptic-oblong, oblong or ovate-oblong, grad-

ually tapering to an acute or acuminate (less often obtuse)

apex, obtuse or acute at the decurrent base, margin entire

and often slightly revolute just above the petiole, drying

stiffly chartaceous and often pale brown or grayish green

above, the upper surface glabrous or sparsely and mi-

nutely puberulent and with a dull flat surface (rarely

slightly lustrous), sericeous beneath in early stages with

thin appressed hairs or very sparsely and minutely pu-

berulent, with (3-)4-6(-7) major secondary veins on each

side, domatia of tufted hairs often present in vein axils

beneath. Inflorescences axillary to distal leaves, 5-12 cm

long, paniculate, few flowered, primary peduncle to 9

cm long and much longer than the flowering rachis, 0.5-

1 mm thick and densely puberulent, flowers often in

umbellate clusters of cymes, pedicels 2-1 1 mm long.

Flowers to 5 mm long and 12(-15) mm broad, white,

minutely puberulent, tepals ca. 5 mm long and 3 mm

broad, minutely papillate on the inner surfaces; outer

stamens with broad subsessile anthers 1-1.6 mm long

and 1.2-1.8 mm broad, the 4 thecae in a horizontal line

or slight arc at the base of the thick distally expanded

connective, small (0.4-0.9 mm) staminodia with obtuse

apices usually present, floral tube shallow (ca. 1 mm);

pistil 1.2-2 mm long, style very short (0.5 mm) or not

well differentiated on the conical apex of the ovary. Fruits

rarely collected, borne in a cup 5-12 mm long, 8-10 mm

broad and 3-6 mm deep, with entire margin; berry 1-

1.5 cm long and 8-10 mm in diameter, ellipsoid.

Trees of evergreen montane forest formations,

between 600 and 1400 m elevation; apparently

confined to the Pacific slope of the central high-

lands and Cordillera de Talamanca. Flowering

material has been collected mostly in February

near San Ramon, but in December-April else-

where. Fruits have been collected in March-April.

The species is known from the area of San Ramon,

Alajuela, a few collections in the central highlands,

the easternmost highlands of Puntarenas Province

and eastward to the province of Code, Panama.

Nectandra ramonensis is characterized by the

relatively small pale brown leaves with dull flat

upper surface, lamina base often revolute, few-

flowered inflorescences on long peduncles, large

flowers, unusual stamens, and restricted habitat.

This species is closely related to TV. globosa, and

appears to be a montane derivative of that species.

Collections that may be intermediate between the

two species have been made near San Ramon; see

the discussion under N. globosa. While a subspe-

cific rank may be suggested for this taxon, some

populations are strikingly distinctive. This is es-

pecially true of the collections made in and around

BURGER: FLORA COSTARICENSIS

63

the Chiriqui highlands, with their short elliptic-

oblong leaves (with domatia beneath) and the ten-

dency for the inflorescences to have an umbel-like

configuration of terminal cymules.

Nectandra reticulata (Ruiz & Pa von) Mez, Jahrb.

Konigl. Bot. Gart. Berlin 5: 404. 1889. Laurus

reticulata Ruiz & Pavon, Laurografia Peru-

viana t. 3, 71802. Ocotea mollis H.B.K., Nov.

Gen. Sp. 2: 164. 1817. N. mollis (H.B.K.) Nees,

Syst. laur. 287. 1836. Figure 7.

Trees to 28(-40) m tall, trunks to 75 cm d.b.h., usually

dark gray and smooth, leafy branchlets 2-8 m thick,

densely puberulent with short (0.1-0.5) brownish hairs.

Leaves alternate and usually in 4 ranks, petioles 8-23

mm long, 2-3.5 mm thick, densely puberulent; leaf blades

10-25(-36) cm l°ng. 4-1 1(-14) cm broad, ovate-lanceo-

late to ovate-elliptic or elliptic-oblong, usually broadest

below the middle and tapering gradually to the long-

acuminate apex, obtuse to acute at the base, margin en-

tire and with a vein along the edge, the margin often

revolute near the petiole with expanded auriculate flaps

4-16 mm long forming enclosed spaces on both sides of

the lamina base beneath, drying stiffly chartaceous, gray-

ish puberulent on the veins above and smooth to the

touch, brownish tomentulose beneath with slender

crooked hairs ca. 0.5 mm long, with 6-16 major sec-

ondary veins on each side, tertiary venation subparallel

and prominent beneath. Inflorescences 6-18 cm long,

axillary to distal leaves, paniculate with few or many

clusters of closely approximate flowers on prominent

lateral branches, peduncle 3-10 cm long, 1.5-2 mm thick,

densely puberulent, lateral branches 3-5 cm long. Flow-

ers 4-8 mm long and 8-12 mm broad, white at anthesis

but becoming reddish brown, tepals 4-5.5 mm long,

densely papillate-puberulent within; outer stamens broad

and flat, 1.5-1.7 mm long, thecae in a plane or in a partly

superposed arc, connective usually developed distally

beyond the thecae, staminodia slender or absent; pistil

2.5-3.5 mm long, stigma discoid. Fruits not seen from

Central America (the following from high altitude col-

lections in the Andes): fruiting receptacles obconic, 6-

12 mm long and 10-16 mm broad, shallow and saucer-

like or deeper (2-4 mm) and more cupulate; berry 10-

20 mm long, ovoid-oblong, abruptly rounded at the apex.

Trees of both evergreen rain forests and ever-

green formations with a pronounced dry season,

on both the Caribbean and Pacific sides of Costa

Rica, from near sea level to 1200 (71600) m ele-

vation. Flowers have been collected in October-

February and April. The species ranges from

southern Mexico through Central America to Peru

and southern Brazil.

Nectandra reticulata is recognized by its nar-

rowly ovate (almost lanceolate) leaves with long-

acuminate apices and dense brownish puberulence

beneath. This species often has flaplike auriculate

developments on the underside of the lamina near

the petiole. These are bent under to form partly

enclosed spaces, but not all leaves or specimens

have this development. The lack of mature fruiting

collections in Central America is unusual for a

species that is fairly often collected. The early stages

of fruit development seem to have the drupe com-

pletely enclosed within the urceolate hypanthium.

Nectandra salicifolia (H.B.K.) Nees, Syst. laur. 302.

1836. Ocotea salicifolia H.B.K., Nov. Gen. Sp.

2: 166. 1817. Figure 17.

Shrubs and small trees 4-12 (rarely to 25) m tall, leafy

branchlets 1.5-4 mm thick, minutely (0.1-0.2 mm) ap-

pressed puberulent with thin ascending hairs, becoming

(sub)glabrous and brownish to gray. Leaves alternate,

petioles 3-8(-14) mm long, ca. 1 mm thick, usually sul-

cate above; leaf blades 6-15 cm long, 1.5-6 cm broad,

narrowly elliptic, elliptic-lanceolate or elliptic-oblong to

ovate-elliptic, tapering gradually or abruptly to an acu-

minate (rarely acute) apex, the tip 4-16 mm long, acute

to obtuse at the base and slightly decurrent on the petiole,

margin entire, drying stiffly chartaceous to subcoriaceous

and often darker above than below when dried, glabrous

above and usually lustrous, glabrous beneath except for

the tufted hairs (domatia) in the axils of proximal sec-

ondary veins in some collections, with 4-7 major sec-

ondary veins on each side, the secondaries usually weak-

ly loop-connected near the margin in the distal half of

the lamina, minor venation raised on both surfaces when

dry. Inflorescences solitary in the axils of distal leaves

or pseudoterminal (sometimes appearing to be several

in the axil due to a very short peduncle), broadly pani-

culate and many-flowered or rarely racemose to umbel-

late in few-flowered panicles, 4-ll(-17) cm long, pri-

mary peduncle (2-)6-60 mm long, reddish to whitish

and sparsely to densely puberulent with slender minute

(0. 1 mm) hairs, pedicels 2-3(-4) mm long. Flowers white,

3-4 mm long, 5-6(-8) mm broad, tepals 2.5-3.8 mm

long and 1.2-1.7 mm broad, spreading or reflexed at

anthesis, puberulent on the outside and papillate within;

outer stamens subsessile or the filaments to 0.3 mm long,

outer anthers 0.4-0.6 mm long, 0.5-0.7 mm broad, rect-

angular to reniform and convex to emarginate distally,

the thecae in a single plane or the lower lateral to the

upper in an arc, inner stamens ca. 0.8 mm long and with

large glands, staminodes 0.4-0.8 mm long and slender

or slightly thickened apically (sometimes absent or mi-

nute); pistil 1-1 .7 mm long, ovary 0.6-1 mm in diameter

(rarely puberulent), style 0.3-0.6 mm long, stigma dis-

coid or simple. Fruits borne on a short-conical or saucer-

like receptacle 3-5 mm long, (5-)8-10 mm broad and

1-3 mm deep, the margin entire or slightly undulate,

becoming reddish or purple; berry 1-1.6 cm long and 1-

1.5 cm in diameter, subglobose to ellipsoid-oblong, be-

coming black (red?).

Shrubs and small trees of evergreen and partly

deciduous forest formations, between sea level and

64

FIELDIANA: BOTANY

600 (900) m elevation in the Caribbean lowlands,

the General Valley and the Golfo Dulce region in

Costa Rica. Flowering material has been collected

in January-August in Costa Rica and January-

June in Guatemala; mature fruits have been col-

lected in August-November in Central America.

This species (in a wide sense) ranges from north-

eastern Mexico through Central America to Costa

Rica and Panama.

Nectandra salicifolia is recognized by the rela-

tively small, stiff, essentially glabrous leaves with

the minor venation raised on both surfaces and

the upper surface usually lustrous when dried. The

usual presence of tufted domatia and the weakly

loop-connected secondary veins are associated with

small puberulent flowers with very small anthers,

short styles, small shallow fruiting receptacles, and

usually subglobose fruits. These characters vary

greatly and can make some populations appear

quite different from others. Collections from

southwestern Costa Rica tend to have more cau-

date-acuminate leaves with conspicuous "drip

tips," while material from the Caribbean slope is

often narrowly elliptic. Nectandra salicina appears

to be a highland derivative of N. salicifolia, and

there appear to be collections intermediate be-

tween the two in the Cordillera de Guanacaste; see

the discussion under N. salicina. Nectandra sa-

vannarum (Standl. & Steyerm.) C. K. Allen of

Guatemala and Belize is probably an unusual form

of N. salicifolia with small umbellate inflores-

cences and small ovate leaves with prominent bas-

al secondary veins. Trees referred to as N. salici-

folia in Paul Allen's book (1956) are actually N.

turbacensis. Some material placed here was earlier

identified as N. coriacea (Sw.) Griseb., which is

probably restricted to the West Indies and Yucatan

peninsula (Rohwer, pers. comm.).

Nectandra salicina C. K. Allen, J. Arnold Arbor.

26: 385. 1945. N. smithii C. K. Allen, loc. cit.

370. 1945. Figure 4.

Trees 5-10 m tall, leafy branchlets l-4(-5) mm thick,

at first minutely puberulent with appressed yellowish

brown hairs, becoming glabrous, smooth, and grayish.

Leaves alternate and often densely clustered at the ends

of branches, petioles 4-12 mm long, ca. 1 mm thick, flat

or slightly sulcate above and with lateral margins; leaf

blades 5-9(-14) cm long, l-3(-4) cm broad, narrowly

elliptic to lanceolate or elliptic-oblong, tapering gradu-

ally to the acuminate apex, the narrowed tip 0.5-2 cm

long, tapering gradually to the attenuate base, margin

entire or undulate, drying stiffly chartaceous or subcor-

iaceous and often greenish brown, glabrous and lustrous

above with the tertiary veins slightly raised, glabrous or

very sparsely puberulent beneath, rarely with tufts of

hairs (domatia) in the vein axils, with (3-)4-6(-7) major

secondary veins on each side, the tertiary venation al-

ways forming a raised but irregular reticulum on the

upper surface of the dried leaf. Inflorescences axillary or

extra-axillary (nodes with undeveloped leaves?), to 12

cm long, paniculate but with few lateral branches, pe-

duncles (2-)4-8 cm long, minutely puberulent, pedicels

ca. 4 mm long and articulate in the middle. Flowers ca.

3.5 mm long and 6-7 mm broad, yellowish white, dense-

ly and minutely puberulent, tepals 2.5-3 mm long, densely

papillate-puberulent within; androecium ca. 3 mm in

diameter, outer stamens with short (0.2 mm) filaments,

outer anthers 0.5-0.8 mm long and 0.7-1 mm broad,

thecae usually in an arc with the lower lateral to the

upper, inner stamens narrow, staminodes to 1 mm long

or absent; pistil 1-1.8 mm long, ovary globose, 0.7-1.1

mm in diameter, style narrow, stigma discoid or simple.

Fruits borne in a conical or broadly campanulate and

very shallow (1-2 mm) cup, 5-10 mm long above the

thickened pedicel and 4-1 1 mm broad at the top, margin

entire or undulate, becoming red; berry 10-18 mm long

and 7-16 mm thick (dry), ellipsoid during development

but becoming globose at full maturity.

Trees of evergreen forests on both the Caribbean

and Pacific slopes, from about 600 to 1700 m

elevation. Flowering collections have been made

in January-May and July-August; fruits have been

collected in March-November. The species, as here

defined, ranges from the westernmost parts of the

Cordillera de Guanacaste through the Cordillera

de Tilaran to the eastern part of the central vol-

canic highlands (near Turrialba, Cartage).

Nectandra salicina is recognized by its small

lustrous narrowly elliptic and acuminate leaves

with the tertiary venation usually prominent on

both surfaces. The small few-flowered inflores-

cences with reddish rachises and puberulent little

flowers, and the ellipsoid to globose fruits borne

in shallow cups also distinguish this species. The

wood and leaves are said to be sweet-smelling.

Nectandra salicina appears to intergrade with very

similar smaller-leaved specimens of N. salicifolia

at lower elevations, and it would appear that it

should really be called a subspecies of N. salici-

folia. However, N. salicifolia encompasses much

geographic variation in the lowlands, and its re-

lationship with material called N. latifolia is not

clear. Until these matters are clarified, we prefer

to think of these taxa as species; see the discussion

under N. latifolia. Specimens intermediate be-

tween typical N. salicina and N. salicifolia have

been collected at lower elevations (about 400 to

900 m) in the Cordillera de Guanacaste, but they

lack fruits, which are generally larger in N. salicina.

Nectandra smithii was based on Austin Smith

BURGER: FLORA COSTARICENSIS

65

//. 541 (AA,F) from near Zarcero at 1600 m ele-

vation. The leaves resemble those of some collec-

tions ofN. salicifolia but are more elliptic-oblong,

and a few of the leaves have distinctive slightly

rounded caudate-acuminate tips. Brenes 6825 (F)

was cited in the description ofN. smithiiand seems

to be intermediate between the type collection and

typical material of N. salicina; it is for this reason

that N. smithii appears to be an unusual variant

of N. salicina. Rohwer (p. 81, 1986) points out

that the flowers have some Ocotea-like character-

istics, both as regards pubescence and thecae ar-

rangement in the outer anthers. This suggests that

the type may be of hybrid origin; however, these

characteristics are also sometimes seen in N. sal-

icina.

Nectandra sinuata Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 402. 1889. Figure 7.

Small or medium-sized trees to 20 m tall, trunk to 1

m d.b.h., leafy branchlets 2-8 mm thick, densely villous-

tomentulose with slender yellowish brown or grayish

hairs ca. 0.5 mm long. Leaves alternate, petioles 8-32(-40)

mm long, 1-4 mm thick, densely puberulent and ob-

scurely sulcate above; leaf blades 10-22(-30) cm long,

3.5-1 2(-l 9) cm broad, oblong-elliptic to slightly obovate

or oblong-obovate, usually abruptly narrowed to the ob-

tuse or short-acuminate apex, usually abruptly narrowed

to the obtuse or truncated base (rarely acute), the larger

leaves often rounded and subcordate at the base, laminae

drying stiffly chartaceous, sparsely puberulent above and

densely tomentulose to sericeous beneath with slender

straight or curved hairs 0.3-1 mm long, with 10-12 ma-

jor secondary veins on each side, the central secondaries

arising at angles of 40°-75°. Inflorescences axillary to

distal leaves, to 30 cm long, open panicles puberulent in

all parts, peduncle to 1 6 cm long (often more than half

the length of the inflorescence), ca. 2 mm thick, pedicels

5-12 mm long. Flowers ca. 10 mm long, 10-20 mm

broad, tepals 5-8 mm long and ca. 4 mm broad (dry),

pale rose, pink or white (reddish brown when dry), dense-

ly tomentulose on the outside and papillate-puberulent

within; androecium ca. 5 mm in diameter, outer anthers

2-3 mm long on short (0.5 mm) thick filaments, con-

nective often expanded and the stamen tepal-like in form,

thecae in an arc or sometimes superposed, staminodes

absent; pistil 2-3 mm long, style ca. 1 mm long with

lobed disclike stigma. Fruit cups ca. 20 mm long and 16

mm broad, entire or lobed at the margin, ca. 5 mm deep;

berry 2-2.5 cm long and 1-1.8 cm thick, ellipsoid-ob-

long, sparsely to densely sericeous in the lower fourth,

glabrous above except at the apex, becoming black.

Trees of evergreen and partly deciduous forest

formations of the central highlands and Pacific

slope in Costa Rica, between 300 and 1500 m

elevation. (A few collections have been made on

the Caribbean slope near Turrialba, but these may

have been planted.) Flowering collections have

been made in January-April in Costa Rica, and

fruits have been collected in April-June. The

species ranges from southern Mexico along the

Pacific side of Central America to the western part

of the Cordillera de Talamanca (near Sta. Maria

de Dota).

Nectandra sinuata is distinguished by the dense

puberulence of its long-peduncled inflorescences

and lower leaf surfaces, the large densely puber-

ulent flowers, and the ellipsoid fruits with ap-

pressed hairs near the base. The oblong leaves

abruptly narrowed at both apex and base, pref-

erence for a seasonally dry habitat, and frequent

pink flowers further characterize this species. The

tepal-like anthers are unusual and sometimes have

the valves in a superposed Ocotea-like arrange-

ment. Rohwer (pers. comm.) has suggested trans-

ferring this species to Ocotea.

Nectandra turbacensis (H.B.K.) Nees, Hufeland.

ill. 14. 1833; Syst. laur. 316. 1836. Ocotea tur-

bacensis H.B.K., Nov. Gen. Sp. 2: 162. 1817.

AT. concinna Nees, Syst. laur. 322. 1836. N. ner-

vosa Mez & J. D. Smith ex Mez, Bull. Herb.

Boissier, ser. 2, 3: 235. 1903. Figure 18.

Trees to 20(30) m tall, to 80 cm d.b.h., leafy branchlets

1.5-5 mm thick, glabrous or sparsely puberulent with

thin minute (0.2 mm) appressed ascending hairs, becom-

ing dark and smooth. Leaves alternate, petioles 5-12 mm

long, 0.7-1.2 mm thick, usually glabrous and dark, flat

or slightly sulcate between the adaxial ridges; leaf blades

5-17 cm long, 2-4.5(-6) cm broad, lanceolate to nar-

rowly ovate or narrowly oblong, tapering gradually to

the acute or acuminate apex, acute (rarely obtuse) at the

base, margin entire or undulate, stiffly chartaceous and

often grayish green above, glabrous above and below,

the midvein and minor venation slightly raised above,

with (3-)5-7 major secondary veins on each side (rarely

with a basal pair of secondaries strongly ascending and

almost tripliveined, as in Allen 5710), ciliate pit domatia

often present in proximal vein axils beneath. Inflores-

cences in the axils of distal leaves or pseudoterminal, to

1 7 cm long, paniculate with spreading lateral branches,

peduncle (0.5-)2-6 cm long, 0.5-1.5 mm thick, glabrous

(rarely sparsely and minutely puberulent), pedicels 1-4

mm long. Flowers ca. 3 mm long and 5-8 mm broad,

white, sparsely puberulent on the outside, tepals 2.2-3

mm long, 2-2.6 mm broad, densely papillate-puberulent

within; androecium 3 mm in diameter, outer stamens

subsessile, outer anthers 0.6-0.7 mm long, 0.8-1.1 mm

broad, thecae in a horizontal row, the connective slightly

expanded distally and papillate, inner stamens 0.9-1.4

mm long, staminodes minute or absent; pistil 1.5-2 mm

long, ovary 1-1.4 mm in diameter, style short (ca. 0.8

mm) and thick, stigma discoid. Fruits borne on a short

(4-10 mm) cupulate receptacle 8-12 mm broad at the

apex and (2-)3-4 mm deep, margin entire and often with

66

FIELDIANA: BOTANY

2 concentric ridges, greenish or bluish; berry 10-14 mm

long and 7-10 mm thick (dry), oblong-ellipsoid, becom-

ing black.

Trees usually found in secondary associations

in evergreen and partly deciduous forest forma-

tions, from near sea level to 1200(1700) m. Flow-

ers have been collected in September-March (May)

in Costa Rica; fruiting is said to occur in March-

April in southwestern Costa Rica (P. H. Allen,

1956). The species ranges from Veracruz, Mexico,

to Colombia, Venezuela, Peru, and Bolivia.

Nectandra turbacensis is recognized by the gla-

brous leaves that dry dull grayish with smooth

surfaces and obscure minor venation (as in the

type) or with the minor venation slightly raised

on a darker background (as in the populations of

southwestern Costa Rica). The outer subsessile an-

thers with their slightly prolonged (but truncated

or obtuse) and papillate connective and their the-

cae in a single plane are distinctive. The entire

fruit cup enclosing about '/4 of the fruit and a

general lack of puberulence (except for distal parts

of the inflorescence and perianth) further distin-

guish this species. The veins are not loop-con-

nected near the margin, and the flowers and sta-

mens are larger than in N. salicifolia and its allies.

Trees with long narrow leaves and prominent

minor venation have been called N. panamensis

Mez, and are characteristic of southwestern Costa

Rica and western Panama in the Pacific lowlands.

It may be that the populations of the Pacific low-

lands deserve subspecific recognition. The species

is rarely collected in Central America and appears

to be restricted to the Caribbean lowlands in

northern Central America. A population with ab-

normal flowers was sampled by Standley and Val-

erio (45469, 45906, 45909, 46192, all us) near

Tilaran in 1926.

Ocotea Aublet

REFERENCE— J. G. Rohwer, Prodromus einer

Monographic der Gattung Ocotea Aubl. (Laura-

ceae), sensu lato. Mitt. Inst. Allg. Bot. Hamburg

20: 1-278. 1986.

Trees or shrubs, bisexual or rarely unisexual (dioe-

cious), glabrous to densely puberulent. Leaves alternate,

rarely subopposite or verticellate, petiolate or sessile but

narrowed near the base, the leaf blades entire, charta-

ceous to coriaceous when dry, with pinnate venation

(rarely with strongly developed basal secondaries and

subtripliveined), with or without domatia. Inflores-

cences usually solitary and axillary to distal leaves, some-

times pseudoterminal, lacking a subtending involucre of

bracts, paniculate or occasionally few-flowered and ra-

cemose, pedicels usually well developed, bracts and brac-

teoles usually deciduous. Flowers bisexual or unisexual,

(male flowers usually with a pistillode and female flowers

with staminodes when unisexual), generally small (2-5

mm long), white to yellowish or green, the floral tube

(hypanthium) well developed, small or absent but always

accrescent in fruits, perianth 6-parted with 2 whorls of

3 tepals each, imbricate, equal or subequal, the tepals

thick to thin and glabrous to densely puberulent; fertile

stamens 9 in 3 series, the outer 2 series (often appearing

as a single whorl) similar in size and shape and with

introrse dehiscence, filaments present or absent, anthers

variously shaped but usually as broad as long or longer

than broad (flat and triangular to laminar or tepal-like

in some species), the connective rarely prolonged beyond

the thecae, each anther with 4 thecae arranged in 2 planes

(superposed) or with the lower thecae slightly lateral to

the upper (this character can vary within the same flower

in a few species and makes differentiation from Nectan-

dra difficult), opening by flaps attached at the top, the

inner 3 stamens also 4-thecous, usually with longer fil-

aments and narrower anthers that open extrorsely or with

the upper thecae opening laterally and the lower extrorse,

each inner stamen with 2 conspicuous sessile (rarely stip-

itate) glands at the base of the filament (6 glands per

flower), the glands thick and bluntly rounded or angular,

an inner whorl of staminodes (series IV) usually absent

or poorly developed (less than 0.6 mm long); pistil with

globose, ovoid, or turbinate ovary, usually slightly nar-

rowed at the base, rarely puberulent, style slender or

thick, stigma simple to discoid or capitate (rarely 2- or

3-lobed). Fruits borne on a flattened discoid to cupulate

or hemispheric receptacle, perianth usually deciduous

and the cup with a single-margined edge (with a double

margined rim in O. dendrodaphne, O. veraguensis and

some flat disclike receptacles), the margin sometimes

undulate (rarely 6-lobed), pedicel thickened in fruits and

erect or pendulous; fruits a single-seeded berry, ovoid to

ellipsoid, oblong or globose, usually narrowed and slight-

ly flattened at the base, abruptly rounded at the apex,

style or stigma rarely persisting.

