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

7
121
188
47
	,
	atae
	Laura;
	Legum i ; ,
101
	•
u> O -J

171 i

-

,

	Rani.
86
	Rhamn
94
	•
134
141

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