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Full text of "Flora Costaricensis"

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 











































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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 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 34 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 612 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 cm long, elliptic to narrowly oblong; from above 1 000 m elevation . 

Fig. 4 

4b. Leaves usually becoming more than 1 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 30S 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 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 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; 0400 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 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) c m l n g. 4-1 1(-14) cm broad, ovate-lanceo- 
late to ovate-elliptic or elliptic-oblong, usually broadest 
below the middle and tapering gradually to the long- 
acuminate apex, obtuse to acute at the base, margin en- 
tire and with a vein along the edge, the margin often 
revolute near the petiole with expanded auriculate flaps 
4-16 mm long forming enclosed spaces on both sides of 
the lamina base beneath, drying stiffly chartaceous, gray- 
ish puberulent on the veins above and smooth to the 
touch, brownish tomentulose beneath with slender 
crooked hairs ca. 0.5 mm long, with 6-16 major sec- 
ondary veins on each side, tertiary venation subparallel 
and prominent beneath. Inflorescences 6-18 cm long, 
axillary to distal leaves, paniculate with few or many 
clusters of closely approximate flowers on prominent 
lateral branches, peduncle 3-10 cm long, 1.5-2 mm thick, 
densely puberulent, lateral branches 3-5 cm long. Flow- 
ers 4-8 mm long and 8-12 mm broad, white at anthesis 
but becoming reddish brown, tepals 4-5.5 mm long, 
densely papillate-puberulent within; outer stamens broad 
and flat, 1.5-1.7 mm long, thecae in a plane or in a partly 
superposed arc, connective usually developed distally 
beyond the thecae, staminodia slender or absent; pistil 
2.5-3.5 mm long, stigma discoid. Fruits not seen from 
Central America (the following from high altitude col- 
lections in the Andes): fruiting receptacles obconic, 6- 
12 mm long and 10-16 mm broad, shallow and saucer- 
like or deeper (2-4 mm) and more cupulate; berry 10- 
20 mm long, ovoid-oblong, abruptly rounded at the apex. 

Trees of both evergreen rain forests and ever- 
green formations with a pronounced dry season, 
on both the Caribbean and Pacific sides of Costa 
Rica, from near sea level to 1200 (71600) m ele- 
vation. Flowers have been collected in October- 
February and April. The species ranges from 
southern Mexico through Central America to Peru 
and southern Brazil. 

Nectandra reticulata is recognized by its nar- 
rowly ovate (almost lanceolate) leaves with long- 
acuminate apices and dense brownish puberulence 



beneath. This species often has flaplike auriculate 
developments on the underside of the lamina near 
the petiole. These are bent under to form partly 
enclosed spaces, but not all leaves or specimens 
have this development. The lack of mature fruiting 
collections in Central America is unusual for a 
species that is fairly often collected. The early stages 
of fruit development seem to have the drupe com- 
pletely enclosed within the urceolate hypanthium. 



Nectandra salicifolia (H.B.K.) Nees, Syst. laur. 302. 
1836. Ocotea salicifolia H.B.K., Nov. Gen. Sp. 
2: 166. 1817. Figure 17. 

Shrubs and small trees 4-12 (rarely to 25) m tall, leafy 
branchlets 1.5-4 mm thick, minutely (0.1-0.2 mm) ap- 
pressed puberulent with thin ascending hairs, becoming 
(sub)glabrous and brownish to gray. Leaves alternate, 
petioles 3-8(-14) mm long, ca. 1 mm thick, usually sul- 
cate above; leaf blades 6-15 cm long, 1.5-6 cm broad, 
narrowly elliptic, elliptic-lanceolate or elliptic-oblong to 
ovate-elliptic, tapering gradually or abruptly to an acu- 
minate (rarely acute) apex, the tip 4-16 mm long, acute 
to obtuse at the base and slightly decurrent on the petiole, 
margin entire, drying stiffly chartaceous to subcoriaceous 
and often darker above than below when dried, glabrous 
above and usually lustrous, glabrous beneath except for 
the tufted hairs (domatia) in the axils of proximal sec- 
ondary veins in some collections, with 4-7 major sec- 
ondary veins on each side, the secondaries usually weak- 
ly loop-connected near the margin in the distal half of 
the lamina, minor venation raised on both surfaces when 
dry. Inflorescences solitary in the axils of distal leaves 
or pseudoterminal (sometimes appearing to be several 
in the axil due to a very short peduncle), broadly pani- 
culate and many-flowered or rarely racemose to umbel- 
late in few-flowered panicles, 4-ll(-17) cm long, pri- 
mary peduncle (2-)6-60 mm long, reddish to whitish 
and sparsely to densely puberulent with slender minute 
(0. 1 mm) hairs, pedicels 2-3(-4) mm long. Flowers white, 
3-4 mm long, 5-6(-8) mm broad, tepals 2.5-3.8 mm 
long and 1.2-1.7 mm broad, spreading or reflexed at 
anthesis, puberulent on the outside and papillate within; 
outer stamens subsessile or the filaments to 0.3 mm long, 
outer anthers 0.4-0.6 mm long, 0.5-0.7 mm broad, rect- 
angular to reniform and convex to emarginate distally, 
the thecae in a single plane or the lower lateral to the 
upper in an arc, inner stamens ca. 0.8 mm long and with 
large glands, staminodes 0.4-0.8 mm long and slender 
or slightly thickened apically (sometimes absent or mi- 
nute); pistil 1-1 .7 mm long, ovary 0.6-1 mm in diameter 
(rarely puberulent), style 0.3-0.6 mm long, stigma dis- 
coid or simple. Fruits borne on a short-conical or saucer- 
like receptacle 3-5 mm long, (5-)8-10 mm broad and 
1-3 mm deep, the margin entire or slightly undulate, 
becoming reddish or purple; berry 1-1.6 cm long and 1- 
1.5 cm in diameter, subglobose to ellipsoid-oblong, be- 
coming black (red?). 

Shrubs and small trees of evergreen and partly 
deciduous forest formations, between sea level and 



64 



FIELDIANA: BOTANY 



600 (900) m elevation in the Caribbean lowlands, 
the General Valley and the Golfo Dulce region in 
Costa Rica. Flowering material has been collected 
in January-August in Costa Rica and January- 
June in Guatemala; mature fruits have been col- 
lected in August-November in Central America. 
This species (in a wide sense) ranges from north- 
eastern Mexico through Central America to Costa 
Rica and Panama. 

Nectandra salicifolia is recognized by the rela- 
tively small, stiff, essentially glabrous leaves with 
the minor venation raised on both surfaces and 
the upper surface usually lustrous when dried. The 
usual presence of tufted domatia and the weakly 
loop-connected secondary veins are associated with 
small puberulent flowers with very small anthers, 
short styles, small shallow fruiting receptacles, and 
usually subglobose fruits. These characters vary 
greatly and can make some populations appear 
quite different from others. Collections from 
southwestern Costa Rica tend to have more cau- 
date-acuminate leaves with conspicuous "drip 
tips," while material from the Caribbean slope is 
often narrowly elliptic. Nectandra salicina appears 
to be a highland derivative of N. salicifolia, and 
there appear to be collections intermediate be- 
tween the two in the Cordillera de Guanacaste; see 
the discussion under N. salicina. Nectandra sa- 
vannarum (Standl. & Steyerm.) C. K. Allen of 
Guatemala and Belize is probably an unusual form 
of N. salicifolia with small umbellate inflores- 
cences and small ovate leaves with prominent bas- 
al secondary veins. Trees referred to as N. salici- 
folia in Paul Allen's book (1956) are actually N. 
turbacensis. Some material placed here was earlier 
identified as N. coriacea (Sw.) Griseb., which is 
probably restricted to the West Indies and Yucatan 
peninsula (Rohwer, pers. comm.). 



Nectandra salicina C. K. Allen, J. Arnold Arbor. 
26: 385. 1945. N. smithii C. K. Allen, loc. cit. 
370. 1945. Figure 4. 

Trees 5-10 m tall, leafy branchlets l-4(-5) mm thick, 
at first minutely puberulent with appressed yellowish 
brown hairs, becoming glabrous, smooth, and grayish. 
Leaves alternate and often densely clustered at the ends 
of branches, petioles 4-12 mm long, ca. 1 mm thick, flat 
or slightly sulcate above and with lateral margins; leaf 
blades 5-9(-14) cm long, l-3(-4) cm broad, narrowly 
elliptic to lanceolate or elliptic-oblong, tapering gradu- 
ally to the acuminate apex, the narrowed tip 0.5-2 cm 
long, tapering gradually to the attenuate base, margin 
entire or undulate, drying stiffly chartaceous or subcor- 
iaceous and often greenish brown, glabrous and lustrous 



above with the tertiary veins slightly raised, glabrous or 
very sparsely puberulent beneath, rarely with tufts of 
hairs (domatia) in the vein axils, with (3-)4-6(-7) major 
secondary veins on each side, the tertiary venation al- 
ways forming a raised but irregular reticulum on the 
upper surface of the dried leaf. Inflorescences axillary or 
extra-axillary (nodes with undeveloped leaves?), to 12 
cm long, paniculate but with few lateral branches, pe- 
duncles (2-)4-8 cm long, minutely puberulent, pedicels 
ca. 4 mm long and articulate in the middle. Flowers ca. 
3.5 mm long and 6-7 mm broad, yellowish white, dense- 
ly and minutely puberulent, tepals 2.5-3 mm long, densely 
papillate-puberulent within; androecium ca. 3 mm in 
diameter, outer stamens with short (0.2 mm) filaments, 
outer anthers 0.5-0.8 mm long and 0.7-1 mm broad, 
thecae usually in an arc with the lower lateral to the 
upper, inner stamens narrow, staminodes to 1 mm long 
or absent; pistil 1-1.8 mm long, ovary globose, 0.7-1.1 
mm in diameter, style narrow, stigma discoid or simple. 
Fruits borne in a conical or broadly campanulate and 
very shallow (1-2 mm) cup, 5-10 mm long above the 
thickened pedicel and 4-1 1 mm broad at the top, margin 
entire or undulate, becoming red; berry 10-18 mm long 
and 7-16 mm thick (dry), ellipsoid during development 
but becoming globose at full maturity. 



Trees of evergreen forests on both the Caribbean 
and Pacific slopes, from about 600 to 1700 m 
elevation. Flowering collections have been made 
in January-May and July-August; fruits have been 
collected in March-November. The species, as here 
defined, ranges from the westernmost parts of the 
Cordillera de Guanacaste through the Cordillera 
de Tilaran to the eastern part of the central vol- 
canic highlands (near Turrialba, Cartage). 

Nectandra salicina is recognized by its small 
lustrous narrowly elliptic and acuminate leaves 
with the tertiary venation usually prominent on 
both surfaces. The small few-flowered inflores- 
cences with reddish rachises and puberulent little 
flowers, and the ellipsoid to globose fruits borne 
in shallow cups also distinguish this species. The 
wood and leaves are said to be sweet-smelling. 
Nectandra salicina appears to intergrade with very 
similar smaller-leaved specimens of N. salicifolia 
at lower elevations, and it would appear that it 
should really be called a subspecies of N. salici- 
folia. However, N. salicifolia encompasses much 
geographic variation in the lowlands, and its re- 
lationship with material called N. latifolia is not 
clear. Until these matters are clarified, we prefer 
to think of these taxa as species; see the discussion 
under N. latifolia. Specimens intermediate be- 
tween typical N. salicina and N. salicifolia have 
been collected at lower elevations (about 400 to 
900 m) in the Cordillera de Guanacaste, but they 
lack fruits, which are generally larger in N. salicina. 

Nectandra smithii was based on Austin Smith 



BURGER: FLORA COSTARICENSIS 



65 



//. 541 (AA,F) from near Zarcero at 1600 m ele- 
vation. The leaves resemble those of some collec- 
tions ofN. salicifolia but are more elliptic-oblong, 
and a few of the leaves have distinctive slightly 
rounded caudate-acuminate tips. Brenes 6825 (F) 
was cited in the description ofN. smithiiand seems 
to be intermediate between the type collection and 
typical material of N. salicina; it is for this reason 
that N. smithii appears to be an unusual variant 
of N. salicina. Rohwer (p. 81, 1986) points out 
that the flowers have some Ocotea-like character- 
istics, both as regards pubescence and thecae ar- 
rangement in the outer anthers. This suggests that 
the type may be of hybrid origin; however, these 
characteristics are also sometimes seen in N. sal- 
icina. 



Nectandra sinuata Mez, Jahrb. Konigl. Bot. Gart. 
Berlin 5: 402. 1889. Figure 7. 

Small or medium-sized trees to 20 m tall, trunk to 1 
m d.b.h., leafy branchlets 2-8 mm thick, densely villous- 
tomentulose with slender yellowish brown or grayish 
hairs ca. 0.5 mm long. Leaves alternate, petioles 8-32(-40) 
mm long, 1-4 mm thick, densely puberulent and ob- 
scurely sulcate above; leaf blades 10-22(-30) cm long, 
3.5-1 2(-l 9) cm broad, oblong-elliptic to slightly obovate 
or oblong-obovate, usually abruptly narrowed to the ob- 
tuse or short-acuminate apex, usually abruptly narrowed 
to the obtuse or truncated base (rarely acute), the larger 
leaves often rounded and subcordate at the base, laminae 
drying stiffly chartaceous, sparsely puberulent above and 
densely tomentulose to sericeous beneath with slender 
straight or curved hairs 0.3-1 mm long, with 10-12 ma- 
jor secondary veins on each side, the central secondaries 
arising at angles of 40-75. Inflorescences axillary to 
distal leaves, to 30 cm long, open panicles puberulent in 
all parts, peduncle to 1 6 cm long (often more than half 
the length of the inflorescence), ca. 2 mm thick, pedicels 
5-12 mm long. Flowers ca. 10 mm long, 10-20 mm 
broad, tepals 5-8 mm long and ca. 4 mm broad (dry), 
pale rose, pink or white (reddish brown when dry), dense- 
ly tomentulose on the outside and papillate-puberulent 
within; androecium ca. 5 mm in diameter, outer anthers 
2-3 mm long on short (0.5 mm) thick filaments, con- 
nective often expanded and the stamen tepal-like in form, 
thecae in an arc or sometimes superposed, staminodes 
absent; pistil 2-3 mm long, style ca. 1 mm long with 
lobed disclike stigma. Fruit cups ca. 20 mm long and 16 
mm broad, entire or lobed at the margin, ca. 5 mm deep; 
berry 2-2.5 cm long and 1-1.8 cm thick, ellipsoid-ob- 
long, sparsely to densely sericeous in the lower fourth, 
glabrous above except at the apex, becoming black. 

Trees of evergreen and partly deciduous forest 
formations of the central highlands and Pacific 
slope in Costa Rica, between 300 and 1500 m 
elevation. (A few collections have been made on 
the Caribbean slope near Turrialba, but these may 



have been planted.) Flowering collections have 
been made in January-April in Costa Rica, and 
fruits have been collected in April-June. The 
species ranges from southern Mexico along the 
Pacific side of Central America to the western part 
of the Cordillera de Talamanca (near Sta. Maria 
de Dota). 

Nectandra sinuata is distinguished by the dense 
puberulence of its long-peduncled inflorescences 
and lower leaf surfaces, the large densely puber- 
ulent flowers, and the ellipsoid fruits with ap- 
pressed hairs near the base. The oblong leaves 
abruptly narrowed at both apex and base, pref- 
erence for a seasonally dry habitat, and frequent 
pink flowers further characterize this species. The 
tepal-like anthers are unusual and sometimes have 
the valves in a superposed Ocotea-like arrange- 
ment. Rohwer (pers. comm.) has suggested trans- 
ferring this species to Ocotea. 



Nectandra turbacensis (H.B.K.) Nees, Hufeland. 
ill. 14. 1833; Syst. laur. 316. 1836. Ocotea tur- 
bacensis H.B.K., Nov. Gen. Sp. 2: 162. 1817. 
AT. concinna Nees, Syst. laur. 322. 1836. N. ner- 
vosa Mez & J. D. Smith ex Mez, Bull. Herb. 
Boissier, ser. 2, 3: 235. 1903. Figure 18. 

Trees to 20(30) m tall, to 80 cm d.b.h., leafy branchlets 
1.5-5 mm thick, glabrous or sparsely puberulent with 
thin minute (0.2 mm) appressed ascending hairs, becom- 
ing dark and smooth. Leaves alternate, petioles 5-12 mm 
long, 0.7-1.2 mm thick, usually glabrous and dark, flat 
or slightly sulcate between the adaxial ridges; leaf blades 
5-17 cm long, 2-4.5(-6) cm broad, lanceolate to nar- 
rowly ovate or narrowly oblong, tapering gradually to 
the acute or acuminate apex, acute (rarely obtuse) at the 
base, margin entire or undulate, stiffly chartaceous and 
often grayish green above, glabrous above and below, 
the midvein and minor venation slightly raised above, 
with (3-)5-7 major secondary veins on each side (rarely 
with a basal pair of secondaries strongly ascending and 
almost tripliveined, as in Allen 5710), ciliate pit domatia 
often present in proximal vein axils beneath. Inflores- 
cences in the axils of distal leaves or pseudoterminal, to 
1 7 cm long, paniculate with spreading lateral branches, 
peduncle (0.5-)2-6 cm long, 0.5-1.5 mm thick, glabrous 
(rarely sparsely and minutely puberulent), pedicels 1-4 
mm long. Flowers ca. 3 mm long and 5-8 mm broad, 
white, sparsely puberulent on the outside, tepals 2.2-3 
mm long, 2-2.6 mm broad, densely papillate-puberulent 
within; androecium 3 mm in diameter, outer stamens 
subsessile, outer anthers 0.6-0.7 mm long, 0.8-1.1 mm 
broad, thecae in a horizontal row, the connective slightly 
expanded distally and papillate, inner stamens 0.9-1.4 
mm long, staminodes minute or absent; pistil 1.5-2 mm 
long, ovary 1-1.4 mm in diameter, style short (ca. 0.8 
mm) and thick, stigma discoid. Fruits borne on a short 
(4-10 mm) cupulate receptacle 8-12 mm broad at the 
apex and (2-)3-4 mm deep, margin entire and often with 



66 



FIELDIANA: BOTANY 



2 concentric ridges, greenish or bluish; berry 10-14 mm 
long and 7-10 mm thick (dry), oblong-ellipsoid, becom- 
ing black. 

Trees usually found in secondary associations 
in evergreen and partly deciduous forest forma- 
tions, from near sea level to 1200(1700) m. Flow- 
ers have been collected in September-March (May) 
in Costa Rica; fruiting is said to occur in March- 
April in southwestern Costa Rica (P. H. Allen, 
1956). The species ranges from Veracruz, Mexico, 
to Colombia, Venezuela, Peru, and Bolivia. 

Nectandra turbacensis is recognized by the gla- 
brous leaves that dry dull grayish with smooth 
surfaces and obscure minor venation (as in the 
type) or with the minor venation slightly raised 
on a darker background (as in the populations of 
southwestern Costa Rica). The outer subsessile an- 
thers with their slightly prolonged (but truncated 
or obtuse) and papillate connective and their the- 
cae in a single plane are distinctive. The entire 
fruit cup enclosing about '/4 of the fruit and a 
general lack of puberulence (except for distal parts 
of the inflorescence and perianth) further distin- 
guish this species. The veins are not loop-con- 
nected near the margin, and the flowers and sta- 
mens are larger than in N. salicifolia and its allies. 

Trees with long narrow leaves and prominent 
minor venation have been called N. panamensis 
Mez, and are characteristic of southwestern Costa 
Rica and western Panama in the Pacific lowlands. 
It may be that the populations of the Pacific low- 
lands deserve subspecific recognition. The species 
is rarely collected in Central America and appears 
to be restricted to the Caribbean lowlands in 
northern Central America. A population with ab- 
normal flowers was sampled by Standley and Val- 
erio (45469, 45906, 45909, 46192, all us) near 
Tilaran in 1926. 



Ocotea Aublet 

REFERENCE J. G. Rohwer, Prodromus einer 
Monographic der Gattung Ocotea Aubl. (Laura- 
ceae), sensu lato. Mitt. Inst. Allg. Bot. Hamburg 
20: 1-278. 1986. 