A genus of perhaps 300 to 400 species centered

in the American tropics, with a few species in Af-

rica and a group of species in Madagascar. In Costa

Rica there appear to be a few species intermediate

between Ocotea and Nectandra, and between Oco-

tea and Phoebe. The usual lack of three well-de-

veloped staminodes help distinguish Ocotea from

our species of Phoebe. The character of superposed

(2-ranked) thecae valves, which separates Ocotea

from Nectandra, displays considerable variation

with some intermediate states among a few species.

While it is easy to say that these relatively rare

intermediate states invalidate current generic con-

cepts, there does not seem to be any way of de-

veloping better generic concepts at this time.

Lumping these genera together produces nomen-

clatural confusion with no increased understand-

BURGER: FLORA COSTARICENSIS

67

ing of how the species should be arranged. The

present grouping of species in Ocotea appears to

form a central assemblage from which other lin-

eages may have been derived (cf. Rohwer & Ku-

bitzki, 1985; Rohwer, 1986). For example, one

population of O. insularis appears to be in the

process of losing the upper thecae in each anther,

and this may have been how Aiouea costaricensis

originated (see the discussion under these species

and under Aiouea, Nectandra, and Phoebe).

Ocotea is recognized by the 6-parted flowers with

equal or subequal tepals, the nine stamens with

four superposed valves in each, the three inner

stamens with two conspicuous glands, the absence

or poor development of staminodes, and the re-

ceptacle forming a cup or disc in fruits. There is

great morphological variation within our species.

The stamens range from sessile lamina-like struc-

tures to those with well-developed slender fila-

ments and clearly differentiated anthers. Compar-

ison with other genera and with related species

within Ocotea suggests that laminar stamens are

not a primitive condition in Lauraceae (cf. Rohwer

& Kubitzki, 1985). Likewise, the fruiting recep-

tacle ranges from a small disc to a deep hemi-

spheric cup.

Key to Species of Ocotea in Costa Rica

la. Distal branchlets hollow and often harboring small ants; leaves usually more than 1 5 cm long and

narrowly oblong to elliptic-oblong or narrowly obovate, always glabrous beneath; small trees of

wet evergreen forest understory 2a

1 b. Distal branchlets solid, the center with wood or pith and not consistently hollow 6a

2a. Flowers 4-6 mm long; anthers 2-3 mm long, flat and tepal-like; fruit cups ca. 1 5 mm broad

and 3-7 mm deep; leaves drying grayish green and subcoriaceous, usually elliptic-oblong; 0-

1 500 m elevation O. dendrodaphne

2b. Flowers 2-3 mm long; anthers to 1 mm long and not tepal-like; fruit cups 6-12 mm broad

and only 1-2 mm deep 3a

3a. Leaves usually drying chartaceous and usually very dark in color; 0-1000 m elevation . . .

4a

3b. Leaves usually drying subcoriaceous and grayish green to orange brown or dull brown ....

5a

4a. Leaves becoming 30-50 cm long, often narrowly obovate; flowers essentially glabrous,

outer filaments glabrous; fruits 20-35 mm long O. atirrensis

4b. Leaves not exceeding 30 cm in length, usually narrowly elliptic-oblong; flowers minutely

puberulent, filaments minutely puberulent; fruits 1 2-20 mm long O. wedeliana

5a. Distal stems strongly angled with 3-5 prominent longitudinal ridges; leaves to 40(-55) cm

long, often narrowly obovate; fruit cups often with persisting perianth bases; 0-1 100 m . .

O. nicaraguensis

5b. Distal stems without prominent ridges, not strongly angled in cross section; leaves 10-30 cm

long, often narrowly oblong; fruit cups with entire margins; 600-2300 m elevation

O. paulii

6a. Leaves usually becoming more than 30 cm long, obovate and gradually narrowed to the base,

drying stiffly chartaceous 7a

6b. Leaves rarely more than 30 cm in length (sprout shoots from nonflowering basal branches may

be larger) 9a

7a. Leaves to 50 cm long and 25 cm broad, glabrous beneath; flowers 3 mm long and 3 mm

broad, outer anthers ca. 0.6 mm long; Golfo Dulce area O. rivularis

7b. Leaves to 40 cm long, conspicuously puberulent beneath; flowers 5-7 mm long and 6-10

mm broad; Caribbean slope and lowlands 8a

8a. Leaves often narrowly obovate, to 1 6 cm broad; inflorescence rachis sparsely hirsute; outer

anthers 1 .2-1 .4 mm long; ca. 1 400 m elevation O. lentil

8b. Leaves often broadly obovate, to 22 cm broad; inflorescence rachis densely yellowish pu-

berulent; outer anthers 1-1 .2 mm long; 0-900 m O. valerioides

68

FIELDIANA: BOTANY

9a. Leaves densely puberulent beneath with longer (0.3-1 mm) hairs, the hairs spreading or appressed,

puberulence on the lower surface soft or slightly rough to the touch 1 Oa

9b. Leaves glabrous to sparsely puberulent beneath (lower surface), the hairs small (0.05-0.2 mm) or

inconspicuous and difficult to see with a 1 OX hand lens, puberulence of the lower leaf surface not

discernable to the touch 25a

lOa. Lower leaf surface sericeous with lustrous silvery or reddish hairs, the leaves drying stiffly

coriaceous and decurrent on a very short petiole; flowers 4-7 mm long and densely puberulent;

only from above 2000 m O. calophylla

lOb. Lower leaf surfaces not densely sericeous with lustrous silvery or reddish hairs; petioles well

developed 1 la

1 la. Largest leaves rarely more than 1 0 cm long 1 2a

1 Ib. Largest leaves usually more than 1 3 cm long 1 5a

12a. Flowers glabrous to sparsely puberulent, often with well-developed staminodes; lamina

base not decurrent on the petiole, leaves usually drying chartaceous 1 3a

12b. Flowers minutely and densely puberulent, staminodes absent; lamina base decurrent

on the petiole, leaves often drying subcoriaceous 1 4a

1 3a. Fruit cups 10-1 5 mm broad; pubescence on the lower leaf surfaces brownish and

slightly rough to the touch; 1000-3200 m elevation O. pittieri

1 3b. Fruit cups ca. 6 mm broad; pubescence on the lower leaf surfaces grayish and

soft to the touch; 1400-2300 m elevation O. mollicella

14a. Leaves usually elliptic and 1.5-4 cm broad, with an acute apex; flowers 3—4 mm long,

outer anthers 0.6-0.9 mm long; western highlands, 1400-1600 m . O. monteverdensis

14b. Leaves usually oblong and 1.5-6 cm broad, with an obtuse apex; flowers 4-5 mm long,

outer anthers ca. 1 mm long; central and eastern highlands, 1 700-3000 m elevation

O. austinii

1 5a. Base of the leaf blades decurrent on the petiole and the petiole often poorly denned, margin

of the lamina often revolute near the petiole 1 6a

1 5b. Base of the leaf blade not consistently decurrent on the petiole 1 9a

16a. Leaves usually narrowly elliptic to oblong, 1.5-6 cm broad; montane species rarely

encountered below 1 200 m elevation 1 4a

16b. Leaves often obovate or slightly obovate, 3-1 1 cm broad; rarely found above 1 100 m

17a

17a. Petioles clearly differentiated from the narrowed lamina base, leaves drying stiffly

chartaceous, with 4-8 pairs of major secondary veins; fruit cups 10-16 mm broad . .

O. hartshorniana

17b. Petioles not clearly differentiated from the narrowed lamina base, leaves drying sub-

coriaceous, with 9-12 pairs of major secondary veins; fruit cups 5-12 mm broad . . .

18a

1 8a. Fruit cups flat and saucer-like, without dentate lobes O. stenoneura

1 8b. Fruit cups deeply cupulate and with persisting perianth lobes O. dentata

19a. Leaves broadly obovate to broadly elliptic, usually rounded or abruptly narrowed at the

apex, usually drying dark brown above, densely puberulent beneath; outer anthers 0.7-1.5

mm long; Caribbean slope and central Cordilleras 20a

1 9b. Leaves rarely obovate or broadly elliptic, not rounded at the apex, rarely drying dark brown,

sparsely puberulent beneath 25a

20a. Trees of montane formations, (8 00-) 1000-2 5 00 m elevation; leaves often drying sub-

coriaceous, oblong to broadly elliptic, obovate or suborbicular; flowers 6-1 5 mm broad

21a

20b. Trees not known from above 800 m elevation in Costa Rica; laminae usually drying

stiffly chartaceous, usually obovate to oblong or pandurate; flowers 4-12 mm broad

23a

2 la. Flowers glabrous on the outside, outer anthers subsessile or with very short

filaments; fruits borne in a deep (5-8 mm) cup 10-18 mm broad . O. valeriana

BURGER: FLORA COSTARICENSIS 69

2 1 b. Flowers densely puberulent on the outside, outer anthers with prominent filaments

22a

22a. Leaves usually elliptic to oblong, 3-9 cm broad; flowers 6-8 mm broad; fruits in

a shallow entire cup; 1 500-2500 m elevation O. pseudopalmana

22b. Leaves usually broadly elliptic to suborbicular, 6-14 cm broad; flowers ca. 12

mm broad; fruits in a deep lobed cup; 800-1400 m O. gomezii

23a. Leaves 10-22 cm broad, obovate to pandurate, on short thick (3-6 mm) petioles;

inflorescences apparently erect; perianth sparsely puberulent O. valerioides

23b. Leaves 4-14 cm broad, petioles rarely more than 3 mm thick 24a

24a. Leaves often obovate to pandurate; inflorescences pendant with long thin peduncles,

flowers often borne on umbellate lateral branches; perianth usually glabrous on the

outside and drying dark O. helicterifolia

24b. Leaves obovate to broadly oblong; inflorescences usually erect and broadly paniculate

with racemose or branched lateral branches; perianth puberulent on the outside and

drying brownish O. mollifolia

25a. Distal branchlets strongly winged (alate) with narrow longitudinal wings 2-3 mm high, the stems

3-5-angled in cross section; leaves narrowly oblong to narrowly obovate and coriaceous, with 9-

14 pairs of major secondary veins; Pacific slope below 1000 m elevation O. aurantiodora

25b. Distal branches without conspicuous longitudinal wings, the stems sometimes bluntly angled in

cross section but not with 2-3 mm high ridges 26a

26a. Leaf base usually decurrent on the petiole, the leaf base and petiole often poorly differentiated,

lamina margin often revolute near the petiole, the leaves petiolate to sessile 27a

26b. Leaf base not usually decurrent on the petiole, lamina acute to obtuse or rounded at the base and

usually clearly differentiated from the petioles, the leaves never sessile or subsessile 36a

27a. Leaf base broadly revolute (auriculate) and forming 2 broad flaps on the underside, petiole

little developed and the leaves subsessile, broadly obovate and often coriaceous; (700-) 1 1 00-

2300 m elevation O. endresiana

27b. Leaf base flat or revolute but not forming broad flaps beneath, never auriculate and usually

petiolate 28a

28a. Mature leaves usually with slender appressed ascending hairs parallel with the secondary

veins over the lower surface; trees often with prop roots at the base of the trunk 29a

28b. Mature leaves glabrous or variously puberulent but not with slender appressed hairs paral-

leling the secondary veins on the lower surfaces; trees without prop roots(?) 3 la

29a. Leaves elliptic to oblanceolate or obovate, 2-5 cm broad, leaf base usually long-de-

current (to 5 cm long), with 5-9 pairs of major secondary veins; fruits ellipsoid; Pacific

slope at 600-1400 m (if higher elevation cf. O. whitei) O. skutchii

29b. Leaves broadly obovate, 4.5-1 1 cm broad, long- or short-decurrent at the base, with

7-12 pairs of major secondary veins; fruits globose to ovoid and 2-4 cm long . . 30a

30a. Fruits ellipsoid to ovoid, 3-4 cm long in large deep ( 1 2 mm) cups; leaves drying smooth

and dull above with the minor venation obscure; Caribbean lowlands

O. sp. aff. caracasana

30b. Fruits globose to oblong, 2-3 cm long, often in conical cups with reflexed edges; leaves

drying with a lustrous sheen and the minor venation raised above; 1500-2500 m

elevation O. glaucosericea

3 1 a. Leaves often rounded at the apex or bluntly obtuse (rarely acute to short-acuminate in smaller

leaves), often obovate, (2-)5-9(-l 2) cm broad, drying coriaceous or subcoriaceous and often

yellowish brown to pale reddish brown; fruit cups 6-9 mm broad and often with persisting

perianth bases on the edge, fruits ellipsoid, 1-2 cm long; 0-2000 m O. insularis

3 Ib. Leaves bluntly acute to acuminate at the apex, sometimes bluntly obtuse but never rounded,

often oblanceolate to narrowly elliptic-obovate or oblong, 1.5-5(-6) cm broad; fruit cups

without persisting perianth 32a

32a. Flowers 2-5 mm long, functionally unisexual; fruits subglobose, 6-16 mm in diameter; leaves

drying grayish green or yellowish green, elliptic-oblong to obovate 33a

32b. Flowers 1.5-3 mm long, bisexual; fruits globose or oblong, but never with a persisting style;

70 FIELDIANA: BOTANY

leaves often drying very dark in Costa Rica but variable in O. whitei, often oblanceolate to

very narrowly elliptical 34a

33a. Fruits with a persisting style base at the top and borne on a thick flat double-rimmed

disclike receptacle 6-10 mm broad, berry 10-16 mm in diameter; leaves often obovate

and drying grayish; 0-1 900 m O. floribunda

33b. Fruits without a persisting style base, receptacle 4-8 mm broad, berry 6-8 mm in

diameter (dried); leaves often elliptic-oblong and drying yellowish; 0-500 m

O. puberula

34a. Fruits 1 5-38 mm long and 11-18 mm thick (dry), ellipsoid and borne in cupulate receptacles

8-14 mm broad; leaves often narrowly elliptic; 1300-2500 m elevation O. whitei

34b. Fruits rarely exceeding 1 5 mm in length, globose to oblong, subtended by shallow receptacles

4-6 mm broad; leaves usually oblanceolate to narrowly elliptic-obovate 35a

35a. Flowers 2-3 mm long, outer anthers ca. 0.6 mm long; fruits globose, 5-8 mm in diameter

(dry); leaves without domatia; Caribbean lowlands O. sp. aff. O. bijuga

35b. Flowers 1.5-2.5 mm long, outer anthers ca. 1 mm long; fruits oblong, 9-18 mm long and

7-9 mm in diameter; pit domatia often present on leaf undersurfaces; (0-)600-1000 m . . .

O. oblonga

36a. Leaves drying very dark (almost black) and thin-chartaceous; fruiting receptacle gradually ex-

panding (obconic) to the 1 cm broad apex, concave or slightly cupulate, fruits 2-3 cm long and 1-

2 cm thick; flowers glabrous and small 37a

36b. Leaves drying grayish to dark brown, stiffly chartaceous to coriaceous if drying very dark . . 38a

37a. Flowers 1.5-3 mm long, outer anthers ca. 0.7 mm long and the connective not expanded;

leaves elliptic to elliptic-oblong, with 4-7 pairs of major secondary veins, glabrous; 0-1700

m elevation O. tenera

37b. Flowers 3-5 mm long, outer anthers ca. 1 mm long and with the connective expanded distally;

leaves ovate to broadly elliptic, with 3-5 pairs of major secondary veins, sparsely puberulent;

900-2000 m O. brenesii

38a. Minor venation raised on the lower leaf surface and forming a reticulum of small areolae 0.3-1

mm broad, the areolae often with subequal sides; the leaves often drying yellowish green or grayish

green, glabrous or glabrescent on the lower surface 39a

38b. Minor venation not raised and not forming a definite reticulum on the lower (abaxial) leaf surfaces,

areolae very irregular if present; glabrous to puberulent beneath 43a

39a. Leaves gradually narrowed to the base, usually narrowly obovate and drying grayish green,

domatia lacking; fruits globose with short persisting style at the apex and subtended by a

small (6-10 mm) flat disclike receptacle; 0-1900 m elevation O. floribunda

39b. Leaves abruptly narrowed at the base (obtuse to acute), pit domatia present; fruiting recep-

tacles cupulate, 8-14 mm broad and 2-5 mm deep; fruits mostly ellipsoid, 2-3 cm long and

the style not persisting 40a

40a. Leaves broadly obovate and large (14-30 cm x 7-18 cm), rounded to bluntly obtuse at the

apex, drying yellowish brown and subcoriaceous; fruit cup with an irregular margin; flowers

unknown; 500-1000 m elevation O. sp. A aff. O. laetevirens

40b. Leaves never broadly obovate and not more than 10 cm broad, usually acuminate at the

apex; flowers 2-4 mm long, anthers 0.7-1 mm long; fruit cups with an entire margin . 41a

4 la. Leaves usually drying yellowish green and often subcoriaceous, often broadly elliptic, retic-

ulum of minor venation poorly defined; fruit cup 10-16 mm broad, 3-5 mm deep

O. laetevirens

41b. Leaves usually drying stiffly chartaceous; fruit cups only 2-3 mm deep 42a

42a. Leaves usually drying grayish green and with a reticulum of small (0.5 mm) areolae beneath;

fruit cup 8-14 mm broad with flaring edges; inflorescences usually paniculate; 0-2100 m

elevation O. meziana

42b. Leaves usually drying yellowish brown and lustrous above, reticulation of veins not forming

small areolae beneath; fruit cup ca. 10 mm broad and with persisting lobes; inflorescences

often umbellate or racemose; Pacific Highlands, 1 600-2000 m O. viridiflora

BURGER: FLORA COSTARICENSIS 71

43a. Flowers to 7 mm long and 5-10 mm broad, outer anthers 1.5-2.5 mm long and usually flattened

44a

43b. Flowers to 5 mm long, outer anthers less than 1 mm long; evergreen forests, 0-1800 m elevation

45a

44a. Flowers minutely puberulent; leaves drying grayish, with 5-8 pairs of major secondary veins;

deciduous and partly deciduous forest formations, 0-1600 m O. veraguensis

44b. Flowers glabrous; leaves drying dark brown with 3-6 pairs of major secondary veins; ever-

green forests, 1 600-2200 m O. holdridgeiana

45a. Flowers 2-2.5 mm long, glabrous and drying black; leaves usually oblong and caudate-acuminate

at the apex, drying grayish and chartaceous, glabrous or very sparsely puberulent; fruit cups ca. 1

cm broad; 0-300(-800) m elevation O. cernua

45b. Flowers 2-5 mm long and minutely puberulent; leaves never caudate-acuminate, often drying

grayish green and subcoriaceous; fruit cups 5-8 mm broad, dark and with pale lenticels; 0-1800

m 46a

46a. Leaves usually glabrous beneath, the major veins not impressed above, often elliptic-oblong; distal

branchlets strongly 2- or 3-angled in cross section; flowers unisexual O. leucoxylon

46b. Leaves usually puberulent beneath, the major veins impressed above and the leaves somewhat

bullate, usually broadly elliptic to slightly obovate; distal branchlets with longitudinal ridges but

not strongly angled in cross section; flowers not known in Costa Rica O. babosa

Ocotea atirrensis Mez & J. D. Smith, Bot. Jahrb.

Syst. 30, Beibl. 67: 1 8. 1 90 1 . O. pedalifolia Mez,

loc. cit. 19. 1901. Figure 2.

Shrubs or small and slender trees to 5(-8) m tall, leafy

branchlets 3-6 mm thick, glabrous and smooth, strongly

ridged or terete, distal stems hollow and often inhabited

by small ants. Leaves alternate, petioles 6-18 mm long,

1.5-3.5 mm thick, with adaxial margins forming a deep

sulcus above and partly overlapping; leaf blades 12-

35(-55) cm long, 5-1 3(-22) cm broad, narrowly obovate

to oblong-obovate or oblanceolate, usually abruptly nar-

rowed and caudate-acuminate at the apex, the tip often

1-3 cm long and 2-4 mm broad, gradually narrowed to

the acute or obtuse base, margin entire and slightly un-

dulate, drying chartaceous and dark gray, glabrous above

and below, with 7-12 major secondary veins on each

side, venation usually flat above and more prominent

beneath, the tertiary veins often subparallel between the

secondaries. Inflorescences axillary to distal leaves (rare-

ly pseudoterminal), 10-18 cm long (to 30 cm in fruits),

paniculate with distal branches much shorter than the

basal branches, glabrous, peduncles 1-6 cm long, be-

coming pendulous, pedicels 1-3 mm long. Flowers buds

ca. 2 mm long and 2 mm broad, yellow, glabrous or with

a few hairs on the outside, tepals ca. 1.5 mm long and

1 .2 mm broad near the base, ovate; outer anthers ca. 0.6

mm long, narrow with superposed and slightly overlap-

ping thecae, filaments 0.1-0.5 mm long, staminodes ab-

sent; pistil ca. 2 mm long, style ca. 1 mm long, stigma

simple or slightly discoid. Fruits borne on a receptacle

8-12 mm long, 6-10 mm broad and 1-3 mm deep, ur-

ceolate (but solid) to oblong and warty, bright red; fruits

2-3.5 cm long, 1-2 cm thick (dry), oblong-ellipsoid to

narrowly ovoid, narrowed at the base, becoming black.

Usually found as understory treelets in ever-

green wet forest formations, from near sea level

to 1100 m elevation. Most flowering collections

have been made in January-April, with a few in

August-October. Mature fruits have been collect-

ed in January-April and July-December. The

species is to be expected in southeastern Nicaragua

and has been collected along the Caribbean side

of Costa Rica, in the General Valley, Osa Penin-

sula, and adjacent Panama.

Ocotea atirrensis is distinguished by the large

thin obovate leaves with caudate-acuminate tips,

lack of puberulence, hollow distal stems, small

stature in the understory of usually lowland wet

forests, small flowers, and unusual fruiting recep-

tacle (a deep cup is not developed). This species

is closely related to O. paulii and O. wedeliana but

is distinguished by the large thin obovate leaves

with caudate tips, tertiary veins more often par-

allel, and different habitats. This species is similar

to O. nicaraguensis, but that species has thicker

leaves that do not dry dark and are not caudate-

acuminate at the apex and its stems are more

prominently ridged. Some collections between 400

and 1 000 m elevation on the Caribbean slope have

smaller, narrower, stiffer leaves that do not dry so

dark and resemble O. paulii. Whether this is just

clinal variation or reflects intergradation and gene

flow with O. paulii should be investigated. See the

discussion under Aiouea vexatrix by van der Werff

in Ann. Missouri Bot. Card. 75: 405. 1988.

Ocotea aurantiodora (Ruiz & Pavon) Mez, Jahrb.

Konigl. Bot. Gart. Berlin 5: 295. 1889. Laurus

72

FIELDIANA: BOTANY

aurantiodora Ruiz & Pavon, Laurografia Pe-

ruviana t. 15, 71802. Mespilodaphne aurantio-

dora Meissn. in DC., Prodr. 15(1): 101. 1864.

Small trees to 12 m tall and ca. 25 cm d.b.h., leafy

branchlets 6-14 mm thick, puberulent with short (0.4

mm) thin ascending brownish hairs, becoming glabres-

cent and dark in color, distal stems strongly angled with

prominent (ca. 1 mm) longitudinal ridges, usually with

5 ridges (or wings) in cross section. Leaves alternate,

petioles ca. 10 mm long and 3 mm thick, flat above or

the 2 adaxial ridges forming a shallow sulcus; leaf blades

13-35 cm long, 4-10 cm broad, very narrowly obovate

to oblanceolate or oblong-elliptic, usually tapering grad-

ually to a short acuminate apex, tapering gradually to

the cuneate or decurrent base, margin slightly revolute,

drying subcoriaceous, glabrous above with the slightly

raised minor venation forming a reticulum of small ar-

eolae (0.5-1 mm broad), lower surface with slender ap-

pressed straight yellowish hairs 0.2-0.7 mm long and

usually parallel with the secondary veins, with 9-14 ma-

jor secondary veins on each side and arising at angles of

45°-50°. Inflorescences axillary to distal leaves, 10-22

cm long, becoming 30 cm long in fruits, peduncles 3-9

cm long and 2-4 mm thick, sparsely puberulent and with

longitudinal ridges, pedicels ca. 1 mm long. Flowers ap-

parently unisexual, ca. 2 mm long and 3 mm broad,

yellowish white and minutely puberulent, tepals 1.5-2

mm long and 1.5 mm broad; outer stamens with short

(0.4 mm) filaments and narrow anthers ca. 0.8 mm long

with superposed thecae, inner stamens ca. 1.5 mm long,

glands closely appressed and resembling an annular disc,

staminodes absent; pistil/pistillode ca. 1 .8 mm long, stig-

ma disclike. Fruits borne in a receptacle ca. 6 mm long

and 8 mm broad, rounded and cupulate in form, the

opening ca. 5 mm in diameter and 3-4 mm deep, margin

rounded and entire; berry ca. 8 mm long and 5-6 mm

thick, oblong and bluntly rounded at both ends.

Trees of evergreen and partly deciduous forest

formations, at elevations from 200 to 900 m. A

single flowering collection has been made in July

in Costa Rica, and fruiting material has been col-

lected in January-April. This species is known from

southern Nicaragua, and from the General Valley

in southern Costa Rica. The species ranges from

southern Nicaragua and Costa Rica to Peru and

the Amazon Basin.

Ocotea aurantiodora is recognized by its thick,

strongly winged stems, long, stiff, narrowly oblong

or obovate leaves with slender appressed hairs be-

neath, small functionally unisexual flowers, and

small oblong fruits half immersed in a thick round-

ed cup. This species is similar to O. nicaraguensis,

but lacks the hollow ant-inhabited stems, and the

flowers and fruits are very different. This very dis-

tinctive species has been interpreted by Rohwer

(1986) to include O. longifolia H.B.K., O. gran-

difolia (Nees) Mez, and O. opifera Martius, among

others. However, in his monograph, Rohwer ( 1 986)

considered the Ruiz and Pavon names nomina

nuda because only their illustrations were pub-

lished. He has since changed his mind and we agree

that the Ruiz and Pavon names, based on very

fine illustrations, should be considered as effec-

tively published. The text for these illustrations

was published in Anal. Inst. Bot. Cavanilles 13:

21. 1954.

Ocotea austinii C. K. Allen, J. Arnold Arbor. 26:

350. 1945. O. irazuensis Lundell, Wrightia 5:

339. 1977. Figures.

Trees to 25 m tall and 65 cm d.b.h., leafy branchlets

1.8-5 mm thick, very minutely puberulent with brown-

ish hairs in early stages, becoming (sub)glabrous, dark

grayish and longitudinally ridged. Leaves alternate and

usually clustered at the ends of branchlets, petioles 5-

20 mm long, 1.5-3 mm broad, flat or sulcate above,

lateral margins continuous with the lamina margins; leaf

blades 3-12(-16) cm long, 1.5-4(-6) cm broad, oblong

to elliptic-oblong, abruptly narrowed at the short-acute

or obtuse apex, obtuse at the base with the margin re-

volute and decurrent on the petiole, drying subcoria-

ceous, upper surfaces usually lustrous and often with the

tertiary venation forming a raised reticulum, densely

subsericeous beneath with slender straight appressed yel-

lowish or brownish hairs 0.2-0.5 mm long and oriented

parallel with the major veins, becoming glabrescent, with

(5-)6-9(-l 1) major veins on each side, the central sec-

ondaries arising at angles of 40°-60°. Inflorescences ax-

illary to distal leaves, 6-15 cm long, paniculate, pedun-

cles 2.5-7 cm long, 1-3 mm thick, minutely puberulent

with slender appressed yellowish brown hairs, flowers in

short-stalked clusters, pedicels 1-3 mm long. Flowers 4-

5 mm broad, densely and minutely puberulent on the

outside, perianth-lobes 2.5-3 mm long, 2 mm broad;

outer stamens with anthers ca. 1 mm long and 1 mm

broad, the thecae superposed or the lower slightly lateral

to the upper, filaments ca. 0.5 mm long, inner anthers

narrow, staminodes absent; pistil ca. 2.5 mm long, gla-

brous, style to 1 mm long and slender, stigma discoid

or simple. Fruits borne in an obconic cup ca. 12 mm

long, 10-15 mm broad and 1-5 mm deep; berry 2-3.5

cm long, 1-2.5 cm in diameter, ellipsoid to ovoid, be-

coming purple.

Trees of higher montane wet evergreen forest

formations, from 1 700 to 3000 m elevation, rang-

ing from the northern edge of the Meseta Central

(near Palmira, Alajuela) through the central vol-

canic chain to the western part of the Cordillera

de Talamanca (near Empalme and Sta. Maria de

Dota, San Jose). Both flowers and fruits have been

collected in May-June, August, and December-

March. This species is endemic to central Costa

Rica but has closely related taxa in the Cordillera

de Tilaran and the Chiriqui highlands of Panama.

Ocotea austinii is characterized by its high-al-

BURGER: FLORA COSTARICENSIS

73

titude habitat, small stiff oblong leaves with lus-

trous upper surfaces and reticulate tertiary vena-

tion, the decurrent and revolute lamina base, silky

yellowish or orange brown pubescence on younger

parts and young leaves beneath, and the small pu-

berulent flowers. This species is very closely re-

lated to O. whitei which usually has oblanceolate

leaves, and varies from densely sericeous to almost

glabrous (see the discussion under O. whitei). Oco-

tea austinii appears to be part of a group of species

including O. sp. aff. caracasana, O. glaucosericea,

O. whitei, and O. skutchii. A Costa Rican name is

ira rosa.

Ocotea babosa C. K. Allen, Mem. New York Bot.

Gard. 15: 82. 1966, sensu lato. Figure 8.

Trees 15-35 m tall, to 1 m d.b.h., bark gray or mar-

bled, leafy branchlets 2-6 mm thick, puberulent with

crooked or straight brownish tomentulose hairs 0.2-0.5

mm long, soon becoming (sub)glabrous and grayish or

black. Leaves alternate, petioles 8-22(-30) mm long, 1.5-

2.3 mm thick, minutely puberulent or glabrescent, sul-

cate above; leaf blades 14-26 cm long, 8-12.5 cm broad,

broadly elliptic to ovate-elliptic or slightly obovate, acute

to short-acuminate at the apex, obtuse at the base, mar-

gin often somewhat undulate, drying stiffly chartaceous

to subcoriaceous, minutely puberulent on the veins above

but glabrescent between the veins, lustrous and some-

what bullate between major veins above, hirsutulous or

tomentulose with spreading slender straight brownish

hairs to 0.5 mm long beneath, with 4-7 major secondary

veins on each side, central secondaries arising at angles

of 35°-55° and loop-connected near the margin distally,

secondary veins slightly impressed above and very

prominent beneath, tertiary venation prominent be-

neath. Inflorescences axillary to distal leaves, 7-10 cm

long and few-flowered, becoming 10-12 cm long in fruits.

Flowers said to be 3.5-4.5 mm and campanulate, tepals

ca. 2.5 mm long; outer stamens ca. 1 mm long and the

outer anthers slightly longer than their filaments, stam-

inodes absent; pistil ca. 2.5 mm long, ovary ellipsoid to

obovoid, stigma papillate. Fruits borne on a campanu-

late-cylindrical pedicel gradually or abruptly expanded

to the flat or saucer-like receptacle, 4-10 mm long and

ca. 6 mm broad, only 1-2 mm deep, margin often 6-

lobed with persisting perianth bases or entire, pedicel

often with lenticels and verrucous; berry globose 7-10

mm in diameter (dried), dark green at maturity, drying

black and lustrous.

Trees of lowland evergreen forest formations in

Costa Rica. The species appears to flower in De-

cember-February in Costa Rica; fruits have been

collected in April and June. This species is known

from the La Selva area (Heredia) at about 100 m

elevation and Pejibaye (Cartago) at 500 m, both

on the Caribbean slope of Costa Rica. The species

(in a wide sense) ranges from Costa Rica to Ven-

ezuela.

Ocotea babosa is recognized by the distinctive

leaves with prominent venation and distinctive

pubescence beneath. The major veins are im-

pressed above and often give a broadly bullate

appearance to the upper surface. The small inflo-

rescences and small globose fruits on slightly ex-

panded receptacles, often with persisting perianth

bases, also distinguish this species. Herbarium

material resembles some species ofPersea. In Cos-

ta Rica, this species is only known from the fol-

lowing fruiting collections: Hammel & Trainer

10871, Hammel 11530, and McDowell 854 (all

DUKE), from La Selva; and Stork 2807 (F), from

Pejibaye (Pejivalle). While it is not unusual for a

species to have a lower altitudinal range in isth-

mian Central America than in continental South

America, the difference in altitudinal range of the

Costa Rican collections and those from northern

South America (at 1 800-2200 m) is greater than

normally seen, and there are differences in fruit

dimensions and upper leaf surfaces. Thus, C. K.

Allen's name O. babosa (1966) may prove to be

incorrect for our material. We are following Ham-

mel's usage (1986).

Ocotea sp. aff. O. bijuga (Rottb.) Bernard!, Can-

dollea 22: 59. 1967. Nectandra bijuga Rottboel,

PI. Surinam. Rar. 12. 1776. Figure 11.

Trees to 30 m tall and 70 cm d.b.h., trunks whitish

to pale gray, leafy branchlets, 1.3-5 mm thick, minutely

(0. 1 mm) and sparsely appressed puberulent or glabrous,

dark brown and longitudinally ridged, becoming terete

and grayish. Leaves alternate, petioles 5-15 mm long,

1-2 mm broad, sparsely and minutely puberulent, flat

or sulcate above; leaf blades 6-15 cm long, 1.5-5 cm

broad, narrowly elliptic-obovate to oblanceolate or nar-

rowly oblong, tapering to an obtuse or bluntly short-

acuminate apex, tapering gradually to the acute or at-

tenuate base and decurrent on the petiole, margin entire

and often slightly revolute (especially near the base),

drying stiffly chartaceous to subcoriaceous, glabrous and

dull or slightly lustrous above when dry with the minor

venation obscure, glabrous or very sparsely appressed-

puberulent beneath, with 5-9 major secondary veins on

each side, central secondaries arising at angles of 35°-

50°. Inflorescences axillary to distal leaves or extra-ax-

illary, 6-1 5 cm long, paniculate with short (2 cm) lateral

branches, peduncle 2-5 cm long, ca. 1.3 mm thick, gla-

brous or minutely puberulent, drying dark, pedicels 0-

2 mm long. Flowers 2-3 mm long, ca. 4 mm broad, very

minutely appressed puberulent on the outside, tepals 1.2-

1.7 mm long and ca. 1.2 mm broad, pellucid punctate;

outer stamens with short (0.5 mm) puberulent filaments,

outer anthers ca. 0.6 mm long and equally broad, thecae

superposed, inner stamens 1.4 mm long with large (0.5

74

F1ELDIANA: BOTANY

mm) glands, staminodes absent; pistil ca. 2 mm long,

style narrow and equalling the length of the ovary, stigma

simple or discoid. Fruit cup poorly developed, funnel-

form in life (fide Hammel) but drying flat or saucer-

shaped and about 5 mm broad, abruptly expanded above

the slightly thickened (1.5 mm) pedicel, red; berry glo-

bose, 5-8 mm in diameter when dry (to 1 2 mm in life),

green to greenish purple.

Trees of ridge tops in primary evergreen lowland

Caribbean rain forest in Costa Rica. Flowering is

in October, and fruiting material has been col-

lected in February and April. Costa Rican material

is only known from La Selva. The species with

which our material is provisionally placed occurs

in Colombia, Venezuela, and Brazil (see below).

Ocotea sp. aff. O. bijuga is recognized by its

glabrous narrowly elliptic to oblanceolate leaves

with decurrent leaf base and the small globose

fruits borne on a small flat or slightly concave

receptacle. The leaves tend to dry a dark olive

green above and brownish beneath. The associa-

tion of this material with Bernardi's concept of O.

bijuga, including O. florulenta (Meissner) Mez,

should be considered tentative. Rohwer (pers.

comm.) believes that O. bijuga is part of the Oco-

tea cernua complex. Also, Ocotea bijuga is said to

be dioecious, while ours appear to have bisexual

flowers; this also suggests that O. bijuga may not

be closely related to the Costa Rican material placed

here. The form and texture of the leaves are very

similar to Costa Rican material of O. oblonga and

the westernmost populations of 0. whitei(ai Mon-

teverde), but those species have very different fruits,

and prefer higher-elevation habitats. At present

this taxon is known from only five collections in

Costa Rica: Folsom 9112 (DUKE); Hammel 10229

(DUKE), 11663 (F, DUKE); Hartshorn 1585 (F, CR);

and Ocampo 3646 (CR).

Ocotea brenesii Standl., Field Mus. Nat. Hist., Bot.

Sen 18: 454. 1937. Nectandra brenesii (Standl.)

C. K. Allen, J. Arnold Arbor. 26: 370. 1945.

Figure 13.

Small to large trees, 6-28 m tall, trunks to 2 m broad

at the base, leafy branchlets 1-3.5 mm thick, appressed

puberulent at first with minute (0.2 mm) straight yel-

lowish hairs, quickly becoming (sub)glabrous and dark

in color, terete. Leaves alternate, petioles 6-1 3 mm long,

ca. 1 mm thick, minutely appressed puberulent or gla-

brescent, with a narrow adaxial sulcus; leaf blades 6-

12(-15) cm long, 2.5-6(-8) cm broad, ovate to ovate-

elliptic or broadly elliptic to ovate-oblong, short-acu-

minate to long-acuminate at the apex, acute to obtuse

or rounded at the base, drying thin chartaceous and very

dark in color, often lustrous above, appressed puberulent

on both surfaces but glabrescent with the minute (0.1-

0.3 mm) thin straight hairs persisting on the larger veins,

with 3-5 major secondaries on each side, central sec-

ondaries arising at angles of 35°-60°. Inflorescences ax-

illary to distal leaves, to 10 cm long, flowers few in an

open racemose panicle, peduncles to 5 cm long and 0.5

mm thick (dry), puberulent with straight appressed as-

cending hairs, pedicels ca. 3 mm long. Flowers 3-5 mm

long, 5-10 mm broad, yellowish or white, perianth gla-

brous or papillate-puberulent on the outer surfaces, te-

pals ca. 2 mm broad, glabrous within; outer stamens

subsessile or on short (0.3-0.4 mm) filaments, outer an-

thers ca. 1 mm long, ovate with superposed thecae, the

connective slightly developed beyond the thecae, inner

stamens with narrowly oblong anthers, staminodes ab-

sent, the floral tube with straight lustrous hairs; pistil

1.5-1.7 mm long, glabrous, ovary ovoid or angular, style

ca. 0.8 mm long, stigma slightly discoid. Fruits borne

on fruiting receptacles gradually expanded from the

thickened pedicel and conic, ca. 10 mm long and 8 mm

broad; berry poorly known, apparently to 24 mm long

and 1 6 mm in diameter, ovoid-oblong (based on A. Smith

517, 4102, F).

Trees of evergreen montane forest formations,

between 900 and 2000 m elevation in central Cos-

ta Rica. Flowering collections have been made in

February-May; mature fruits were collected in

April. This species is endemic and known only

from the northern region of the central highlands,

from near the Reserva Forestal de San Ramon to

the area between Poas and Barva (Barba) volcan-

oes in Alajuela Province.

Ocotea brenesii is distinguished by its relatively

broad leaves with few secondary veins that dry

thin in texture and very dark in color, rather large

flowers on open racemose panicles, and its re-

stricted habitat. It resembles O. tenera but that

species is glabrous in all its parts, has smaller flow-

ers, and narrower leaves. Ocotea brenesii resem-

bles the type of Ocotea pittieri (Tonduz 1 1893, us);

see the discussion under that species. Ocotea hold-

ridgeiana with larger flowers and stiffer leaves may

also be related to O. brenesii.

Ocotea brenesii has been placed into synonymy

under O. heydeana (Mez & J. D. Smith) Bernardi,

by Bernardi (1967, p. 93) and by Rohwer (1986,

p. 63). Collections of 0. heydeana differ in having

stiffly chartaceous leaves that usually turn olive

green when dried, and possess four to six pairs of

major secondary veins; also flowers and floral parts

are larger than in O. brenesii. Nevertheless, these

species are closely related. Ocotea heydeana (often

referred to as Nectandra heydeana) ranges from

Honduras to Chiapas at elevations of 700 to

1600m.

BURGER: FLORA COSTARICENSIS

75

Ocotea calophylla Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 298. 1889. Pleurothyrium velutinum

Meissn. in DC., Prodr. 15, pt. 1: 170. 1864, not

O. velutina Martius. O. fulvescens Slandl. & L.

O. Williams, Ceiba 1: 237. 1951. Figure 5.

Trees to 20 m tall, leafy branchlets 5-12 mm thick,

densely sericeous with reddish brown hairs in early stages

but the hairs becoming dark gray and matted. Leaves

alternate and subsessile, petioles short (0-10 mm) and

poorly differentiated from the cuneate lamina base; leaf

blades 10-19 cm long, 4-6 cm broad, narrowly elliptic

to elliptic-oblong, obtuse at the apex, attenuate at the

base with the margin revolute but not decurrent on the

stem, drying subcoriaceous, densely and minutely ap-

pressed puberulent above but becoming dull grayish and

the pubescence obscure, very densely and conspicuously

sericeous beneath with lustrous reddish brown or silvery

brown appressed ascending hairs, with 10-13 major sec-

ondary veins on each side, the central secondaries arising

at angles of 40°-70°, tertiary venation obscure. Inflores-

cences to 25 cm long, axillary to distal leaves, flowers

crowded in a compact panicle with lateral branches ca.

3 cm long, peduncles about half the length of the inflo-

rescence (4-12 cm) and 4-5 mm thick, reddish brown

puberulent. Flowers 4-7 mm long and 6-10 mm broad,

densely puberulent or tomentulose on the outside, outer

tepals ca. 4 mm long and 3 mm broad; outer anthers

1 .6-1 .8 mm long on slender glabrous filaments of nearly

equal length, thecae superposed but the upper introrse

and the lower latrorse (in the outer series), staminodes

present and small; pistil slender, to 4.5 mm long, style

1-3 mm long. Fruit cups ca. 1 cm broad and 3 mm deep

(immature?); berry ca. 20 mm long and 17 mm in di-

ameter, globose-ellipsoid.

Trees of wet evergreen montane forest forma-

tions, between 2600 and 3000 m elevation. Flow-

ers have been collected in January-February, and

fruiting inflorescences have been collected in Sep-

tember in Costa Rica. This species is also known

from Colombia and Venezuela at elevations from

2000 to 3 100m.

Ocotea calophylla is one of Costa Rica's most

distinctive species of Ocotea; the thick leaves with

dense lustrous pubescence beneath are unique

among Costa Rica's Lauraceae, and resemble some

Sapotaceae. The subsessile leaves with revolute

lamina base, thick stems, reddish brown inflores-

cence, and high montane habitat further distin-

guish this species. We have only seen the following

collections: Lems 5352 (F, NY); Leon 2166 (CR, us,

the type of O fulvescens); and Madriz 12 (CR, F).

The local names are reported as ira zoncho and

yema huevo. Many South American specimens of

this species have dull reddish pubescence, but some

do have the lustrous hairs that makes our Costa

Rican material so striking. The leaves of this species

are folded in an unusual way in bud, leaving a

diagonal line across the upper left of the lamina

and a short line on the right side near the base

(viewing the lamina undersurface with the apex

up and the base down). Ocotea guianensis Aublet

of lowland formations in South America has sim-

ilar lustrous pubescence, and the same pattern of

folding in leaf vernation, and must be related.

Closely related Andean species are O. micans Mez

and O. sericea H.B.K.

Ocotea sp. aff. caracasana (Nees) Mez, Jahrb. Kon-

igl. Bot. Gart. Berlin 5: 292. 1889. Oreodaphne

caracasana Nees, Linnaea 2 1 : 522. 1 849. Figure

10.

Trees to 30 m tall and 60 cm d.b.h., occasionally with

prop roots from near the base, dioecious or apparently

bisexual in ours, leafy branchlets 2.5-8 mm thick, ap-

pressed puberulent with minute (0.1-0.4 mm) slender

ascending pale yellowish to golden hairs, often with lon-

gitudinal ridges but becoming terete and grayish. Leaves

alternate, petioles 0-1 5 mm long but poorly defined, with

lateral margins continuous with the decurrent lamina

base, minutely appressed puberulent, narrowed portion

of the lamina base 1-4 cm long; leaf blades 12-20(-24)

cm long, 4.5-8(-ll) cm broad, elliptic-obovate to ob-

ovate, oblong-obovate or oblanceolate, abruptly nar-

rowed or rounded to a short-acuminate or obtuse apex,

tapering gradually to the cuneate and decurrent base,

margin usually revolute near the base, drying subcoria-

ceous to coriaceous, glabrous above or minutely ap-

pressed puberulent on the flat or impressed midvein near

the base, usually dull above and slightly glaucous be-

neath, minutely and often obscurely puberulent beneath

with thin straight appressed yellowish hairs 0.1-0.3 mm

long and usually parallel with the secondary veins or

glabrescent, with (6-) 7-1 2 major secondary veins on each

side, central secondaries arising at angles of 35°-50°, ter-

tiary venation subparallcl and slightly raised beneath.

Inflorescences axillary to distal leaves or distal leafless

nodes, 7-12 cm long, panicles broadly paniculate, many-

flowered, peduncles 2-8 cm long, 2-3 mm thick, densely

puberulent and silvery to reddish brown in color, ped-

icels 0-4 mm long. Flowers 3.5-5 mm long, 4-6 mm

broad, campanulate, densely puberulent on the outside,

tepals 1.5-2.8 mm long, 1.3-2.3 mm broad; outer sta-

mens on short (0.4 mm) broad glabrous filaments, outer

anthers 0.5-1 mm long, usually narrower than long, the-

cae superposed but occasionally quite variable, inner

stamens 1.3-2 mm long, often with large glands, stam-

inodes to 1 . 1 mm long and variable or absent; pistil 1 .6-

2.5 mm long, the slender style 1-1.5 mm long, stigma

discoid or simple. Fruits borne in a large hemispherical

cup 18-28 mm broad and 8-15 mm deep, margin often

with teethlike projections from the persisting perianth;

berry 3—4 cm long and 2-2.5 cm in diameter, ovoid.

Trees of evergreen rain forest formations in the

Caribbean lowlands of Costa Rica. Flowers and

fruits were both collected in May. This taxon may

76

FIELDIANA: BOTANY

be endemic to Costa Rica if it proves to be different

from true O. caracasana, which occurs in Colom-

bia, Venezuela, and the Guianas.

Ocotea sp. aff. caracasana is recognized by the

larger slightly obovate leaves, abruptly short-acu-

minate at the apex and decurrent and revolute at

the base. The large fruits and the stiff leaves with

characteristic appressed hairs are also helpful in

recognizing this species and its relatives. The pu-

berulent flowers were described as unisexual but

the Costa Rican material appears to be bisexual.

Our material is derived from only a single tree at

La Selva (Hammel p. 225, 1986: Hammel 12450,

DUKE, F). The Hammel collection differs from the

type photo (Karsten 32, F) in its lack of the elongate

racemiform inflorescences, its secondary veins

arising at a narrower angle, and its revolute lamina

base being not as long. But there are also many

similarities between the Costa Rican material and

the type collection. This species, whatever its final

name, is related to O. glaucosericea and O. skutchii

of Costa Rica's highland forests; see the discus-

sions under those species. While the foliage of these

species resembles that of O. insularis and its allies,

the fruits, parallel-appressed hairs, and prop roots

indicate that they may not be closely related.

Ocotea cernua (Nees) Mez, Jahrb. Konigl. Bot.

Gart. Berlin 5: 377. 1889. Oreodaphne cernua

Nees, Syst. laur. 424. 1836. Figure 14.

ca. 2.5 mm long and urceolate, with a deep tube, tepals

1-1.8 mm long and minutely papillate near the tips,

nonfunctional anthers ca. 0.5 mm long, pistil ca. 1.5 mm

long, stigma subsessile on the narrow apex of the ovary.

Fruits borne in a cup 6-8 mm long and 9-12 mm broad,

4-5 mm deep, margin entire, thickened part of the ped-

icel 5-10 mm long, becoming red and enclosing the lower

'/? of the fruits; berry 1.5-2 cm long, ovoid, becoming

black (rare in herbaria).

Small trees of lowland evergreen forest forma-

tions on both the Caribbean and southern Pacific

slopes, below 900 m elevation in Costa Rica.

Flowering occurs in February-May, and fruits have

been collected in August-November in Costa Rica.

The species ranges from southern Mexico and the

West Indies through Central America to South

America.

Ocotea cernua is distinguished by its elliptic-

oblong leaves that nearly always dry grayish or

yellowish and often have a caudate-acuminate

apex, and the slender inflorescences that are ra-

cemiform or open panicles with racemose lateral

branches. The lack of puberulence on almost all

parts, the ebracteate inflorescences, and the small

unisexual flowers that usually dry black are ad-

ditional features of this unusual species. Ocotea

cernua, O. meziana, and O. tenera are very similar

in the field but differ in their dried appearance (cf.

Hammel, 1986). Licaria cufodontisii and L. sar-

apiquensis are also similar to O. cernua, especially

in the appearance of leafy twigs and inflorescences.

Shrubs or small to medium-sized trees 5-10(-20) m

tall, dioecious (unisexual), leafy branchlets 1.2-3 mm

thick, apical buds slightly puberulent but stems essen-

tially glabrous, terete or slightly ridged. Leaves alternate,

approximate but not congested distally, petioles 6-20

mm long, ca. 1 mm thick, glabrous and with a shallow

adaxial sulcus; leaf blades 6-14(-18) cm long, 3-6(-8)

cm broad, oblong to elliptic-oblong, caudate acuminate

or short- to long-acuminate at the apex, often with a

narrow (ca. 3 mm) tip to 2 cm long, acute to obtuse at

the base, margin often undulate, drying chartaceous to

stiffly chartaceous, grayish and glabrous above, glabrous

or sparsely puberulent along the major veins beneath,

the primary veins flat above, with 3-6 major secondary

veins on each side, central secondaries arising at angles

of 30°-70°. Inflorescences axillary or extra-axillary on

distal twigs (sometimes pseudoterminal and stemlike),

3.5-15 cm long, paniculate and usually with racemose

lateral branches or small and raceme-like, glabrous or

with very few appressed hairs, often drying very dark

(both flowers and inflorescence), peduncle to 5 cm long,

ca. 0.5 mm thick, pedicels 1-4 mm long, slender. Flowers

unisexual, drying dark, glabrous on the exterior; male

flowers ca. 2.2 mm long and 3-4 mm broad, campan-

ulate, tepals 1.2-1.8 mm long, outer anthers 0.6-0.8 mm

long, slightly narrower than long, subsessile, thecae su-

perposed, a pistilode present or absent; female flowers

BURGER: FLORA COSTARICENSIS

Ocotea dendrodaphne Mez, Jahrb. Konigl. Bot.