Trees or shrubs, bisexual or rarely unisexual (dioe- 
cious), glabrous to densely puberulent. Leaves alternate, 
rarely subopposite or verticellate, petiolate or sessile but 
narrowed near the base, the leaf blades entire, charta- 
ceous to coriaceous when dry, with pinnate venation 
(rarely with strongly developed basal secondaries and 
subtripliveined), with or without domatia. Inflores- 



cences usually solitary and axillary to distal leaves, some- 
times pseudoterminal, lacking a subtending involucre of 
bracts, paniculate or occasionally few-flowered and ra- 
cemose, pedicels usually well developed, bracts and brac- 
teoles usually deciduous. Flowers bisexual or unisexual, 
(male flowers usually with a pistillode and female flowers 
with staminodes when unisexual), generally small (2-5 
mm long), white to yellowish or green, the floral tube 
(hypanthium) well developed, small or absent but always 
accrescent in fruits, perianth 6-parted with 2 whorls of 
3 tepals each, imbricate, equal or subequal, the tepals 
thick to thin and glabrous to densely puberulent; fertile 
stamens 9 in 3 series, the outer 2 series (often appearing 
as a single whorl) similar in size and shape and with 
introrse dehiscence, filaments present or absent, anthers 
variously shaped but usually as broad as long or longer 
than broad (flat and triangular to laminar or tepal-like 
in some species), the connective rarely prolonged beyond 
the thecae, each anther with 4 thecae arranged in 2 planes 
(superposed) or with the lower thecae slightly lateral to 
the upper (this character can vary within the same flower 
in a few species and makes differentiation from Nectan- 
dra difficult), opening by flaps attached at the top, the 
inner 3 stamens also 4-thecous, usually with longer fil- 
aments and narrower anthers that open extrorsely or with 
the upper thecae opening laterally and the lower extrorse, 
each inner stamen with 2 conspicuous sessile (rarely stip- 
itate) glands at the base of the filament (6 glands per 
flower), the glands thick and bluntly rounded or angular, 
an inner whorl of staminodes (series IV) usually absent 
or poorly developed (less than 0.6 mm long); pistil with 
globose, ovoid, or turbinate ovary, usually slightly nar- 
rowed at the base, rarely puberulent, style slender or 
thick, stigma simple to discoid or capitate (rarely 2- or 
3-lobed). Fruits borne on a flattened discoid to cupulate 
or hemispheric receptacle, perianth usually deciduous 
and the cup with a single-margined edge (with a double 
margined rim in O. dendrodaphne, O. veraguensis and 
some flat disclike receptacles), the margin sometimes 
undulate (rarely 6-lobed), pedicel thickened in fruits and 
erect or pendulous; fruits a single-seeded berry, ovoid to 
ellipsoid, oblong or globose, usually narrowed and slight- 
ly flattened at the base, abruptly rounded at the apex, 
style or stigma rarely persisting. 

A genus of perhaps 300 to 400 species centered 
in the American tropics, with a few species in Af- 
rica and a group of species in Madagascar. In Costa 
Rica there appear to be a few species intermediate 
between Ocotea and Nectandra, and between Oco- 
tea and Phoebe. The usual lack of three well-de- 
veloped staminodes help distinguish Ocotea from 
our species of Phoebe. The character of superposed 
(2-ranked) thecae valves, which separates Ocotea 
from Nectandra, displays considerable variation 
with some intermediate states among a few species. 
While it is easy to say that these relatively rare 
intermediate states invalidate current generic con- 
cepts, there does not seem to be any way of de- 
veloping better generic concepts at this time. 
Lumping these genera together produces nomen- 
clatural confusion with no increased understand- 



BURGER: FLORA COSTARICENSIS 



67 



ing of how the species should be arranged. The 
present grouping of species in Ocotea appears to 
form a central assemblage from which other lin- 
eages may have been derived (cf. Rohwer & Ku- 
bitzki, 1985; Rohwer, 1986). For example, one 
population of O. insularis appears to be in the 
process of losing the upper thecae in each anther, 
and this may have been how Aiouea costaricensis 
originated (see the discussion under these species 
and under Aiouea, Nectandra, and Phoebe). 

Ocotea is recognized by the 6-parted flowers with 
equal or subequal tepals, the nine stamens with 
four superposed valves in each, the three inner 



stamens with two conspicuous glands, the absence 
or poor development of staminodes, and the re- 
ceptacle forming a cup or disc in fruits. There is 
great morphological variation within our species. 
The stamens range from sessile lamina-like struc- 
tures to those with well-developed slender fila- 
ments and clearly differentiated anthers. Compar- 
ison with other genera and with related species 
within Ocotea suggests that laminar stamens are 
not a primitive condition in Lauraceae (cf. Rohwer 
& Kubitzki, 1985). Likewise, the fruiting recep- 
tacle ranges from a small disc to a deep hemi- 
spheric cup. 



Key to Species of Ocotea in Costa Rica 

la. Distal branchlets hollow and often harboring small ants; leaves usually more than 1 5 cm long and 
narrowly oblong to elliptic-oblong or narrowly obovate, always glabrous beneath; small trees of 

wet evergreen forest understory 2a 

1 b. Distal branchlets solid, the center with wood or pith and not consistently hollow 6a 

2a. Flowers 4-6 mm long; anthers 2-3 mm long, flat and tepal-like; fruit cups ca. 1 5 mm broad 
and 3-7 mm deep; leaves drying grayish green and subcoriaceous, usually elliptic-oblong; 0- 

1 500 m elevation O. dendrodaphne 

2b. Flowers 2-3 mm long; anthers to 1 mm long and not tepal-like; fruit cups 6-12 mm broad 

and only 1-2 mm deep 3a 

3a. Leaves usually drying chartaceous and usually very dark in color; 0-1000 m elevation . . . 

4a 

3b. Leaves usually drying subcoriaceous and grayish green to orange brown or dull brown .... 

5a 

4a. Leaves becoming 30-50 cm long, often narrowly obovate; flowers essentially glabrous, 

outer filaments glabrous; fruits 20-35 mm long O. atirrensis 

4b. Leaves not exceeding 30 cm in length, usually narrowly elliptic-oblong; flowers minutely 

puberulent, filaments minutely puberulent; fruits 1 2-20 mm long O. wedeliana 

5a. Distal stems strongly angled with 3-5 prominent longitudinal ridges; leaves to 40(-55) cm 
long, often narrowly obovate; fruit cups often with persisting perianth bases; 0-1 100 m . . 

O. nicaraguensis 

5b. Distal stems without prominent ridges, not strongly angled in cross section; leaves 10-30 cm 

long, often narrowly oblong; fruit cups with entire margins; 600-2300 m elevation 

O. paulii 

6a. Leaves usually becoming more than 30 cm long, obovate and gradually narrowed to the base, 

drying stiffly chartaceous 7a 

6b. Leaves rarely more than 30 cm in length (sprout shoots from nonflowering basal branches may 

be larger) 9a 

7a. Leaves to 50 cm long and 25 cm broad, glabrous beneath; flowers 3 mm long and 3 mm 

broad, outer anthers ca. 0.6 mm long; Golfo Dulce area O. rivularis 

7b. Leaves to 40 cm long, conspicuously puberulent beneath; flowers 5-7 mm long and 6-10 

mm broad; Caribbean slope and lowlands 8a 

8a. Leaves often narrowly obovate, to 1 6 cm broad; inflorescence rachis sparsely hirsute; outer 

anthers 1 .2-1 .4 mm long; ca. 1 400 m elevation O. lentil 

8b. Leaves often broadly obovate, to 22 cm broad; inflorescence rachis densely yellowish pu- 
berulent; outer anthers 1-1 .2 mm long; 0-900 m O. valerioides 



68 



FIELDIANA: BOTANY 



9a. Leaves densely puberulent beneath with longer (0.3-1 mm) hairs, the hairs spreading or appressed, 

puberulence on the lower surface soft or slightly rough to the touch 1 Oa 

9b. Leaves glabrous to sparsely puberulent beneath (lower surface), the hairs small (0.05-0.2 mm) or 
inconspicuous and difficult to see with a 1 OX hand lens, puberulence of the lower leaf surface not 

discernable to the touch 25a 

lOa. Lower leaf surface sericeous with lustrous silvery or reddish hairs, the leaves drying stiffly 
coriaceous and decurrent on a very short petiole; flowers 4-7 mm long and densely puberulent; 

only from above 2000 m O. calophylla 

lOb. Lower leaf surfaces not densely sericeous with lustrous silvery or reddish hairs; petioles well 

developed 1 la 

1 la. Largest leaves rarely more than 1 cm long 1 2a 

1 Ib. Largest leaves usually more than 1 3 cm long 1 5a 

12a. Flowers glabrous to sparsely puberulent, often with well-developed staminodes; lamina 

base not decurrent on the petiole, leaves usually drying chartaceous 1 3a 

12b. Flowers minutely and densely puberulent, staminodes absent; lamina base decurrent 

on the petiole, leaves often drying subcoriaceous 1 4a 

1 3a. Fruit cups 10-1 5 mm broad; pubescence on the lower leaf surfaces brownish and 

slightly rough to the touch; 1000-3200 m elevation O. pittieri 

1 3b. Fruit cups ca. 6 mm broad; pubescence on the lower leaf surfaces grayish and 

soft to the touch; 1400-2300 m elevation O. mollicella 

14a. Leaves usually elliptic and 1.5-4 cm broad, with an acute apex; flowers 34 mm long, 

outer anthers 0.6-0.9 mm long; western highlands, 1400-1600 m . O. monteverdensis 

14b. Leaves usually oblong and 1.5-6 cm broad, with an obtuse apex; flowers 4-5 mm long, 

outer anthers ca. 1 mm long; central and eastern highlands, 1 700-3000 m elevation 

O. austinii 

1 5a. Base of the leaf blades decurrent on the petiole and the petiole often poorly denned, margin 

of the lamina often revolute near the petiole 1 6a 

1 5b. Base of the leaf blade not consistently decurrent on the petiole 1 9a 

16a. Leaves usually narrowly elliptic to oblong, 1.5-6 cm broad; montane species rarely 

encountered below 1 200 m elevation 1 4a 

16b. Leaves often obovate or slightly obovate, 3-1 1 cm broad; rarely found above 1 100 m 

17a 

17a. Petioles clearly differentiated from the narrowed lamina base, leaves drying stiffly 
chartaceous, with 4-8 pairs of major secondary veins; fruit cups 10-16 mm broad . . 

O. hartshorniana 

17b. Petioles not clearly differentiated from the narrowed lamina base, leaves drying sub- 
coriaceous, with 9-12 pairs of major secondary veins; fruit cups 5-12 mm broad . . . 

18a 

1 8a. Fruit cups flat and saucer-like, without dentate lobes O. stenoneura 

1 8b. Fruit cups deeply cupulate and with persisting perianth lobes O. dentata 

19a. Leaves broadly obovate to broadly elliptic, usually rounded or abruptly narrowed at the 
apex, usually drying dark brown above, densely puberulent beneath; outer anthers 0.7-1.5 

mm long; Caribbean slope and central Cordilleras 20a 

1 9b. Leaves rarely obovate or broadly elliptic, not rounded at the apex, rarely drying dark brown, 

sparsely puberulent beneath 25a 

20a. Trees of montane formations, (8 00-) 1000-2 5 00 m elevation; leaves often drying sub- 
coriaceous, oblong to broadly elliptic, obovate or suborbicular; flowers 6-1 5 mm broad 

21a 

20b. Trees not known from above 800 m elevation in Costa Rica; laminae usually drying 
stiffly chartaceous, usually obovate to oblong or pandurate; flowers 4-12 mm broad 

23a 

2 la. Flowers glabrous on the outside, outer anthers subsessile or with very short 
filaments; fruits borne in a deep (5-8 mm) cup 10-18 mm broad . O. valeriana 



BURGER: FLORA COSTARICENSIS 69 



2 1 b. Flowers densely puberulent on the outside, outer anthers with prominent filaments 

22a 

22a. Leaves usually elliptic to oblong, 3-9 cm broad; flowers 6-8 mm broad; fruits in 

a shallow entire cup; 1 500-2500 m elevation O. pseudopalmana 

22b. Leaves usually broadly elliptic to suborbicular, 6-14 cm broad; flowers ca. 12 

mm broad; fruits in a deep lobed cup; 800-1400 m O. gomezii 

23a. Leaves 10-22 cm broad, obovate to pandurate, on short thick (3-6 mm) petioles; 

inflorescences apparently erect; perianth sparsely puberulent O. valerioides 

23b. Leaves 4-14 cm broad, petioles rarely more than 3 mm thick 24a 

24a. Leaves often obovate to pandurate; inflorescences pendant with long thin peduncles, 
flowers often borne on umbellate lateral branches; perianth usually glabrous on the 

outside and drying dark O. helicterifolia 

24b. Leaves obovate to broadly oblong; inflorescences usually erect and broadly paniculate 
with racemose or branched lateral branches; perianth puberulent on the outside and 

drying brownish O. mollifolia 

25a. Distal branchlets strongly winged (alate) with narrow longitudinal wings 2-3 mm high, the stems 
3-5-angled in cross section; leaves narrowly oblong to narrowly obovate and coriaceous, with 9- 

14 pairs of major secondary veins; Pacific slope below 1000 m elevation O. aurantiodora 

25b. Distal branches without conspicuous longitudinal wings, the stems sometimes bluntly angled in 

cross section but not with 2-3 mm high ridges 26a 

26a. Leaf base usually decurrent on the petiole, the leaf base and petiole often poorly differentiated, 

lamina margin often revolute near the petiole, the leaves petiolate to sessile 27a 

26b. Leaf base not usually decurrent on the petiole, lamina acute to obtuse or rounded at the base and 

usually clearly differentiated from the petioles, the leaves never sessile or subsessile 36a 

27a. Leaf base broadly revolute (auriculate) and forming 2 broad flaps on the underside, petiole 
little developed and the leaves subsessile, broadly obovate and often coriaceous; (700-) 1 1 00- 

2300 m elevation O. endresiana 

27b. Leaf base flat or revolute but not forming broad flaps beneath, never auriculate and usually 

petiolate 28a 

28a. Mature leaves usually with slender appressed ascending hairs parallel with the secondary 

veins over the lower surface; trees often with prop roots at the base of the trunk 29a 

28b. Mature leaves glabrous or variously puberulent but not with slender appressed hairs paral- 
leling the secondary veins on the lower surfaces; trees without prop roots(?) 3 la 

29a. Leaves elliptic to oblanceolate or obovate, 2-5 cm broad, leaf base usually long-de- 
current (to 5 cm long), with 5-9 pairs of major secondary veins; fruits ellipsoid; Pacific 

slope at 600-1400 m (if higher elevation cf. O. whitei) O. skutchii 

29b. Leaves broadly obovate, 4.5-1 1 cm broad, long- or short-decurrent at the base, with 

7-12 pairs of major secondary veins; fruits globose to ovoid and 2-4 cm long . . 30a 

30a. Fruits ellipsoid to ovoid, 3-4 cm long in large deep ( 1 2 mm) cups; leaves drying smooth 

and dull above with the minor venation obscure; Caribbean lowlands 

O. sp. aff. caracasana 

30b. Fruits globose to oblong, 2-3 cm long, often in conical cups with reflexed edges; leaves 
drying with a lustrous sheen and the minor venation raised above; 1500-2500 m 

elevation O. glaucosericea 

3 1 a. Leaves often rounded at the apex or bluntly obtuse (rarely acute to short-acuminate in smaller 
leaves), often obovate, (2-)5-9(-l 2) cm broad, drying coriaceous or subcoriaceous and often 
yellowish brown to pale reddish brown; fruit cups 6-9 mm broad and often with persisting 

perianth bases on the edge, fruits ellipsoid, 1-2 cm long; 0-2000 m O. insularis 

3 Ib. Leaves bluntly acute to acuminate at the apex, sometimes bluntly obtuse but never rounded, 
often oblanceolate to narrowly elliptic-obovate or oblong, 1.5-5(-6) cm broad; fruit cups 

without persisting perianth 32a 

32a. Flowers 2-5 mm long, functionally unisexual; fruits subglobose, 6-16 mm in diameter; leaves 

drying grayish green or yellowish green, elliptic-oblong to obovate 33a 

32b. Flowers 1.5-3 mm long, bisexual; fruits globose or oblong, but never with a persisting style; 

70 FIELDIANA: BOTANY 



leaves often drying very dark in Costa Rica but variable in O. whitei, often oblanceolate to 

very narrowly elliptical 34a 

33a. Fruits with a persisting style base at the top and borne on a thick flat double-rimmed 
disclike receptacle 6-10 mm broad, berry 10-16 mm in diameter; leaves often obovate 

and drying grayish; 0-1 900 m O. floribunda 

33b. Fruits without a persisting style base, receptacle 4-8 mm broad, berry 6-8 mm in 

diameter (dried); leaves often elliptic-oblong and drying yellowish; 0-500 m 

O. puberula 

34a. Fruits 1 5-38 mm long and 11-18 mm thick (dry), ellipsoid and borne in cupulate receptacles 
8-14 mm broad; leaves often narrowly elliptic; 1300-2500 m elevation O. whitei 

34b. Fruits rarely exceeding 1 5 mm in length, globose to oblong, subtended by shallow receptacles 
4-6 mm broad; leaves usually oblanceolate to narrowly elliptic-obovate 35a 

35a. Flowers 2-3 mm long, outer anthers ca. 0.6 mm long; fruits globose, 5-8 mm in diameter 
(dry); leaves without domatia; Caribbean lowlands O. sp. aff. O. bijuga 

35b. Flowers 1.5-2.5 mm long, outer anthers ca. 1 mm long; fruits oblong, 9-18 mm long and 
7-9 mm in diameter; pit domatia often present on leaf undersurfaces; (0-)600-1000 m . . . 

O. oblonga 

36a. Leaves drying very dark (almost black) and thin-chartaceous; fruiting receptacle gradually ex- 
panding (obconic) to the 1 cm broad apex, concave or slightly cupulate, fruits 2-3 cm long and 1- 

2 cm thick; flowers glabrous and small 37a 

36b. Leaves drying grayish to dark brown, stiffly chartaceous to coriaceous if drying very dark . . 38a 

37a. Flowers 1.5-3 mm long, outer anthers ca. 0.7 mm long and the connective not expanded; 
leaves elliptic to elliptic-oblong, with 4-7 pairs of major secondary veins, glabrous; 0-1700 
m elevation O. tenera 

37b. Flowers 3-5 mm long, outer anthers ca. 1 mm long and with the connective expanded distally; 
leaves ovate to broadly elliptic, with 3-5 pairs of major secondary veins, sparsely puberulent; 

900-2000 m O. brenesii 

38a. Minor venation raised on the lower leaf surface and forming a reticulum of small areolae 0.3-1 

mm broad, the areolae often with subequal sides; the leaves often drying yellowish green or grayish 

green, glabrous or glabrescent on the lower surface 39a 

38b. Minor venation not raised and not forming a definite reticulum on the lower (abaxial) leaf surfaces, 

areolae very irregular if present; glabrous to puberulent beneath 43a 

39a. Leaves gradually narrowed to the base, usually narrowly obovate and drying grayish green, 
domatia lacking; fruits globose with short persisting style at the apex and subtended by a 
small (6-10 mm) flat disclike receptacle; 0-1900 m elevation O. floribunda 

39b. Leaves abruptly narrowed at the base (obtuse to acute), pit domatia present; fruiting recep- 
tacles cupulate, 8-14 mm broad and 2-5 mm deep; fruits mostly ellipsoid, 2-3 cm long and 
the style not persisting 40a 

40a. Leaves broadly obovate and large (14-30 cm x 7-18 cm), rounded to bluntly obtuse at the 
apex, drying yellowish brown and subcoriaceous; fruit cup with an irregular margin; flowers 
unknown; 500-1000 m elevation O. sp. A aff. O. laetevirens 

40b. Leaves never broadly obovate and not more than 10 cm broad, usually acuminate at the 
apex; flowers 2-4 mm long, anthers 0.7-1 mm long; fruit cups with an entire margin . 41a 

4 la. Leaves usually drying yellowish green and often subcoriaceous, often broadly elliptic, retic- 
ulum of minor venation poorly defined; fruit cup 10-16 mm broad, 3-5 mm deep 

O. laetevirens 

41b. Leaves usually drying stiffly chartaceous; fruit cups only 2-3 mm deep 42a 

42a. Leaves usually drying grayish green and with a reticulum of small (0.5 mm) areolae beneath; 
fruit cup 8-14 mm broad with flaring edges; inflorescences usually paniculate; 0-2100 m 
elevation O. meziana 

42b. Leaves usually drying yellowish brown and lustrous above, reticulation of veins not forming 
small areolae beneath; fruit cup ca. 10 mm broad and with persisting lobes; inflorescences 
often umbellate or racemose; Pacific Highlands, 1 600-2000 m O. viridiflora 

BURGER: FLORA COSTARICENSIS 71 



43a. Flowers to 7 mm long and 5-10 mm broad, outer anthers 1.5-2.5 mm long and usually flattened 
44a 

43b. Flowers to 5 mm long, outer anthers less than 1 mm long; evergreen forests, 0-1800 m elevation 

45a 

44a. Flowers minutely puberulent; leaves drying grayish, with 5-8 pairs of major secondary veins; 

deciduous and partly deciduous forest formations, 0-1600 m O. veraguensis 

44b. Flowers glabrous; leaves drying dark brown with 3-6 pairs of major secondary veins; ever- 
green forests, 1 600-2200 m O. holdridgeiana 

45a. Flowers 2-2.5 mm long, glabrous and drying black; leaves usually oblong and caudate-acuminate 
at the apex, drying grayish and chartaceous, glabrous or very sparsely puberulent; fruit cups ca. 1 
cm broad; 0-300(-800) m elevation O. cernua 

45b. Flowers 2-5 mm long and minutely puberulent; leaves never caudate-acuminate, often drying 
grayish green and subcoriaceous; fruit cups 5-8 mm broad, dark and with pale lenticels; 0-1800 
m 46a 

46a. Leaves usually glabrous beneath, the major veins not impressed above, often elliptic-oblong; distal 
branchlets strongly 2- or 3-angled in cross section; flowers unisexual O. leucoxylon 

46b. Leaves usually puberulent beneath, the major veins impressed above and the leaves somewhat 
bullate, usually broadly elliptic to slightly obovate; distal branchlets with longitudinal ridges but 
not strongly angled in cross section; flowers not known in Costa Rica O. babosa 



Ocotea atirrensis Mez & J. D. Smith, Bot. Jahrb. 
Syst. 30, Beibl. 67: 1 8. 1 90 1 . O. pedalifolia Mez, 
loc. cit. 19. 1901. Figure 2. 