Gart. Berlin 5: 238. 1889. O. quisara Mez & J.

D. Smith, Bot. Gaz. 33: 259. 1902. O. ovan-

densis Lundell, Contr. Univ. Mich. Herb. 6: 16.

1941. Figure 3.

Shrubs or small treelets to 6 m tall, leafy branchlets

3-7 mm thick, minutely (0.1-0.3 mm) appressed pu-

berulent with straight ascending hairs in early stages,

soon becoming glabrous and pale gray, lenticellate and

ridged but becoming terete in age, distal stems hollow

and often with small ants. Leaves alternate, petioles 6-

36 mm long, 1.5-3 mm thick, sulcate above, drying dark

in color; leaf blades 14-36 cm long, 5-14 cm broad,

elliptic to narrowly elliptic-oblong, widest at or slightly

below the middle, acute or short to long acuminate at

the apex (occasionally rounded/obtuse), acute to obtuse

at the base, drying very stiffly chartaceous to subcoria-

ceous and grayish green, glabrous and slightly lustrous

on both surfaces, with 5-10 major secondary veins on

each side, the central secondaries arising at angles of 40°-

70° tertiary venation slightly raised and reticulate be-

neath. Inflorescences to 1 5 cm long, borne close together

at the branchlet tips in the axils of what appear to be

leaf scars (perhaps early caducous leaf scales), paniculate

77

in form with basal branches larger than the distal, pri-

mary peduncle to 5 cm long, very minutely puberulent

in early stages. Flowers 4-6 mm long, to 8 mm broad,

white and sweet-scented, tepals to 5 mm long, densely

and minutely papillate-puberulent; outer anthers 2-3 mm

long, broadly flattened and somewhat tepal-like, ovate-

triangular to lanceolate in outline with the thecae often

on the basal half of the inner face, minutely puberulent,

filaments very short (0.5 mm) and stout, staminodes not

seen; pistil 1.9-2.4 mm long, style 1.1-1.5 mm long,

slender, stigma slightly discoid. Fruit cups 8-10 mm

long, ca. 1 5 mm broad and 3-7 mm deep, enclosing the

basal 'A of the fruits, conical to cupulate in form, often

with a second ridge around the rim; berry 15-25 mm

long, 10-12 mm thick, bluntly rounded at the apex, nar-

rowly ovoid to oblong, black at maturity.

Understory plants of wet evergreen forest for-

mations, between sea level and 1 500 m elevation,

along the Caribbean slopes and in the central high-

lands. Flowering collections have been made in

January-June, October, and December; fruiting

occurs throughout the year, but most collections

have been made in July-October in Costa Rica.

The species ranges from central Mexico to central

Panama.

Ocotea dendrodaphne is recognized by its hol-

low stems often inhabited by ants, the large gla-

brous elliptic-oblong leaves that dry very stiff and

gray-green, the large flowers with tepaloid sta-

mens, and distinctive deep cups. This species is

often found on slopes and ridges. Two unusual

features of this species are the way in which the

inflorescences originate from leafless nodes, and

the frequently inflated distal branchlet-tips with

longitudinal slitlike openings into the hollow stems.

The latter are associated with ants. The large flow-

ers with broadly flattened outer stamens and the

fruit cups with ridged rims (resembling Licaria,

but not always developed) indicate a relationship

with O. veraguensis. However, specimens may be

more easily confused with O. paulii and its allies,

which also have hollow stems, similar leaves, and

are treelets of wet forest understory.

Ocotea dentata van der Werff, sp. nov. Figure 20

Arbor ad 25 m. Ramuli juniores angulares, adpresse

pubescentes. vetustiores teretes glabrescentesque. Folia

obovata, 20-35 x 8- 12 cm, apice rotundata velbreviter

acuminata, basi decurrente et reflexa, supra praeter cos-

tarn nervosque leviter pubescentes laevia glabraque, sub-

tus sparse pubescentia, pilis brevibus et plerumque er-

ectis; costa, nervis elevata, venatione reticulata et parum

elevata; nervis 9-12. Petioli sulcati, 0.5-1.5 cm longi.

Inflorescentiae pubescentes, in axillis foliorum vel brac-

tearum deciduarum, foliis breviores vel aequales, ram-

ificatione divaricata. Flores fragrantes, hermaphroditi;

tepala magnitudine aequalia, ca. 1.5 mm longa, ovata,

omnina intus glabra, 3 exterioribus extus dense puberulis

sed 3 interioribus minus puberulis; stamina 9, 4-locel-

lata, 6 exterioribus ca. 1.1 mm longis, antheris glabris,

ca. 0.6 mm longis, thecis introrsis, filament is ca. 0.5 mm

longis, propre apicem pilis minutis muni t is: 3 interior-

ibus ca. 1 .2 mm longis, antheris glabris, thecis inferior-

ibus extrorsis, thecis superioribus lateralibus, filamentis

ca. 0.5 mm longis, 2 glandulis munitis; pagina interiore

filamentorum propre apicem pubescentia densa munita;

staminodia nulla. Ovarium ovoideum, glabrum, in sicco

0.5 mm longum, stylo ca. 1 mm longo, glabro. Fructus

viridis, ellipsoideus, 10x6 mm; cupula tepalis persis-

tentibus munita.

Trees to 25 m tall, leafy branchlets 2.5-8 mm thick,

angular when young but becoming terete, at first ap-

pressed puberulent but glabrescent in age. Leaves alter-

nate, petioles difficult to delimit because of the decurrent

lamina-base, 5-15 mm long, 2.5-7 mm broad and sul-

cate above; leaf blade (15-)20-35 cm long, (5-)8-12 cm

broad, obovate to elliptic-obovate or elliptic-oblong,

usually broadest at or above the middle, abruptly nar-

rowed and often rounded at the obtuse to short-acu-

minate apex, tapering to the cuneate and decurrent base,

margin somewhat revolute near the base and decurrent

on the petiole, drying stiffly chartaceous to subcoria-

ceous, glabrous above or with some hairs above the ma-

jor veins, lower surface usually sparsely puberulent with

erect or spreading hairs 0.2-0.5 mm long, with 9-12

major secondary veins on each side, central secondaries

arising at angles of 30°-50°. Inflorescences axillary to

distal leaves or deciduous bracts, solitary, 12-25 cm long,

robust and many-flowered panicles, peduncle 3-12 cm

long, 1.7-3.5 mm thick and with grayish or brownish

puberulence, pedicels to 2 mm long, flowers in cymose

groups. Flowers 3-5 mm long and 4-5 mm broad, yel-

lowish green, tepals 1.5-2.5 mm long, subequal, outer

tepals densely puberulent on the outer surfaces, glabrous

within; outer stamens with short (0.2-0.4 mm) filaments,

outer anthers 0.5-0.8 mm long and equally broad or

narrower, thecae superposed or the lower slightly lateral,

inner stamens 1 .2-1 .4 mm long, with a tuft of hairs below

the anthers, staminodes absent; pistil 1.5-2.6 mm long,

ovary ovoid, style 1-1.5 mm long, slender, stigma slightly

discoid. Fruits borne on a short (8-10 mm) cupulate

receptacle 5-12 mm broad, the margin usually with per-

sisting perianth bases; berry ellipsoid or oblong, 10 mm

long and 6 mm in diameter (in the type) but perhaps

becoming 3 cm long and 2 cm in diameter.

TYPE— Costa Rica, Prov. Limon, Reserva Biol-

ogica Hitoy Cerere, 100-125 m, 29 Aug. 1985, L.

D. Gomez & G. Herrera 23653 (holotype, MO; is-

otypes, CR, F). Additional Collections: Nicaragua,

Dept. Matagalpa, Neil I 77/5, Stevens & Riviera

20899, Dept. Zelaya, Araquistain 3165, Grijalva

264, Stevens 8005, 12444 (all MO); Costa Rica,

Prov. Alajuela, Haber 5557, 6178 (MO); Hartshorn

1543 (CR, F); Panama Prov. Bocas del Toro, Lao

94, von Wedel 720, 1382 (all MO).

78

FIELDIANA: BOTANY

Trees of evergreen rain forest formations, from

near sea level to 1000 m elevation. Flowers have

been collected in August and October-December;

young fruits have been collected in February, ma-

ture fruits in March. This species ranges along the

Caribbean slope and lowlands, from Nicaragua to

Panama, and in the evergreen forests of the south-

ern Pacific slope in Costa Rica.

Ocotea dentata is recognized by the larger stiff

puberulent obovate leaves with decurrent and re-

volute leaf bases, and the fruit cups often with

persisting perianth parts. This species is closely

related to O. insularis (in a wide sense). The pu-

bescence helps separate this species from O. in-

sularis, O. ira, and O. rivularis. The dense pub-

erulence of the flowers, and the cupules with

persistent tepals (hence the dentate appearance and

specific name) are also distinctive. Earlier, some of

the material placed here was identified as O. sten-

oneura. Unfortunately, Ocotea stenoneura is very

poorly understood at present, but it has very dif-

ferent fruits (q.v.). Additional material was used

to amplify the English description beyond the

measurements cited for the type in the Latin de-

scription.

Ocotea endresiana Mez, Jahrb. Konigl. Bot. Gart.

Berlin 5: 257. 1889. Figure 10.

Trees to 25 m tall, leafy branchlets 2-6 mm thick,

glabrous or minutely appressed puberulent at first, with

prominent ridges but soon becoming terete, dark brown

to dark gray. Leaves alternate, petioles poorly denned or

absent, narrowed portion of the lamina base 0-2(-3) cm

long, flat above, with strongly revolute margins; leaf

blades 6-17 cm long, 3-9 cm broad (25 x 18 cm in

sprout shoots), obovate to elliptic-obovate, obtuse to

abruptly rounded at the apex, tapering gradually to the

decurrent base, revolute with the margin curled under

near the base, drying subcoriaceous or coriaceous, gla-

brous and often lustrous above, glabrous or minutely

(0. 1-0.2 mm) puberulent beneath, rarely with longer tufts

of hairs (domatia) in the vein axils and along the midvein

beneath, with 5-9 major secondary veins on each side,

central secondaries arising at angles of 40°-60°, tertiary

venation slightly raised on both surfaces but usually ob-

scure, often subparallel between the secondaries, major

veins often reddish in early stages. Inflorescences axillary

to distal leaves or apparently terminal, to 1 7 cm long,

broadly paniculate but few-branched, peduncle to 6 cm

long, 2-3 mm thick, glabrous and drying dark brown to

black, pedicels ca. 2 mm long. Flowers 2.5-5 mm long,

3-6 mm broad, yellowish, glabrous to densely and mi-

nutely puberulent on the outer surfaces, minutely pap-

illate-puberulent within, tepals ca. 2.5 mm long; outer

stamens with short (0.5 mm) filaments, outer anthers

0.5-0.8(-1.1) mm long and equally broad or narrower,

staminodes absent (rarely present?); pistil 1 .5-2 mm long,

style narrow and usually equalling the ovary in length,

stigma simple or discoid. Fruits borne in cupulate re-

ceptacles 8-1 5 mm long and 7-10 mm broad, rim entire

and the surfaces smooth, becoming red; berry 1 5-1 8 mm

long, 8-10 mm thick (dried), oblong-ellipsoid or elliptic-

cylindrical.

Trees of montane evergreen forest formations,

between (700) 1 100 and 2300 m elevation, along

the Caribbean slope and continental divide. Flow-

ering material has been collected in August-Oc-

tober and February; mature fruits have been col-

lected in February-March and June. The species

ranges from the Cordillera de Tilaran (Monte-

verde area), along the northern and eastern edge

of the central highlands, to the area of Cartago and

Volcan Turrialba. This taxon of doubtful status is

endemic to Costa Rica and adjacent Panama (see

below).

Ocotea endresiana is recognized by the stiff

bluntly obovate leaves, with decurrent leaf bases

which have broadly revolute margins on the un-

derside of a poorly defined petiole/leaf base. The

long narrow fruits in smooth entire-rimmed cups

only 1 cm broad also distinguishes this species. It

appears to be common only north of San Ramon

and near Vara Blanca de Sarapiqui. The very re-

stricted range is an argument for considering these

plants no more than a subspecific element of O.

insularis in a wide sense. However, the leaf bases

are very distinctive, as are the fruiting stages. See

the discussion under O. insularis. Ocotea endre-

siana has sometimes been confused with O. skut-

chii, which also has long-decurrent leaf bases but

different flowers, and a characteristically fine in-

dumentum on the undersides of the leaves. We

have not seen the type and are following earlier

usage in the application of this name.

Ocotea floribunda (Sw.) Mez, Jahrb. Konigl. Bot.

Gart. Berlin 5: 325. 1889. Laurus floribunda

Sw., Prodr. 65. 1788. O. wachenheimh ' Benoist,

Bull. Mus. Hist. Nat. Paris 30: 510. 1924. Fig-

ures 14.

Trees 1 5-30 m tall, trunk 30-80 cm in diameter, dioe-

cious (unisexual), leafy branchlets 1.5-5 mm thick, very

minutely (0. 1 mm) puberulent with inconspicuous hairs

near the apex but quickly becoming glabrous and dark

in color, smooth and terete. Leaves alternate or subop-

posite, petioles 4-14 mm long, 0.8-1.9 mm thick, gla-

brous and flat above, drying dark in color; leaf blades

7-1 5(-19) cm long, 2-6(-8) cm broad, elliptic to oblong-

obovate, narrowly obovate or obovate, usually broadest

BURGER: FLORA COSTARICENSIS

79

above the middle, acute to short-acuminate (rarely

rounded) at the apex, acute to cuneate at the base and

slightly decurrent on the petiole, margin entire and slightly

recurved when dry, drying stiffly chartaceous to subcor-

iaceous and pale to dark grayish green, essentially gla-

brous above and below, with (3-)5-8 major secondary

veins on each side, with a few weakly denned loop-con-

nections near the margin, central secondaries arising at

angles of 40°-60°, the larger veins often drying reddish

brown, the smallest veins usually forming a fine (0.1-

0.6 mm) reticulum usually visible on both surfaces but

more prominent beneath. Inflorescences axillary, extra-

axillary or pseudoterminal, 3-14 cm long, racemose or

paniculate, few to many-flowered, peduncle 1-4 cm long,

glabrous or very minutely puberulent, drying dark, ped-

icels 1-3 mm long, ca. 0.8 mm thick. Flowers function-

ally unisexual, 5-7 mm broad, greenish to cream-white,

outer tepals broadly imbricate, 2-3(-4) mm long and 2-

3 mm broad, minutely puberulent on both surfaces, be-

coming reflexed; male flowers with stamens on promi-

nent (0.9 mm) filaments, anthers ca. 1.3 mm long and

1.2 mm broad, the lower thecae slightly lateral to the

upper, inner stamens ca. 2 mm long, staminodia absent,

pistillode ca. 3 mm long; female flowers with the outer

stamens on short (0.2 mm) filaments and with smaller

(0.7 mm) anthers, pistil ca. 3 mm long, ovary ovoid and

equalling the style in length, stigma discoid. Fruits borne

a flat disci ike receptacle 6-10 mm in diameter and 1.5-

2 mm thick, often slightly recurved, margin rounded in

life (fide Hammel) but with upper and lower ridges when

dry, sometimes with the bases of persisting floral parts,

green; berry subglobose, 13-18 mm long and 10-15 mm

in diameter, green and glaucous at maturity, usually with

short ( 1 mm) persisting style base at the apex of the fruits.

Trees of wet evergreen forest formations, from

near sea level to 1 500 m elevation. In Costa Rica,

flowers have been collected in October and De-

cember-February. Young fruits have been col-

lected in June, and mature fruits in August-No-

vember and March (near Panama). The species

has been collected in the Caribbean lowlands (La

Selva), near Turrialba, at Monteverde, and on the

Pacific slope near Panama at 1 300 m elevation. It

has not been collected on the Pacific slope below

1200 m in Costa Rica. The species ranges from

Nicaragua southward to Peru, Venezuela, the

Guianas, and the West Indies.

Ocotea floribunda is recognized by its very dark

(almost black) stems, stiff leaves drying grayish

green and with a finely reticulated surface, uni-

sexual flowers, globose fruits with persisting style

base, and the flattened fruiting receptacle with

ridged edge. Herbarium material of this species

resembles O. meziana, O. veraguensis, and some

species of Licaria. This species appears to be un-

common and has been collected in Costa Rica only

recently. Costa Rican material differs from the West

Indian collections in a number of ways and may

prove to be worthy of subspecific recognition. There

is an unusual population in the southern part of

the Cordillera de Talamanca, where the base of

the lamina is both decurrent and revolute (Davidse

24359, CR, MO; Hartshorn 2165, Lent 2705 CR, F);

flowers of the Davidse collection make the place-

ment of these collections under O. floribunda quite

certain.

Ocotea glaucosericea Rohwer, Mitt. Inst. Allg. Hot.

Hamburg 20: 144. 1986, nomen novum forNec-

tandra hypoglauca Standl. ex C. K. Allen, J.

Arnold Arbor. 26: 399. 1945, not O. hypoglauca

(Nees) Mez. Figure 12.

Trees to over 30 m tall, leafy branchlets 3-8 mm thick,

densely and minutely (0.2 mm) puberulent with ascend-

ing or appressed hairs, yellowish brown to grayish brown,

longitudinally ridged. Leaves alternate in a spiral, peti-

oles poorly defined and continuous with the decurrent

lamina base, 1-3 cm long, with broad revolute lateral

margins; leaf blades 9-17 cm long, 4-10 cm broad, ob-

ovate to narrowly obovate, elliptic-obovate or oblong-

obovate, abruptly rounded at the apex or obtuse (bluntly

acute in narrow leaves), tapering gradually to the cuneate

and decurrent base, margin entire and usually revolute

at the base, drying subcoriaceous and lustrous above with

the minor venation slightly raised, glabrous except on

the midvein basally, paler in color beneath and with

minute (0.1-0.2 mm) slender appressed hairs usually

lying parallel with the secondary veins, with (5-)6-9 ma-

jor secondary veins on each side, the central secondaries

arising at angles of 30°-50°. Inflorescences axillary to

distal leaves, 8-18 cm long, paniculate, peduncles 3-9

cm long, densely appressed puberulent and yellowish,

pedicels 2-4 mm long. Flowers ca. 4 mm long, 4-6 mm

broad, densely grayish or yellowish brown puberulent

on the outside, tepals ca. 2.8 mm long and 2.2 mm broad;

outer stamens 1-1.5 mm long with short (0.2-0.6 mm)

broad filaments, outer anthers 0.9-1 mm long and 0.7-

0.8 mm broad, the thecae superposed or the lower some-

what lateral to the upper, staminodes absent or small (to

0.6 mm); pistil ca. 3 mm long, ovary ellipsoid, style ca.

1 mm long, stigma capitate. Fruits borne in a conical

receptacle about 2 cm long, 1.2-1.8 cm broad and ca. 5

mm deep; berry 2.5-3 cm long, 2-2.5 cm in diameter,

globose or oblong.

Trees of evergreen montane forest formations,

from 1500 to 2500 m elevation. Flowering ma-

terial has been collected in April-May and July;

fruits have been collected in March-June. The

species is endemic to the Cordillera de Talamanca

in Costa Rica (El Cedral de Sta. Maria de Dota,

La Cangreja, and Las Tablas near the Rio Coton-

cito), and appears to be common in the Chiriqui

highlands of Panama.

Ocotea glaucosericea is recognized by its stiff

obovate leaves, lustrous on the upper surface and

decurrent and revolute at the base. The highland

80

FIELDIANA: BOTANY

habitat, upper leaf surface, and smaller fruits on

shallower receptacles distinguish this species from

O. sp. aff. caracasana, with which this species has

been considered synonymous by Hammel (1986).

This species is also closely related to O. skutchii,

and there are collections that appear to be inter-

mediate between them. Ocotea austinii and O.

whitei are also part of this complex. All these taxa

appear to be part of a closely related species group,

united by characters of the flowers, large fruits,

leaf form, and the slender appressed hairs on the

lower leaf surfaces. These species resemble O. in-

sularis and its allies, but those differ in pubescence

and fruits, and are not as closely related.

Ocotea gomezii W. Burger, sp. nov. Figure 6.

Arbor 10 m alta, ramulis foliiferis 3-6 mm crassis,

ferrugineo-tomentosis. Folia alterna, petiolis 1 2-26 mm

longis, tomentosis; laminis 10-18(-24) cm longis, 6-

11 (-14. 5) cm latis, late ellipticis vel suborbicularibus,

apice obtuso vel breviter acuminato, subtus ferrugineo

puberulis, nervis secondariis 5-8(-10) paribus. Inflores-

centiae 10-18 cm longae, paniculatae, floribus paucis,

pedunculis 6-13 cm longis, tomentosis. Flores ca. 10

mm longi et 1 2 mm lati, tepalis ca. 6 mm longis, pub-

erulis; stamina ser. I-II ca. 3 mm longa, antheris ca. 1 .6

mm longis; gynoecium 2.5-3.7 mm longum. Fructus ig-

notus; cupula 1 5-20 mm longa et 1 5-20 mm lata, mar-

gine lobata.

Trees 6-10 m tall, leafy branchlets 3-6 mm thick and

densely ferrugineous-tomentulose with soft curved and

straight hairs ca. 0.5 mm long, remaining tomentulose

but becoming grayish. Leaves alternate and not crowded

distally, petioles 12-26 mm long, 2.5-3.5 mm thick,

densely tomentulose; leaf blades 10-18(-24) cm long, 6-

11 (-14. 5) cm broad, broadly elliptic to suborbicular,

usually broadest near the middle and rarely somewhat

ovate or obovate, bluntly obtuse to short-acuminate with

a tip to 1 5 mm long, obtuse to rounded and subtruncate

at the base, drying stiffly chartaceous and dark brown to

olive green, slightly lustrous above and minutely puber-

ulent above the major veins, densely tomentulose be-

neath with ferruginous or yellowish brown hairs ca. 0.7

mm long, with 5-8(-10) major secondary veins on each

side, central secondaries arising at angles of 45°-60°, ma-

jor veins very prominent beneath and forming an arcuate

submarginal vein distally, tertiary venation also slightly

raised. Inflorescences few-branched panicles in axils of

distal leaves or pseudoterminal, 10-18 cm long, pedun-

cles 6-13 cm long, 1.7-3 mm thick, densely ferruginous

puberulent, flowers borne in groups of 1-3 on short (1

cm) lateral branches subtended by a bract 5-6 mm long.

Flowers ca. 10mm long and 12mm broad, campanulate,

yellowish white, densely puberulent on the outside and

papillate-puberulent within, tepals to 6 mm long and 3

mm wide with the inner series smaller than the outer;

outer stamens to 3 mm long, outer anthers ca. 1 .6 mm

long and 1 mm broad, oblong, valves superposed, fila-

ments with long trichomes, inner stamens with large

glands abaxially, staminodes absent; pistil 2.5-3.7 mm

long, ovary narrowed to the base, style ca. 2 mm long,

slender, stigma slightly lobed. Fruits borne in a conical

cup 1 5-20 mm long and 1 5-20 mm broad, the cup sur-

face densely hirsutulous, the perianth parts persisting

and forming large (3-5 mm) lobes on the margin of the

cup; young berry with hairs over the basal surface, ma-

ture berry not seen.

TYPE— Costa Rica, Alajuela Province, valley of

the Rio Lorencito in the Reserva Forestal de San

Ramon, 14 March 1987. Gomez- Laurito, Burger,

Mora, Ortiz & Antonio 11450 (holotype, CR; iso-

typCS, CAS, F, DUKE, HBG, MO, NY, USj).

Trees of very moist cloud forest and premontane

rain forest formations of the Caribbean slope and

continental divide, from 800 to 1400 m elevation.

Flowers have been collected in January-April and

July. Fruits have been reported in August and Jan-

uary. The species is endemic to Costa Rica, and

ranges from near Volcan Rincon de la Vieja in the

west to Moravia de Chirripo in the east, and prob-

ably further eastward along the Caribbean slope

of the Cordillera de Talamanca.

Ocotea gomezii is a very unusual species, dis-

tinguished by its ferruginous puberulence, broadly

rounded leaves, large puberulent flowers, and

fruiting receptacles with large perianth lobes. The

puberulence and lobed fruiting receptacles resem-

ble O. mollifolia, but that species has very different

inflorescences, and thinner, more obovate leaves.

Ocotea pseudopalmana is similar but has smaller

flowers, entire fruit cups, and grows at higher el-

evations. The large glands of the new species are

reminiscent of Pleurothyriumpalmanum, and there

are other similarities that should be looked at fur-

ther. The first collection of this species was by

Brenes (328(685)/6092, F, NY); other collections

are Bermudez 268 (usj), Dryer 1612 (CR, F), Gom-

ez-L. 4590 (CR, usj), Hammel & Trainer 15044

(MO), Holdridge 6827, 6829 (CR, NY), and Poveda

et al. 947 (CR, F), 75/9 (usj). The best flowering

collection of this species was made by Jorge Gom-

ez-Laurito, whose collections are adding signifi-

cantly to our knowledge of Costa Rica's rich flora.

Ocotea hartshorniana Hammel, J. Arnold Arbor.

67: 128. 1986. Figure 8.

Trees 10-30 m tall, to 1 m diameter d.b.h., with but-

tresses to 2 m high and 1 m broad or with prop roots,

bark pinkish tan to orange, leafy branchlets 2.5-6 cm

thick, at first densely reddish brown or yellowish brown

puberulent with hairs 0. 1-0.5 mm long, often ascending

BURGER: FLORA COSTARICENSIS

81

and slightly strigulose, longitudinal ridges usually pres-

ent. Leaves alternate, petioles 4-1 8(-24) mm long, densely

puberulent or the hairs minute and apparently glabrous,

flat above or slightly sulcate; leaf blades 1 2-1 7(-20) cm

long, 3-6(-8) cm broad, oblong-obovate to obovate or

elliptic-oblong, usually broadest above the middle,

abruptly narrowed to a short acuminate or caudate-acu-

minate apex, gradually narrowed to a decurrent base,

margins entire and usually recurved at the base, drying

stiffly chartaceous, puberulent on the mid vein above or

glabrescent, sparsely to densely puberulent on the veins

beneath, hairs 0.2-0.5 mm long and often reddish brown

in color, sometimes glaucous beneath, with 4-8 major

secondaries on each side, central secondaries arising at

angles of 25°-55°, tertiary veins usually subparallel and

at right angles to the secondaries. Inflorescences 5-15

cm long, axillary to distal leaves or occasionally pseu-

doterminal, broadly (8 cm) paniculate, many flowered

and yellowish brown to reddish brown puberulent, pe-

duncle 1-5 cm long, distal branches of the panicle much

shorter than the basal. Flowers 3-4 mm long, ca. 4 mm

broad, densely short-puberulent, tepals 1.5-2 mm long

and erect at anthesis; outer stamens ca. 1 .2 mm long with

anthers 0.6-0.8 mm long and equally broad or slightly

narrower, thecae superposed, staminodes absent; pistil

ca. 2.5 mm long, ovary 1.5 mm long and narrowed at

the base, stigma discoid. Fruits enclosed within a cup

10-15 mm long and 10-16 mm broad, ca. 6 mm deep

and enclosing '/z-Vj of the fruits, margin entire or with

broad blunt lobes, cuplike and rounded at the base,

brownish, pedicels 1.5-4 mm thick; berry ca. 2 cm long

and 1 cm thick (dried), ovoid-cylindrical or oblong, ob-

tuse at the apex and apiculate, green at maturity.

Trees of wet evergreen forests of the Caribbean

slope and lowlands, from 50 to 1200 m elevation.

Flowers have been collected in April-July; young

fruits have been collected in September-February,

while mature fruits have been collected in April-

May. This species is only known from La Selva,

the Caribbean slopes of the central highlands in

Costa Rica, and a collection from Esmeraldas

Province in Ecuador.

Ocotea hartshorniana is recognized by its large

stature, the obovate-oblong laminae with subpar-

allel tertiary veins, and the usually reddish brown

or yellowish brown puberulence on flowers, inflo-

rescences, and leaves. The stilt roots or buttresses

are also unusual. The young globose "cups" have

a very small opening and enclose nearly all of the

developing fruits. This species is related to Ocotea

sp. aff. caracasana and its allies, a species group

with stilt roots, decurrent lamina bases, and very

similar flowers.

Ocotea helicterifolia (Meissn.) Hemsl., Biol. centr.

amer., Hot. 3: 73. 1882. Oreodaphne helicteri-

folia Meissn. in DC., Prodr. 1 5, pt. 1: 123. 1864.

O. mexicana Meissn., loc. cit. 118.1 864. Phoebe

helicterifolia (Meissn.) Mez, Jahrb. Konigl. Bot.

Gait. Berlin 5: 193. 1889. Phoebe betazensis Mez,

loc. cit. 192. 1889. Figure 8.

Small to medium-sized trees 4-15(-25) m tall, leafy

branchlets 2-5 mm thick, hirsute with slender stiff yel-

lowish hairs to 2.5 mm long, becoming glabrous and

terete. Leaves alternate or occasionally congested at the

ends of branchlets, petioles 4—12 mm long, ca. 1.5 mm

thick, densely hirsutulous; leaf blades (9-)l 2-25(-30) cm

long, (2.5-)4.5-10(-l 2) cm broad, elliptic-obovate to ob-

lanceolate, obovate or pandurate, usually broadest above

the middle, gradually to abruptly narrowed at the apex

and acuminate (rarely rounded or obtuse) at the apex,

the narrowed tip 0-1 5 mm long, rounded to subcordate

or obtuse at the base, often unequal at the petiole, margin

entire, drying membranaceous to stiffly chartaceous and

yellowish green to brown, puberulent on the major veins

above, conspicuously puberulent beneath with slender

brownish hairs to 2-3 mm long, with 6-10(-12) major

secondary veins on each side, central secondaries arising

at angles of 45°-66°, tertiary veins elevated beneath and

often subparallel. Inflorescence axillary from distal leaves

or from leafless nodes, to 18(-25) cm long and pendu-

lous, few-flowered panicles with a long (to 1 3 cm) hir-

sutulous peduncle in ours (occasionally glabrous else-

where), flowers often in umbellate clusters on slender

lateral branches, pedicels 3-6 mm long and very slender.

Flowers 3-6 mm long and 3-5 mm broad, white or yel-

lowish, perianth usually glabrous on the outside and

drying dark; outer anthers 0.8-1.6 m long, 0.7-1.3 mm

broad, elliptic-oblong to ovate, flat, thecae superposed,

the connective sometimes prolonged beyond the thecae,

filament short or absent, inner stamens narrow, stami-

nodes slender to ovate or absent, to 0.8 mm long; pistil

ca. 2 mm long with globose-ellipsoid ovary and thick

style ca. 1 mm long, stigma subcapitate or discoid. Fruits

borne on a long pendulous infructescence, pedicels to 2

cm long and 3 mm thick below the abruptly expanded

cupulate receptacle, cup 3-4 mm long, 7-14 mm broad

and 2-3 mm deep, becoming red; berry ellipsoid 2-2.4

cm long and 1.2-1.7 cm in diameter (dry), becoming

black.

Trees of lowland evergreen rain forests in Costa

Rica, from 20 to 500 m elevation on the Caribbean

slopes, and in the Osa peninsula (as interpreted

here, see below). Flowering material has been col-

lected in February-March; fruits have been col-

lected in August. The species ranges from eastern

and southern Mexico through Guatemala and

Honduras to Costa Rica.

Ocotea helicterifolia (often called Phoebe helic-

terifolia) is a very variable species but usually eas-

ily identified because of the short-petiolate leaves

with long (ca. 2 mm) slender hairs, laminae broad-

est above the middle, acuminate at the apex and

slightly rounded at the base, and the long-pendant

inflorescences with puberulent peduncles, but with

glabrous or sparsely puberulent perianth that dries

black. The relatively large subsessile outer stamens

82

FIELDIANA: BOTANY

are flat and sometimes have their connective pro-

longed distally. The very shallow fruiting recep-

tacle, and usual presence of staminodes (in north-

ern populations) further distinguish this species.

However, there are many problems associated with

the specimens placed here. First, they encompass

an unusual spectrum of variation, especially in

Mexico and Guatemala. Secondly, O. valeriana

may prove to be a higher altitude subspecies, since

it seems to parallel the variation seen in the Gua-

temalan and Honduran highlands of material re-

tained in O. helicterifolia. See the discussion under

O. valeriana and O. valerioides. Nectandra beli-

zensis may also be related, though the form of the

anthers is different.

Ocotea holdridgeiana W. Burger, sp. nov. Figure

12.

Arbor 5-1 5 m alta, ramulis foliiferis 1.5-3.5 mm eras-

sis. Folia alterna, petiolis 8-18 mm longis; laminis 8-

1 5(-l 8) cm longis et 2-5(-6.5) cm latis, ellipticis, ovatis,

elliptico-ovatis vel lanceolatis, apice acuminate, subtus

glabris vel puberulis in nervis, saepe domatiis praeditis,

nervis secondariis 3-6 paribus. Inflorescentiae panicu-

latae racemiformae, 3-9 cm longae, pedunculis usque 4

cm longis. Flores usque 7 mm longi et 10 mm lati, extus

glabri, periantho intus papillato-puberulo; stamina ser

I-II 2-2.8 mm longa, antheris 1.5-2.4 mm longis, stam-

inodiis gracilibus vel nullis; gynoecio 1 .6-2.7 mm longo,

stylo usque 1.2 mm longo. Fructus oblongus vel ellip-

soidus, ca. 25 mm longus et 13 mm crassus; cupula 1

cm lata et 1-2 mm profunda.

Trees 5-15 m tall, leafy branchlets 1.5-3.5 mm thick,

minutely (0.1-0.2 mm) appressed puberulent with slen-

der brownish hairs, becoming longitudinally striate and

dark gray. Leaves alternate, petioles 8-1 8 mm long, 0.8-

1.5 mm thick, sulcate above, sparsely and very minutely

appressed puberulent; leaf blades 8-1 5(-l 8) cm long, 2-

5(-6.5) cm broad, elliptic to elliptic-ovate, elliptic-ob-

long or lanceolate, tapering gradually to the acuminate

apex, the narrowed tip 0.2-1 .2 cm long, acute at the base

and slightly decurrent on the petiole, drying stiffly char-

taceous and dark or yellowish brown, the upper surface

often lustrous and with the minor venation easily visible,

glabrous or minutely puberulent above the major veins,

glabrous or appressed puberulent along the veins be-

neath, with 3-6 major secondary veins on each side, the

basal veins usually strongly ascending (subtripliveined)

or the venation pinnate, central secondaries arising at

angles of 30°-50°, conspicuous pit domatia usually pres-

ent in the axils of the proximal veins beneath, the dom-

atia often raised and visible on the upper surface (dry).

Inflorescences axillary to distal leaves or pseudotermin-

al, 3-9 cm long, few-flowered and racemose, primary

peduncle to 4 cm long, ca. 1 mm thick, minutely ap-

pressed puberulent, pedicels 2-4 mm long. Flowers to 7

mm long and 10 mm broad, white, glabrous on the out-

side, tepals 2.5-5.6 mm long, 1 .4-4 mm broad, papillate-

puberulent on the inner surface; outer stamens 2-2.8 mm

long and with a short broad filament, outer anthers 1.5-

2.4 mm long, oblong, connective often slightly prolonged

beyond the superposed thecae, inner stamens to 3 mm

long, staminodes small (1 mm) and usually slender or

absent; pistil 1.6-2.7 mm long, style to 1.2 mm long,

ovary globose to obovoid, stigma capitate. Fruits borne

on an expanded (1 cm broad) receptacle only 1-2 mm

deep, fruiting pedicels ca. 15 mm long and 2.5-4 mm

thick, gradually expanded at the apex into the shallow

receptacle, red; berry ca. 25 mm long and 13 mm in

diameter, oblong-ellipsoid or slightly oblong-obovoid,

green or tinged with red.

TYPE— Costa Rica, San Jose Province, 0.9 km

above La Chonta and 1 00 m from the Pan Amer-

ican highway, elevation 2460 m, 11 May 1969,

Roy W. Lent 7677(holotype, F 1958283; negative,

6 1 1 24; isotypes, CR, DUKE, MO). Additional Costa

Rican collections are: Brown 17393 (F, LSU), Da-

vidse & G. Herrera 29106 (CR, MO), Gomez et al.

21496 (CR, MO), Gomez- Laurito 1 1077 (CR, F, usj),

Holdridge 6570 (CR, NY), Madriz 34 (CR, F), Poveda

753 (CR, F), and Zamora et al. 879 (CR, F).

Trees of evergreen montane rain forest and

montane wet forest formations, between ( 1 1 00)

1600 and 2700 m elevation. Flowering collections

have been made in February-April, July-Septem-

ber, and November-December. Fruiting material

has been collected in February, April, and No-

vember. This species is only known from the Ca-

ribbean slopes and near the continental divide in

the central highlands and the Cordillera de Tala-

manca; it also occurs in westernmost Panama

(McPherson 9444, MO).

Ocotea holdridgeiana is recognized by the lus-

trous narrow leaves, often somewhat tripliveined,

and with conspicuous pit domatia in basal vein

axils on the lower surface. The large flowers, gla-

brous on the outside, long (2 mm) stamens with

short (0.2-0.4 mm) filaments, connective often

prolonged a little beyond the thecae, and the sub-

capitate stigma are also unusual features. These

characteristics define a very distinctive species,

which appears to be related to O. brenesii. Some

specimens had earlier been identified as Phoebe

tonduzii, but the type of that name appears to be

an unusual variant of P. cinnamomifolia. While

O. holdridgeiana has staminodes, it does not have

three well-developed staminodes in each flower,

as is characteristic of Phoebe. The many collec-

tions of Leslie Holdridge and his astute identifi-

cations have been extremely helpful in under-

standing the Lauraceae of Costa Rica.

BURGER: FLORA COSTARICENSIS

83

Ocotea insularis (Meissn.) Mez, Jahrb. Konigl. Bot.

Gart. Berlin 5: 271. 1889. Phoebe insularis

Meissn. in DC., Prodr. 15, pt. 1: 33. 1864. O.

cuneala Mez, Bot. Jahrb. Syst. 30, Beibl. 67: 17.

1901, non O. cuneata (Griseb.) Gomez, 1894.

O. ira Mez & Pittier, Bull. Herb. Boissier, ser.

2, 3: 232. 1903. 0. tonduzii 'Standley, Publ. Field

Mus. Nat. Hist., Bot. Ser. 18: 456. 1937 (based

on O. cuneata Mez). Aiouea lundelliana C. K.

Allen, J. Arnold Arbor. 26: 419. 1945. Figure

10.

Trees 10-25 m tall, trunks sometimes with short but-

tresses, leafy branchlets 2.5-7(-12) mm thick, glabrous

or minutely (0.1-0.3 mm) appressed puberulent with

slender straight yellowish brown hairs (at least near the

shoot apex), longitudinally ridged but becoming terete,

dark brown to grayish brown. Leaves alternate and usu-

ally close together distally, petioles 0-12 mm long and

often poorly differentiated, flat or slightly sulcate above,

usually with revolute margins; leaf blades (4-)8-20(-28)

cm long, (2-)5-9(-12) cm broad, obovate to spatulate,

elliptic-obovate or obovate-oblong, abruptly rounded at

the bluntly obtuse or short-acuminate apex (sometimes

acute on small leaves), tapering gradually to the cuneate

or attenuate and decurrent base, the margin entire and

slightly revolute (especially at the base), narrowed por-

tion of the lamina base to 2 cm long, drying subcoria-

ceous and dark brown to pale grayish brown (often red-

dish brown in lower-elevation populations), glabrous and

the minor veins obscure above, lower surfaces glabrous

or with thin inconspicuous appressed hairs 0.2-0.3 mm

long, domatia of tufted hairs often present in the vein

axils beneath, with 6-10 major secondary veins on each

side, central secondaries arising at angles of 30°-60°, ter-

tiary venation subparallel but usually obscure. Inflores-

cences 1 0-20 cm long, axillary to distal leaves, broadly

paniculate with flowers usually clustered at the ends of

short (1-3 cm) lateral branches, peduncle 4-10 cm long

and 2-3 mm thick, reddish brown, glabrous or puber-

ulent, pedicels 1-4 mm long and sparsely puberulent.

Flowers densely to sparsely puberulent on the outside,

tepals 1.5-2 mm long, ca. 1.3 mm broad; outer stamens

with short (0.2-0.6 mm) filaments, outer anthers 0.5-1

mm long and equally broad or narrower, ovate to oblong,

thecae superposed, staminodes very rarely present; pistil

1 .5-2.8 mm long, ovary half the length of the pistil, style

slender, stigma discoid. Fruits borne on a short (4-7 mm)

shallow (2-4 mm deep) conical or cupulate receptacle

6-9(-12) mm broad at the top, margin entire or with

persisting perianth parts, the cup abruptly expanded above

the thickened (2-3 mm) pedicel, cup and pedicel ca. 1.5

cm long, becoming red; berry ellipsoid-oblong, 12-20

mm long and 6-10 mm in diameter (dry), dark green to

black at maturity.

Trees of wet evergreen rain forest and cloud

forest formations, from near sea level to 1600

(2000) m elevation. Flowering collections have

been made in all months except October. Fruits

have been collected in all months except Novem-

ber. This species (in a wide sense) probably ranges

from southeastern Nicaragua to Panama, and pos-

sibly as far as coastal Ecuador. It was first collected

on Cocos Island.

Ocotea insularis is recognized by its stiff, essen-

tially glabrous, obovate leaves usually bluntly

rounded at the apex and decurrent on the petiole,

with revolute margins basally. The relatively small

ellipsoid-oblong fruits, borne in small cups with

perianth parts usually persisting on the margin,

also help to identify members of this species. Un-

fortunately, this species shows considerable local

divergence (see below) and appears to be closely

related to Aiouea costaricensis. Ocotea endresiana

may be no more than an unusual form of O. in-

sularis. Difficulties can also be encountered in dif-

ferentiating material of Ocotea sp. aff. caracasana

and its allies (including O. skutchii and O. whitei),

all of which have decurrent leaf bases and rather

similar floral morphology. See the discussions un-

der these species. Ocotea rivularis is also closely

related but it has very large leaves.

The highland populations of Ocotea insularis

seem to be common around Monteverde in the

Cordillera de Tilaran, and a number of specimens

come from the northwestern edge of the Meseta

Central (San Ramon and Zarcero). Only a very

few collections have been made near Volcan Poas

and Volcan Barva (Barba) and on the western side

of Irazu. A few collections have been made south

of Cartago, from Muneco, Orosi, and Cachi, in

very wet forests. The species is apparently more

common in the western part of the Cordillera de

Talamanca, in the Chiriqui highlands, and east-

ward into Code Province, Panama. Among the

collections from the Chiriqui highlands are several

in which the upper thecae of the outer anthers are

reduced or absent. It was such a collection (P.

White 225, MO) which was the basis of the name

Aiouea lundelliana. A similar situation seems to

be found in the flowers of Williams 16344 (us,

usj) from near La Congreja, Cartago. These spec-

imens suggest a trend in loss of the upper thecae,

and they may indicate how Aiouea costaricensis

(q.v.) originated from Ocotea insularis-like ances-

tors. In this regard it is interesting to note that

Aiouea costaricensis and O. insularis appear to be

sympatric only between Volcan Barva (Barba) and

Zarcero, despite living in rather similar forest for-

mations elsewhere. These two species and their

interrelatedness deserve closer study.

The name Ocotea insularis is based on the Cocos

Island population, which differs from most of our

highland material in its larger more elliptic leaves

with acute/obtuse apices and a tendency to dry

84

FIELDIANA: BOTANY

dark. The tufts of reddish hairs along the midvein

on the undersides of the Cocos Island specimens

are not found in any mainland material, though

somewhat similar hairs may be found in the vein

axils. Some collections from the General Valley

closely resemble the Cocos Island material as re-

gards leaf form and color after drying. The lower-

elevation collections from the Caribbean slope also

have larger ( 1 2-28 cm) leaves, but they are often

slightly lustrous above, and are more reddish brown

beneath; in addition, the fruit cups are often entire.

The following key summarizes some of this vari-

ation. It is clear that the problems in this group

of species can only be solved after further study.

We have decided to treat this group conservatively

and place collections possibly belonging to O. ira

and O. tonduzii under O. insularis sensu lato. We

feel that without a thorough study of collections

from Panama, coastal Colombia, and coastal Ec-

uador no better classification can be presented at

this time.

Key to Possible Subdivision of Ocotea insularis

la. Leaves elliptic to obovate and often bluntly acute or acuminate at the apex, usually drying dark

brown, with tufts of reddish brown hairs along the midrib on the lower leaf surface; a dominant

tree of Cocos Island O. insularis s.s.

Ib. Leaves usually obovate to oblanceolate with the larger laminae usually rounded at the apex, never

with tufts of reddish hairs along the length of the midvein (but sometimes in the vein axils); trees

of the mainland 2a

2a. Leaves drying darker brown or reddish brown, 15-25 cm long; fruits 15-25 mm long and borne in

a cup with an entire margin; from below 1 000 m elevation O. ira

2b. Leaves usually drying pale grayish brown, 5-20 cm long; fruits 12-18 mm long and usually borne

in a cup with some persisting perianth lobes; above 1000 m in the Cordillera de Tilaran and around

the central highlands O. tonduzii

Ocotea laetevirens Standl. & Steyerm., Publ. Field

Mus. Nat. Hist., Bot. Ser. 23: 1 14. 1944. Figure

12.

Trees 4-18 m tall, leafy branchlets 2-8 mm thick,

strongly 1-3-ridged from early stages (strongly angular

in cross section) but becoming terete, glabrous or with

minute (0.2 mm) thin ascending hairs near the shoot

apex, greenish (dry) or brown. Leaves alternate, petioles

8-22 mm long, 1.2-2.5 mm thick, with adaxial ridges

forming a sulcus; leaf blades 7-1 8 cm long, 3-8 cm broad,

elliptic to elliptic-obovate or elliptic-oblong, acute to

short-acuminate (occasionally bluntly obtuse) at the apex,

tapering gradually to the obtuse or acute base and slightly

decurrent on the petiole, margin slightly revolute when

dry, drying stiffly chartaceous to subcoriaceous and yel-

lowish green to greenish brown, glabrous and slightly

lustrous with the tertiary venation visible on the upper

surface, glabrescent and duller beneath with the minor

venation raised and forming a poorly denned reticulum,

with 3-7 major secondary veins on each side, the central

secondaries arising at angles of 30°-60°, the basal sec-

ondaries occasionally prominent and arcuate-ascending,

domatia-like developments of tufted hairs or pits often

present in the axils of proximal secondary veins beneath.

Inflorescences axillary to both distal and proximal leaves,

to 20 cm long, an open many-branched panicle, peduncle

3-12 cm long and ca. 1.3 mm thick, glabrous, pedicels

ca. 2 mm long. Flowers 2.5-4 mm long, 2.5-4 mm broad,

campanulate, white but drying dark, glabrous on the

outside or minutely papillate distally, tepals ca. 2 mm

long and 1.7 mm broad; outer stamens 1.4-1.8 mm long

with oblong or ovate anthers 0.7-0.8(-1 . 1) mm long and

0.6-0.8(-0.9 ) mm broad, filaments usually prominent

(0.5-1 mm) and broad, thecae clearly superposed, inner

stamens 1.5-2 mm long and with large glands, stami-

nodes absent or very slender (to 0.8 mm long); pistil 1.7-

2.5 mm long, ovary 1-1.3 mm in diameter, style 0.7-

1 . 1 mm long, stigma simple or oblique. Fruits borne in

a cupulate or conic receptacle 1 .2-2.5 cm long, 1-1 .6 cm

broad and 3-6 mm deep, gradually expanded above the

thickened pedicel, surface rough but lacking prominent

lenticels, becoming red; berry 2.2-3 cm long, 1.3-1.8 cm

in diameter (dry), ellipsoid, becoming black.

Trees of montane evergreen forest formations,

from 1200-2000 m elevation. In Costa Rica flow-

ering material has been collected in March-April,

July, and December; fruits have been collected in

March-April, June-August, and December-Jan-

uary. This species has only been collected from

the northeastern portion of the Meseta Central

(Volcan Barva to Tablazo) and the western part

of the Cordillera de Talamanca in Costa Rica. The

species is also known from Guatemala.

Ocotea laetevirens is distinguished by its stiff

BURGER: FLORA COSTARICENSIS

85

glabrous leaves that often turn yellowish green or

pale brown when dried, the slightly decurrent lam-

ina base, the almost glabrous flowers, the outer

stamens with prominent filaments, poorly devel-

oped stigma on a conspicuous style, shallow fruit

cup, and ellipsoid fruits. We follow C. K. Allen

(1945) and Hammel (1986) in using this name for

these Costa Rican plants. Even though the leaves

of the type (Steyermark 49189, F, from Guate-

mala) are thinner and much more acuminate than

in Costa Rican material, the flowers are nearly

identical. Other highland species with stiff gla-

brous pale-drying leaves that are often glossy above

and can be confused with this species are Ocotea

meziana, Nectandra cufodontisii , and Phoebe

hammeliana.

This species is poorly understood at present. It

may be much more variable than described above,

with some individuals having smaller, more co-

riaceous leaves. Two larger-leaved (20-30 cm) col-

lections, with longer more open pendant infruc-

tescences, from the Osa peninsula (Primack 96,

MO) and Mastatal de Puriscal (Gomez- Laurito

11237, CR, F, usj) are tentatively placed under this

species. (Note that these last two collections are

both from lower elevation habitats, and they may

not be conspecific.) It seems best to place these

latter anomalous collections under O. laetevirens

until we have better samples of the populations

they represent. Ocotea verapazensis Standl. &

Steyerm. and O. standleyi C. K. Allen (based on

Phoebe macrophylla Standl. & Steyerm.) may prove

to be synonymous with an expanded concept of

O. laetevirens; all the relevant types are from ev-

ergreen forests at about 1 500 m elevation in Gua-

temala. See also Ocotea sp. aff. O. laetevirens at

the end of the species descriptions.