Shrubs or small and slender trees to 5(-8) m tall, leafy 
branchlets 3-6 mm thick, glabrous and smooth, strongly 
ridged or terete, distal stems hollow and often inhabited 
by small ants. Leaves alternate, petioles 6-18 mm long, 
1.5-3.5 mm thick, with adaxial margins forming a deep 
sulcus above and partly overlapping; leaf blades 12- 
35(-55) cm long, 5-1 3(-22) cm broad, narrowly obovate 
to oblong-obovate or oblanceolate, usually abruptly nar- 
rowed and caudate-acuminate at the apex, the tip often 
1-3 cm long and 2-4 mm broad, gradually narrowed to 
the acute or obtuse base, margin entire and slightly un- 
dulate, drying chartaceous and dark gray, glabrous above 
and below, with 7-12 major secondary veins on each 
side, venation usually flat above and more prominent 
beneath, the tertiary veins often subparallel between the 
secondaries. Inflorescences axillary to distal leaves (rare- 
ly pseudoterminal), 10-18 cm long (to 30 cm in fruits), 
paniculate with distal branches much shorter than the 
basal branches, glabrous, peduncles 1-6 cm long, be- 
coming pendulous, pedicels 1-3 mm long. Flowers buds 
ca. 2 mm long and 2 mm broad, yellow, glabrous or with 
a few hairs on the outside, tepals ca. 1.5 mm long and 
1 .2 mm broad near the base, ovate; outer anthers ca. 0.6 
mm long, narrow with superposed and slightly overlap- 
ping thecae, filaments 0.1-0.5 mm long, staminodes ab- 
sent; pistil ca. 2 mm long, style ca. 1 mm long, stigma 
simple or slightly discoid. Fruits borne on a receptacle 
8-12 mm long, 6-10 mm broad and 1-3 mm deep, ur- 
ceolate (but solid) to oblong and warty, bright red; fruits 
2-3.5 cm long, 1-2 cm thick (dry), oblong-ellipsoid to 
narrowly ovoid, narrowed at the base, becoming black. 

Usually found as understory treelets in ever- 
green wet forest formations, from near sea level 



to 1100 m elevation. Most flowering collections 
have been made in January-April, with a few in 
August-October. Mature fruits have been collect- 
ed in January-April and July-December. The 
species is to be expected in southeastern Nicaragua 
and has been collected along the Caribbean side 
of Costa Rica, in the General Valley, Osa Penin- 
sula, and adjacent Panama. 

Ocotea atirrensis is distinguished by the large 
thin obovate leaves with caudate-acuminate tips, 
lack of puberulence, hollow distal stems, small 
stature in the understory of usually lowland wet 
forests, small flowers, and unusual fruiting recep- 
tacle (a deep cup is not developed). This species 
is closely related to O. paulii and O. wedeliana but 
is distinguished by the large thin obovate leaves 
with caudate tips, tertiary veins more often par- 
allel, and different habitats. This species is similar 
to O. nicaraguensis, but that species has thicker 
leaves that do not dry dark and are not caudate- 
acuminate at the apex and its stems are more 
prominently ridged. Some collections between 400 
and 1 000 m elevation on the Caribbean slope have 
smaller, narrower, stiffer leaves that do not dry so 
dark and resemble O. paulii. Whether this is just 
clinal variation or reflects intergradation and gene 
flow with O. paulii should be investigated. See the 
discussion under Aiouea vexatrix by van der Werff 
in Ann. Missouri Bot. Card. 75: 405. 1988. 

Ocotea aurantiodora (Ruiz & Pavon) Mez, Jahrb. 
Konigl. Bot. Gart. Berlin 5: 295. 1889. Laurus 



72 



FIELDIANA: BOTANY 



aurantiodora Ruiz & Pavon, Laurografia Pe- 
ruviana t. 15, 71802. Mespilodaphne aurantio- 
dora Meissn. in DC., Prodr. 15(1): 101. 1864. 

Small trees to 12 m tall and ca. 25 cm d.b.h., leafy 
branchlets 6-14 mm thick, puberulent with short (0.4 
mm) thin ascending brownish hairs, becoming glabres- 
cent and dark in color, distal stems strongly angled with 
prominent (ca. 1 mm) longitudinal ridges, usually with 
5 ridges (or wings) in cross section. Leaves alternate, 
petioles ca. 10 mm long and 3 mm thick, flat above or 
the 2 adaxial ridges forming a shallow sulcus; leaf blades 
13-35 cm long, 4-10 cm broad, very narrowly obovate 
to oblanceolate or oblong-elliptic, usually tapering grad- 
ually to a short acuminate apex, tapering gradually to 
the cuneate or decurrent base, margin slightly revolute, 
drying subcoriaceous, glabrous above with the slightly 
raised minor venation forming a reticulum of small ar- 
eolae (0.5-1 mm broad), lower surface with slender ap- 
pressed straight yellowish hairs 0.2-0.7 mm long and 
usually parallel with the secondary veins, with 9-14 ma- 
jor secondary veins on each side and arising at angles of 
45-50. Inflorescences axillary to distal leaves, 10-22 
cm long, becoming 30 cm long in fruits, peduncles 3-9 
cm long and 2-4 mm thick, sparsely puberulent and with 
longitudinal ridges, pedicels ca. 1 mm long. Flowers ap- 
parently unisexual, ca. 2 mm long and 3 mm broad, 
yellowish white and minutely puberulent, tepals 1.5-2 
mm long and 1.5 mm broad; outer stamens with short 
(0.4 mm) filaments and narrow anthers ca. 0.8 mm long 
with superposed thecae, inner stamens ca. 1.5 mm long, 
glands closely appressed and resembling an annular disc, 
staminodes absent; pistil/pistillode ca. 1 .8 mm long, stig- 
ma disclike. Fruits borne in a receptacle ca. 6 mm long 
and 8 mm broad, rounded and cupulate in form, the 
opening ca. 5 mm in diameter and 3-4 mm deep, margin 
rounded and entire; berry ca. 8 mm long and 5-6 mm 
thick, oblong and bluntly rounded at both ends. 



Trees of evergreen and partly deciduous forest 
formations, at elevations from 200 to 900 m. A 
single flowering collection has been made in July 
in Costa Rica, and fruiting material has been col- 
lected in January-April. This species is known from 
southern Nicaragua, and from the General Valley 
in southern Costa Rica. The species ranges from 
southern Nicaragua and Costa Rica to Peru and 
the Amazon Basin. 

Ocotea aurantiodora is recognized by its thick, 
strongly winged stems, long, stiff, narrowly oblong 
or obovate leaves with slender appressed hairs be- 
neath, small functionally unisexual flowers, and 
small oblong fruits half immersed in a thick round- 
ed cup. This species is similar to O. nicaraguensis, 
but lacks the hollow ant-inhabited stems, and the 
flowers and fruits are very different. This very dis- 
tinctive species has been interpreted by Rohwer 
(1986) to include O. longifolia H.B.K., O. gran- 
difolia (Nees) Mez, and O. opifera Martius, among 
others. However, in his monograph, Rohwer ( 1 986) 



considered the Ruiz and Pavon names nomina 
nuda because only their illustrations were pub- 
lished. He has since changed his mind and we agree 
that the Ruiz and Pavon names, based on very 
fine illustrations, should be considered as effec- 
tively published. The text for these illustrations 
was published in Anal. Inst. Bot. Cavanilles 13: 
21. 1954. 



Ocotea austinii C. K. Allen, J. Arnold Arbor. 26: 
350. 1945. O. irazuensis Lundell, Wrightia 5: 
339. 1977. Figures. 

Trees to 25 m tall and 65 cm d.b.h., leafy branchlets 
1.8-5 mm thick, very minutely puberulent with brown- 
ish hairs in early stages, becoming (sub)glabrous, dark 
grayish and longitudinally ridged. Leaves alternate and 
usually clustered at the ends of branchlets, petioles 5- 
20 mm long, 1.5-3 mm broad, flat or sulcate above, 
lateral margins continuous with the lamina margins; leaf 
blades 3-12(-16) cm long, 1.5-4(-6) cm broad, oblong 
to elliptic-oblong, abruptly narrowed at the short-acute 
or obtuse apex, obtuse at the base with the margin re- 
volute and decurrent on the petiole, drying subcoria- 
ceous, upper surfaces usually lustrous and often with the 
tertiary venation forming a raised reticulum, densely 
subsericeous beneath with slender straight appressed yel- 
lowish or brownish hairs 0.2-0.5 mm long and oriented 
parallel with the major veins, becoming glabrescent, with 
(5-)6-9(-l 1) major veins on each side, the central sec- 
ondaries arising at angles of 40-60. Inflorescences ax- 
illary to distal leaves, 6-15 cm long, paniculate, pedun- 
cles 2.5-7 cm long, 1-3 mm thick, minutely puberulent 
with slender appressed yellowish brown hairs, flowers in 
short-stalked clusters, pedicels 1-3 mm long. Flowers 4- 
5 mm broad, densely and minutely puberulent on the 
outside, perianth-lobes 2.5-3 mm long, 2 mm broad; 
outer stamens with anthers ca. 1 mm long and 1 mm 
broad, the thecae superposed or the lower slightly lateral 
to the upper, filaments ca. 0.5 mm long, inner anthers 
narrow, staminodes absent; pistil ca. 2.5 mm long, gla- 
brous, style to 1 mm long and slender, stigma discoid 
or simple. Fruits borne in an obconic cup ca. 12 mm 
long, 10-15 mm broad and 1-5 mm deep; berry 2-3.5 
cm long, 1-2.5 cm in diameter, ellipsoid to ovoid, be- 
coming purple. 

Trees of higher montane wet evergreen forest 
formations, from 1 700 to 3000 m elevation, rang- 
ing from the northern edge of the Meseta Central 
(near Palmira, Alajuela) through the central vol- 
canic chain to the western part of the Cordillera 
de Talamanca (near Empalme and Sta. Maria de 
Dota, San Jose). Both flowers and fruits have been 
collected in May-June, August, and December- 
March. This species is endemic to central Costa 
Rica but has closely related taxa in the Cordillera 
de Tilaran and the Chiriqui highlands of Panama. 

Ocotea austinii is characterized by its high-al- 



BURGER: FLORA COSTARICENSIS 



73 



titude habitat, small stiff oblong leaves with lus- 
trous upper surfaces and reticulate tertiary vena- 
tion, the decurrent and revolute lamina base, silky 
yellowish or orange brown pubescence on younger 
parts and young leaves beneath, and the small pu- 
berulent flowers. This species is very closely re- 
lated to O. whitei which usually has oblanceolate 
leaves, and varies from densely sericeous to almost 
glabrous (see the discussion under O. whitei). Oco- 
tea austinii appears to be part of a group of species 
including O. sp. aff. caracasana, O. glaucosericea, 
O. whitei, and O. skutchii. A Costa Rican name is 
ira rosa. 



Ocotea babosa C. K. Allen, Mem. New York Bot. 
Gard. 15: 82. 1966, sensu lato. Figure 8. 

Trees 15-35 m tall, to 1 m d.b.h., bark gray or mar- 
bled, leafy branchlets 2-6 mm thick, puberulent with 
crooked or straight brownish tomentulose hairs 0.2-0.5 
mm long, soon becoming (sub)glabrous and grayish or 
black. Leaves alternate, petioles 8-22(-30) mm long, 1.5- 
2.3 mm thick, minutely puberulent or glabrescent, sul- 
cate above; leaf blades 14-26 cm long, 8-12.5 cm broad, 
broadly elliptic to ovate-elliptic or slightly obovate, acute 
to short-acuminate at the apex, obtuse at the base, mar- 
gin often somewhat undulate, drying stiffly chartaceous 
to subcoriaceous, minutely puberulent on the veins above 
but glabrescent between the veins, lustrous and some- 
what bullate between major veins above, hirsutulous or 
tomentulose with spreading slender straight brownish 
hairs to 0.5 mm long beneath, with 4-7 major secondary 
veins on each side, central secondaries arising at angles 
of 35-55 and loop-connected near the margin distally, 
secondary veins slightly impressed above and very 
prominent beneath, tertiary venation prominent be- 
neath. Inflorescences axillary to distal leaves, 7-10 cm 
long and few-flowered, becoming 10-12 cm long in fruits. 
Flowers said to be 3.5-4.5 mm and campanulate, tepals 
ca. 2.5 mm long; outer stamens ca. 1 mm long and the 
outer anthers slightly longer than their filaments, stam- 
inodes absent; pistil ca. 2.5 mm long, ovary ellipsoid to 
obovoid, stigma papillate. Fruits borne on a campanu- 
late-cylindrical pedicel gradually or abruptly expanded 
to the flat or saucer-like receptacle, 4-10 mm long and 
ca. 6 mm broad, only 1-2 mm deep, margin often 6- 
lobed with persisting perianth bases or entire, pedicel 
often with lenticels and verrucous; berry globose 7-10 
mm in diameter (dried), dark green at maturity, drying 
black and lustrous. 

Trees of lowland evergreen forest formations in 
Costa Rica. The species appears to flower in De- 
cember-February in Costa Rica; fruits have been 
collected in April and June. This species is known 
from the La Selva area (Heredia) at about 100 m 
elevation and Pejibaye (Cartago) at 500 m, both 
on the Caribbean slope of Costa Rica. The species 



(in a wide sense) ranges from Costa Rica to Ven- 
ezuela. 

Ocotea babosa is recognized by the distinctive 
leaves with prominent venation and distinctive 
pubescence beneath. The major veins are im- 
pressed above and often give a broadly bullate 
appearance to the upper surface. The small inflo- 
rescences and small globose fruits on slightly ex- 
panded receptacles, often with persisting perianth 
bases, also distinguish this species. Herbarium 
material resembles some species ofPersea. In Cos- 
ta Rica, this species is only known from the fol- 
lowing fruiting collections: Hammel & Trainer 
10871, Hammel 11530, and McDowell 854 (all 
DUKE), from La Selva; and Stork 2807 (F), from 
Pejibaye (Pejivalle). While it is not unusual for a 
species to have a lower altitudinal range in isth- 
mian Central America than in continental South 
America, the difference in altitudinal range of the 
Costa Rican collections and those from northern 
South America (at 1 800-2200 m) is greater than 
normally seen, and there are differences in fruit 
dimensions and upper leaf surfaces. Thus, C. K. 
Allen's name O. babosa (1966) may prove to be 
incorrect for our material. We are following Ham- 
mel's usage (1986). 



Ocotea sp. aff. O. bijuga (Rottb.) Bernard!, Can- 
dollea 22: 59. 1967. Nectandra bijuga Rottboel, 
PI. Surinam. Rar. 12. 1776. Figure 11. 

Trees to 30 m tall and 70 cm d.b.h., trunks whitish 
to pale gray, leafy branchlets, 1.3-5 mm thick, minutely 
(0. 1 mm) and sparsely appressed puberulent or glabrous, 
dark brown and longitudinally ridged, becoming terete 
and grayish. Leaves alternate, petioles 5-15 mm long, 
1-2 mm broad, sparsely and minutely puberulent, flat 
or sulcate above; leaf blades 6-15 cm long, 1.5-5 cm 
broad, narrowly elliptic-obovate to oblanceolate or nar- 
rowly oblong, tapering to an obtuse or bluntly short- 
acuminate apex, tapering gradually to the acute or at- 
tenuate base and decurrent on the petiole, margin entire 
and often slightly revolute (especially near the base), 
drying stiffly chartaceous to subcoriaceous, glabrous and 
dull or slightly lustrous above when dry with the minor 
venation obscure, glabrous or very sparsely appressed- 
puberulent beneath, with 5-9 major secondary veins on 
each side, central secondaries arising at angles of 35- 
50. Inflorescences axillary to distal leaves or extra-ax- 
illary, 6-1 5 cm long, paniculate with short (2 cm) lateral 
branches, peduncle 2-5 cm long, ca. 1.3 mm thick, gla- 
brous or minutely puberulent, drying dark, pedicels 0- 
2 mm long. Flowers 2-3 mm long, ca. 4 mm broad, very 
minutely appressed puberulent on the outside, tepals 1.2- 
1.7 mm long and ca. 1.2 mm broad, pellucid punctate; 
outer stamens with short (0.5 mm) puberulent filaments, 
outer anthers ca. 0.6 mm long and equally broad, thecae 
superposed, inner stamens 1.4 mm long with large (0.5 



74 



F1ELDIANA: BOTANY 



mm) glands, staminodes absent; pistil ca. 2 mm long, 
style narrow and equalling the length of the ovary, stigma 
simple or discoid. Fruit cup poorly developed, funnel- 
form in life (fide Hammel) but drying flat or saucer- 
shaped and about 5 mm broad, abruptly expanded above 
the slightly thickened (1.5 mm) pedicel, red; berry glo- 
bose, 5-8 mm in diameter when dry (to 1 2 mm in life), 
green to greenish purple. 

Trees of ridge tops in primary evergreen lowland 
Caribbean rain forest in Costa Rica. Flowering is 
in October, and fruiting material has been col- 
lected in February and April. Costa Rican material 
is only known from La Selva. The species with 
which our material is provisionally placed occurs 
in Colombia, Venezuela, and Brazil (see below). 

Ocotea sp. aff. O. bijuga is recognized by its 
glabrous narrowly elliptic to oblanceolate leaves 
with decurrent leaf base and the small globose 
fruits borne on a small flat or slightly concave 
receptacle. The leaves tend to dry a dark olive 
green above and brownish beneath. The associa- 
tion of this material with Bernardi's concept of O. 
bijuga, including O. florulenta (Meissner) Mez, 
should be considered tentative. Rohwer (pers. 
comm.) believes that O. bijuga is part of the Oco- 
tea cernua complex. Also, Ocotea bijuga is said to 
be dioecious, while ours appear to have bisexual 
flowers; this also suggests that O. bijuga may not 
be closely related to the Costa Rican material placed 
here. The form and texture of the leaves are very 
similar to Costa Rican material of O. oblonga and 
the westernmost populations of 0. whitei(ai Mon- 
teverde), but those species have very different fruits, 
and prefer higher-elevation habitats. At present 
this taxon is known from only five collections in 
Costa Rica: Folsom 9112 (DUKE); Hammel 10229 
(DUKE), 11663 (F, DUKE); Hartshorn 1585 (F, CR); 
and Ocampo 3646 (CR). 



Ocotea brenesii Standl., Field Mus. Nat. Hist., Bot. 
Sen 18: 454. 1937. Nectandra brenesii (Standl.) 
C. K. Allen, J. Arnold Arbor. 26: 370. 1945. 
Figure 13. 

Small to large trees, 6-28 m tall, trunks to 2 m broad 
at the base, leafy branchlets 1-3.5 mm thick, appressed 
puberulent at first with minute (0.2 mm) straight yel- 
lowish hairs, quickly becoming (sub)glabrous and dark 
in color, terete. Leaves alternate, petioles 6-1 3 mm long, 
ca. 1 mm thick, minutely appressed puberulent or gla- 
brescent, with a narrow adaxial sulcus; leaf blades 6- 
12(-15) cm long, 2.5-6(-8) cm broad, ovate to ovate- 
elliptic or broadly elliptic to ovate-oblong, short-acu- 
minate to long-acuminate at the apex, acute to obtuse 
or rounded at the base, drying thin chartaceous and very 



dark in color, often lustrous above, appressed puberulent 
on both surfaces but glabrescent with the minute (0.1- 
0.3 mm) thin straight hairs persisting on the larger veins, 
with 3-5 major secondaries on each side, central sec- 
ondaries arising at angles of 35-60. Inflorescences ax- 
illary to distal leaves, to 10 cm long, flowers few in an 
open racemose panicle, peduncles to 5 cm long and 0.5 
mm thick (dry), puberulent with straight appressed as- 
cending hairs, pedicels ca. 3 mm long. Flowers 3-5 mm 
long, 5-10 mm broad, yellowish or white, perianth gla- 
brous or papillate-puberulent on the outer surfaces, te- 
pals ca. 2 mm broad, glabrous within; outer stamens 
subsessile or on short (0.3-0.4 mm) filaments, outer an- 
thers ca. 1 mm long, ovate with superposed thecae, the 
connective slightly developed beyond the thecae, inner 
stamens with narrowly oblong anthers, staminodes ab- 
sent, the floral tube with straight lustrous hairs; pistil 
1.5-1.7 mm long, glabrous, ovary ovoid or angular, style 
ca. 0.8 mm long, stigma slightly discoid. Fruits borne 
on fruiting receptacles gradually expanded from the 
thickened pedicel and conic, ca. 10 mm long and 8 mm 
broad; berry poorly known, apparently to 24 mm long 
and 1 6 mm in diameter, ovoid-oblong (based on A. Smith 
517, 4102, F). 



Trees of evergreen montane forest formations, 
between 900 and 2000 m elevation in central Cos- 
ta Rica. Flowering collections have been made in 
February-May; mature fruits were collected in 
April. This species is endemic and known only 
from the northern region of the central highlands, 
from near the Reserva Forestal de San Ramon to 
the area between Poas and Barva (Barba) volcan- 
oes in Alajuela Province. 

Ocotea brenesii is distinguished by its relatively 
broad leaves with few secondary veins that dry 
thin in texture and very dark in color, rather large 
flowers on open racemose panicles, and its re- 
stricted habitat. It resembles O. tenera but that 
species is glabrous in all its parts, has smaller flow- 
ers, and narrower leaves. Ocotea brenesii resem- 
bles the type of Ocotea pittieri (Tonduz 1 1893, us); 
see the discussion under that species. Ocotea hold- 
ridgeiana with larger flowers and stiffer leaves may 
also be related to O. brenesii. 