Ocotea lentil W. Burger, sp. nov. Figure 2.

Arbor 4-8 m alta, ramulis foliiferis 5-10 mm crassis.

Folia alterna, petiolis 8-30 mm longis et 2.5-4 mm cras-

sis; laminis 18-40 cm longis et 8-16 cm latis, obovatis

vel elliptico-obovatis, apice breviter acuminato vel ro-

tundato, basi cuneato, subtus strigosis in nervis, pilis

0.2-1 mm longis, nervis secondariis 7-1 1 paribus. In-

florescentiae paniculatae, 8-20 cm longae, saepe racem-

iformae, pedunculis 3-12 cm longis. Flores 5-7 mm lon-

gi et 6-10 mm lati, extus glahri vel minute puberuli,

tepalis intus papillato-puberulis; stamina ser. I-II an-

theris 1 .2-1 .4 mm longis, staminodiis nullis vel parvulis

(0.6 mm); gynoecium ca. 2.5 mm longum, ovario ellip-

soideo, 1.5-1.9 mm longo. Fructus ellipsoideus vel ob-

longoideus, 3-6 cm longus et 2-3 cm crassus; cupula 8-

25 mm lata, 2-8 mm profunda.

Small trees 4-8 m tall, leafy branchlets 5- 1 0 mm thick,

hirsute with straight stiff yellowish hairs 0.5-1.3 mm

long and persisting on the terete stems. Leaves alternate

or crowded together distally, petioles 8-30 mm long, 2.5-

4 mm thick, densely hirsute with stiff hairs, narrowly

sulcate above; leaf blades 1 8-40 cm long, 8-16 cm broad,

obovate to elliptic-obovate, abruptly narrowed to the

short-acuminate or rounded apex, gradually tapering to

the cuneate base, margin often undulate (dry), the lam-

inae drying chartaceous to stiffly chartaceous and dark

greenish or brown above, densely and minutely (0.3 mm)

puberulent on the major veins above and with longer

(ca. 1 mm) hairs between the veins, lower surface with

stiff straight hairs 0.2-0.8(-1.2) long on both the major

and minor veins, with 7-1 1 major secondary veins on

each side, central secondaries arising at angles of 40°-

70°, secondary and tertiary venation prominent beneath,

tertiary veins often subparallel and perpendicular to the

secondaries. Inflorescences axillary to distal leaves or

from leafless nodes, 8-20 cm long, elongate panicles with

short lateral branches (almost racemiform), primary pe-

duncle 3-12 cm long, slender (1.5 mm) and sparsely

hirsute, pedicels to 10 mm long. Flowers 5-7 mm long,

6-10 mm broad, perianth glabrous or sparsely puberu-

lent on the outside, papillate-puberulent on the inside;

outer stamens subsessile, anthers 1.2-1.4 mm long, ca.

1.2 mm broad, thecae superposed but the lower often

somewhat lateral, staminodes absent or small (0.6 mm)

and clavate, with erect straight hairs around the rim of

the floral tube and base of the stamens; pistil ca. 2.5 mm

long, ovary 1.5-1.9 mm long, ellipsoid, style slender,

stigma capitate. Fruits borne on a flat, conical or broadly

cupulate receptacle, 4-10 mm long, 8-25 mm broad and

2-8 mm deep, pedicels becoming 2-4 cm long and 2.5-

6 mm thick; berry 3-6 cm long, 2-3 cm in diameter,

ellipsoid, the base not tightly enclosed in the cup when

a cup is present (dry).

TYPE— Costa Rica, Cartago Province, hillside

overlooking the Rio Grande de Orosi, 3.5 km SE

of Tapanti, elevation 1400 m, 9 April 1967, Roy

W. Lent 794 (holotype, F 1749021; negative,

61119; isotypes, CR, MO).

Small trees of the very wet cloud forest for-

mations of the Caribbean slope, at about 1400 m

elevation. Rowers have been collected in April,

while fruits have been collected in April and Au-

gust. At present the species is known from only

three collections (Gomez 18814, CR, MO, Lent 794,

2070, CR, F), all from near Tapanti, along the Rio

Grande de Orosi.

Ocotea lentii is recognized by its very large but

thin-textured obovate leaves with short straight

stiff hairs, long pedunculate inflorescences with

relatively few flowers, and large ellipsoid fruits.

The upper edge of the floral tube with straight

lustrous hairs, and the subsessile anthers about as

broad as long, are also distinctive features. The

general form of the flowers and vegetative parts

relate this species to Ocotea valeriana, but the very

86

FIELDIANA: BOTANY

large leaves and fruits of O. lentil are amply dis-

tinct. Ocotea valerioides, with large leaves and very

similar floral structure, is more closely related. It

too, has a tendency for the outer anthers to become

broad and develop a Nectandra-\ike configuration.

Ocotea lentii and O. valerioides differ in elevation,

leaf texture, and color (dried), as well as vesture,

inflorescence-form, and size of the fruits. See the

discussions under O. valerioides and O. mollifolia.

Roy Lent's Costa Rican collections number about

4,000 and include many rare and important finds,

of which this unusual species is an example.

Ocotea leucoxylon (Sw.) Laness., PI. util. colon,

franc. 156. 1886. Laurus leucoxylon Sw., Prodr.

65. 1 788. 0. subsericea Standl., Publ. Field Mus.

Nat. Hist., Hot. Ser. 18: 456. 1937. O. lenticel-

lata Lundell, Wrightia 5: 54. 1974. Figure 14.

Small or medium-sized trees, 3-15(-25 m) tall, dioe-

cious (unisexual), leafy branchlets 1.7-6.5 m thick, gla-

brous or with very minute (0.05-0.1 mm) grayish ap-

pressed ascending hairs, often with longitudinal ridges

but becoming terete. Leaves alternate and evenly spaced,

petioles 10-22 mm long, 1.5-2.5 mm thick, with ele-

vated adaxial ridges usually forming a shallow sulcus

above; leaf blades 10-23(-27) cm long, 3-9 cm broad,

elliptic-oblong to oblong or elliptic-lanceolate, usually

acuminate at the apex with a narrow tip 0.5-2 cm long,

acute to obtuse at the base, margin usually slightly re-

volute, the laminae drying very stiffly chartaceous to

subcoriaceous and grayish green to dark olive green, gla-

brous above and the midvein flat, essentially glabrous

(in ours) or very sparsely and minutely puberulent be-

neath, with 4-8 major secondary veins on each side,

primary and secondary veins prominent beneath, central

secondaries arising at angles of 30°-50°(-600), epidermal

cells somewhat convex on the lower surface and giving

an uneven or scale-like effect (10 x). Inflorescences sol-

itary from the axils of leaves or leafless nodes throughout

the length of new twigs, 4-9(-13) cm long, paniculate

with well-spaced lateral branches, basal branches longer

than the distal, pedicels 1-3 mm long. Male flowers ca.

2.5 mm long and 2.5 mm broad (to 3 x 3 mm elsewhere),

white to yellowish, tepals 1.5(-2) mm long and 1 mm

broad, very minutely papillate puberulent on the edges;

outer stamens with anthers 0.5-1 mm long and 0.5-0.9

mm broad, with short (0.2-0.3 mm) broad filaments,

thecae superposed or the lower slightly lateral to the

upper, inner stamens ca. 1.5 mm long with anthers 0.9

mm long, staminodes absent; pistillode ca. 2 mm long

and 0.2-0. 5 mm thick, stigma discoid. Female flowers

not seen (probably very similar to the male). Fruits sub-

tended by a broadly obconic or saucer-like receptacle 4-

10 mm long, 5-8 mm broad, and 1-3 mm deep, reddish

brown, often with distinctive wart like lenticels, rim en-

tire; berry 8-10 mm in diameter (dry, to 1 5 mm in life),

globose to globose-obovoid, green to purple, the fruiting

inflorescences rarely more than 8 cm long.

Trees of tropical wet to premontane rain forest

formations on the Caribbean slope and adjacent

continental divide, from 20 to 1 800 m elevation.

The trees flower in February-July at La Selva

(Hammel 1986, p. 229), and fruits have been col-

lected throughout the year, except November, in

Costa Rica. The species ranges from southern

Mexico and the West Indies to South America.

Ocotea leucoxylon is recognized by it stiffoblong

glabrous leaves, strongly angular young stems,

small functionally unisexual flowers, and small

globose fruits, subtended by a small cupulate or

obconic warty receptacle. Hammel (1986) men-

tions the foul-smelling foliage (when crushed), the

distantly spaced inflorescences, grayish green leaves

and scaly lower leaf surface as additional distin-

guishing features. The "scaly effect" on the lower

leaf surface may simply be the reflections of light

from the convex surfaces of epidermal cells (in

life); there are no scales under the microscope

( 1 00 x ). We follow Hammel ( 1 986), who included

O. lenticellata from Belize under this species. This

species has also been cited as O. leucoxylon (Sw.)

Gomez de la Maza (Dice. Nombres Vulg. Cub.

Puerto Riq. 12.), but Howard (1981) states that

this publication was issued in 1889, and the day

and the month are unknown. The type of Stan-

dley's O. subsericea (Brenes 6789, CR, F) has im-

mature foliage and larger more puberulent flowers;

it probably belongs in synonymy here.

Ocotea meziana C. K. Allen, J. Arnold. Arbor. 26:

360-361. 1945. Figure 14.

Trees 2-1 5 m tall, trunk to 80 cm d.b.h., bark grayish

or brown, leafy branchlets 1.3-4 mm thick, at first with

short (0.2 mm) ascending yellowish hairs but quickly

becoming glabrous and pale grayish to yellowish green.

Leaves alternate, larger in low elevation plants than in

montane plants (see below), petioles 5-1 6(-30) mm long,

1-2 mm broad, flat or slightly sulcate above; leaf blades

5-15 or 1 1-22 cm long, 2-6 or 4-10 cm broad, elliptic

to narrowly elliptic or elliptic-oblong (broadly elliptic at

lower elevations), tapering gradually to the short-acu-

minate apex and acute base (tapering more abruptly in

the broader leaves of lower elevations), drying stiffly

chartaceous and gray or greenish gray, smooth and gla-

brous above with the venation flat, essentially glabrous

beneath, with 4-6 major secondary veins on each side,

the central secondaries arising at angles of 30°-60°, with

pit domatia in the axils of major veins or along the

secondary veins, the smallest veins forming a distinct

reticulum on the lower surfaces with areolae about 0.5

mm broad. Inflorescences axillary or extra-axillary on

distal twigs, to 10(-15) cm long, few-branched panicles,

peduncle 1.5-6 cm long, glabrous or very minutely pu-

berulent, lateral branches with few (3-7) flowers, pedicels

BURGER: FLORA COSTARICENSIS

87

2-5 mm long and articulate near the base, drying dark.

Flowers ca. 2-3 mm long, 4 mm broad, yellowish green

but drying black, essentially glabrous, tepals 1 .2-2 mm

long; outer stamens with prominent (ca. 1 mm) filaments,

outer anthers 0.7-1 mm long, narrow with superposed

thecae, staminodes absent; pistil 1 .5-2.5 long, ovary tur-

binate or narrowed near the base, style ca. 1 mm long,

stigma simple. Fruits borne on a shallow (ca. 2-3 mm

cup) cup 8-14 mm broad, 1.5-3 cm long, funnelform

and often abruptly flared at the apex, red; berry 2-3 cm

long, 1.3-2 cm in diameter (dried), becoming purple or

black.

Trees of the wet evergreen forest formations of

the Caribbean slope and the continental divide,

from 50 to 2300 m elevation. Flowering collec-

tions have been made in February-July, Septem-

ber-October, and December. Fruiting material has

been collected in July-December. The species, as

presently understood, ranges from northern Costa

Rica to Code Province in Panama.

Ocotea meziana is recognized by the few-flow-

ered and essentially glabrous inflorescences with

small flowers that dry blackish, the glabrous (or

very minutely puberulent) leaves that dry grayish

or greenish and slightly lustrous beneath, and the

shallow funnelform cupules about 1 cm broad at

the apex. This treatment follows Hammel's ( 1 986)

interpretation, and includes the larger-leaved

specimens from La Selva, together with the small-

er-leaved highland (800 to 2100 m) material. Pit

domatia along the secondary veins are often pres-

ent in highland material, as in the type collection

(A. Smith 359, F) from about 1600 m elevation.

Ocotea meziana is closely related to O. laetevirens

(q.v.). Herbarium material ofO. meziana may re-

semble Ocotea cernua, O. tenera, Licaria cufo-

dontisii, and L. sarapiquensis , but the fine retic-

ulum formed by the minor venation on the lower

surface when dry helps to separate O. meziana

from all of these other species.

on each side, the central secondaries arising at angles of

30°-50°. Inflorescences small and axillary to older or

fallen leaves, racemose panicles to 6 cm long, peduncles

to 4 cm long, slender (0.5 mm) and grayish puberulent,

flowers solitary or 2-3 on short branches of the rachis,

pedicels 1-5 mm long. Flowers 4-6 mm broad, buds 2-

3 mm long, tepals ca. 3 mm long, obtuse, glabrous dis-

tally and within, pellucid punctate; outer stamens ca. 1 .3

mm long with anthers ca. 0.8 mm long and 0.9 mm

broad, ovate with superposed thecae, inner stamens with

oblong anthers ca. 0.7 mm long, staminodes small (0.5-

0.8 mm) and with or without a glandlike apex; pistil ca.

2 mm long, ovary globose and glabrous, style ca. 0.6 mm

long, stigma simple or slightly discoid. Fruits borne on

a short cup ca. 6 mm broad, usually with the broad but

short tepals persisting at the edge, becoming red; berry

ca. 2 cm long and 1.3 cm in diameter, ellipsoid, becoming

black.

Trees of wet evergreen montane forest forma-

tions, between 1400 and 2300 m elevation in cen-

tral Costa Rica. Flowering collections have been

made in December-February; fruits have been

collected in May-August. This species is only

known from the region around Zarcero, the slopes

of Volcan Irazu, and the western parts of the Cor-

dillera de Talamanca (Copey and Sta. Maria de

Dota) in Costa Rica.

Ocotea mollicella is recognized by its small nar-

row leaves with soft grayish puberulence beneath,

small inflorescences that become dark on drying,

presence of staminodes, and restricted highland

habitat. There is little good fruiting material, and

the fruiting description may require revision. This

species is known as quizarra amarillo. This species

is very closely related to Ocotea pittieri, and there

is the possibility that O. mollicella is only an un-

usual subspecific element of that species. How-

ever, the soft grayish puberulence is quite char-

acteristic and intermediate collections are not

apparent. The two species are here transferred from

Phoebe to Ocotea; see the discussion under O. pit-

tieri.

Ocotea mollicella (Blake) van der Werff, comb,

nov. Phoebe mollicella Blake, Contr. Gray Herb.

52: 64. 1917. Figure 4.

Trees 6-16 m tall, distal branchlets 0.7-3 mm thick,

usually pale grayish or yellowish brown tomentulose with

thin hairs 0.1-0.3 mm long, terete, becoming slightly

striate. Leaves alternate, petioles 6-12 mm long, ca. 1

mm thick, flat above, puberulent; leaf blades 3-7(-9) cm

long, l-2.3(-3) cm broad, narrowly elliptic to elliptic-

lanceolate or elliptic-oblong, tapering gradually to the

acute or acuminate apex, acute at the base, drying char-

taceous, usually dark and slightly lustrous above, gla-

brescent above, grayish puberulent below with slender

hairs 0. 1-0.4 mm long, with 3-6 major secondary veins

Ocotea mollifolia Mez & Pittier, Bull. Herb. Bois-

sier, ser. 2, 3: 233. 1903. Figure 7.

Small and medium-sized trees 4-20 m tall, prop roots

often present (fide Hammel), leafy branchlets 2-6 mm

thick, densely puberulent with yellowish brown to orange

brown hairs 0.2-0.5 mm long. Leaves alternate or oc-

casionally subopposite, petioles 4-12(-18) mm long, 1.5-

3.5 mm thick, densely puberulent; leaf blades (9-) 12-

24 cm long, 6-14 cm broad, obovate to broadly elliptic-

obovate or oblong, nearly always broadest above the

middle, usually rounded and abruptly narrowed to an

acute or short-acuminate apex with a tip 0. 5- 1 . 5 cm long,

narrowed gradually to an acute or obtuse base (never

88

FIELDIANA: BOTANY

decurrent), drying chartaceous to stiffly chartaceous, pu-

berulent on the larger veins above, puberulent beneath

with straight or crooked slender hairs to 0.5 mm long,

yellowish brown to dull brown in color, with 5-9 major

secondary veins on each side, central secondaries arising

at angles of 30°-60°, tertiary veins prominent beneath

and often subparallel and perpendicular to the second-

aries. Inflorescences axillary or apparently extra-axillary,

6-20(-30) cm long, an open panicle, primary peduncle

to 14 cm long and 1 .5-2.5 mm thick, densely puberulent,

pedicels 2-9 mm long. Flowers 3-4(-5) mm long, 3-5(-7)

mm broad, cream white but becoming brownish on

drying, tepals 1.5-2.2(-3) mm long, densely puberulent

on the outside; outer stamens subsessile with ovate to

oblong anthers 0.7-1 .4 mm long, thecae superposed and

the connective sometimes prolonged apically, inner sta-

mens 1-1.8 mm long, staminodes present and slender

(ca. 0.8 mm long) or absent; pistil 1.5-2.5 mm long,

ovary turbinate or ellipsoid, style short to long ( 1 .4 mm)

and glabrous or puberulent (as in the type), stigma simple

or slightly discoid. Fruits borne on a conical cup 10-18

mm long and ca. 1 5 mm broad, shallow or deep (2-5

mm), rim of the cup usually with broad (3 mm) persisting

perianth parts 1-2 mm long, becoming red; berry 2-3(-4)

cm long, 1.5-2 cm in diameter, oblong-cylindrical.

Trees of stream sides and forest understory usu-

ally on hilly sites, on the lower slopes of the Ca-

ribbean escarpment between 50 and 900 m ele-

vation, and near Palmar Sur de Osa on the Pacific

slope. Flowering material has been collected in

January-March, and flowering has been observed

in February-June at La Selva (Hammel 1986).

Mature fruits have been collected in September-

January. This species is only known from Costa

Rica.

Ocotea mollifolia is recognized by the conspic-

uously puberulent and usually obovate lamina that

are thin in texture and, while narrowed gradually

to the base, are not decurrent on the petiole. The

open inflorescences with widely spaced flowers,

dense puberulence on most parts, fruit cups with

persistent lobes on the margins, and prop roots

near the base of the trunk are further distinguishing

characters. Despite obvious differences between

typical members of each species, this species may

be related to O. gomezii; collections from around

500 to 700 m exhibit some shared characteristics

(but they are not clearly intermediate as one would

expect from a cline or hybridization). In this regard

Burger et al. 11730 (CR, F, NY) from about 600 m

has much larger flowers and floral parts than our

other, lower-elevation, collections of O. mollifolia.

While these flowers approach those of O. gomezii

in size, the large open inflorescences are more typ-

ical O. mollifolia. The prop roots and other char-

acteristics suggest a relationship with O. hart-

shorniana and O. sp. aff. caracasana.

Ocotea monteverdensis W. Burger, sp. nov. Fig-

ure 4.

Arbor usque 25 m alta, ramulis foliiferis 2-5.5 mm

crassis. Folia alterna, petiolis 2-8 mm longis; laminis 4—

12 cm longis et 1.5-4 cm latis, ellipticis vel anguste

elliptico-oblongis, apice saepe acuto, basi acuto vel cu-

neato, decurrente, subtus adpresse puberulis, nervis se-

condariis 3-8 paribus. Inflorescentiae paniculatae, 6-12

cm longae, pedunculis 2-5 cm longis. Flores 3-3.5 mm

longi et 4-5 mm lati, extus dense puberuli, tepalis intus

papillato-puberulis; stamina ser. I-II antheris 0.6-0.9

mm longis, staminodiis nullis; gynoecium 1.5-2.2 mm

longum. Fructus ellipsoideus, usque 3 cm longus et 1.5

cm crassus; cupula 12-16 mm lata et 1-3 mm profunda.

Trees to 25 m tall, to 55 cm d.b.h., leafy branchlets

2-5.5 mm thick, at first yellowish brown or orange brown

tomentulose with dense short (ca. 0.2 mm) tomentulose

hairs, with prominent longitudinal ridges but becoming

terete and grayish. Leaves alternate, often closely clus-

tered distally, petioles not clearly differentiated, 2-8 mm

long, with lateral margins and flat above; leaf blades 4-

12 cm long, 1.5-4 cm broad, elliptic to narrowly elliptic

or narrowly elliptic-oblong, tapering gradually to an acute

(rarely obtuse) apex, tapering gradually to the acute (ob-

tuse or cuneate) base and decurrent on the petiole, mar-

gin slightly revolute near the base, the laminae drying

stiffly chartaceous or subcoriaceous, grayish green or

grayish brown above with the tertiary veins paler and

distinct, smooth above and essentially glabrous except

above the midvein, grayish beneath with a dense cov-

ering of small (0.2 mm) slender appressed ascending hairs,

glabrescent in age, with 3-8 major secondary veins on

each side, central secondaries arising at angles of 30°-

60°. Inflorescences axillary to distal leaves or pseudo-

terminal, 6-12 cm long, paniculate and many-flowered,

peduncles 2-5 cm long, ca. 2 mm thick, brownish pu-

berulent, pedicels 1-3 mm long. Flowers 3-3.5 mm long,

4-5 mm broad, densely ferruginous puberulent on the

exterior, tepals 1.5-2 mm long and ca. 1.5 mm broad,

papillate puberulent within; outer stamens 1.3-1.5 mm

long, subsessile or with short (0.3 mm) puberulent fila-

ments, outer anthers 0.6-0.9 mm long, 0.6-0.7 mm broad,

rectangular, thecae superposed or the lower slightly lat-

eral, inner stamens ca. 1.6 mm long with anthers ca. 0.8

mm long, glands large, staminodes not seen; pistil 1.5-

2.2 mm long, style 0.5-0.7 mm long and slender, stigma

simple. Fruits borne in a shallow (1-3 mm) cup 12-16

mm broad, expanded gradually from the pedicel and

obconic or somewhat saucer-shaped, red, fruiting pedicel

4-10 mm long and ca. 3 mm thick; berry becoming as

much as 3 cm long and 1.5 cm in diameter (dry), ellip-

soid, green.

TYPE— Costa Rica, Puntarenas Province, Mon-

teverde, in Figuerola's pasture, elevation about

1450 m, 27 July 1977, Gary S. Hartshorn 1900

(holotype, CR 91924; isotypes, F 1866719; nega-

tive, 61120, MO).

Trees of montane cloud forest formations in the

Cordillera de Tilaran and westernmost slopes of

BURGER: FLORA COSTARICENSIS

89

the Meseta Central, from about 800 to 900 m near

San Ramon to 1400 to 1600 m at Monteverde.

Flowering material has been collected in June-

August, and fruiting material has been collected

in July-August. This species is only known from

west-central Costa Rica.

Ocotea monteverdensis is recognized by the

smaller elliptic leaves that are densely appressed

puberulent beneath, the decurrent lamina base, the

ferruginous puberulence on stems and inflores-

cences, the larger ellipsoid fruits borne in shallow

cups, and the restricted range. Specimens of this

species may resemble Ocotea mollicella, but that

species lacks the dense pubescence, the decurrent

leaf base, and differs in flowers and fruits. Ocotea

monteverdensis appears to be most closely related

to O. skutchii., O. hartshorniana, and their allies.

The following collections are placed here: Brenes

3849 (F, NY), Dryer 1332, 1591 (CR, F), Gentry &

Haber 48736 (MO), Poveda 1108 (CR, F), J. & K.

Utley 5407 (DUKE, F), and N. Wheelwright 1, BOB,

33, 75 (all MO).

Ocotea nicaraguensis Mez, Jahrb. Konigl. Bot.

Gait. Berlin 5: 238. 1889. O. pentagona Mez,

Bot. Jahrb. Syst. 30, Beibl. 67: 17. 1901. Fig-

ure 3.

Slender trees 6-8(-18) m tall, trunk 10-20 cm d.b.h.,

leafy branchlets 4-8 mm thick, strongly angled in cross

section with 3-5 prominent longitudinal ridges, very mi-

nutely appressed puberulent or glabrous, distal stems

usually hollow and harboring ants. Leaves alternate, pet-

ioles 5-1 2(-20) mm long, 2-4 mm thick, slightly or deep-

ly sulcate above, often drying dark; leaf blades 1 4-40(-5 5)

cm long, 4.5-12(-15) cm broad, very narrowly obovate

to oblong-obovate, oblanceolate or oblong-elliptic,

rounded to bluntly obtuse or short-acuminate at the apex,

tapering gradually to the obtuse (rounded) to cuneate

(but not decurrent) base, drying subcoriaceous to cori-

aceous and often pale grayish brown, glabrous above and

below, midvein up to 5 mm broad beneath, with 7-

10(-12) major secondary veins on each side, central sec-

ondaries arising at angles of 30°-50°(-650), tertiary veins

often perpendicular to the secondaries and subparallel,

slightly raised on the lower surface, domatia absent. In-

florescences axillary to distal leaves, 1 5-30 cm long, pan-

iculate but with short (2-3 cm) lateral branches and

somewhat racemose in form, peduncle 7-12 cm long, 2-

3 mm thick, very minutely puberulent or glabrous, ped-

icels 1.5-5 mm long. Flowers 2.2-3 mm long, 3-5 mm

broad, very minutely puberulent or glabrous, tepals ovate

and thin, usually glabrous within; outer stamens with

short (0.5 mm) filaments and narrow anthers ca. 0.9 mm

long, thecae superposed, inner stamens 1 .4-1 .8 mm long,

staminodia absent; pistil 2-2.4 mm long, the style nar-

rowed gradually and equalling the ovary in length, stigma

slightly discoid. Fruits borne on a shallow (1-2 mm deep)

receptacle ca. 8 mm broad, with the perianth bases (2-

3 mm broad and 1-2 mm long) usually persisting, pedicel

becoming 5-1 5 mm long and 3-4 mm thick at the apex,

becoming reddish; berry 12-25 mm long and 6-12 mm

thick (dried), ellipsoid to ovoid, becoming purple or black.