Ocotea brenesii has been placed into synonymy 
under O. heydeana (Mez & J. D. Smith) Bernardi, 
by Bernardi (1967, p. 93) and by Rohwer (1986, 
p. 63). Collections of 0. heydeana differ in having 
stiffly chartaceous leaves that usually turn olive 
green when dried, and possess four to six pairs of 
major secondary veins; also flowers and floral parts 
are larger than in O. brenesii. Nevertheless, these 
species are closely related. Ocotea heydeana (often 
referred to as Nectandra heydeana) ranges from 
Honduras to Chiapas at elevations of 700 to 
1600m. 



BURGER: FLORA COSTARICENSIS 



75 



Ocotea calophylla Mez, Jahrb. Konigl. Bot. Gart. 
Berlin 5: 298. 1889. Pleurothyrium velutinum 
Meissn. in DC., Prodr. 15, pt. 1: 170. 1864, not 
O. velutina Martius. O. fulvescens Slandl. & L. 
O. Williams, Ceiba 1: 237. 1951. Figure 5. 

Trees to 20 m tall, leafy branchlets 5-12 mm thick, 
densely sericeous with reddish brown hairs in early stages 
but the hairs becoming dark gray and matted. Leaves 
alternate and subsessile, petioles short (0-10 mm) and 
poorly differentiated from the cuneate lamina base; leaf 
blades 10-19 cm long, 4-6 cm broad, narrowly elliptic 
to elliptic-oblong, obtuse at the apex, attenuate at the 
base with the margin revolute but not decurrent on the 
stem, drying subcoriaceous, densely and minutely ap- 
pressed puberulent above but becoming dull grayish and 
the pubescence obscure, very densely and conspicuously 
sericeous beneath with lustrous reddish brown or silvery 
brown appressed ascending hairs, with 10-13 major sec- 
ondary veins on each side, the central secondaries arising 
at angles of 40-70, tertiary venation obscure. Inflores- 
cences to 25 cm long, axillary to distal leaves, flowers 
crowded in a compact panicle with lateral branches ca. 
3 cm long, peduncles about half the length of the inflo- 
rescence (4-12 cm) and 4-5 mm thick, reddish brown 
puberulent. Flowers 4-7 mm long and 6-10 mm broad, 
densely puberulent or tomentulose on the outside, outer 
tepals ca. 4 mm long and 3 mm broad; outer anthers 
1 .6-1 .8 mm long on slender glabrous filaments of nearly 
equal length, thecae superposed but the upper introrse 
and the lower latrorse (in the outer series), staminodes 
present and small; pistil slender, to 4.5 mm long, style 
1-3 mm long. Fruit cups ca. 1 cm broad and 3 mm deep 
(immature?); berry ca. 20 mm long and 17 mm in di- 
ameter, globose-ellipsoid. 

Trees of wet evergreen montane forest forma- 
tions, between 2600 and 3000 m elevation. Flow- 
ers have been collected in January-February, and 
fruiting inflorescences have been collected in Sep- 
tember in Costa Rica. This species is also known 
from Colombia and Venezuela at elevations from 
2000 to 3 100m. 

Ocotea calophylla is one of Costa Rica's most 
distinctive species of Ocotea; the thick leaves with 
dense lustrous pubescence beneath are unique 
among Costa Rica's Lauraceae, and resemble some 
Sapotaceae. The subsessile leaves with revolute 
lamina base, thick stems, reddish brown inflores- 
cence, and high montane habitat further distin- 
guish this species. We have only seen the following 
collections: Lems 5352 (F, NY); Leon 2166 (CR, us, 
the type of O fulvescens); and Madriz 12 (CR, F). 
The local names are reported as ira zoncho and 
yema huevo. Many South American specimens of 
this species have dull reddish pubescence, but some 
do have the lustrous hairs that makes our Costa 
Rican material so striking. The leaves of this species 
are folded in an unusual way in bud, leaving a 



diagonal line across the upper left of the lamina 
and a short line on the right side near the base 
(viewing the lamina undersurface with the apex 
up and the base down). Ocotea guianensis Aublet 
of lowland formations in South America has sim- 
ilar lustrous pubescence, and the same pattern of 
folding in leaf vernation, and must be related. 
Closely related Andean species are O. micans Mez 
and O. sericea H.B.K. 



Ocotea sp. aff. caracasana (Nees) Mez, Jahrb. Kon- 
igl. Bot. Gart. Berlin 5: 292. 1889. Oreodaphne 
caracasana Nees, Linnaea 2 1 : 522. 1 849. Figure 
10. 

Trees to 30 m tall and 60 cm d.b.h., occasionally with 
prop roots from near the base, dioecious or apparently 
bisexual in ours, leafy branchlets 2.5-8 mm thick, ap- 
pressed puberulent with minute (0.1-0.4 mm) slender 
ascending pale yellowish to golden hairs, often with lon- 
gitudinal ridges but becoming terete and grayish. Leaves 
alternate, petioles 0-1 5 mm long but poorly defined, with 
lateral margins continuous with the decurrent lamina 
base, minutely appressed puberulent, narrowed portion 
of the lamina base 1-4 cm long; leaf blades 12-20(-24) 
cm long, 4.5-8(-ll) cm broad, elliptic-obovate to ob- 
ovate, oblong-obovate or oblanceolate, abruptly nar- 
rowed or rounded to a short-acuminate or obtuse apex, 
tapering gradually to the cuneate and decurrent base, 
margin usually revolute near the base, drying subcoria- 
ceous to coriaceous, glabrous above or minutely ap- 
pressed puberulent on the flat or impressed midvein near 
the base, usually dull above and slightly glaucous be- 
neath, minutely and often obscurely puberulent beneath 
with thin straight appressed yellowish hairs 0.1-0.3 mm 
long and usually parallel with the secondary veins or 
glabrescent, with (6-) 7-1 2 major secondary veins on each 
side, central secondaries arising at angles of 35-50, ter- 
tiary venation subparallcl and slightly raised beneath. 
Inflorescences axillary to distal leaves or distal leafless 
nodes, 7-12 cm long, panicles broadly paniculate, many- 
flowered, peduncles 2-8 cm long, 2-3 mm thick, densely 
puberulent and silvery to reddish brown in color, ped- 
icels 0-4 mm long. Flowers 3.5-5 mm long, 4-6 mm 
broad, campanulate, densely puberulent on the outside, 
tepals 1.5-2.8 mm long, 1.3-2.3 mm broad; outer sta- 
mens on short (0.4 mm) broad glabrous filaments, outer 
anthers 0.5-1 mm long, usually narrower than long, the- 
cae superposed but occasionally quite variable, inner 
stamens 1.3-2 mm long, often with large glands, stam- 
inodes to 1 . 1 mm long and variable or absent; pistil 1 .6- 
2.5 mm long, the slender style 1-1.5 mm long, stigma 
discoid or simple. Fruits borne in a large hemispherical 
cup 18-28 mm broad and 8-15 mm deep, margin often 
with teethlike projections from the persisting perianth; 
berry 34 cm long and 2-2.5 cm in diameter, ovoid. 



Trees of evergreen rain forest formations in the 
Caribbean lowlands of Costa Rica. Flowers and 
fruits were both collected in May. This taxon may 



76 



FIELDIANA: BOTANY 



be endemic to Costa Rica if it proves to be different 
from true O. caracasana, which occurs in Colom- 
bia, Venezuela, and the Guianas. 

Ocotea sp. aff. caracasana is recognized by the 
larger slightly obovate leaves, abruptly short-acu- 
minate at the apex and decurrent and revolute at 
the base. The large fruits and the stiff leaves with 
characteristic appressed hairs are also helpful in 
recognizing this species and its relatives. The pu- 
berulent flowers were described as unisexual but 
the Costa Rican material appears to be bisexual. 
Our material is derived from only a single tree at 
La Selva (Hammel p. 225, 1986: Hammel 12450, 
DUKE, F). The Hammel collection differs from the 
type photo (Karsten 32, F) in its lack of the elongate 
racemiform inflorescences, its secondary veins 
arising at a narrower angle, and its revolute lamina 
base being not as long. But there are also many 
similarities between the Costa Rican material and 
the type collection. This species, whatever its final 
name, is related to O. glaucosericea and O. skutchii 
of Costa Rica's highland forests; see the discus- 
sions under those species. While the foliage of these 
species resembles that of O. insularis and its allies, 
the fruits, parallel-appressed hairs, and prop roots 
indicate that they may not be closely related. 



Ocotea cernua (Nees) Mez, Jahrb. Konigl. Bot. 
Gart. Berlin 5: 377. 1889. Oreodaphne cernua 
Nees, Syst. laur. 424. 1836. Figure 14. 



ca. 2.5 mm long and urceolate, with a deep tube, tepals 
1-1.8 mm long and minutely papillate near the tips, 
nonfunctional anthers ca. 0.5 mm long, pistil ca. 1.5 mm 
long, stigma subsessile on the narrow apex of the ovary. 
Fruits borne in a cup 6-8 mm long and 9-12 mm broad, 
4-5 mm deep, margin entire, thickened part of the ped- 
icel 5-10 mm long, becoming red and enclosing the lower 
'/? of the fruits; berry 1.5-2 cm long, ovoid, becoming 
black (rare in herbaria). 

Small trees of lowland evergreen forest forma- 
tions on both the Caribbean and southern Pacific 
slopes, below 900 m elevation in Costa Rica. 
Flowering occurs in February-May, and fruits have 
been collected in August-November in Costa Rica. 
The species ranges from southern Mexico and the 
West Indies through Central America to South 
America. 

Ocotea cernua is distinguished by its elliptic- 
oblong leaves that nearly always dry grayish or 
yellowish and often have a caudate-acuminate 
apex, and the slender inflorescences that are ra- 
cemiform or open panicles with racemose lateral 
branches. The lack of puberulence on almost all 
parts, the ebracteate inflorescences, and the small 
unisexual flowers that usually dry black are ad- 
ditional features of this unusual species. Ocotea 
cernua, O. meziana, and O. tenera are very similar 
in the field but differ in their dried appearance (cf. 
Hammel, 1986). Licaria cufodontisii and L. sar- 
apiquensis are also similar to O. cernua, especially 
in the appearance of leafy twigs and inflorescences. 



Shrubs or small to medium-sized trees 5-10(-20) m 
tall, dioecious (unisexual), leafy branchlets 1.2-3 mm 
thick, apical buds slightly puberulent but stems essen- 
tially glabrous, terete or slightly ridged. Leaves alternate, 
approximate but not congested distally, petioles 6-20 
mm long, ca. 1 mm thick, glabrous and with a shallow 
adaxial sulcus; leaf blades 6-14(-18) cm long, 3-6(-8) 
cm broad, oblong to elliptic-oblong, caudate acuminate 
or short- to long-acuminate at the apex, often with a 
narrow (ca. 3 mm) tip to 2 cm long, acute to obtuse at 
the base, margin often undulate, drying chartaceous to 
stiffly chartaceous, grayish and glabrous above, glabrous 
or sparsely puberulent along the major veins beneath, 
the primary veins flat above, with 3-6 major secondary 
veins on each side, central secondaries arising at angles 
of 30-70. Inflorescences axillary or extra-axillary on 
distal twigs (sometimes pseudoterminal and stemlike), 
3.5-15 cm long, paniculate and usually with racemose 
lateral branches or small and raceme-like, glabrous or 
with very few appressed hairs, often drying very dark 
(both flowers and inflorescence), peduncle to 5 cm long, 
ca. 0.5 mm thick, pedicels 1-4 mm long, slender. Flowers 
unisexual, drying dark, glabrous on the exterior; male 
flowers ca. 2.2 mm long and 3-4 mm broad, campan- 
ulate, tepals 1.2-1.8 mm long, outer anthers 0.6-0.8 mm 
long, slightly narrower than long, subsessile, thecae su- 
perposed, a pistilode present or absent; female flowers 

BURGER: FLORA COSTARICENSIS 



Ocotea dendrodaphne Mez, Jahrb. Konigl. Bot. 
Gart. Berlin 5: 238. 1889. O. quisara Mez & J. 
D. Smith, Bot. Gaz. 33: 259. 1902. O. ovan- 
densis Lundell, Contr. Univ. Mich. Herb. 6: 16. 
1941. Figure 3. 

Shrubs or small treelets to 6 m tall, leafy branchlets 
3-7 mm thick, minutely (0.1-0.3 mm) appressed pu- 
berulent with straight ascending hairs in early stages, 
soon becoming glabrous and pale gray, lenticellate and 
ridged but becoming terete in age, distal stems hollow 
and often with small ants. Leaves alternate, petioles 6- 
36 mm long, 1.5-3 mm thick, sulcate above, drying dark 
in color; leaf blades 14-36 cm long, 5-14 cm broad, 
elliptic to narrowly elliptic-oblong, widest at or slightly 
below the middle, acute or short to long acuminate at 
the apex (occasionally rounded/obtuse), acute to obtuse 
at the base, drying very stiffly chartaceous to subcoria- 
ceous and grayish green, glabrous and slightly lustrous 
on both surfaces, with 5-10 major secondary veins on 
each side, the central secondaries arising at angles of 40- 
70 tertiary venation slightly raised and reticulate be- 
neath. Inflorescences to 1 5 cm long, borne close together 
at the branchlet tips in the axils of what appear to be 
leaf scars (perhaps early caducous leaf scales), paniculate 



77 



in form with basal branches larger than the distal, pri- 
mary peduncle to 5 cm long, very minutely puberulent 
in early stages. Flowers 4-6 mm long, to 8 mm broad, 
white and sweet-scented, tepals to 5 mm long, densely 
and minutely papillate-puberulent; outer anthers 2-3 mm 
long, broadly flattened and somewhat tepal-like, ovate- 
triangular to lanceolate in outline with the thecae often 
on the basal half of the inner face, minutely puberulent, 
filaments very short (0.5 mm) and stout, staminodes not 
seen; pistil 1.9-2.4 mm long, style 1.1-1.5 mm long, 
slender, stigma slightly discoid. Fruit cups 8-10 mm 
long, ca. 1 5 mm broad and 3-7 mm deep, enclosing the 
basal 'A of the fruits, conical to cupulate in form, often 
with a second ridge around the rim; berry 15-25 mm 
long, 10-12 mm thick, bluntly rounded at the apex, nar- 
rowly ovoid to oblong, black at maturity. 

Understory plants of wet evergreen forest for- 
mations, between sea level and 1 500 m elevation, 
along the Caribbean slopes and in the central high- 
lands. Flowering collections have been made in 
January-June, October, and December; fruiting 
occurs throughout the year, but most collections 
have been made in July-October in Costa Rica. 
The species ranges from central Mexico to central 
Panama. 

Ocotea dendrodaphne is recognized by its hol- 
low stems often inhabited by ants, the large gla- 
brous elliptic-oblong leaves that dry very stiff and 
gray-green, the large flowers with tepaloid sta- 
mens, and distinctive deep cups. This species is 
often found on slopes and ridges. Two unusual 
features of this species are the way in which the 
inflorescences originate from leafless nodes, and 
the frequently inflated distal branchlet-tips with 
longitudinal slitlike openings into the hollow stems. 
The latter are associated with ants. The large flow- 
ers with broadly flattened outer stamens and the 
fruit cups with ridged rims (resembling Licaria, 
but not always developed) indicate a relationship 
with O. veraguensis. However, specimens may be 
more easily confused with O. paulii and its allies, 
which also have hollow stems, similar leaves, and 
are treelets of wet forest understory. 



Ocotea dentata van der Werff, sp. nov. Figure 20 

Arbor ad 25 m. Ramuli juniores angulares, adpresse 
pubescentes. vetustiores teretes glabrescentesque. Folia 
obovata, 20-35 x 8- 12 cm, apice rotundata velbreviter 
acuminata, basi decurrente et reflexa, supra praeter cos- 
tarn nervosque leviter pubescentes laevia glabraque, sub- 
tus sparse pubescentia, pilis brevibus et plerumque er- 
ectis; costa, nervis elevata, venatione reticulata et parum 
elevata; nervis 9-12. Petioli sulcati, 0.5-1.5 cm longi. 
Inflorescentiae pubescentes, in axillis foliorum vel brac- 
tearum deciduarum, foliis breviores vel aequales, ram- 



ificatione divaricata. Flores fragrantes, hermaphroditi; 
tepala magnitudine aequalia, ca. 1.5 mm longa, ovata, 
omnina intus glabra, 3 exterioribus extus dense puberulis 
sed 3 interioribus minus puberulis; stamina 9, 4-locel- 
lata, 6 exterioribus ca. 1.1 mm longis, antheris glabris, 
ca. 0.6 mm longis, thecis introrsis, filament is ca. 0.5 mm 
longis, propre apicem pilis minutis muni t is: 3 interior- 
ibus ca. 1 .2 mm longis, antheris glabris, thecis inferior- 
ibus extrorsis, thecis superioribus lateralibus, filamentis 
ca. 0.5 mm longis, 2 glandulis munitis; pagina interiore 
filamentorum propre apicem pubescentia densa munita; 
staminodia nulla. Ovarium ovoideum, glabrum, in sicco 
0.5 mm longum, stylo ca. 1 mm longo, glabro. Fructus 
viridis, ellipsoideus, 10x6 mm; cupula tepalis persis- 
tentibus munita. 

Trees to 25 m tall, leafy branchlets 2.5-8 mm thick, 
angular when young but becoming terete, at first ap- 
pressed puberulent but glabrescent in age. Leaves alter- 
nate, petioles difficult to delimit because of the decurrent 
lamina-base, 5-15 mm long, 2.5-7 mm broad and sul- 
cate above; leaf blade (15-)20-35 cm long, (5-)8-12 cm 
broad, obovate to elliptic-obovate or elliptic-oblong, 
usually broadest at or above the middle, abruptly nar- 
rowed and often rounded at the obtuse to short-acu- 
minate apex, tapering to the cuneate and decurrent base, 
margin somewhat revolute near the base and decurrent 
on the petiole, drying stiffly chartaceous to subcoria- 
ceous, glabrous above or with some hairs above the ma- 
jor veins, lower surface usually sparsely puberulent with 
erect or spreading hairs 0.2-0.5 mm long, with 9-12 
major secondary veins on each side, central secondaries 
arising at angles of 30-50. Inflorescences axillary to 
distal leaves or deciduous bracts, solitary, 12-25 cm long, 
robust and many-flowered panicles, peduncle 3-12 cm 
long, 1.7-3.5 mm thick and with grayish or brownish 
puberulence, pedicels to 2 mm long, flowers in cymose 
groups. Flowers 3-5 mm long and 4-5 mm broad, yel- 
lowish green, tepals 1.5-2.5 mm long, subequal, outer 
tepals densely puberulent on the outer surfaces, glabrous 
within; outer stamens with short (0.2-0.4 mm) filaments, 
outer anthers 0.5-0.8 mm long and equally broad or 
narrower, thecae superposed or the lower slightly lateral, 
inner stamens 1 .2-1 .4 mm long, with a tuft of hairs below 
the anthers, staminodes absent; pistil 1.5-2.6 mm long, 
ovary ovoid, style 1-1.5 mm long, slender, stigma slightly 
discoid. Fruits borne on a short (8-10 mm) cupulate 
receptacle 5-12 mm broad, the margin usually with per- 
sisting perianth bases; berry ellipsoid or oblong, 10 mm 
long and 6 mm in diameter (in the type) but perhaps 
becoming 3 cm long and 2 cm in diameter. 



TYPE Costa Rica, Prov. Limon, Reserva Biol- 
ogica Hitoy Cerere, 100-125 m, 29 Aug. 1985, L. 
D. Gomez & G. Herrera 23653 (holotype, MO; is- 
otypes, CR, F). Additional Collections: Nicaragua, 
Dept. Matagalpa, Neil I 77/5, Stevens & Riviera 
20899, Dept. Zelaya, Araquistain 3165, Grijalva 
264, Stevens 8005, 12444 (all MO); Costa Rica, 
Prov. Alajuela, Haber 5557, 6178 (MO); Hartshorn 
1543 (CR, F); Panama Prov. Bocas del Toro, Lao 
94, von Wedel 720, 1382 (all MO). 



78 



FIELDIANA: BOTANY 



Trees of evergreen rain forest formations, from 
near sea level to 1000 m elevation. Flowers have 
been collected in August and October-December; 
young fruits have been collected in February, ma- 
ture fruits in March. This species ranges along the 
Caribbean slope and lowlands, from Nicaragua to 
Panama, and in the evergreen forests of the south- 
ern Pacific slope in Costa Rica. 

Ocotea dentata is recognized by the larger stiff 
puberulent obovate leaves with decurrent and re- 
volute leaf bases, and the fruit cups often with 
persisting perianth parts. This species is closely 
related to O. insularis (in a wide sense). The pu- 
bescence helps separate this species from O. in- 
sularis, O. ira, and O. rivularis. The dense pub- 
erulence of the flowers, and the cupules with 
persistent tepals (hence the dentate appearance and 
specific name) are also distinctive. Earlier, some of 
the material placed here was identified as O. sten- 
oneura. Unfortunately, Ocotea stenoneura is very 
poorly understood at present, but it has very dif- 
ferent fruits (q.v.). Additional material was used 
to amplify the English description beyond the 
measurements cited for the type in the Latin de- 
scription. 



Ocotea endresiana Mez, Jahrb. Konigl. Bot. Gart. 
Berlin 5: 257. 1889. Figure 10. 

Trees to 25 m tall, leafy branchlets 2-6 mm thick, 
glabrous or minutely appressed puberulent at first, with 
prominent ridges but soon becoming terete, dark brown 
to dark gray. Leaves alternate, petioles poorly denned or 
absent, narrowed portion of the lamina base 0-2(-3) cm 
long, flat above, with strongly revolute margins; leaf 
blades 6-17 cm long, 3-9 cm broad (25 x 18 cm in 
sprout shoots), obovate to elliptic-obovate, obtuse to 
abruptly rounded at the apex, tapering gradually to the 
decurrent base, revolute with the margin curled under 
near the base, drying subcoriaceous or coriaceous, gla- 
brous and often lustrous above, glabrous or minutely 
(0. 1-0.2 mm) puberulent beneath, rarely with longer tufts 
of hairs (domatia) in the vein axils and along the midvein 
beneath, with 5-9 major secondary veins on each side, 
central secondaries arising at angles of 40-60, tertiary 
venation slightly raised on both surfaces but usually ob- 
scure, often subparallel between the secondaries, major 
veins often reddish in early stages. Inflorescences axillary 
to distal leaves or apparently terminal, to 1 7 cm long, 
broadly paniculate but few-branched, peduncle to 6 cm 
long, 2-3 mm thick, glabrous and drying dark brown to 
black, pedicels ca. 2 mm long. Flowers 2.5-5 mm long, 
3-6 mm broad, yellowish, glabrous to densely and mi- 
nutely puberulent on the outer surfaces, minutely pap- 
illate-puberulent within, tepals ca. 2.5 mm long; outer 
stamens with short (0.5 mm) filaments, outer anthers 
0.5-0.8(-1.1) mm long and equally broad or narrower, 



staminodes absent (rarely present?); pistil 1 .5-2 mm long, 
style narrow and usually equalling the ovary in length, 
stigma simple or discoid. Fruits borne in cupulate re- 
ceptacles 8-1 5 mm long and 7-10 mm broad, rim entire 
and the surfaces smooth, becoming red; berry 1 5-1 8 mm 
long, 8-10 mm thick (dried), oblong-ellipsoid or elliptic- 
cylindrical. 

Trees of montane evergreen forest formations, 
between (700) 1 100 and 2300 m elevation, along 
the Caribbean slope and continental divide. Flow- 
ering material has been collected in August-Oc- 
tober and February; mature fruits have been col- 
lected in February-March and June. The species 
ranges from the Cordillera de Tilaran (Monte- 
verde area), along the northern and eastern edge 
of the central highlands, to the area of Cartago and 
Volcan Turrialba. This taxon of doubtful status is 
endemic to Costa Rica and adjacent Panama (see 
below). 

Ocotea endresiana is recognized by the stiff 
bluntly obovate leaves, with decurrent leaf bases 
which have broadly revolute margins on the un- 
derside of a poorly defined petiole/leaf base. The 
long narrow fruits in smooth entire-rimmed cups 
only 1 cm broad also distinguishes this species. It 
appears to be common only north of San Ramon 
and near Vara Blanca de Sarapiqui. The very re- 
stricted range is an argument for considering these 
plants no more than a subspecific element of O. 
insularis in a wide sense. However, the leaf bases 
are very distinctive, as are the fruiting stages. See 
the discussion under O. insularis. Ocotea endre- 
siana has sometimes been confused with O. skut- 
chii, which also has long-decurrent leaf bases but 
different flowers, and a characteristically fine in- 
dumentum on the undersides of the leaves. We 
have not seen the type and are following earlier 
usage in the application of this name. 



Ocotea floribunda (Sw.) Mez, Jahrb. Konigl. Bot. 
Gart. Berlin 5: 325. 1889. Laurus floribunda 
Sw., Prodr. 65. 1788. O. wachenheimh ' Benoist, 
Bull. Mus. Hist. Nat. Paris 30: 510. 1924. Fig- 
ures 14. 

Trees 1 5-30 m tall, trunk 30-80 cm in diameter, dioe- 
cious (unisexual), leafy branchlets 1.5-5 mm thick, very 
minutely (0. 1 mm) puberulent with inconspicuous hairs 
near the apex but quickly becoming glabrous and dark 
in color, smooth and terete. Leaves alternate or subop- 
posite, petioles 4-14 mm long, 0.8-1.9 mm thick, gla- 
brous and flat above, drying dark in color; leaf blades 
7-1 5(-19) cm long, 2-6(-8) cm broad, elliptic to oblong- 
obovate, narrowly obovate or obovate, usually broadest 



BURGER: FLORA COSTARICENSIS 



79 



above the middle, acute to short-acuminate (rarely 
rounded) at the apex, acute to cuneate at the base and 
slightly decurrent on the petiole, margin entire and slightly 
recurved when dry, drying stiffly chartaceous to subcor- 
iaceous and pale to dark grayish green, essentially gla- 
brous above and below, with (3-)5-8 major secondary 
veins on each side, with a few weakly denned loop-con- 
nections near the margin, central secondaries arising at 
angles of 40-60, the larger veins often drying reddish 
brown, the smallest veins usually forming a fine (0.1- 
0.6 mm) reticulum usually visible on both surfaces but 
more prominent beneath. Inflorescences axillary, extra- 
axillary or pseudoterminal, 3-14 cm long, racemose or 
paniculate, few to many-flowered, peduncle 1-4 cm long, 
glabrous or very minutely puberulent, drying dark, ped- 
icels 1-3 mm long, ca. 0.8 mm thick. Flowers function- 
ally unisexual, 5-7 mm broad, greenish to cream-white, 
outer tepals broadly imbricate, 2-3(-4) mm long and 2- 
3 mm broad, minutely puberulent on both surfaces, be- 
coming reflexed; male flowers with stamens on promi- 
nent (0.9 mm) filaments, anthers ca. 1.3 mm long and 
1.2 mm broad, the lower thecae slightly lateral to the 
upper, inner stamens ca. 2 mm long, staminodia absent, 
pistillode ca. 3 mm long; female flowers with the outer 
stamens on short (0.2 mm) filaments and with smaller 
(0.7 mm) anthers, pistil ca. 3 mm long, ovary ovoid and 
equalling the style in length, stigma discoid. Fruits borne 
a flat disci ike receptacle 6-10 mm in diameter and 1.5- 
2 mm thick, often slightly recurved, margin rounded in 
life (fide Hammel) but with upper and lower ridges when 
dry, sometimes with the bases of persisting floral parts, 
green; berry subglobose, 13-18 mm long and 10-15 mm 
in diameter, green and glaucous at maturity, usually with 
short ( 1 mm) persisting style base at the apex of the fruits. 



Trees of wet evergreen forest formations, from 
near sea level to 1 500 m elevation. In Costa Rica, 
flowers have been collected in October and De- 
cember-February. Young fruits have been col- 
lected in June, and mature fruits in August-No- 
vember and March (near Panama). The species 
has been collected in the Caribbean lowlands (La 
Selva), near Turrialba, at Monteverde, and on the 
Pacific slope near Panama at 1 300 m elevation. It 
has not been collected on the Pacific slope below 
1200 m in Costa Rica. The species ranges from 
Nicaragua southward to Peru, Venezuela, the 
Guianas, and the West Indies. 

Ocotea floribunda is recognized by its very dark 
(almost black) stems, stiff leaves drying grayish 
green and with a finely reticulated surface, uni- 
sexual flowers, globose fruits with persisting style 
base, and the flattened fruiting receptacle with 
ridged edge. Herbarium material of this species 
resembles O. meziana, O. veraguensis, and some 
species of Licaria. This species appears to be un- 
common and has been collected in Costa Rica only 
recently. Costa Rican material differs from the West 
Indian collections in a number of ways and may 
prove to be worthy of subspecific recognition. There 



is an unusual population in the southern part of 
the Cordillera de Talamanca, where the base of 
the lamina is both decurrent and revolute (Davidse 
24359, CR, MO; Hartshorn 2165, Lent 2705 CR, F); 
flowers of the Davidse collection make the place- 
ment of these collections under O. floribunda quite 
certain. 



Ocotea glaucosericea Rohwer, Mitt. Inst. Allg. Hot. 
Hamburg 20: 144. 1986, nomen novum forNec- 
tandra hypoglauca Standl. ex C. K. Allen, J. 
Arnold Arbor. 26: 399. 1945, not O. hypoglauca 
(Nees) Mez. Figure 12. 

Trees to over 30 m tall, leafy branchlets 3-8 mm thick, 
densely and minutely (0.2 mm) puberulent with ascend- 
ing or appressed hairs, yellowish brown to grayish brown, 
longitudinally ridged. Leaves alternate in a spiral, peti- 
oles poorly defined and continuous with the decurrent 
lamina base, 1-3 cm long, with broad revolute lateral 
margins; leaf blades 9-17 cm long, 4-10 cm broad, ob- 
ovate to narrowly obovate, elliptic-obovate or oblong- 
obovate, abruptly rounded at the apex or obtuse (bluntly 
acute in narrow leaves), tapering gradually to the cuneate 
and decurrent base, margin entire and usually revolute 
at the base, drying subcoriaceous and lustrous above with 
the minor venation slightly raised, glabrous except on 
the midvein basally, paler in color beneath and with 
minute (0.1-0.2 mm) slender appressed hairs usually 
lying parallel with the secondary veins, with (5-)6-9 ma- 
jor secondary veins on each side, the central secondaries 
arising at angles of 30-50. Inflorescences axillary to 
distal leaves, 8-18 cm long, paniculate, peduncles 3-9 
cm long, densely appressed puberulent and yellowish, 
pedicels 2-4 mm long. Flowers ca. 4 mm long, 4-6 mm 
broad, densely grayish or yellowish brown puberulent 
on the outside, tepals ca. 2.8 mm long and 2.2 mm broad; 
outer stamens 1-1.5 mm long with short (0.2-0.6 mm) 
broad filaments, outer anthers 0.9-1 mm long and 0.7- 
0.8 mm broad, the thecae superposed or the lower some- 
what lateral to the upper, staminodes absent or small (to 
0.6 mm); pistil ca. 3 mm long, ovary ellipsoid, style ca. 
1 mm long, stigma capitate. Fruits borne in a conical 
receptacle about 2 cm long, 1.2-1.8 cm broad and ca. 5 
mm deep; berry 2.5-3 cm long, 2-2.5 cm in diameter, 
globose or oblong. 

Trees of evergreen montane forest formations, 
from 1500 to 2500 m elevation. Flowering ma- 
terial has been collected in April-May and July; 
fruits have been collected in March-June. The 
species is endemic to the Cordillera de Talamanca 
in Costa Rica (El Cedral de Sta. Maria de Dota, 
La Cangreja, and Las Tablas near the Rio Coton- 
cito), and appears to be common in the Chiriqui 
highlands of Panama. 

Ocotea glaucosericea is recognized by its stiff 
obovate leaves, lustrous on the upper surface and 
decurrent and revolute at the base. The highland 



80 



FIELDIANA: BOTANY 



habitat, upper leaf surface, and smaller fruits on 
shallower receptacles distinguish this species from 
O. sp. aff. caracasana, with which this species has 
been considered synonymous by Hammel (1986). 
This species is also closely related to O. skutchii, 
and there are collections that appear to be inter- 
mediate between them. Ocotea austinii and O. 
whitei are also part of this complex. All these taxa 
appear to be part of a closely related species group, 
united by characters of the flowers, large fruits, 
leaf form, and the slender appressed hairs on the 
lower leaf surfaces. These species resemble O. in- 
sularis and its allies, but those differ in pubescence 
and fruits, and are not as closely related. 



Ocotea gomezii W. Burger, sp. nov. Figure 6. 

Arbor 10 m alta, ramulis foliiferis 3-6 mm crassis, 
ferrugineo-tomentosis. Folia alterna, petiolis 1 2-26 mm 
longis, tomentosis; laminis 10-18(-24) cm longis, 6- 
11 (-14. 5) cm latis, late ellipticis vel suborbicularibus, 
apice obtuso vel breviter acuminato, subtus ferrugineo 
puberulis, nervis secondariis 5-8(-10) paribus. Inflores- 
centiae 10-18 cm longae, paniculatae, floribus paucis, 
pedunculis 6-13 cm longis, tomentosis. Flores ca. 10 
mm longi et 1 2 mm lati, tepalis ca. 6 mm longis, pub- 
erulis; stamina ser. I-II ca. 3 mm longa, antheris ca. 1 .6 
mm longis; gynoecium 2.5-3.7 mm longum. Fructus ig- 
notus; cupula 1 5-20 mm longa et 1 5-20 mm lata, mar- 
gine lobata. 

Trees 6-10 m tall, leafy branchlets 3-6 mm thick and 
densely ferrugineous-tomentulose with soft curved and 
straight hairs ca. 0.5 mm long, remaining tomentulose 
but becoming grayish. Leaves alternate and not crowded 
distally, petioles 12-26 mm long, 2.5-3.5 mm thick, 
densely tomentulose; leaf blades 10-18(-24) cm long, 6- 
11 (-14. 5) cm broad, broadly elliptic to suborbicular, 
usually broadest near the middle and rarely somewhat 
ovate or obovate, bluntly obtuse to short-acuminate with 
a tip to 1 5 mm long, obtuse to rounded and subtruncate 
at the base, drying stiffly chartaceous and dark brown to 
olive green, slightly lustrous above and minutely puber- 
ulent above the major veins, densely tomentulose be- 
neath with ferruginous or yellowish brown hairs ca. 0.7 
mm long, with 5-8(-10) major secondary veins on each 
side, central secondaries arising at angles of 45-60, ma- 
jor veins very prominent beneath and forming an arcuate 
submarginal vein distally, tertiary venation also slightly 
raised. Inflorescences few-branched panicles in axils of 
distal leaves or pseudoterminal, 10-18 cm long, pedun- 
cles 6-13 cm long, 1.7-3 mm thick, densely ferruginous 
puberulent, flowers borne in groups of 1-3 on short (1 
cm) lateral branches subtended by a bract 5-6 mm long. 
Flowers ca. 10mm long and 12mm broad, campanulate, 
yellowish white, densely puberulent on the outside and 
papillate-puberulent within, tepals to 6 mm long and 3 
mm wide with the inner series smaller than the outer; 
outer stamens to 3 mm long, outer anthers ca. 1 .6 mm 
long and 1 mm broad, oblong, valves superposed, fila- 
ments with long trichomes, inner stamens with large 



glands abaxially, staminodes absent; pistil 2.5-3.7 mm 
long, ovary narrowed to the base, style ca. 2 mm long, 
slender, stigma slightly lobed. Fruits borne in a conical 
cup 1 5-20 mm long and 1 5-20 mm broad, the cup sur- 
face densely hirsutulous, the perianth parts persisting 
and forming large (3-5 mm) lobes on the margin of the 
cup; young berry with hairs over the basal surface, ma- 
ture berry not seen. 

TYPE Costa Rica, Alajuela Province, valley of 
the Rio Lorencito in the Reserva Forestal de San 
Ramon, 14 March 1987. Gomez- Laurito, Burger, 
Mora, Ortiz & Antonio 11450 (holotype, CR; iso- 

typCS, CAS, F, DUKE, HBG, MO, NY, USj). 

Trees of very moist cloud forest and premontane 
rain forest formations of the Caribbean slope and 
continental divide, from 800 to 1400 m elevation. 
Flowers have been collected in January-April and 
July. Fruits have been reported in August and Jan- 
uary. The species is endemic to Costa Rica, and 
ranges from near Volcan Rincon de la Vieja in the 
west to Moravia de Chirripo in the east, and prob- 
ably further eastward along the Caribbean slope 
of the Cordillera de Talamanca. 

Ocotea gomezii is a very unusual species, dis- 
tinguished by its ferruginous puberulence, broadly 
rounded leaves, large puberulent flowers, and 
fruiting receptacles with large perianth lobes. The 
puberulence and lobed fruiting receptacles resem- 
ble O. mollifolia, but that species has very different 
inflorescences, and thinner, more obovate leaves. 
Ocotea pseudopalmana is similar but has smaller 
flowers, entire fruit cups, and grows at higher el- 
evations. The large glands of the new species are 
reminiscent of Pleurothyriumpalmanum, and there 
are other similarities that should be looked at fur- 
ther. The first collection of this species was by 
Brenes (328(685)/6092, F, NY); other collections 
are Bermudez 268 (usj), Dryer 1612 (CR, F), Gom- 
ez-L. 4590 (CR, usj), Hammel & Trainer 15044 
(MO), Holdridge 6827, 6829 (CR, NY), and Poveda 
et al. 947 (CR, F), 75/9 (usj). The best flowering 
collection of this species was made by Jorge Gom- 
ez-Laurito, whose collections are adding signifi- 
cantly to our knowledge of Costa Rica's rich flora. 



Ocotea hartshorniana Hammel, J. Arnold Arbor. 
67: 128. 1986. Figure 8. 

Trees 10-30 m tall, to 1 m diameter d.b.h., with but- 
tresses to 2 m high and 1 m broad or with prop roots, 
bark pinkish tan to orange, leafy branchlets 2.5-6 cm 
thick, at first densely reddish brown or yellowish brown 
puberulent with hairs 0. 1-0.5 mm long, often ascending 



BURGER: FLORA COSTARICENSIS 



81 



and slightly strigulose, longitudinal ridges usually pres- 
ent. Leaves alternate, petioles 4-1 8(-24) mm long, densely 
puberulent or the hairs minute and apparently glabrous, 
flat above or slightly sulcate; leaf blades 1 2-1 7(-20) cm 
long, 3-6(-8) cm broad, oblong-obovate to obovate or 
elliptic-oblong, usually broadest above the middle, 
abruptly narrowed to a short acuminate or caudate-acu- 
minate apex, gradually narrowed to a decurrent base, 
margins entire and usually recurved at the base, drying 
stiffly chartaceous, puberulent on the mid vein above or 
glabrescent, sparsely to densely puberulent on the veins 
beneath, hairs 0.2-0.5 mm long and often reddish brown 
in color, sometimes glaucous beneath, with 4-8 major 
secondaries on each side, central secondaries arising at 
angles of 25-55, tertiary veins usually subparallel and 
at right angles to the secondaries. Inflorescences 5-15 
cm long, axillary to distal leaves or occasionally pseu- 
doterminal, broadly (8 cm) paniculate, many flowered 
and yellowish brown to reddish brown puberulent, pe- 
duncle 1-5 cm long, distal branches of the panicle much 
shorter than the basal. Flowers 3-4 mm long, ca. 4 mm 
broad, densely short-puberulent, tepals 1.5-2 mm long 
and erect at anthesis; outer stamens ca. 1 .2 mm long with 
anthers 0.6-0.8 mm long and equally broad or slightly 
narrower, thecae superposed, staminodes absent; pistil 
ca. 2.5 mm long, ovary 1.5 mm long and narrowed at 
the base, stigma discoid. Fruits enclosed within a cup 
10-15 mm long and 10-16 mm broad, ca. 6 mm deep 
and enclosing '/z-Vj of the fruits, margin entire or with 
broad blunt lobes, cuplike and rounded at the base, 
brownish, pedicels 1.5-4 mm thick; berry ca. 2 cm long 
and 1 cm thick (dried), ovoid-cylindrical or oblong, ob- 
tuse at the apex and apiculate, green at maturity. 

Trees of wet evergreen forests of the Caribbean 
slope and lowlands, from 50 to 1200 m elevation. 
Flowers have been collected in April-July; young 
fruits have been collected in September-February, 
while mature fruits have been collected in April- 
May. This species is only known from La Selva, 
the Caribbean slopes of the central highlands in 
Costa Rica, and a collection from Esmeraldas 
Province in Ecuador. 

Ocotea hartshorniana is recognized by its large 
stature, the obovate-oblong laminae with subpar- 
allel tertiary veins, and the usually reddish brown 
or yellowish brown puberulence on flowers, inflo- 
rescences, and leaves. The stilt roots or buttresses 
are also unusual. The young globose "cups" have 
a very small opening and enclose nearly all of the 
developing fruits. This species is related to Ocotea 
sp. aff. caracasana and its allies, a species group 
with stilt roots, decurrent lamina bases, and very 
similar flowers. 



Ocotea helicterifolia (Meissn.) Hemsl., Biol. centr. 
amer., Hot. 3: 73. 1882. Oreodaphne helicteri- 
folia Meissn. in DC., Prodr. 1 5, pt. 1: 123. 1864. 
O. mexicana Meissn., loc. cit. 118.1 864. Phoebe 



helicterifolia (Meissn.) Mez, Jahrb. Konigl. Bot. 
Gait. Berlin 5: 193. 1889. Phoebe betazensis Mez, 
loc. cit. 192. 1889. Figure 8. 

Small to medium-sized trees 4-15(-25) m tall, leafy 
branchlets 2-5 mm thick, hirsute with slender stiff yel- 
lowish hairs to 2.5 mm long, becoming glabrous and 
terete. Leaves alternate or occasionally congested at the 
ends of branchlets, petioles 412 mm long, ca. 1.5 mm 
thick, densely hirsutulous; leaf blades (9-)l 2-25(-30) cm 
long, (2.5-)4.5-10(-l 2) cm broad, elliptic-obovate to ob- 
lanceolate, obovate or pandurate, usually broadest above 
the middle, gradually to abruptly narrowed at the apex 
and acuminate (rarely rounded or obtuse) at the apex, 
the narrowed tip 0-1 5 mm long, rounded to subcordate 
or obtuse at the base, often unequal at the petiole, margin 
entire, drying membranaceous to stiffly chartaceous and 
yellowish green to brown, puberulent on the major veins 
above, conspicuously puberulent beneath with slender 
brownish hairs to 2-3 mm long, with 6-10(-12) major 
secondary veins on each side, central secondaries arising 
at angles of 45-66, tertiary veins elevated beneath and 
often subparallel. Inflorescence axillary from distal leaves 
or from leafless nodes, to 18(-25) cm long and pendu- 
lous, few-flowered panicles with a long (to 1 3 cm) hir- 
sutulous peduncle in ours (occasionally glabrous else- 
where), flowers often in umbellate clusters on slender 
lateral branches, pedicels 3-6 mm long and very slender. 
Flowers 3-6 mm long and 3-5 mm broad, white or yel- 
lowish, perianth usually glabrous on the outside and 
drying dark; outer anthers 0.8-1.6 m long, 0.7-1.3 mm 
broad, elliptic-oblong to ovate, flat, thecae superposed, 
the connective sometimes prolonged beyond the thecae, 
filament short or absent, inner stamens narrow, stami- 
nodes slender to ovate or absent, to 0.8 mm long; pistil 
ca. 2 mm long with globose-ellipsoid ovary and thick 
style ca. 1 mm long, stigma subcapitate or discoid. Fruits 
borne on a long pendulous infructescence, pedicels to 2 
cm long and 3 mm thick below the abruptly expanded 
cupulate receptacle, cup 3-4 mm long, 7-14 mm broad 
and 2-3 mm deep, becoming red; berry ellipsoid 2-2.4 
cm long and 1.2-1.7 cm in diameter (dry), becoming 
black. 