Trees of evergreen forest formations below 1 000

(71300) m elevation, on both the Caribbean and

Pacific slopes of Costa Rica. Flowering collections

have been made in August-October, and fruiting

material has been collected in January-April and

November. The species appears to be most com-

mon in the Caribbean lowlands, the Cordillera de

Tilaran (600 to 1000 m), the General Valley, and

the Golfo Dulce region. This species ranges from

southeastern Nicaragua to westernmost Panama.

Ocotea nicaraguensis is recognized by its large,

stiff, often oblanceolate leaves on short thick pet-

ioles, the hollow distal stems, lack of conspicuous

puberulence, and the small fruiting receptacles with

(usually) persisting perianth bases. The prominent

longitudinal ridges on the young stems are also

useful in identification, and separating this species

from O. atirrensis, with much thinner leaves that

are often caudate-acuminate and dry dark in color.

Ocotea paulii of higher elevations also lacks the

prominently ridged stems and has smaller more

oblong leaves. All three of these species have the

distal stems hollow and often inhabited by small

ants. Ocotea aurantiodora has similar leaves and

solid stems with even more prominent ridges, but

the fruits are very different.

Ocotea oblonga (Meissn.) Mez, Jahrb. Konigl. Bot.

Gait. Berlin 5: 366. 1889. Mespilodaphne ob-

longa Meissn. in DC., Prodr. 15, pt. 1: 107.

1864. O. portoricensis Mez, loc. cit. 364. 1889.

Phoebe mayana Lundell, Amer. Midi. Natural-

ist 29: 473. 1943. Figure 11.

Trees 8-30 m tall, 20-45 cm d.b.h., dioecious, often

becoming buttressed in age, sometimes with whorled

branching, leafy branchlets 1.5-4 mm thick, glabrous or

very minutely (0. 1 mm) puberulent near the apex, lon-

gitudinally ridged but becoming terete and lenticellate,

dark or grayish. Leaves alternate, petioles 1-2 cm long

but poorly differentiated, 1-2 mm broad, flat above, gla-

brous or minutely puberulent; leaf blades 5-15(-22) cm

long, 2-4(-6) cm broad, oblanceolate to narrowly ellip-

tic-obovate or narrowly obovate, usually short-acumin-

ate at the apex (bluntly acute to obtuse), tapering very

gradually to the cuneate or decurrent base, drying char-

taceous to stiffly chartaceous and usually very dark above

with the major veins slightly raised and the minor veins

90

FIELDIANA: BOTANY

obscure, grayish or brownish beneath and glabrous or

obscurely appressed puberulent with small (0.2-0.4 mm)

thin hairs, with 5-8 major secondary veins on each side,

the central secondaries arising at angles of 30°-50°, pit

domatia often present near the mid vein beneath. Inflo-

rescences axillary to distal leaves, 5-13 cm long, race-

mose panicles with short (1-2 cm) lateral branches, pe-

duncle 8-30 mm long, 0.7-2 mm thick, sparsely to densely

puberulent, pedicels 1-3 mm long. Flowers functionally

unisexual, 1.7-2.5 mm long, ca. 3.5 mm broad, cam-

panulate, minutely and densely appressed puberulent on

the outside, tepals 1.2-1.5(-2) mm long, 1.2-1.4 mm

broad, papillate puberulent within; male flowers with

outer anthers ca. 1 mm long on filaments ca. 0.4 mm

long, thecae superposed in oblong anthers, inner stamens

ca. 2 mm long with small (0.5 mm) glands, staminodes

absent; female flowers with stamens half the size of those

in the male flowers; pistil 1.5-2 mm long, ovary ovoid

and ca. 0.7 mm in diameter, style thick and ca. 1 mm

long, stigma a flat-topped disc. Fruits borne on a flat or

saucer-like receptacle 4-6 mm broad and only 1-2 mm

long, abruptly expanded above the short (3-8 mm) thick-

ened (3-4 mm) pedicel (the thickened pedicel may look

like a cup in early stages); berry 9-18 mm long, 7-9 mm

in diameter (dry), ellipsoid to ovoid or oblong.

Trees of evergreen forest formations of the Ca-

ribbean slope and partly deciduous formations of

the General Valley, between 600 and 1000 m el-

evation in Costa Rica; from near sea level to 1 000

m elsewhere. Flowers have been collected in May,

August, and October in Costa Rica and July-Au-

gust in central Panama; fruits have been collected

in October-early February. This species ranges

from Belize and the West Indies to Bolivia and

the Guianas.

Ocotea oblonga is recognized by the smaller ob-

lanceolate leaves with decurrent lamina base, ob-

scure pubescence, unusual pit domatia (sometimes

absent), unisexual flowers, and small ellipsoid fruits

on small flat receptacles. Vegetatively, our collec-

tions resemble the lowland O. sp. aff. O. bijuga

and the Monteverde population of O. whitei, but

those species have very different fruits, and the

plants are not dioecious. Costa Rican collections

are from near Volcan Rincon de La Vieja (Guan-

acaste), the region of Buena Vista de San Carlos

(Alajuela), Tucurrique and Juan Vinas (Cartago),

and near the Rio Convento in the General Valley

(San Jose/Puntarenas). This species is similar to

O. cuneifolia (Ruiz & Pavon) Mez, O. neesiana

(Miq.) Kosterm., and O.jlorulenta (Meissn.) Mez,

of South America. Ocotea eucuneata Lundell with

slightly thinner and more acuminate leaves may

be a synonym from Belize.

Ocotea paulii C. K. Allen, J. Arnold Arbor. 26:

345. 1945. Figure3.

Small trees 3-8(-15) m tall, occasionally shrub like

and somewhat scadent, leafy branchlets 3-8 mm thick,

glabrous, with 3-5 prominent longitudinal ridges, smooth

and grayish or brown between the ridges (rarely terete),

distal stems hollow. Leaves alternate or rarely pseudov-

erticillate, petioles 6-15(-22) long, 1.5-3(-4) mm thick,

slightly canaliculate above, glabrous and usually drying

dark; leaf blades 10-23(-33) cm long, 3-7.5 cm broad,

narrowly oblong to elliptic-oblong or very narrowly ovate-

oblong, abruptly short-acuminate at the apex (occasion-

ally rounded or tapering gradually to a longer acuminate

apex), acute to obtuse or slightly rounded at the base,

margin entire and flat or revolute, drying subcoriaceous

to coriaceous and grayish to reddish brown, glabrous

above and below, with 5-1 1 major secondary veins on

each side, major veins usually prominent beneath, cen-

tral secondaries arising at angles of 35°-55°, tertiary veins

often subparallel but not conspicuous. Inflorescences 1 5-

25 cm long, subterminal or axillary to distal leaves, pan-

iculate with lateral branches to 5 cm long, many-flow-

ered, peduncle 5-8 cm long, glabrescent, pedicels 1-2

mm long. Flowers 2-3 mm long and ca. 3 mm broad,

glabrous or with a few minute hairs, tepals ca. 1.5 mm

long and equally broad, ovate; outer stamens with short

(0.4 mm) filaments, outer anthers 0.7-0.8 mm long, nar-

rowly ovate with superposed thecae, inner stamens ca.

1.2 mm long, staminodes absent; pistil 1.5-2 mm long,

style equalling the ovoid ovary in length, stigma slightly

discoid. Fruits borne on short (2-4 mm) shallow recep-

tacles 6-10 mm broad and 1-2 mm deep, margin usually

entire, pedicels becoming 6 mm long and 2-3 mm broad

at the apex; berry ca. 2 cm long and 1 cm thick (dry),

ellipsoid-oblong.

Small trees and clambering shrubs of evergreen

montane forest formations, between 600 and 1 600

(2300) m elevation. Flowers have been collected

in all months except October-November, but fruits

have only been collected in March and May. This

species is only known from the northern part of

the Meseta Central (San Ramon to Zarcero) and

the highlands of Chiriqui, Code, and Panama

provinces in Panama.

Ocotea paulii is recognized by its usually small

stature, hollow distal stems, narrowly oblong leaves

often exceeding 1 5 cm in length, glabrous parts,

and small narrow fruits borne on small shallow

receptacles. This species differs from the closely

related O. atirrensis, O. wedeliana, and O. nicar-

aguensis in its higher-altitude habitat, smaller,

more consistently oblong leaves, and more pro-

fusely branched inflorescences. There appears to

be intergradation between O. atirrensis and O.

paulii on the Caribbean slope at about 1000 m

elevation. The populations of Costa Rica and Pan-

ama differ somewhat in leaf form and texture. More

importantly, while Panamanian collections may

have the upper thecae of the outer anthers reduced

in size, there are Costa Rican collections in which

BURGER: FLORA COSTARICENSIS

91

the outer anthers may be 2-thecous in some flow-

ers (.-I. Smith 633, 907, F).

Ocotea pittieri (Mez) van der Werff, comb. nov.

Phoebe pittieri Mez, Hot. Jahrb. Syst. 30, Beibl.

67: 16. 1901. Figure 4.

Shrubs or small to medium-sized trees, 2-10(-20) m

tall, leafy branchlets 1.3-4 mm thick, at first grayish or

yellowish puberulent with short (0.1-0.3 mm) straight

or crooked hairs, persisting or glabrescent. Leaves alter-

nate, petioles 5-12 mm long, ca. 1 mm thick, puberulent,

lateral margins flat above or slightly grooved (not sul-

cate): leaf blades (2.5-)4-8(-10) cm long, 1.5-4(-5) cm

broad, narrowly to broadly elliptic, elliptic-oblong or

slightly obovate, acute to short-acuminate (obtuse in short

broad leaves) at the apex, acute to obtuse at the base,

drying stiffly chartaceous, pale brown or occasionally

dark, usually lustrous and sparsely puberulent on the

midvein above, very sparsely to densely puberulent be-

neath with slender or crooked hairs, often with tufts of

longer (0.6 mm) hairs in the vein axils or along the

midvein beneath, with 2-6 major secondary veins on

each side, central secondaries arising at angles of 30°-

60°. Inflorescences axillary from distal or older leaves,

5-9 cm long, paniculate, peduncles usually longer than

the flowering rachis, pedicels 4-8 mm long, glabrous or

puberulent. Flowers 3-8 mm broad, tepals 2-3 mm long,

equal, glabrous or puberulent on the outside, glabrous

or papillate-puberulent within; outer stamens with ob-

long anthers 0.7-1 mm long, 0.5-1 mm broad, on fila-

ments 0.5-1 mm long, inner stamens ca. 1.4 mm long

with narrow anthers 0.7 mm long, staminodes 0.5-1 mm

long and slender or capitate (sometimes absent), floral

tube often with hairs at the edge; pistil 2-3 mm long,

glabrous, style ca. 1 mm long, stigma simple or lobed.

Fruits subtended by a shallow cup 10-1 2 mm broad and

2-3 mm deep, margin entire or with short persisting

perianth parts, gradually or abruptly expanded above the

elongated (1-2 cm) thickened (ca. 4 mm) pedicel; berry

15-30 mm long, 10-20 mm thick, ellipsoid, dark green.

Shrubs and trees of evergreen montane forest

formations, between (1000) 1500 and 3200 m el-

evation. Flowers have been collected in September

and December-June; fruits have been collected in

January-August. The species is found in the Cor-

dillera de Tilaran, along the northern edge of the

Meseta Central near Palmira, on Volcan Irazu, and

above Cartago, and from the Cordillera de Tala-

manca to the Chiriqui highlands of Panama.

Ocotea pittieri (formerly Phoebe pittieri) is usu-

ally recognized by the stiff smaller hirsutulous

leaves, with minor venation raised below, the often

lustrous upper surfaces, and a tendency to dry yel-

lowish brown. We interpret this species broadly,

and in the sense that the name has been used by

Standley and others (in herb.). Ocotea pittieri has

not been collected in the area between Volcan Poas

and the western slopes of Volcan Irazu. This pe-

culiar pattern is also found in O. mollicella which

is closely related and easy to confuse with O. pit-

tieri. The leaves appear to be broader and larger

in collections from the eastern part of the Cordi-

llera de Talamanca. Ira amarillo is reported as a

common name in Costa Rica.

Staminodes are often poorly developed in this

species, though they can be well developed in some

individuals. Leaf texture and venation is very dif-

ferent from Phoebe cinnamomifolia and its allies.

Moreover, Oscar Castro of the University of Costa

Rica has found (pers. comm.) that the chemistry

of 0. pittieri is like that of some Costa Rican Oco-

tea spp., and not like that of Phoebe mexicana

(part of P. cinnamomifolia in a wide sense). Thus

the correct placement of Ocotea pittieri and its

close relative O. mollicella is in the genus Ocotea.

Specimens from Mexico and northern Central

America placed under the names Phoebe bour-

geauviana Mez and Phoebe psychotrioides Mez ap-

pear to be closely related to O. pittieri.

The type of Ocotea pittieri (Tonduz 11893, pho-

to of specimen destroyed at B in F, us) has broader

thinner leaves that have dried darker than is usual

for this species. This type material resembles O.

brenesii somewhat, but that species never has such

narrow leaves, and the leaves usually dry very dark

and thin. It appears that some newly flushed leaves

have dried dark and thin (in the material at us),

but other leaves (as in the photo from B) appear

more characteristic of the specimens traditionally

placed under this name in herbaria. The photo,

we believe, justifies the continued use of this name

for the material placed here.

Ocotea pseudopalmana W. Burger, sp. nov. Fig-

ure 6.

Arbor usque 30 m alta, ramulis foliiferis 2.6-9 mm

crassis, ferrugineo-puberulis. Folia alterna, petiolis 6-15

mm longis, puberulis; laminis 8-16(-21) cm longis et 3-

9 cm latis, ellipticis vel oblongo-ellipticis, apice obtuso

vel brevi-acuminato, subtus brunneolo-puberulis, nervis

secondariis 5-8 paribus. Inflorescentiae 6-1 5 cm longae,

pauciflorae, pedunculis 4-1 0 cm longis, puberulis. Flores

5-7 mm longi, 6-8 mm lati, extus puberuli, stamina ser.

I-II ca. 1.5 mm longa, antheris ca. 1 mm longis, stam-

inodiis nullis; gynoecium 2.3-4 mm longum, stylo usque

2.3 mm longo. Fructus ellipsoideus vel oblongus, 2.2-

2.8 cm longus et 1.5-2 cm crassus; cupula 11-14 mm

lata, 3-8 mm longa.

Trees 4-20(-30) m tall, leafy branchlets 2.6-6(-9) mm

thick, densely velutinous to tomentulose with short (0.2-

0.6 mm) yellowish brown or brownish hairs, terete or

92

FIELDIANA: BOTANY

with longitudinal ridges in early stages. Leaves alternate

in a spiral, petioles 6-15 mm long, 2-3 mm thick, flat

or slightly sulcate above, densely puberulent; leaf blades

8-16(-21) cm long, 3-9 cm broad, elliptic to elliptic-

oblong or slightly obovate, tapering abruptly to a short-

acuminate or bluntly obtuse apex, obtuse to cuneate at

the base and often slightly decurrent, margin often slight-

ly revolute (especially near the base), drying subcoria-

ceous or coriaceous, puberulent on the major veins above

but becoming glabrescent and impressed in age, lower

surface with brown or yellowish brown hairs 0.3-0.5 mm

long, with 5-8 major secondary veins on each side, the

central secondaries arising at angles of 45°-70°, tertiary

venation usually prominent beneath. Inflorescences sol-

itary and axillary to leaves, 6-15 cm long, racemose or

narrowly paniculate and with few (4-12) flowers, pe-

duncle 4-10 cm long, 1 .2-2 mm thick, densely brownish

puberulent, pedicels 3-8 mm long. Flowers 5-7 mm long,

6-8 mm broad, campanulate, densely puberulent on the

outside, tepals ca. 3.5 mm long and 3 mm broad, drying

dark and papillate-puberulent within; outer stamens with

prominent (0.6 mm) broad filaments and oblong anthers

ca. 1 mm long and 1 mm broad, inner stamens ca. 2

mm long with glands 0.7-1 mm thick, staminodia ab-

sent; pistil 2.3-4 mm long, ovary narrowed at the base,

style 1.5-2.3 mm long, stigma subcapitate. Fruits borne

on a flat or shallow (0-3 mm) cup 11-14 mm broad, to

8 mm long and conical or as little as 3 mm long and

saucer-like, the cup gradually or abruptly expanded above

the thickened pedicel, red at maturity; berry 2.2-2.8 cm

long and 1.5-2 cm in diameter (dry), broadly ellipsoid

or oblong, becoming black.

TYPE— Costa Rica, San Jose Province, east of

La Chonta, 1 1 May 1969, Roy W. Lent 1679 (ho-

lotype, F 1749018; negative, 61218; isotype, CR).

Trees of wet evergreen montane forest forma-

tions, from 1500 to 2500 m elevation in areas

subjected to moist winds from the Caribbean and

adjacent areas. Flowering material has been col-

lected in February-September; mature fruits have

been collected in April-October. This species is

only known from the slopes of Volcan Irazu and

eastward through the Cordillera de Talamanca as

far as the border with Panama.

Ocotea pseudopalmana is recognized by the

dense pubescence on the underside of the leaves

and exterior of flowers, the stiff leaves that usually

dry dark brown above, the few-flowered inflores-

cences, the larger flower size, and the cloud forest

habitat. This species is related to O. gomezii with

much larger flowers, larger more rounded leaves,

and lower-altitude (800 to 1400 m) habitat. This

species resembles O. mollifolia, but that species

has broader thinner leaves, smaller flowers, and

fruit cups with persisting perianth lobes. This

species also resembles members of the O. helic-

terifolia-O. valeriana complex, but those species

have smaller glabrescent flowers with staminodes

and deeply cupulate fruiting receptacles. The ma-

terial placed here resembles Phoebe amplifolia Mez

& J. D. Smith, from highland Guatemala; but while

that species has rather similar flowers, the small

(0.7 mm) sagittate-capitate staminodes are dis-

tinctive, and the leaves are larger and often with

long (3 cm) petioles. Some of the collections placed

here were earlier identified as Ocotea palmana,

but that species is actually a Pleurothyrium (q.v.).

Material placed here includes: Almeda & Nakai

3834 (CAS, CR, F), Brown 17518 (F, LSU), Davidse

et al. 28550, 28906 (CR, MO), Holdridge6555, 6575

(CR, NY), Rossbach 3171 (GH), Standley 38633,

38848 (us), Wilbur & Almeda 16714 (DUKE).

Ocotea puberula (Rich.) Nees, Syst. Laurin. 472.

1836. Laurus puberula Rich., Actes Soc. Hist.

Nat. Paris 1: 108. 1792. O. pyramidata Blake

ex Brandegee, Univ. Calif. Publ. Bot. 7: 326.

1920.

Trees 7-15(-27) m tall, to 50 cm d.b.h., bark pale

gray, leafy branchlets 1 .7-7 mm thick, flattened or ribbed

with longitudinal ridges extending downward from the

leaf bases, glabrous or sparsely and minutely (0. 1 mm)

puberulent, young stems drying dark, older stems pale

grayish. Leaves with petioles 10-26(-30) mm long, 0.7-

2 mm broad, glabrous distally but with minute (0. 1 mm)

hairs near the base adaxially, distally sulcate above; leaf

blades 7-2 1(-25) cm long, 2.5-8(-l 2) cm broad, elliptic-

oblong to narrowly elliptic or narrowly ovate-elliptic,

tapering gradually to an acuminate apex (or tapering

abruptly to a short-acuminate apex in broader leaves),

acute to obtuse at the base and slightly decurrent on the

petiole, drying stiffly chartaceous and grayish green to

olive green, dull or lustrous above, essentially glabrous

above, glabrous beneath or with minute hairs along the

major veins, with 5-9 major secondary veins, lacking

domatia, tertiary veins elevated on the dried surfaces

above and below and forming a weak reticulum of broad

(1-3 mm) areolae. Inflorescences axillary, 4-7 cm long,

peduncles 1-8 mm long, rachis minutely puberulent, with

a few proximal lateral branches to 15mm long, pedicels

1-3 mm long. Flowers unisexual and dioecious, glabrous

distally on the outside; male flowers 3-5 mm long, peri-

anth lobes 2-2.7 mm long, 1.7-2.2 mm broad; outer

stamens 2-2.5 mm long, filament flattened, anthers ca.

1.2 mm long and 0.7 mm broad, valves superposed,

inner stamens slightly shorter, glands 0.7-1 mm wide,

staminodes absent; pistillode ca. 1.3 mm long and 0.4

mm in diameter. Female flowers 2-3 mm long, 2.5-3.5

mm broad, perianth parts ca. 1.5 mm long, nonfunc-

tional stamens 1-1.2 mm long and with anthers ca. 0.5

mm long; pistil ca. 2.2 mm long, ovary ca. 0.9 mm in

diameter, style ca. 0.8 mm long, stigma flat, disclike, and

drying dark. Fruits borne on a short (2-4 mm) distally

flattened receptacle 4-8 mm broad at the top (but more

cuplike in life), with undulate or 6-lobed margin, pedicels

4-16 mm long; berry 6-8 mm in diameter (dried), glo-

bose, green becoming black and lustrous.

BURGER: FLORA COSTARICENSIS

93

A species of lowland evergreen and partly de-

ciduous forest formations. Flowers have been col-

lected in December-January in Panama; fruits have

been collected in January-February. Though little

collected in Central America, this species ranges

from Mexico to Argentina.

Ocotea puberula is recognized by the function-

ally unisexual flowers on unisexual (dioecious)

trees, small globose fruits subtended by flattened

receptacles or weakly developed cups, and larger

elliptic-oblong Nectandra-like leaves. Fruiting

material closely resembles a number of our species

ofNectandra, but the inflorescences of O. puberula

have fewer branches and the cups are not as well

developed. This species was first collected in Costa

Rica near Palmar Norte in 1988, after the manu-

script was completed; it was not possible to include

this species in the keys. We thank J. Rohwer for

the determination.

Ocotea rivularis Standl. & L. O. Williams, Ceiba

1: 238.1951. Figure 2.

Medium-sized trees to 1 5 m tall, leafy branchlets 4—

12 mm thick. Leaves alternate, petioles ca. 1 cm long

and 5 mm thick, sulcate and winged, glabrous; leaf blade

30-45 cm long, 15-23 cm broad, obovate, abruptly

rounded near the apex to a bluntly short-acuminate apex,

tapering gradually to the attenuate base, margin entire

and slightly revolute, drying stiffly chartaceous, smooth

and glabrous above, essentially glabrous and reddish

brown beneath, with 10-14 major secondary veins on

each side, central secondaries arising at angles of 50°-

65°, tertiary venation subparallel but obscure. Inflores-

cences axillary to distal leaves, 1 8-30 cm long, panicu-

late, peduncles 7-10 cm long, reddish and 2-3.5 mm

thick, pedicels 1-2 mm long, minutely puberulent. Flow-

ers ca. 3 mm long and 3 mm broad, greenish, tepals to

2 mm long and glabrous within; outer stamens with fil-

aments almost equalling the length of the anthers, outer

anthers ca. 0.6 mm long and 0.5 mm broad, staminodes

absent; pistil ca. 2 mm long with slender style 0.8 mm

long. Fruits borne in a small (4 mm long and 6 mm

broad) cupulate receptacle 2-3 mm deep with persisting

small broad perianth bases; berry ca. 1 2 mm long and

ellipsoid.

Trees of the evergreen rain forest formations of

the Pacific lowlands; flowering in July-early Au-

gust. This species is endemic to southern Puntar-

enas Province in the Golfo Dulce area.

Ocotea rivularis is easily recognized by its very

large obovate thin-textured leaves, attenuate lam-

ina base with revolute margins, large inflores-

cences with small flowers, and small lobed cups

with ellipsoid fruits. The leaves are essentially gla-

brous but may have very thin appressed hairs (0.2

mm long) on the underside which are difficult to

see. This species is very closely related to O. in-

sularis, with smaller thicker leaves. Paul Allen

(1956, pp. 276-277) stated that the species was

common around the type locality near Esquinas.

This species is only known from Allen 5590 (F,

MO, us, the type) and Grayum et al. 4068 (CR, MO),

from near Rincon.

Ocotea skutchii C. K. Allen, J. Arnold Arbor. 26:

352. 1945. Nectandra producta C. K. Allen, loc.

cit. 397. 1945. O. williamsii P. A. Allen, The

Rain Forests of Golfo Dulce, 410, pi. 27, 1956,

non O. williamsii (O. C. Schmidt) Kosterm., J.