Trees of lowland evergreen rain forests in Costa 
Rica, from 20 to 500 m elevation on the Caribbean 
slopes, and in the Osa peninsula (as interpreted 
here, see below). Flowering material has been col- 
lected in February-March; fruits have been col- 
lected in August. The species ranges from eastern 
and southern Mexico through Guatemala and 
Honduras to Costa Rica. 

Ocotea helicterifolia (often called Phoebe helic- 
terifolia) is a very variable species but usually eas- 
ily identified because of the short-petiolate leaves 
with long (ca. 2 mm) slender hairs, laminae broad- 
est above the middle, acuminate at the apex and 
slightly rounded at the base, and the long-pendant 
inflorescences with puberulent peduncles, but with 
glabrous or sparsely puberulent perianth that dries 
black. The relatively large subsessile outer stamens 



82 



FIELDIANA: BOTANY 



are flat and sometimes have their connective pro- 
longed distally. The very shallow fruiting recep- 
tacle, and usual presence of staminodes (in north- 
ern populations) further distinguish this species. 
However, there are many problems associated with 
the specimens placed here. First, they encompass 
an unusual spectrum of variation, especially in 
Mexico and Guatemala. Secondly, O. valeriana 
may prove to be a higher altitude subspecies, since 
it seems to parallel the variation seen in the Gua- 
temalan and Honduran highlands of material re- 
tained in O. helicterifolia. See the discussion under 
O. valeriana and O. valerioides. Nectandra beli- 
zensis may also be related, though the form of the 
anthers is different. 



Ocotea holdridgeiana W. Burger, sp. nov. Figure 

12. 

Arbor 5-1 5 m alta, ramulis foliiferis 1.5-3.5 mm eras- 
sis. Folia alterna, petiolis 8-18 mm longis; laminis 8- 
1 5(-l 8) cm longis et 2-5(-6.5) cm latis, ellipticis, ovatis, 
elliptico-ovatis vel lanceolatis, apice acuminate, subtus 
glabris vel puberulis in nervis, saepe domatiis praeditis, 
nervis secondariis 3-6 paribus. Inflorescentiae panicu- 
latae racemiformae, 3-9 cm longae, pedunculis usque 4 
cm longis. Flores usque 7 mm longi et 10 mm lati, extus 
glabri, periantho intus papillato-puberulo; stamina ser 
I-II 2-2.8 mm longa, antheris 1.5-2.4 mm longis, stam- 
inodiis gracilibus vel nullis; gynoecio 1 .6-2.7 mm longo, 
stylo usque 1.2 mm longo. Fructus oblongus vel ellip- 
soidus, ca. 25 mm longus et 13 mm crassus; cupula 1 
cm lata et 1-2 mm profunda. 

Trees 5-15 m tall, leafy branchlets 1.5-3.5 mm thick, 
minutely (0.1-0.2 mm) appressed puberulent with slen- 
der brownish hairs, becoming longitudinally striate and 
dark gray. Leaves alternate, petioles 8-1 8 mm long, 0.8- 
1.5 mm thick, sulcate above, sparsely and very minutely 
appressed puberulent; leaf blades 8-1 5(-l 8) cm long, 2- 
5(-6.5) cm broad, elliptic to elliptic-ovate, elliptic-ob- 
long or lanceolate, tapering gradually to the acuminate 
apex, the narrowed tip 0.2-1 .2 cm long, acute at the base 
and slightly decurrent on the petiole, drying stiffly char- 
taceous and dark or yellowish brown, the upper surface 
often lustrous and with the minor venation easily visible, 
glabrous or minutely puberulent above the major veins, 
glabrous or appressed puberulent along the veins be- 
neath, with 3-6 major secondary veins on each side, the 
basal veins usually strongly ascending (subtripliveined) 
or the venation pinnate, central secondaries arising at 
angles of 30-50, conspicuous pit domatia usually pres- 
ent in the axils of the proximal veins beneath, the dom- 
atia often raised and visible on the upper surface (dry). 
Inflorescences axillary to distal leaves or pseudotermin- 
al, 3-9 cm long, few-flowered and racemose, primary 
peduncle to 4 cm long, ca. 1 mm thick, minutely ap- 
pressed puberulent, pedicels 2-4 mm long. Flowers to 7 
mm long and 10 mm broad, white, glabrous on the out- 



side, tepals 2.5-5.6 mm long, 1 .4-4 mm broad, papillate- 
puberulent on the inner surface; outer stamens 2-2.8 mm 
long and with a short broad filament, outer anthers 1.5- 
2.4 mm long, oblong, connective often slightly prolonged 
beyond the superposed thecae, inner stamens to 3 mm 
long, staminodes small (1 mm) and usually slender or 
absent; pistil 1.6-2.7 mm long, style to 1.2 mm long, 
ovary globose to obovoid, stigma capitate. Fruits borne 
on an expanded (1 cm broad) receptacle only 1-2 mm 
deep, fruiting pedicels ca. 15 mm long and 2.5-4 mm 
thick, gradually expanded at the apex into the shallow 
receptacle, red; berry ca. 25 mm long and 13 mm in 
diameter, oblong-ellipsoid or slightly oblong-obovoid, 
green or tinged with red. 

TYPE Costa Rica, San Jose Province, 0.9 km 
above La Chonta and 1 00 m from the Pan Amer- 
ican highway, elevation 2460 m, 11 May 1969, 
Roy W. Lent 7677(holotype, F 1958283; negative, 
6 1 1 24; isotypes, CR, DUKE, MO). Additional Costa 
Rican collections are: Brown 17393 (F, LSU), Da- 
vidse & G. Herrera 29106 (CR, MO), Gomez et al. 
21496 (CR, MO), Gomez- Laurito 1 1077 (CR, F, usj), 
Holdridge 6570 (CR, NY), Madriz 34 (CR, F), Poveda 
753 (CR, F), and Zamora et al. 879 (CR, F). 



Trees of evergreen montane rain forest and 
montane wet forest formations, between ( 1 1 00) 
1600 and 2700 m elevation. Flowering collections 
have been made in February-April, July-Septem- 
ber, and November-December. Fruiting material 
has been collected in February, April, and No- 
vember. This species is only known from the Ca- 
ribbean slopes and near the continental divide in 
the central highlands and the Cordillera de Tala- 
manca; it also occurs in westernmost Panama 
(McPherson 9444, MO). 

Ocotea holdridgeiana is recognized by the lus- 
trous narrow leaves, often somewhat tripliveined, 
and with conspicuous pit domatia in basal vein 
axils on the lower surface. The large flowers, gla- 
brous on the outside, long (2 mm) stamens with 
short (0.2-0.4 mm) filaments, connective often 
prolonged a little beyond the thecae, and the sub- 
capitate stigma are also unusual features. These 
characteristics define a very distinctive species, 
which appears to be related to O. brenesii. Some 
specimens had earlier been identified as Phoebe 
tonduzii, but the type of that name appears to be 
an unusual variant of P. cinnamomifolia. While 
O. holdridgeiana has staminodes, it does not have 
three well-developed staminodes in each flower, 
as is characteristic of Phoebe. The many collec- 
tions of Leslie Holdridge and his astute identifi- 
cations have been extremely helpful in under- 
standing the Lauraceae of Costa Rica. 



BURGER: FLORA COSTARICENSIS 



83 



Ocotea insularis (Meissn.) Mez, Jahrb. Konigl. Bot. 
Gart. Berlin 5: 271. 1889. Phoebe insularis 
Meissn. in DC., Prodr. 15, pt. 1: 33. 1864. O. 
cuneala Mez, Bot. Jahrb. Syst. 30, Beibl. 67: 17. 
1901, non O. cuneata (Griseb.) Gomez, 1894. 
O. ira Mez & Pittier, Bull. Herb. Boissier, ser. 
2, 3: 232. 1903. 0. tonduzii 'Standley, Publ. Field 
Mus. Nat. Hist., Bot. Ser. 18: 456. 1937 (based 
on O. cuneata Mez). Aiouea lundelliana C. K. 
Allen, J. Arnold Arbor. 26: 419. 1945. Figure 
10. 

Trees 10-25 m tall, trunks sometimes with short but- 
tresses, leafy branchlets 2.5-7(-12) mm thick, glabrous 
or minutely (0.1-0.3 mm) appressed puberulent with 
slender straight yellowish brown hairs (at least near the 
shoot apex), longitudinally ridged but becoming terete, 
dark brown to grayish brown. Leaves alternate and usu- 
ally close together distally, petioles 0-12 mm long and 
often poorly differentiated, flat or slightly sulcate above, 
usually with revolute margins; leaf blades (4-)8-20(-28) 
cm long, (2-)5-9(-12) cm broad, obovate to spatulate, 
elliptic-obovate or obovate-oblong, abruptly rounded at 
the bluntly obtuse or short-acuminate apex (sometimes 
acute on small leaves), tapering gradually to the cuneate 
or attenuate and decurrent base, the margin entire and 
slightly revolute (especially at the base), narrowed por- 
tion of the lamina base to 2 cm long, drying subcoria- 
ceous and dark brown to pale grayish brown (often red- 
dish brown in lower-elevation populations), glabrous and 
the minor veins obscure above, lower surfaces glabrous 
or with thin inconspicuous appressed hairs 0.2-0.3 mm 
long, domatia of tufted hairs often present in the vein 
axils beneath, with 6-10 major secondary veins on each 
side, central secondaries arising at angles of 30-60, ter- 
tiary venation subparallel but usually obscure. Inflores- 
cences 1 0-20 cm long, axillary to distal leaves, broadly 
paniculate with flowers usually clustered at the ends of 
short (1-3 cm) lateral branches, peduncle 4-10 cm long 
and 2-3 mm thick, reddish brown, glabrous or puber- 
ulent, pedicels 1-4 mm long and sparsely puberulent. 
Flowers densely to sparsely puberulent on the outside, 
tepals 1.5-2 mm long, ca. 1.3 mm broad; outer stamens 
with short (0.2-0.6 mm) filaments, outer anthers 0.5-1 
mm long and equally broad or narrower, ovate to oblong, 
thecae superposed, staminodes very rarely present; pistil 
1 .5-2.8 mm long, ovary half the length of the pistil, style 
slender, stigma discoid. Fruits borne on a short (4-7 mm) 
shallow (2-4 mm deep) conical or cupulate receptacle 
6-9(-12) mm broad at the top, margin entire or with 
persisting perianth parts, the cup abruptly expanded above 
the thickened (2-3 mm) pedicel, cup and pedicel ca. 1.5 
cm long, becoming red; berry ellipsoid-oblong, 12-20 
mm long and 6-10 mm in diameter (dry), dark green to 
black at maturity. 

Trees of wet evergreen rain forest and cloud 
forest formations, from near sea level to 1600 
(2000) m elevation. Flowering collections have 
been made in all months except October. Fruits 
have been collected in all months except Novem- 
ber. This species (in a wide sense) probably ranges 



from southeastern Nicaragua to Panama, and pos- 
sibly as far as coastal Ecuador. It was first collected 
on Cocos Island. 

Ocotea insularis is recognized by its stiff, essen- 
tially glabrous, obovate leaves usually bluntly 
rounded at the apex and decurrent on the petiole, 
with revolute margins basally. The relatively small 
ellipsoid-oblong fruits, borne in small cups with 
perianth parts usually persisting on the margin, 
also help to identify members of this species. Un- 
fortunately, this species shows considerable local 
divergence (see below) and appears to be closely 
related to Aiouea costaricensis. Ocotea endresiana 
may be no more than an unusual form of O. in- 
sularis. Difficulties can also be encountered in dif- 
ferentiating material of Ocotea sp. aff. caracasana 
and its allies (including O. skutchii and O. whitei), 
all of which have decurrent leaf bases and rather 
similar floral morphology. See the discussions un- 
der these species. Ocotea rivularis is also closely 
related but it has very large leaves. 

The highland populations of Ocotea insularis 
seem to be common around Monteverde in the 
Cordillera de Tilaran, and a number of specimens 
come from the northwestern edge of the Meseta 
Central (San Ramon and Zarcero). Only a very 
few collections have been made near Volcan Poas 
and Volcan Barva (Barba) and on the western side 
of Irazu. A few collections have been made south 
of Cartago, from Muneco, Orosi, and Cachi, in 
very wet forests. The species is apparently more 
common in the western part of the Cordillera de 
Talamanca, in the Chiriqui highlands, and east- 
ward into Code Province, Panama. Among the 
collections from the Chiriqui highlands are several 
in which the upper thecae of the outer anthers are 
reduced or absent. It was such a collection (P. 
White 225, MO) which was the basis of the name 
Aiouea lundelliana. A similar situation seems to 
be found in the flowers of Williams 16344 (us, 
usj) from near La Congreja, Cartago. These spec- 
imens suggest a trend in loss of the upper thecae, 
and they may indicate how Aiouea costaricensis 
(q.v.) originated from Ocotea insularis-like ances- 
tors. In this regard it is interesting to note that 
Aiouea costaricensis and O. insularis appear to be 
sympatric only between Volcan Barva (Barba) and 
Zarcero, despite living in rather similar forest for- 
mations elsewhere. These two species and their 
interrelatedness deserve closer study. 

The name Ocotea insularis is based on the Cocos 
Island population, which differs from most of our 
highland material in its larger more elliptic leaves 
with acute/obtuse apices and a tendency to dry 



84 



FIELDIANA: BOTANY 



dark. The tufts of reddish hairs along the midvein 
on the undersides of the Cocos Island specimens 
are not found in any mainland material, though 
somewhat similar hairs may be found in the vein 
axils. Some collections from the General Valley 
closely resemble the Cocos Island material as re- 
gards leaf form and color after drying. The lower- 
elevation collections from the Caribbean slope also 
have larger ( 1 2-28 cm) leaves, but they are often 
slightly lustrous above, and are more reddish brown 
beneath; in addition, the fruit cups are often entire. 



The following key summarizes some of this vari- 
ation. It is clear that the problems in this group 
of species can only be solved after further study. 
We have decided to treat this group conservatively 
and place collections possibly belonging to O. ira 
and O. tonduzii under O. insularis sensu lato. We 
feel that without a thorough study of collections 
from Panama, coastal Colombia, and coastal Ec- 
uador no better classification can be presented at 
this time. 



Key to Possible Subdivision of Ocotea insularis 

la. Leaves elliptic to obovate and often bluntly acute or acuminate at the apex, usually drying dark 
brown, with tufts of reddish brown hairs along the midrib on the lower leaf surface; a dominant 
tree of Cocos Island O. insularis s.s. 

Ib. Leaves usually obovate to oblanceolate with the larger laminae usually rounded at the apex, never 
with tufts of reddish hairs along the length of the midvein (but sometimes in the vein axils); trees 
of the mainland 2a 

2a. Leaves drying darker brown or reddish brown, 15-25 cm long; fruits 15-25 mm long and borne in 
a cup with an entire margin; from below 1 000 m elevation O. ira 

2b. Leaves usually drying pale grayish brown, 5-20 cm long; fruits 12-18 mm long and usually borne 
in a cup with some persisting perianth lobes; above 1000 m in the Cordillera de Tilaran and around 
the central highlands O. tonduzii 



Ocotea laetevirens Standl. & Steyerm., Publ. Field 
Mus. Nat. Hist., Bot. Ser. 23: 1 14. 1944. Figure 
12. 

Trees 4-18 m tall, leafy branchlets 2-8 mm thick, 
strongly 1-3-ridged from early stages (strongly angular 
in cross section) but becoming terete, glabrous or with 
minute (0.2 mm) thin ascending hairs near the shoot 
apex, greenish (dry) or brown. Leaves alternate, petioles 
8-22 mm long, 1.2-2.5 mm thick, with adaxial ridges 
forming a sulcus; leaf blades 7-1 8 cm long, 3-8 cm broad, 
elliptic to elliptic-obovate or elliptic-oblong, acute to 
short-acuminate (occasionally bluntly obtuse) at the apex, 
tapering gradually to the obtuse or acute base and slightly 
decurrent on the petiole, margin slightly revolute when 
dry, drying stiffly chartaceous to subcoriaceous and yel- 
lowish green to greenish brown, glabrous and slightly 
lustrous with the tertiary venation visible on the upper 
surface, glabrescent and duller beneath with the minor 
venation raised and forming a poorly denned reticulum, 
with 3-7 major secondary veins on each side, the central 
secondaries arising at angles of 30-60, the basal sec- 
ondaries occasionally prominent and arcuate-ascending, 
domatia-like developments of tufted hairs or pits often 
present in the axils of proximal secondary veins beneath. 
Inflorescences axillary to both distal and proximal leaves, 
to 20 cm long, an open many-branched panicle, peduncle 
3-12 cm long and ca. 1.3 mm thick, glabrous, pedicels 
ca. 2 mm long. Flowers 2.5-4 mm long, 2.5-4 mm broad, 



campanulate, white but drying dark, glabrous on the 
outside or minutely papillate distally, tepals ca. 2 mm 
long and 1.7 mm broad; outer stamens 1.4-1.8 mm long 
with oblong or ovate anthers 0.7-0.8(-1 . 1) mm long and 
0.6-0.8(-0.9 ) mm broad, filaments usually prominent 
(0.5-1 mm) and broad, thecae clearly superposed, inner 
stamens 1.5-2 mm long and with large glands, stami- 
nodes absent or very slender (to 0.8 mm long); pistil 1.7- 
2.5 mm long, ovary 1-1.3 mm in diameter, style 0.7- 
1 . 1 mm long, stigma simple or oblique. Fruits borne in 
a cupulate or conic receptacle 1 .2-2.5 cm long, 1-1 .6 cm 
broad and 3-6 mm deep, gradually expanded above the 
thickened pedicel, surface rough but lacking prominent 
lenticels, becoming red; berry 2.2-3 cm long, 1.3-1.8 cm 
in diameter (dry), ellipsoid, becoming black. 

Trees of montane evergreen forest formations, 
from 1200-2000 m elevation. In Costa Rica flow- 
ering material has been collected in March-April, 
July, and December; fruits have been collected in 
March-April, June-August, and December-Jan- 
uary. This species has only been collected from 
the northeastern portion of the Meseta Central 
(Volcan Barva to Tablazo) and the western part 
of the Cordillera de Talamanca in Costa Rica. The 
species is also known from Guatemala. 

Ocotea laetevirens is distinguished by its stiff 



BURGER: FLORA COSTARICENSIS 



85 



glabrous leaves that often turn yellowish green or 
pale brown when dried, the slightly decurrent lam- 
ina base, the almost glabrous flowers, the outer 
stamens with prominent filaments, poorly devel- 
oped stigma on a conspicuous style, shallow fruit 
cup, and ellipsoid fruits. We follow C. K. Allen 
(1945) and Hammel (1986) in using this name for 
these Costa Rican plants. Even though the leaves 
of the type (Steyermark 49189, F, from Guate- 
mala) are thinner and much more acuminate than 
in Costa Rican material, the flowers are nearly 
identical. Other highland species with stiff gla- 
brous pale-drying leaves that are often glossy above 
and can be confused with this species are Ocotea 
meziana, Nectandra cufodontisii , and Phoebe 
hammeliana. 

This species is poorly understood at present. It 
may be much more variable than described above, 
with some individuals having smaller, more co- 
riaceous leaves. Two larger-leaved (20-30 cm) col- 
lections, with longer more open pendant infruc- 
tescences, from the Osa peninsula (Primack 96, 
MO) and Mastatal de Puriscal (Gomez- Laurito 
11237, CR, F, usj) are tentatively placed under this 
species. (Note that these last two collections are 
both from lower elevation habitats, and they may 
not be conspecific.) It seems best to place these 
latter anomalous collections under O. laetevirens 
until we have better samples of the populations 
they represent. Ocotea verapazensis Standl. & 
Steyerm. and O. standleyi C. K. Allen (based on 
Phoebe macrophylla Standl. & Steyerm.) may prove 
to be synonymous with an expanded concept of 
O. laetevirens; all the relevant types are from ev- 
ergreen forests at about 1 500 m elevation in Gua- 
temala. See also Ocotea sp. aff. O. laetevirens at 
the end of the species descriptions. 



Ocotea lentil W. Burger, sp. nov. Figure 2. 

Arbor 4-8 m alta, ramulis foliiferis 5-10 mm crassis. 
Folia alterna, petiolis 8-30 mm longis et 2.5-4 mm cras- 
sis; laminis 18-40 cm longis et 8-16 cm latis, obovatis 
vel elliptico-obovatis, apice breviter acuminato vel ro- 
tundato, basi cuneato, subtus strigosis in nervis, pilis 
0.2-1 mm longis, nervis secondariis 7-1 1 paribus. In- 
florescentiae paniculatae, 8-20 cm longae, saepe racem- 
iformae, pedunculis 3-12 cm longis. Flores 5-7 mm lon- 
gi et 6-10 mm lati, extus glahri vel minute puberuli, 
tepalis intus papillato-puberulis; stamina ser. I-II an- 
theris 1 .2-1 .4 mm longis, staminodiis nullis vel parvulis 
(0.6 mm); gynoecium ca. 2.5 mm longum, ovario ellip- 
soideo, 1.5-1.9 mm longo. Fructus ellipsoideus vel ob- 
longoideus, 3-6 cm longus et 2-3 cm crassus; cupula 8- 
25 mm lata, 2-8 mm profunda. 



Small trees 4-8 m tall, leafy branchlets 5- 1 mm thick, 
hirsute with straight stiff yellowish hairs 0.5-1.3 mm 
long and persisting on the terete stems. Leaves alternate 
or crowded together distally, petioles 8-30 mm long, 2.5- 
4 mm thick, densely hirsute with stiff hairs, narrowly 
sulcate above; leaf blades 1 8-40 cm long, 8-16 cm broad, 
obovate to elliptic-obovate, abruptly narrowed to the 
short-acuminate or rounded apex, gradually tapering to 
the cuneate base, margin often undulate (dry), the lam- 
inae drying chartaceous to stiffly chartaceous and dark 
greenish or brown above, densely and minutely (0.3 mm) 
puberulent on the major veins above and with longer 
(ca. 1 mm) hairs between the veins, lower surface with 
stiff straight hairs 0.2-0.8(-1.2) long on both the major 
and minor veins, with 7-1 1 major secondary veins on 
each side, central secondaries arising at angles of 40- 
70, secondary and tertiary venation prominent beneath, 
tertiary veins often subparallel and perpendicular to the 
secondaries. Inflorescences axillary to distal leaves or 
from leafless nodes, 8-20 cm long, elongate panicles with 
short lateral branches (almost racemiform), primary pe- 
duncle 3-12 cm long, slender (1.5 mm) and sparsely 
hirsute, pedicels to 10 mm long. Flowers 5-7 mm long, 
6-10 mm broad, perianth glabrous or sparsely puberu- 
lent on the outside, papillate-puberulent on the inside; 
outer stamens subsessile, anthers 1.2-1.4 mm long, ca. 
1.2 mm broad, thecae superposed but the lower often 
somewhat lateral, staminodes absent or small (0.6 mm) 
and clavate, with erect straight hairs around the rim of 
the floral tube and base of the stamens; pistil ca. 2.5 mm 
long, ovary 1.5-1.9 mm long, ellipsoid, style slender, 
stigma capitate. Fruits borne on a flat, conical or broadly 
cupulate receptacle, 4-10 mm long, 8-25 mm broad and 
2-8 mm deep, pedicels becoming 2-4 cm long and 2.5- 
6 mm thick; berry 3-6 cm long, 2-3 cm in diameter, 
ellipsoid, the base not tightly enclosed in the cup when 
a cup is present (dry). 

TYPE Costa Rica, Cartago Province, hillside 
overlooking the Rio Grande de Orosi, 3.5 km SE 
of Tapanti, elevation 1400 m, 9 April 1967, Roy 
W. Lent 794 (holotype, F 1749021; negative, 
61119; isotypes, CR, MO). 

Small trees of the very wet cloud forest for- 
mations of the Caribbean slope, at about 1400 m 
elevation. Rowers have been collected in April, 
while fruits have been collected in April and Au- 
gust. At present the species is known from only 
three collections (Gomez 18814, CR, MO, Lent 794, 
2070, CR, F), all from near Tapanti, along the Rio 
Grande de Orosi. 

Ocotea lentii is recognized by its very large but 
thin-textured obovate leaves with short straight 
stiff hairs, long pedunculate inflorescences with 
relatively few flowers, and large ellipsoid fruits. 
The upper edge of the floral tube with straight 
lustrous hairs, and the subsessile anthers about as 
broad as long, are also distinctive features. The 
general form of the flowers and vegetative parts 
relate this species to Ocotea valeriana, but the very 



86 



FIELDIANA: BOTANY 



large leaves and fruits of O. lentil are amply dis- 
tinct. Ocotea valerioides, with large leaves and very 
similar floral structure, is more closely related. It 
too, has a tendency for the outer anthers to become 
broad and develop a Nectandra-\ike configuration. 
Ocotea lentii and O. valerioides differ in elevation, 
leaf texture, and color (dried), as well as vesture, 
inflorescence-form, and size of the fruits. See the 
discussions under O. valerioides and O. mollifolia. 
Roy Lent's Costa Rican collections number about 
4,000 and include many rare and important finds, 
of which this unusual species is an example. 



Ocotea leucoxylon (Sw.) Laness., PI. util. colon, 
franc. 156. 1886. Laurus leucoxylon Sw., Prodr. 
65. 1 788. 0. subsericea Standl., Publ. Field Mus. 
Nat. Hist., Hot. Ser. 18: 456. 1937. O. lenticel- 
lata Lundell, Wrightia 5: 54. 1974. Figure 14. 

Small or medium-sized trees, 3-15(-25 m) tall, dioe- 
cious (unisexual), leafy branchlets 1.7-6.5 m thick, gla- 
brous or with very minute (0.05-0.1 mm) grayish ap- 
pressed ascending hairs, often with longitudinal ridges 
but becoming terete. Leaves alternate and evenly spaced, 
petioles 10-22 mm long, 1.5-2.5 mm thick, with ele- 
vated adaxial ridges usually forming a shallow sulcus 
above; leaf blades 10-23(-27) cm long, 3-9 cm broad, 
elliptic-oblong to oblong or elliptic-lanceolate, usually 
acuminate at the apex with a narrow tip 0.5-2 cm long, 
acute to obtuse at the base, margin usually slightly re- 
volute, the laminae drying very stiffly chartaceous to 
subcoriaceous and grayish green to dark olive green, gla- 
brous above and the midvein flat, essentially glabrous 
(in ours) or very sparsely and minutely puberulent be- 
neath, with 4-8 major secondary veins on each side, 
primary and secondary veins prominent beneath, central 
secondaries arising at angles of 30-50(-60 ), epidermal 
cells somewhat convex on the lower surface and giving 
an uneven or scale-like effect (10 x). Inflorescences sol- 
itary from the axils of leaves or leafless nodes throughout 
the length of new twigs, 4-9(-13) cm long, paniculate 
with well-spaced lateral branches, basal branches longer 
than the distal, pedicels 1-3 mm long. Male flowers ca. 
2.5 mm long and 2.5 mm broad (to 3 x 3 mm elsewhere), 
white to yellowish, tepals 1.5(-2) mm long and 1 mm 
broad, very minutely papillate puberulent on the edges; 
outer stamens with anthers 0.5-1 mm long and 0.5-0.9 
mm broad, with short (0.2-0.3 mm) broad filaments, 
thecae superposed or the lower slightly lateral to the 
upper, inner stamens ca. 1.5 mm long with anthers 0.9 
mm long, staminodes absent; pistillode ca. 2 mm long 
and 0.2-0. 5 mm thick, stigma discoid. Female flowers 
not seen (probably very similar to the male). Fruits sub- 
tended by a broadly obconic or saucer-like receptacle 4- 
10 mm long, 5-8 mm broad, and 1-3 mm deep, reddish 
brown, often with distinctive wart like lenticels, rim en- 
tire; berry 8-10 mm in diameter (dry, to 1 5 mm in life), 
globose to globose-obovoid, green to purple, the fruiting 
inflorescences rarely more than 8 cm long. 



Trees of tropical wet to premontane rain forest 
formations on the Caribbean slope and adjacent 
continental divide, from 20 to 1 800 m elevation. 
The trees flower in February-July at La Selva 
(Hammel 1986, p. 229), and fruits have been col- 
lected throughout the year, except November, in 
Costa Rica. The species ranges from southern 
Mexico and the West Indies to South America. 

Ocotea leucoxylon is recognized by it stiffoblong 
glabrous leaves, strongly angular young stems, 
small functionally unisexual flowers, and small 
globose fruits, subtended by a small cupulate or 
obconic warty receptacle. Hammel (1986) men- 
tions the foul-smelling foliage (when crushed), the 
distantly spaced inflorescences, grayish green leaves 
and scaly lower leaf surface as additional distin- 
guishing features. The "scaly effect" on the lower 
leaf surface may simply be the reflections of light 
from the convex surfaces of epidermal cells (in 
life); there are no scales under the microscope 
( 1 00 x ). We follow Hammel ( 1 986), who included 
O. lenticellata from Belize under this species. This 
species has also been cited as O. leucoxylon (Sw.) 
Gomez de la Maza (Dice. Nombres Vulg. Cub. 
Puerto Riq. 12.), but Howard (1981) states that 
this publication was issued in 1889, and the day 
and the month are unknown. The type of Stan- 
dley's O. subsericea (Brenes 6789, CR, F) has im- 
mature foliage and larger more puberulent flowers; 
it probably belongs in synonymy here. 



Ocotea meziana C. K. Allen, J. Arnold. Arbor. 26: 
360-361. 1945. Figure 14. 

Trees 2-1 5 m tall, trunk to 80 cm d.b.h., bark grayish 
or brown, leafy branchlets 1.3-4 mm thick, at first with 
short (0.2 mm) ascending yellowish hairs but quickly 
becoming glabrous and pale grayish to yellowish green. 
Leaves alternate, larger in low elevation plants than in 
montane plants (see below), petioles 5-1 6(-30) mm long, 
1-2 mm broad, flat or slightly sulcate above; leaf blades 
5-15 or 1 1-22 cm long, 2-6 or 4-10 cm broad, elliptic 
to narrowly elliptic or elliptic-oblong (broadly elliptic at 
lower elevations), tapering gradually to the short-acu- 
minate apex and acute base (tapering more abruptly in 
the broader leaves of lower elevations), drying stiffly 
chartaceous and gray or greenish gray, smooth and gla- 
brous above with the venation flat, essentially glabrous 
beneath, with 4-6 major secondary veins on each side, 
the central secondaries arising at angles of 30-60, with 
pit domatia in the axils of major veins or along the 
secondary veins, the smallest veins forming a distinct 
reticulum on the lower surfaces with areolae about 0.5 
mm broad. Inflorescences axillary or extra-axillary on 
distal twigs, to 10(-15) cm long, few-branched panicles, 
peduncle 1.5-6 cm long, glabrous or very minutely pu- 
berulent, lateral branches with few (3-7) flowers, pedicels 



BURGER: FLORA COSTARICENSIS 



87 



2-5 mm long and articulate near the base, drying dark. 
Flowers ca. 2-3 mm long, 4 mm broad, yellowish green 
but drying black, essentially glabrous, tepals 1 .2-2 mm 
long; outer stamens with prominent (ca. 1 mm) filaments, 
outer anthers 0.7-1 mm long, narrow with superposed 
thecae, staminodes absent; pistil 1 .5-2.5 long, ovary tur- 
binate or narrowed near the base, style ca. 1 mm long, 
stigma simple. Fruits borne on a shallow (ca. 2-3 mm 
cup) cup 8-14 mm broad, 1.5-3 cm long, funnelform 
and often abruptly flared at the apex, red; berry 2-3 cm 
long, 1.3-2 cm in diameter (dried), becoming purple or 
black. 

Trees of the wet evergreen forest formations of 
the Caribbean slope and the continental divide, 
from 50 to 2300 m elevation. Flowering collec- 
tions have been made in February-July, Septem- 
ber-October, and December. Fruiting material has 
been collected in July-December. The species, as 
presently understood, ranges from northern Costa 
Rica to Code Province in Panama. 

Ocotea meziana is recognized by the few-flow- 
ered and essentially glabrous inflorescences with 
small flowers that dry blackish, the glabrous (or 
very minutely puberulent) leaves that dry grayish 
or greenish and slightly lustrous beneath, and the 
shallow funnelform cupules about 1 cm broad at 
the apex. This treatment follows Hammel's ( 1 986) 
interpretation, and includes the larger-leaved 
specimens from La Selva, together with the small- 
er-leaved highland (800 to 2100 m) material. Pit 
domatia along the secondary veins are often pres- 
ent in highland material, as in the type collection 
(A. Smith 359, F) from about 1600 m elevation. 
Ocotea meziana is closely related to O. laetevirens 
(q.v.). Herbarium material ofO. meziana may re- 
semble Ocotea cernua, O. tenera, Licaria cufo- 
dontisii, and L. sarapiquensis , but the fine retic- 
ulum formed by the minor venation on the lower 
surface when dry helps to separate O. meziana 
from all of these other species. 



on each side, the central secondaries arising at angles of 
30-50. Inflorescences small and axillary to older or 
fallen leaves, racemose panicles to 6 cm long, peduncles 
to 4 cm long, slender (0.5 mm) and grayish puberulent, 
flowers solitary or 2-3 on short branches of the rachis, 
pedicels 1-5 mm long. Flowers 4-6 mm broad, buds 2- 
3 mm long, tepals ca. 3 mm long, obtuse, glabrous dis- 
tally and within, pellucid punctate; outer stamens ca. 1 .3 
mm long with anthers ca. 0.8 mm long and 0.9 mm 
broad, ovate with superposed thecae, inner stamens with 
oblong anthers ca. 0.7 mm long, staminodes small (0.5- 
0.8 mm) and with or without a glandlike apex; pistil ca. 
2 mm long, ovary globose and glabrous, style ca. 0.6 mm 
long, stigma simple or slightly discoid. Fruits borne on 
a short cup ca. 6 mm broad, usually with the broad but 
short tepals persisting at the edge, becoming red; berry 
ca. 2 cm long and 1.3 cm in diameter, ellipsoid, becoming 
black. 

Trees of wet evergreen montane forest forma- 
tions, between 1400 and 2300 m elevation in cen- 
tral Costa Rica. Flowering collections have been 
made in December-February; fruits have been 
collected in May-August. This species is only 
known from the region around Zarcero, the slopes 
of Volcan Irazu, and the western parts of the Cor- 
dillera de Talamanca (Copey and Sta. Maria de 
Dota) in Costa Rica. 

Ocotea mollicella is recognized by its small nar- 
row leaves with soft grayish puberulence beneath, 
small inflorescences that become dark on drying, 
presence of staminodes, and restricted highland 
habitat. There is little good fruiting material, and 
the fruiting description may require revision. This 
species is known as quizarra amarillo. This species 
is very closely related to Ocotea pittieri, and there 
is the possibility that O. mollicella is only an un- 
usual subspecific element of that species. How- 
ever, the soft grayish puberulence is quite char- 
acteristic and intermediate collections are not 
apparent. The two species are here transferred from 
Phoebe to Ocotea; see the discussion under O. pit- 
tieri. 



Ocotea mollicella (Blake) van der Werff, comb, 
nov. Phoebe mollicella Blake, Contr. Gray Herb. 
52: 64. 1917. Figure 4. 

Trees 6-16 m tall, distal branchlets 0.7-3 mm thick, 
usually pale grayish or yellowish brown tomentulose with 
thin hairs 0.1-0.3 mm long, terete, becoming slightly 
striate. Leaves alternate, petioles 6-12 mm long, ca. 1 
mm thick, flat above, puberulent; leaf blades 3-7(-9) cm 
long, l-2.3(-3) cm broad, narrowly elliptic to elliptic- 
lanceolate or elliptic-oblong, tapering gradually to the 
acute or acuminate apex, acute at the base, drying char- 
taceous, usually dark and slightly lustrous above, gla- 
brescent above, grayish puberulent below with slender 
hairs 0. 1-0.4 mm long, with 3-6 major secondary veins 



Ocotea mollifolia Mez & Pittier, Bull. Herb. Bois- 
sier, ser. 2, 3: 233. 1903. Figure 7. 

Small and medium-sized trees 4-20 m tall, prop roots 
often present (fide Hammel), leafy branchlets 2-6 mm 
thick, densely puberulent with yellowish brown to orange 
brown hairs 0.2-0.5 mm long. Leaves alternate or oc- 
casionally subopposite, petioles 4-12(-18) mm long, 1.5- 
3.5 mm thick, densely puberulent; leaf blades (9-) 12- 
24 cm long, 6-14 cm broad, obovate to broadly elliptic- 
obovate or oblong, nearly always broadest above the 
middle, usually rounded and abruptly narrowed to an 
acute or short-acuminate apex with a tip 0. 5- 1 . 5 cm long, 
narrowed gradually to an acute or obtuse base (never 



88 



FIELDIANA: BOTANY 



decurrent), drying chartaceous to stiffly chartaceous, pu- 
berulent on the larger veins above, puberulent beneath 
with straight or crooked slender hairs to 0.5 mm long, 
yellowish brown to dull brown in color, with 5-9 major 
secondary veins on each side, central secondaries arising 
at angles of 30-60, tertiary veins prominent beneath 
and often subparallel and perpendicular to the second- 
aries. Inflorescences axillary or apparently extra-axillary, 
6-20(-30) cm long, an open panicle, primary peduncle 
to 14 cm long and 1 .5-2.5 mm thick, densely puberulent, 
pedicels 2-9 mm long. Flowers 3-4(-5) mm long, 3-5(-7) 
mm broad, cream white but becoming brownish on 
drying, tepals 1.5-2.2(-3) mm long, densely puberulent 
on the outside; outer stamens subsessile with ovate to 
oblong anthers 0.7-1 .4 mm long, thecae superposed and 
the connective sometimes prolonged apically, inner sta- 
mens 1-1.8 mm long, staminodes present and slender 
(ca. 0.8 mm long) or absent; pistil 1.5-2.5 mm long, 
ovary turbinate or ellipsoid, style short to long ( 1 .4 mm) 
and glabrous or puberulent (as in the type), stigma simple 
or slightly discoid. Fruits borne on a conical cup 10-18 
mm long and ca. 1 5 mm broad, shallow or deep (2-5 
mm), rim of the cup usually with broad (3 mm) persisting 
perianth parts 1-2 mm long, becoming red; berry 2-3(-4) 
cm long, 1.5-2 cm in diameter, oblong-cylindrical. 



Trees of stream sides and forest understory usu- 
ally on hilly sites, on the lower slopes of the Ca- 
ribbean escarpment between 50 and 900 m ele- 
vation, and near Palmar Sur de Osa on the Pacific 
slope. Flowering material has been collected in 
January-March, and flowering has been observed 
in February-June at La Selva (Hammel 1986). 
Mature fruits have been collected in September- 
January. This species is only known from Costa 
Rica. 

Ocotea mollifolia is recognized by the conspic- 
uously puberulent and usually obovate lamina that 
are thin in texture and, while narrowed gradually 
to the base, are not decurrent on the petiole. The 
open inflorescences with widely spaced flowers, 
dense puberulence on most parts, fruit cups with 
persistent lobes on the margins, and prop roots 
near the base of the trunk are further distinguishing 
characters. Despite obvious differences between 
typical members of each species, this species may 
be related to O. gomezii; collections from around 
500 to 700 m exhibit some shared characteristics 
(but they are not clearly intermediate as one would 
expect from a cline or hybridization). In this regard 
Burger et al. 11730 (CR, F, NY) from about 600 m 
has much larger flowers and floral parts than our 
other, lower-elevation, collections of O. mollifolia. 
While these flowers approach those of O. gomezii 
in size, the large open inflorescences are more typ- 
ical O. mollifolia. The prop roots and other char- 
acteristics suggest a relationship with O. hart- 
shorniana and O. sp. aff. caracasana. 



Ocotea monteverdensis W. Burger, sp. nov. Fig- 
ure 4. 

Arbor usque 25 m alta, ramulis foliiferis 2-5.5 mm 
crassis. Folia alterna, petiolis 2-8 mm longis; laminis 4 
12 cm longis et 1.5-4 cm latis, ellipticis vel anguste 
elliptico-oblongis, apice saepe acuto, basi acuto vel cu- 
neato, decurrente, subtus adpresse puberulis, nervis se- 
condariis 3-8 paribus. Inflorescentiae paniculatae, 6-12 
cm longae, pedunculis 2-5 cm longis. Flores 3-3.5 mm 
longi et 4-5 mm lati, extus dense puberuli, tepalis intus 
papillato-puberulis; stamina ser. I-II antheris 0.6-0.9 
mm longis, staminodiis nullis; gynoecium 1.5-2.2 mm 
longum. Fructus ellipsoideus, usque 3 cm longus et 1.5 
cm crassus; cupula 12-16 mm lata et 1-3 mm profunda. 