Sci. Res. (Jakarta) 1: 122. 1952. Figure 11.

Trees to 30 m tall, often with prop roots near the base,

leafy branchlets 2.5-5 mm thick, minutely (0.1-0.3) pu-

berulent with brownish or silvery appressed hairs but

quickly becoming glabrous, terete. Leaves alternate, pet-

ioles 0.5-5 cm long but difficult to delimit because of

the decurrent and revolute lamina base, narrowed por-

tion of the leaf base to 5 cm long, minutely puberulent

or glabrescent, flat above; leaf blades 8-13 cm long, 2.5-

5 cm broad, elliptic-obovate to elliptic-oblanceolate or

narrowly elliptic-oblong, usually short-acuminate at the

apex, tapering gradually to a cuneate or acute base and

decurrent on the petiole, margin strongly revolute near

the base, drying stiffly chartaceous to subcoriaceous, gla-

brous except for minute hairs above the midvein near

the petiole, often lustrous above and glaucous beneath,

minutely and obscurely puberulent beneath with thin

appressed hairs ca. 0.3 mm long and parallel with the

secondary veins, with 5-9 major secondary veins on each

side, central secondaries arising at angles of 35°-55°. In-

florescences axillary to distal leaves, 8-18 cm long, open-

ly branched panicles, peduncles to 6 cm long, ca. 1.5

mm thick and sparsely appressed puberulent, pedicels

l-3(-5) mm long. Flowers 2.5-3.5 mm long, 3.5-5.5 mm

broad, campanulate, tepals ca. 2 mm long and 1.5 mm

broad, obtuse, densely puberulent on the outside and

minutely papillate within; outer stamens 0.6-1.3 mm

long, filaments 0.1-0.7 mm long, outer anthers 0.5-0.8

mm long and equally broad or broader, thecae super-

posed or occasionally the lower somewhat lateral to the

upper (Nectandra-\ike), inner stamens ca. 1.5 mm long

and with large glands, staminodes not seen; pistil 1.5-

2.5 mm long, style slender and equalling the ovary in

length, stigma slightly discoid. Fruits borne in a broadly

cupulate receptacle ca. 1 cm long and 10-15 mm broad,

red; berry 3-4 cm long, 13-18 mm in diameter, ellipsoid-

oblong.

Trees of evergreen lower montane forest for-

mations in the central volcanic highlands and the

Cordillera de Talamanca, from (30) 600 to 1400 m

elevation. Flowering material has been collected

in December (Skutch 3062, GH, NY, the type ofO.

skutchii) and January (Skutch 3906, AA, NY, the

94

FIELDIANA: BOTANY

type of N. productd). Fruits were collected in April

(Solas 392, usj). This species is only known from

the central highlands and the General Valley in

Costa Rica.

Ocotea skutchii is recognized by the average-

sized elliptic-obovate leaves with long-decurrent

and revolute lamina base, the slender appressed

hairs on the lower leaf surfaces, and the puberulent

flowers with variable stamens. The leaves are often

lustrous above and glaucous beneath. This species

is part of a group of taxa related to O. glaucoser-

icea, which often become tall trees with basal prop

roots and have the same characteristic pubescence

on the lower leaf surfaces in early stages. There is

the possibility that the plants placed here may in-

tergrade with O. glaucosericea at slightly higher

elevations. A few collections of O. austinii (such

as Solano 3 CR, F) also appear as if they could

represent intermediates with O. skutchii. There is

also the possibility that this species intergrades

with O. whitei of somewhat higher elevations.

Whether collections which appear intermediate are

simply extremes of variation within their species,

or whether this is actually a single polymorphic

complex of intergrading populations, are ques-

tions that cannot be resolved without further study

and more collections.

Caroline Allen (1966) placed Paul Allen's Oco-

tea williamsii (1956) into synonymy under O.

skutchii, and this may be correct. The long narrow

fruits (4.8 x 2 cm) on small cupular receptacles

are distinctive. The flowers of the type (Allen 5983,

us) are abnormally developed, lacking both sta-

mens and pistils. The tufts of hairs along the mid-

vein on the lower leaf surface may also be abnor-

mal. Regardless, the name O. williamsii is a later

homonym and cannot be used as a specific epithet.

Ocotea stenoneura Mez & Pittier, Bull. Herb. Bois-

sier, ser. 2, 3: 233. 1903, emend. C. K. Allen,

J. Arnold Arbor. 26: 334-335. 1945.

Trees to 20 m tall, leafy branchlets 3-7 mm thick,

densely reddish brown tomentulose, becoming grayish

and terete. Leaves alternate, petioles difficult to delimit

because of the strongly revolute lamina margin, nar-

rowed basal portion of the leaf 1.5-3 cm long and 2.5-

7 mm broad, flat or broadly sulcate above; leaf blades

ca. 1 8-20 cm long, 9 cm broad, broadest at or near the

middle, elliptic-oblong, abruptly narrowed at the acute

to short-acuminate apex, tapering and slightly rounded

above the revolute base, drying subcoriaceous, glabrous

above or with short hairs above the midvein and with

the suhparallcl tertiary venation often slightly raised

above, densely ferruginous-pilose (or tomentulose?) be-

neath with appressed hairs 0.1-0.5 mm long, with 10-

1 2 major secondary veins on each side, the central sec-

ondaries arising at angles of 35°-60°. Inflorescences ax-

illary to distal leaves, somewhat shorter than the leaves,

peduncles 4-8 cm long to 4 mm thick and densely pu-

berulent. Flowers unknown, dried stamens (on the fruit-

ing cupules) resembling those of O. insularis (fide Roh-

wer, 1986, p. 145). Fruits borne on flat saucer-like

receptacles, ca. 1 cm broad and 2-3 mm deep, often with

triangular perianth lobes persisting on the margin; berry

(perhaps not mature) globose, 10-13 mm in diameter.

A poorly known species of evergreen forests,

from near sea level to 1000 m elevation. Collected

near Las Vueltas, Tucurrique, Cartage Province;

fruiting in February (Tonduz 13377, photo of B at

F, the lectotype; C. K. Allen, 1945). Skutch 3014

(MO, NY), flowering in December in the General

Valley, is provisionally placed here. The species

is endemic to Costa Rica.

Ocotea stenoneura is distinguished by its larger

puberulent leaves with short strongly revolute

lamina base, and the small globose fruits on rel-

atively flat receptacles. Whether this material rep-

resents a very rare species or aberrant individuals

of a population placed under another name in this

treatment is difficult to say. Some specimens ear-

lier identified as Ocotea stenoneura are now placed

under O. dentata, a member of the O. insularis

alliance, with very different fruits and more ob-

ovate leaves.

Ocotea tenera Mez & J. D. Smith ex Mez, Bull.

Herb. Boissier, ser. 2, 3: 234. 1903. Figure 13.

Shrubs or small trees 3-8(-12) m tall, to 15(-20) cm

d.b.h., leafy branchlets 1.5-3.8 mm thick, glabrous from

the earliest stages, drying terete and dark. Leaves alter-

nate, usually distichous in a single plane and not crowd-

ed, petioles 6-12 mm long, ca. 1 mm thick, lateral mar-

gins forming a sulcus adaxially, glabrous; leaf blades

(5-)7-16(-20) cm long, (1.5-)3-6(-8) cm broad, elliptic

to elliptic-oblong or ovate-elliptic, tapering gradually to

the acute or acuminate apex, acute to obtuse at the base,

usually drying very dark and thin-chartaceous in texture,

glabrous above and below, minor veins slightly raised

above, with 4-7 major secondary veins on each side,

central secondaries arising at angles of 50°-70°, often

loop-connected near the margin distally, often with pit

domatia. Inflorescences 4-8(-12) cm long, axillary or

extra-axillary on distal stems, paniculate with relatively

few widely spaced flowers, glabrous, peduncles to 4 cm

long, pedicels slender, ca. 3.5 mm long. Flowers greenish

white to yellowish (but drying black), 1.5-3 mm long,

glabrous, tepals ca. 1.5 mm long; outer stamens 0.6-1.2

mm long, with broad short filaments, outer anthers ca.

0.7 mm long, narrowed apically and with superposed

thecae, staminodes absent or minute, glabrous or with

a few hairs around the edge of the floral tube within;

BURGER: FLORA COSTARICENSIS

95

pistil 1.5-2.3 mm long, ovary globose or ovoid, style

usually equalling the ovary in length, stigma simple. Fruits

borne on the flattened apex of the thickened pedicel, a

cuplike receptacle absent or shallow (2-3 mm), 1 cm

broad and with an undulate margin, usually pendant,

becoming rose red or bright red, thickened portion of

the pedicel 1-3 cm long; berry 2-3 cm long and 1-2 cm

thick, ovoid or oblong, becoming purple or black.

Trees usually found at forest edges and in sec-

ondary vegetation at lower elevations and in forest

understory at higher elevation, in evergreen wet

forest formations of the Caribbean slope and cen-

tral highlands, between 50 and 1500 (1700) m

elevation. Flowering collections have been made

in August-March, and fruits have been collected

in August-September. This species is endemic to

Costa Rica, ranging from the Cordillera de Tilaran

to the valley of the Rio Reventazon and Puerto

Limon.

Ocotea tenera is a distinctive species of Laura-

ceae with its small stature, limited range, thin el-

liptic leaves that dry very dark, minute flowers

which almost become black when dry, ellipsoid

fruits on a flattened receptacle at the apex of a

thickened pedicel, and the lack of puberulence.

The species has been reported to be dioecious (C.

K. Allen, 1945, p. 357) but this may be an error,

as most plants appear to have bisexual flowers.

Ocotea tenera is closely related to O. bernoulliana

Mez of Guatemala, but that species has a more

deeply cupulate fruiting receptacle and the leaves

do not become as dark when dry. Ocotea brenesii,

with broader leaves, larger flowers, and the floral

cup hairy within, appears to be closely related to

O. tenera and may be difficult to separate. Ocotea

cernua and O. meziana are also similar, but their

leaves do not become so dark when dried.

Ocotea valeriana (Standl.) W. Burger, comb. nov.

Phoebe valeriana Standl., Field Mus. Nat. Hist.,

Hot. Ser. 18: 460. 1937. P. smithii C. K. Allen,

J. Arnold Arbor. 26: 317-318. 1945. Figure 6.

Trees 3-12(-20) m tall, leafy branchlets 2.5-8 mm

thick, densely strigulose to tomentulose with yellowish

brown to dark brown hairs 0.1-0.5 mm long. Leaves

alternate, differing greatly in size and shape on different

trees, petioles 4-22(-28) mm long, 1.7-3 mm thick, sul-

cate above, tomentulose to strigulose; leaf blades 8-25

cm long, 3.5-14 cm broad, broadly obovate to oblong

(usually widest above the middle), abruptly narrowed to

the short-acuminate, obtuse or rounded apex, obtuse to

rounded (cordulate) at the base, equal or subequal, drying

subcoriaceous and yellowish brown to dark brown, mi-

nutely puberulent on the vein above and sparsely pu-

berulent to glabrous between the veins, dull or lustrous,

strigulose on the veins beneath with slender hairs ca. 0.3

mm long, with (4-)5-10 major secondary veins on each

side, central secondaries arising at angles of 35°-60°. In-

florescences 5-25 cm long, distal and with the subtending

leaves often caducous, peduncles (l-)3-9 cm long, 1-3

mm thick, densely strigulose, dark or with yellowish

brown hairs, lateral branches of the rachis with (l-)3-7

flowers and not further branched, pedicels slender, 2-10

mm long. Flowers 4-6 mm long and 6-15 mm broad,

essentially glabrous outside and within, perianth pellucid

punctate, tepals ca. 3 mm long; outer stamens with an-

thers 0.9-1. 5(-2) mm long, 1-1.3(-1.7) mm broad, ob-

long to slightly broader than long, filaments very short,

staminodes absent or small or up to 1 mm long; pistil

ca. 2.5 mm long, ovary ca. 1.5 mm in diameter, narrowed

at the base, style ca. 1 mm long, stigma discoid. Fruits

borne in a cup 9-14 mm long, 10-18 mm broad, 5-8

mm deep, cupulate or hemispheric, margin entire or with

the persisting perianth bases, red; berry ca. 3 cm long

and 1.8 cm in diameter, obovoid to ellipsoid, obtusely

apiculate at the apex, black.

Trees of evergreen montane forest formations,

from (1000) 1400 to 2200 m elevation. Flowering

in January-May; fruits have been collected in De-

cember-February and May-June. The species

ranges from the Cordillera de Tilaran, along the

Caribbean escarpment and continental divide of

the central volcanic highlands as far east as the

western slopes of the General Valley in the Cor-

dillera de Talamanca. Endemic to Costa Rica (but

see below).

Ocotea valeriana (formerly Phoebe valeriana) is

a very variable species characterized by the stiffly

chartaceous to subcoriaceous laminae that are pu-

berulent beneath and usually broadest above the

middle. The glabrous flowers drying black, pel-

lucid-punctate perianth, distinctive puberulent in-

florescences with lateral branches usually bearing

long-pediceled flowers (and not further branched),

and middle elevation habitat further distinguish

this species. Staminodes may be present or absent,

but there are rarely three well-developed stami-

nodes, as in species of Phoebe. Ocotea valeriana

may prove to be a high elevation derivative of O.

helicterifolia, but the fruit cups are larger and deep-

er than in Costa Rican material of that species.

There are highland specimens from Guatemala

that are very similar to O. valeriana, but these are

placed in O. helicterifolia and seem to intergrade

with more typical lowland forms of O. helicteri-

folia in Guatemala and Mexico. There is little evi-

dence for such intergradation in Costa Rica, but

it may be present (see the discussion under O.

helicterifolia).

Phoebe smithii (based on A. Smith P.C. 367, F)

has very short petioles, short inflorescences, and

96

FIELDIANA: BOTANY

more oblong laminae slightly rounded at the base.

In addition, the anthers are more often broad (their

length equaling their width), and the staminodes

are smaller or more often absent. Specimens here

placed under Ocotea valeriana, but more like the

type of P. smithii appear to predominate in col-

lections made from the Cordillera de Tilaran.

However, there are many intermediates through-

out the range of the species, and populations at

several localities have both the extreme forms and

the intermediates.

Ocotea valerioides W. Burger, sp. nov. Figure 2.

Arbor 5-1 5 m alta, ramulis foliiferis 5-1 2 mm crassis.

Folia alterna, saepe aggregata. petiolis 4-14 mm longis,

3-6 mm crassis; laminis 13-38 cm longis et 10-22 cm

latis, obovatis vel oblongo-obovatis, subtus tomentosis,

nervis secondariis 8-12 paribus. Inflorescentiae pani-

culatae, 9-30 cm longae, pedunculis usque 1 8 cm longis.

Flores 4-5(-7) mm longi et 8-10 mm lati, extus parce

puberuli, tepalis intus minute papillatis; stamina ser. I-

II antheris 1-1.2 mm longis, staminodiis nullis. Fructus

ellipsoideus, 28-36 mm longus.

Small trees 5-1 5 m tall, to 40 cm d.b.h., leafy branch-

lets 5-12 mm thick, densely tomentulose with yellowish

to dark grayish brown hairs to 0.5 mm long. Leaves

usually clustered at the ends of branchlets, petioles 4-

14 mm long, 3-6 mm thick, densely tomentulose, broad-

ly sulcate above; leaf blades 13-38 cm long, 10-22 cm

broad, obovate to oblong-obovate or slightly pandurate,

abruptly rounded and short-acuminate at the apex, ta-

pering gradually to the cuneate base and slightly rounded

at the petiole, drying stiffly chartaceous, dark brown above

and slightly lustrous, minutely puberulent above the ma-

jor veins, lower surface paler in color and with a dense

tomentum of short (0.3 mm) thin straight hairs slightly

rough to the touch, with 8-12 major secondary veins on

each side, central secondaries arising at angles of 40°-

60°, tertiary veins prominent beneath and often subpar-

allel. Inflorescences axillary to distal leaves or scalelike

undeveloped leaves, racemiform panicles with short lat-

eral branches to more openly paniculate with spreading

side branches, (5-)9-30 cm long, primary peduncles to

18 cm long, 1-2.5 mm thick, densely yellowish brown

puberulent, pedicels 3-1 1 mm long. Flowers 8-10 mm

broad, 4-5(-7) mm long, rotate at anthesis, perianth parts

ca. 4 mm long and 2mm broad, sparsely puberulent on

the outside and minutely papillate within; outer stamens

subsessile and sometimes broader than long, outer an-

thers 1-1.2 mm long and 0.9-1.2 mm broad, thecae

usually superposed but the lower occasionally lateral (re-

sembling Nectandra), inner stamens biglandular, stam-

inodes not seen, the edge of the floral tube with straight

erect hairs around the base of the stamens; pistil 1.7-

2.5 mm long, ovary rounded, style slender, 0.7-1 . 1 mm

long, stigma discoid. Fruits borne on a broad (12-16

mm), shallow (ca. 5 mm deep) cup gradually expanded

from a thickened (3-5 mm) pedicel, longitudinal ridges

sometimes present and giving the margin of the cup an

undulate appearance, becoming red; fruits 28-36 mm

long and 14-22 mm in diameter, ellipsoid or slightly

obovoid.

TYPE— Costa Rica, Alajuela Province, 1 km S

of Cariblanco along the Sarapiqui road, elevation

about 850 m, 15 Sept. 1974, Gary S. Hartshorn

1530 (holotype, CR 71922; isotypes, F [negative,

61118], MO).

Trees of the very wet Caribbean slope and low-

lands, between 40 and 850 m elevation. Flowers

have been collected in July and September; fruits

were collected March-May and July. This species

has only been collected in central Costa Rica (from

Cariblanco, Alajuela, to between Siquirres and

Turrialba, Limon) and in the province of Bocas

del Toro, Panama.

Ocotea valerioides is recognized by the large ob-

ovate puberulent leaves with short thick petioles,

the larger flowers with subsessile anthers, and ra-

cemiform panicles with densely yellowish brown

puberulent rachis. This material had been filed

under Phoebe valeriana (now Ocotea valeriana) in

herbaria, but O. valerioides has much larger leaves,

a lower elevation habitat, more puberulent flow-

ers, and stamens with the anthers often developing

a Nectandra-like configuration. In fact, this species

may represent a tendency fully expressed in Nec-

tandra belizensis, which appears to be related, as

does O. helicterifolia. All these species are part of

a complex that deserves serious study. Within our

material the lower elevation collections have larg-

er leaves (Gomez & Herrera, 23619, CR, MO; Gra-

yum et al. 3519, CR, F, MO; Zamora & Elizondo

674, CR, F), and those around 800 m have some-

what smaller leaves (Hartshorn 1530, CR; Hold-

ridge 6779, CR, NY; Poveda & Castro 3920, CR, F).

The altitudinal separation of O. valerioides and O.

valeriana is consistent with patterns seen in many

genera and families; a clinal connection is unlikely.

There may be a close relationship with O. lentii,

and the possibility of intermediates with that

species should be explored. The leaves of a fruiting

collection from Panama (Kirkbride & Duke 486,

MO) differs from Costa Rican collections in vesture

and prominence of the tertiary veins, while another

collection from Panama (von Wedel 2264, MO) is

more typical.

Ocotea veraguensis (Meissn.) Mez, Jahrb. Konigl.

Bot. Gart. Berlin 5: 240. 1889. Sassafridium

veraguense Meissn. in DC., Prodr. 1 5, pt. 1 : 171.

1864. O. paradoxa Mez, Bot. Jahrb. Syst. 30,

BURGER: FLORA COSTARICENSIS

97

Beibl. 67: 16. 1901. O. bakeri Blake, Contrib.

Gray Herb. 52: 65. 1917. Figure 14.

Small trees 4-10 m tall, leafy branchlets 1.2-5 mm

thick, at first minutely puberulent with grayish appressed

hairs 0.1-0.3 mm long, surfaces becoming grayish or

dark and smooth to striate. Leaves separate along the

stem or somewhat clustered distally, petioles (3-)5-

10(-1 5) mm long, ca. 1 mm thick, lateral margins form-

ing an adaxial sulcus and the midvein prominent within

the sulcus; leaf blades 6-14 cm long, 2-5 cm broad,

elliptic to elliptic-oblong, elliptic-obovate or narrowly

ovate-lanceolate, obtuse to bluntly acute at the apex

(sometimes rounded and rarely acute), acute to obtuse

at the base, drying stiffly chartaceous to subcoriaceous,

smooth and glabrous above and below (rarely minutely

puberulent beneath), often drying grayish and slightly

lustrous above, with 5-8 major secondary veins on each

side and the central secondaries arising at angles of 30°-

60°. Inflorescences axillary to distal leaves or pseudo-

terminal, 5-16 cm long, basal branches of the inflores-

cence longer than the distal and forming a rounded pan-

icle of 8-50 flowers, peduncle 3-9 cm long, very minutely

puberulent. Flowers 5-10 mm broad, white turning

brown, with the odor of jasmine, tepals imbricate, to 3

mm broad and rounded at the apex, minutely papillate

puberulent; outer anthers 1.5-2.5 mm long and sessile

or subsessile, ovate-triangular to lanceolate and flat, pap-

illate puberulent, staminodes inconspicuous or absent;

pistil 1.5-2 mm long. Fruits borne in a shallow cup 1-

1.6 cm broad and 1-5 mm deep, usually with an outer

undulate ridge and an interior entire margin, the cup

abruptly expanded above the thickened pedicel, red at

maturity; berry up to 2.5 cm long and 1.8 cm thick,

ellipsoid, becoming black.

A species of deciduous and partly deciduous

forest formations, from near sea level to about

1200 (1600) m elevation along the Pacific slope

and Meseta Central. The principal flowering time

is in December-March, and fruiting is mostly in

February-May. This species ranges from central

Mexico to Panama.

Ocotea veraguensis is recognized by its stiff gla-

brous leaves that often dry grayish (or yellowish),

inflorescences with long peduncles, relatively large

flowers with flattened papillate puberulent outer

stamens, and the short broad fruit cup with 2-

ridged margin. The restriction to deciduous or sea-

sonally very dry forests along the Pacific slope is

another distinction. The margins of the fruit cups

resemble those found in Licaria, and collections

from moister forests, such as on the Osa peninsula,

resemble L. cufodontisii in their darker more acu-

minate leaves.

Ocotea viridiflora Lundell, Wrightia 5: 36. 1974.

Trees 3-8(-20) m tall, leafy branchlets 1-3 mm thick,

glabrous, longitudinally ridged in early stages but soon

becoming terete and dark or grayish. Leaves alternate in

a spiral, petioles (4-)5-12 mm long, 0.7-1.2 mm thick,

glabrous, with adaxial ridges forming a deep narrow sul-

cus above near the stem; leaf blades 3-1 0(-1 2) cm long,

1-4 cm broad, narrowly elliptic to elliptic-oblong or

slightly oblanceolate, gradually narrowed to the acumin-

ate or caudate-acuminate apex, the narrow (2-3 mm) tip

1-2 cm long on some leaves, acute at the base and slightly

decurrent on the petiole, drying stiffly chartaceous, gla-

brous and slightly lustrous above with the venation

slightly elevated, glabrous beneath, with (3-)4-7 major

secondary veins on each side, central secondaries arising

at angles of 30°-60°, the minor venation raised beneath

when dry and forming a reticulum of weakly denned

areolae, distinctive pit domatia often present in the vein

axils and on the secondary veins beneath. Inflorescences

axillary to distal leaves, small (2-5 cm) and racemose or

umbellate (rarely paniculate), peduncles 1-3 cm long and

ca. 0.5 mm thick, glabrous, pedicels 2.5-7 mm long.

Flowers 2.5-4 mm long and 3-4 mm broad, glabrous on

the outside, tepals ca. 2 mm long and glabrous within;

outer stamens 1.3-2 mm long, outer anthers 0.8-1 mm

long and 0.6-0.8 mm broad, rectangular with superposed

thecae, inner stamens ca. 2.2 mm long with large (2.2

mm) glands and strigulose filaments, staminodes slender

and small (0.5 mm), slightly strigulose; pistil 2-3 mm

long, ovary ellipsoid, style half the length of the pistil,

stigma slightly discoid. Fruits borne in a small deep cup

ca. 7 mm long and 10 mm broad, perianth parts per-

sisting or deciduous and the cup entire, red; berry ellip-

soid to oblong, 12-25 mm long and 6-12 mm in di-

ameter, green (becoming black?).

Trees of cloud forest formations at elevations

of 1400 to 2000 m, along the central Cordilleras

and in the Chiriqui highlands. Flowers have been

collected in April-May; fruits have been collected

in March and May-June. A poorly understood

species of western Panama, possibly also repre-

sented in Costa Rica (see below).

Ocotea viridiflora is recognized by its small nar-

rowly elliptic glabrous leaves with elevated ve-

nation (when dry) and distinctive pit domatia, long-

acuminate leaf tips, small few-flowered glabrous

inflorescences, and oblong fruits in well-developed

cups. This species appears to be a close relative of

the larger-leaved O. meziana, and perhaps O. lae-

tevirens. The type of O. viridiflora differs from O.

meziana in having completely glabrous terminal

buds. Our use of this name for specimens from

Monteverde may be incorrect: the Panamanian