Trees to 25 m tall, to 55 cm d.b.h., leafy branchlets 
2-5.5 mm thick, at first yellowish brown or orange brown 
tomentulose with dense short (ca. 0.2 mm) tomentulose 
hairs, with prominent longitudinal ridges but becoming 
terete and grayish. Leaves alternate, often closely clus- 
tered distally, petioles not clearly differentiated, 2-8 mm 
long, with lateral margins and flat above; leaf blades 4- 
12 cm long, 1.5-4 cm broad, elliptic to narrowly elliptic 
or narrowly elliptic-oblong, tapering gradually to an acute 
(rarely obtuse) apex, tapering gradually to the acute (ob- 
tuse or cuneate) base and decurrent on the petiole, mar- 
gin slightly revolute near the base, the laminae drying 
stiffly chartaceous or subcoriaceous, grayish green or 
grayish brown above with the tertiary veins paler and 
distinct, smooth above and essentially glabrous except 
above the midvein, grayish beneath with a dense cov- 
ering of small (0.2 mm) slender appressed ascending hairs, 
glabrescent in age, with 3-8 major secondary veins on 
each side, central secondaries arising at angles of 30- 
60. Inflorescences axillary to distal leaves or pseudo- 
terminal, 6-12 cm long, paniculate and many-flowered, 
peduncles 2-5 cm long, ca. 2 mm thick, brownish pu- 
berulent, pedicels 1-3 mm long. Flowers 3-3.5 mm long, 
4-5 mm broad, densely ferruginous puberulent on the 
exterior, tepals 1.5-2 mm long and ca. 1.5 mm broad, 
papillate puberulent within; outer stamens 1.3-1.5 mm 
long, subsessile or with short (0.3 mm) puberulent fila- 
ments, outer anthers 0.6-0.9 mm long, 0.6-0.7 mm broad, 
rectangular, thecae superposed or the lower slightly lat- 
eral, inner stamens ca. 1.6 mm long with anthers ca. 0.8 
mm long, glands large, staminodes not seen; pistil 1.5- 
2.2 mm long, style 0.5-0.7 mm long and slender, stigma 
simple. Fruits borne in a shallow (1-3 mm) cup 12-16 
mm broad, expanded gradually from the pedicel and 
obconic or somewhat saucer-shaped, red, fruiting pedicel 
4-10 mm long and ca. 3 mm thick; berry becoming as 
much as 3 cm long and 1.5 cm in diameter (dry), ellip- 
soid, green. 

TYPE Costa Rica, Puntarenas Province, Mon- 
teverde, in Figuerola's pasture, elevation about 
1450 m, 27 July 1977, Gary S. Hartshorn 1900 
(holotype, CR 91924; isotypes, F 1866719; nega- 
tive, 61120, MO). 

Trees of montane cloud forest formations in the 
Cordillera de Tilaran and westernmost slopes of 



BURGER: FLORA COSTARICENSIS 



89 



the Meseta Central, from about 800 to 900 m near 
San Ramon to 1400 to 1600 m at Monteverde. 
Flowering material has been collected in June- 
August, and fruiting material has been collected 
in July-August. This species is only known from 
west-central Costa Rica. 

Ocotea monteverdensis is recognized by the 
smaller elliptic leaves that are densely appressed 
puberulent beneath, the decurrent lamina base, the 
ferruginous puberulence on stems and inflores- 
cences, the larger ellipsoid fruits borne in shallow 
cups, and the restricted range. Specimens of this 
species may resemble Ocotea mollicella, but that 
species lacks the dense pubescence, the decurrent 
leaf base, and differs in flowers and fruits. Ocotea 
monteverdensis appears to be most closely related 
to O. skutchii., O. hartshorniana, and their allies. 
The following collections are placed here: Brenes 
3849 (F, NY), Dryer 1332, 1591 (CR, F), Gentry & 
Haber 48736 (MO), Poveda 1108 (CR, F), J. & K. 
Utley 5407 (DUKE, F), and N. Wheelwright 1, BOB, 
33, 75 (all MO). 



Ocotea nicaraguensis Mez, Jahrb. Konigl. Bot. 
Gait. Berlin 5: 238. 1889. O. pentagona Mez, 
Bot. Jahrb. Syst. 30, Beibl. 67: 17. 1901. Fig- 
ure 3. 

Slender trees 6-8(-18) m tall, trunk 10-20 cm d.b.h., 
leafy branchlets 4-8 mm thick, strongly angled in cross 
section with 3-5 prominent longitudinal ridges, very mi- 
nutely appressed puberulent or glabrous, distal stems 
usually hollow and harboring ants. Leaves alternate, pet- 
ioles 5-1 2(-20) mm long, 2-4 mm thick, slightly or deep- 
ly sulcate above, often drying dark; leaf blades 1 4-40(-5 5) 
cm long, 4.5-12(-15) cm broad, very narrowly obovate 
to oblong-obovate, oblanceolate or oblong-elliptic, 
rounded to bluntly obtuse or short-acuminate at the apex, 
tapering gradually to the obtuse (rounded) to cuneate 
(but not decurrent) base, drying subcoriaceous to cori- 
aceous and often pale grayish brown, glabrous above and 
below, midvein up to 5 mm broad beneath, with 7- 
10(-12) major secondary veins on each side, central sec- 
ondaries arising at angles of 30-50(-65 ), tertiary veins 
often perpendicular to the secondaries and subparallel, 
slightly raised on the lower surface, domatia absent. In- 
florescences axillary to distal leaves, 1 5-30 cm long, pan- 
iculate but with short (2-3 cm) lateral branches and 
somewhat racemose in form, peduncle 7-12 cm long, 2- 
3 mm thick, very minutely puberulent or glabrous, ped- 
icels 1.5-5 mm long. Flowers 2.2-3 mm long, 3-5 mm 
broad, very minutely puberulent or glabrous, tepals ovate 
and thin, usually glabrous within; outer stamens with 
short (0.5 mm) filaments and narrow anthers ca. 0.9 mm 
long, thecae superposed, inner stamens 1 .4-1 .8 mm long, 
staminodia absent; pistil 2-2.4 mm long, the style nar- 
rowed gradually and equalling the ovary in length, stigma 



slightly discoid. Fruits borne on a shallow (1-2 mm deep) 
receptacle ca. 8 mm broad, with the perianth bases (2- 
3 mm broad and 1-2 mm long) usually persisting, pedicel 
becoming 5-1 5 mm long and 3-4 mm thick at the apex, 
becoming reddish; berry 12-25 mm long and 6-12 mm 
thick (dried), ellipsoid to ovoid, becoming purple or black. 



Trees of evergreen forest formations below 1 000 
(71300) m elevation, on both the Caribbean and 
Pacific slopes of Costa Rica. Flowering collections 
have been made in August-October, and fruiting 
material has been collected in January-April and 
November. The species appears to be most com- 
mon in the Caribbean lowlands, the Cordillera de 
Tilaran (600 to 1000 m), the General Valley, and 
the Golfo Dulce region. This species ranges from 
southeastern Nicaragua to westernmost Panama. 

Ocotea nicaraguensis is recognized by its large, 
stiff, often oblanceolate leaves on short thick pet- 
ioles, the hollow distal stems, lack of conspicuous 
puberulence, and the small fruiting receptacles with 
(usually) persisting perianth bases. The prominent 
longitudinal ridges on the young stems are also 
useful in identification, and separating this species 
from O. atirrensis, with much thinner leaves that 
are often caudate-acuminate and dry dark in color. 
Ocotea paulii of higher elevations also lacks the 
prominently ridged stems and has smaller more 
oblong leaves. All three of these species have the 
distal stems hollow and often inhabited by small 
ants. Ocotea aurantiodora has similar leaves and 
solid stems with even more prominent ridges, but 
the fruits are very different. 



Ocotea oblonga (Meissn.) Mez, Jahrb. Konigl. Bot. 
Gait. Berlin 5: 366. 1889. Mespilodaphne ob- 
longa Meissn. in DC., Prodr. 15, pt. 1: 107. 
1864. O. portoricensis Mez, loc. cit. 364. 1889. 
Phoebe mayana Lundell, Amer. Midi. Natural- 
ist 29: 473. 1943. Figure 11. 

Trees 8-30 m tall, 20-45 cm d.b.h., dioecious, often 
becoming buttressed in age, sometimes with whorled 
branching, leafy branchlets 1.5-4 mm thick, glabrous or 
very minutely (0. 1 mm) puberulent near the apex, lon- 
gitudinally ridged but becoming terete and lenticellate, 
dark or grayish. Leaves alternate, petioles 1-2 cm long 
but poorly differentiated, 1-2 mm broad, flat above, gla- 
brous or minutely puberulent; leaf blades 5-15(-22) cm 
long, 2-4(-6) cm broad, oblanceolate to narrowly ellip- 
tic-obovate or narrowly obovate, usually short-acumin- 
ate at the apex (bluntly acute to obtuse), tapering very 
gradually to the cuneate or decurrent base, drying char- 
taceous to stiffly chartaceous and usually very dark above 
with the major veins slightly raised and the minor veins 



90 



FIELDIANA: BOTANY 



obscure, grayish or brownish beneath and glabrous or 
obscurely appressed puberulent with small (0.2-0.4 mm) 
thin hairs, with 5-8 major secondary veins on each side, 
the central secondaries arising at angles of 30-50, pit 
domatia often present near the mid vein beneath. Inflo- 
rescences axillary to distal leaves, 5-13 cm long, race- 
mose panicles with short (1-2 cm) lateral branches, pe- 
duncle 8-30 mm long, 0.7-2 mm thick, sparsely to densely 
puberulent, pedicels 1-3 mm long. Flowers functionally 
unisexual, 1.7-2.5 mm long, ca. 3.5 mm broad, cam- 
panulate, minutely and densely appressed puberulent on 
the outside, tepals 1.2-1.5(-2) mm long, 1.2-1.4 mm 
broad, papillate puberulent within; male flowers with 
outer anthers ca. 1 mm long on filaments ca. 0.4 mm 
long, thecae superposed in oblong anthers, inner stamens 
ca. 2 mm long with small (0.5 mm) glands, staminodes 
absent; female flowers with stamens half the size of those 
in the male flowers; pistil 1.5-2 mm long, ovary ovoid 
and ca. 0.7 mm in diameter, style thick and ca. 1 mm 
long, stigma a flat-topped disc. Fruits borne on a flat or 
saucer-like receptacle 4-6 mm broad and only 1-2 mm 
long, abruptly expanded above the short (3-8 mm) thick- 
ened (3-4 mm) pedicel (the thickened pedicel may look 
like a cup in early stages); berry 9-18 mm long, 7-9 mm 
in diameter (dry), ellipsoid to ovoid or oblong. 

Trees of evergreen forest formations of the Ca- 
ribbean slope and partly deciduous formations of 
the General Valley, between 600 and 1000 m el- 
evation in Costa Rica; from near sea level to 1 000 
m elsewhere. Flowers have been collected in May, 
August, and October in Costa Rica and July-Au- 
gust in central Panama; fruits have been collected 
in October-early February. This species ranges 
from Belize and the West Indies to Bolivia and 
the Guianas. 

Ocotea oblonga is recognized by the smaller ob- 
lanceolate leaves with decurrent lamina base, ob- 
scure pubescence, unusual pit domatia (sometimes 
absent), unisexual flowers, and small ellipsoid fruits 
on small flat receptacles. Vegetatively, our collec- 
tions resemble the lowland O. sp. aff. O. bijuga 
and the Monteverde population of O. whitei, but 
those species have very different fruits, and the 
plants are not dioecious. Costa Rican collections 
are from near Volcan Rincon de La Vieja (Guan- 
acaste), the region of Buena Vista de San Carlos 
(Alajuela), Tucurrique and Juan Vinas (Cartago), 
and near the Rio Convento in the General Valley 
(San Jose/Puntarenas). This species is similar to 
O. cuneifolia (Ruiz & Pavon) Mez, O. neesiana 
(Miq.) Kosterm., and O.jlorulenta (Meissn.) Mez, 
of South America. Ocotea eucuneata Lundell with 
slightly thinner and more acuminate leaves may 
be a synonym from Belize. 



Ocotea paulii C. K. Allen, J. Arnold Arbor. 26: 
345. 1945. Figure3. 



Small trees 3-8(-15) m tall, occasionally shrub like 
and somewhat scadent, leafy branchlets 3-8 mm thick, 
glabrous, with 3-5 prominent longitudinal ridges, smooth 
and grayish or brown between the ridges (rarely terete), 
distal stems hollow. Leaves alternate or rarely pseudov- 
erticillate, petioles 6-15(-22) long, 1.5-3(-4) mm thick, 
slightly canaliculate above, glabrous and usually drying 
dark; leaf blades 10-23(-33) cm long, 3-7.5 cm broad, 
narrowly oblong to elliptic-oblong or very narrowly ovate- 
oblong, abruptly short-acuminate at the apex (occasion- 
ally rounded or tapering gradually to a longer acuminate 
apex), acute to obtuse or slightly rounded at the base, 
margin entire and flat or revolute, drying subcoriaceous 
to coriaceous and grayish to reddish brown, glabrous 
above and below, with 5-1 1 major secondary veins on 
each side, major veins usually prominent beneath, cen- 
tral secondaries arising at angles of 35-55, tertiary veins 
often subparallel but not conspicuous. Inflorescences 1 5- 
25 cm long, subterminal or axillary to distal leaves, pan- 
iculate with lateral branches to 5 cm long, many-flow- 
ered, peduncle 5-8 cm long, glabrescent, pedicels 1-2 
mm long. Flowers 2-3 mm long and ca. 3 mm broad, 
glabrous or with a few minute hairs, tepals ca. 1.5 mm 
long and equally broad, ovate; outer stamens with short 
(0.4 mm) filaments, outer anthers 0.7-0.8 mm long, nar- 
rowly ovate with superposed thecae, inner stamens ca. 
1.2 mm long, staminodes absent; pistil 1.5-2 mm long, 
style equalling the ovoid ovary in length, stigma slightly 
discoid. Fruits borne on short (2-4 mm) shallow recep- 
tacles 6-10 mm broad and 1-2 mm deep, margin usually 
entire, pedicels becoming 6 mm long and 2-3 mm broad 
at the apex; berry ca. 2 cm long and 1 cm thick (dry), 
ellipsoid-oblong. 



Small trees and clambering shrubs of evergreen 
montane forest formations, between 600 and 1 600 
(2300) m elevation. Flowers have been collected 
in all months except October-November, but fruits 
have only been collected in March and May. This 
species is only known from the northern part of 
the Meseta Central (San Ramon to Zarcero) and 
the highlands of Chiriqui, Code, and Panama 
provinces in Panama. 

Ocotea paulii is recognized by its usually small 
stature, hollow distal stems, narrowly oblong leaves 
often exceeding 1 5 cm in length, glabrous parts, 
and small narrow fruits borne on small shallow 
receptacles. This species differs from the closely 
related O. atirrensis, O. wedeliana, and O. nicar- 
aguensis in its higher-altitude habitat, smaller, 
more consistently oblong leaves, and more pro- 
fusely branched inflorescences. There appears to 
be intergradation between O. atirrensis and O. 
paulii on the Caribbean slope at about 1000 m 
elevation. The populations of Costa Rica and Pan- 
ama differ somewhat in leaf form and texture. More 
importantly, while Panamanian collections may 
have the upper thecae of the outer anthers reduced 
in size, there are Costa Rican collections in which 



BURGER: FLORA COSTARICENSIS 



91 



the outer anthers may be 2-thecous in some flow- 
ers (.-I. Smith 633, 907, F). 



Ocotea pittieri (Mez) van der Werff, comb. nov. 
Phoebe pittieri Mez, Hot. Jahrb. Syst. 30, Beibl. 
67: 16. 1901. Figure 4. 

Shrubs or small to medium-sized trees, 2-10(-20) m 
tall, leafy branchlets 1.3-4 mm thick, at first grayish or 
yellowish puberulent with short (0.1-0.3 mm) straight 
or crooked hairs, persisting or glabrescent. Leaves alter- 
nate, petioles 5-12 mm long, ca. 1 mm thick, puberulent, 
lateral margins flat above or slightly grooved (not sul- 
cate): leaf blades (2.5-)4-8(-10) cm long, 1.5-4(-5) cm 
broad, narrowly to broadly elliptic, elliptic-oblong or 
slightly obovate, acute to short-acuminate (obtuse in short 
broad leaves) at the apex, acute to obtuse at the base, 
drying stiffly chartaceous, pale brown or occasionally 
dark, usually lustrous and sparsely puberulent on the 
midvein above, very sparsely to densely puberulent be- 
neath with slender or crooked hairs, often with tufts of 
longer (0.6 mm) hairs in the vein axils or along the 
midvein beneath, with 2-6 major secondary veins on 
each side, central secondaries arising at angles of 30- 
60. Inflorescences axillary from distal or older leaves, 
5-9 cm long, paniculate, peduncles usually longer than 
the flowering rachis, pedicels 4-8 mm long, glabrous or 
puberulent. Flowers 3-8 mm broad, tepals 2-3 mm long, 
equal, glabrous or puberulent on the outside, glabrous 
or papillate-puberulent within; outer stamens with ob- 
long anthers 0.7-1 mm long, 0.5-1 mm broad, on fila- 
ments 0.5-1 mm long, inner stamens ca. 1.4 mm long 
with narrow anthers 0.7 mm long, staminodes 0.5-1 mm 
long and slender or capitate (sometimes absent), floral 
tube often with hairs at the edge; pistil 2-3 mm long, 
glabrous, style ca. 1 mm long, stigma simple or lobed. 
Fruits subtended by a shallow cup 10-1 2 mm broad and 
2-3 mm deep, margin entire or with short persisting 
perianth parts, gradually or abruptly expanded above the 
elongated (1-2 cm) thickened (ca. 4 mm) pedicel; berry 
15-30 mm long, 10-20 mm thick, ellipsoid, dark green. 

Shrubs and trees of evergreen montane forest 
formations, between (1000) 1500 and 3200 m el- 
evation. Flowers have been collected in September 
and December-June; fruits have been collected in 
January-August. The species is found in the Cor- 
dillera de Tilaran, along the northern edge of the 
Meseta Central near Palmira, on Volcan Irazu, and 
above Cartago, and from the Cordillera de Tala- 
manca to the Chiriqui highlands of Panama. 

Ocotea pittieri (formerly Phoebe pittieri) is usu- 
ally recognized by the stiff smaller hirsutulous 
leaves, with minor venation raised below, the often 
lustrous upper surfaces, and a tendency to dry yel- 
lowish brown. We interpret this species broadly, 
and in the sense that the name has been used by 
Standley and others (in herb.). Ocotea pittieri has 
not been collected in the area between Volcan Poas 



and the western slopes of Volcan Irazu. This pe- 
culiar pattern is also found in O. mollicella which 
is closely related and easy to confuse with O. pit- 
tieri. The leaves appear to be broader and larger 
in collections from the eastern part of the Cordi- 
llera de Talamanca. Ira amarillo is reported as a 
common name in Costa Rica. 

Staminodes are often poorly developed in this 
species, though they can be well developed in some 
individuals. Leaf texture and venation is very dif- 
ferent from Phoebe cinnamomifolia and its allies. 
Moreover, Oscar Castro of the University of Costa 
Rica has found (pers. comm.) that the chemistry 
of 0. pittieri is like that of some Costa Rican Oco- 
tea spp., and not like that of Phoebe mexicana 
(part of P. cinnamomifolia in a wide sense). Thus 
the correct placement of Ocotea pittieri and its 
close relative O. mollicella is in the genus Ocotea. 
Specimens from Mexico and northern Central 
America placed under the names Phoebe bour- 
geauviana Mez and Phoebe psychotrioides Mez ap- 
pear to be closely related to O. pittieri. 

The type of Ocotea pittieri (Tonduz 11893, pho- 
to of specimen destroyed at B in F, us) has broader 
thinner leaves that have dried darker than is usual 
for this species. This type material resembles O. 
brenesii somewhat, but that species never has such 
narrow leaves, and the leaves usually dry very dark 
and thin. It appears that some newly flushed leaves 
have dried dark and thin (in the material at us), 
but other leaves (as in the photo from B) appear 
more characteristic of the specimens traditionally 
placed under this name in herbaria. The photo, 
we believe, justifies the continued use of this name 
for the material placed here. 



Ocotea pseudopalmana W. Burger, sp. nov. Fig- 
ure 6. 

Arbor usque 30 m alta, ramulis foliiferis 2.6-9 mm 
crassis, ferrugineo-puberulis. Folia alterna, petiolis 6-15 
mm longis, puberulis; laminis 8-16(-21) cm longis et 3- 
9 cm latis, ellipticis vel oblongo-ellipticis, apice obtuso 
vel brevi-acuminato, subtus brunneolo-puberulis, nervis 
secondariis 5-8 paribus. Inflorescentiae 6-1 5 cm longae, 
pauciflorae, pedunculis 4-1 cm longis, puberulis. Flores 
5-7 mm longi, 6-8 mm lati, extus puberuli, stamina ser. 
I-II ca. 1.5 mm longa, antheris ca. 1 mm longis, stam- 
inodiis nullis; gynoecium 2.3-4 mm longum, stylo usque 
2.3 mm longo. Fructus ellipsoideus vel oblongus, 2.2- 
2.8 cm longus et 1.5-2 cm crassus; cupula 11-14 mm 
lata, 3-8 mm longa. 

Trees 4-20(-30) m tall, leafy branchlets 2.6-6(-9) mm 
thick, densely velutinous to tomentulose with short (0.2- 
0.6 mm) yellowish brown or brownish hairs, terete or 



92 



FIELDIANA: BOTANY 



with longitudinal ridges in early stages. Leaves alternate 
in a spiral, petioles 6-15 mm long, 2-3 mm thick, flat 
or slightly sulcate above, densely puberulent; leaf blades 
8-16(-21) cm long, 3-9 cm broad, elliptic to elliptic- 
oblong or slightly obovate, tapering abruptly to a short- 
acuminate or bluntly obtuse apex, obtuse to cuneate at 
the base and often slightly decurrent, margin often slight- 
ly revolute (especially near the base), drying subcoria- 
ceous or coriaceous, puberulent on the major veins above 
but becoming glabrescent and impressed in age, lower 
surface with brown or yellowish brown hairs 0.3-0.5 mm 
long, with 5-8 major secondary veins on each side, the 
central secondaries arising at angles of 45-70, tertiary 
venation usually prominent beneath. Inflorescences sol- 
itary and axillary to leaves, 6-15 cm long, racemose or 
narrowly paniculate and with few (4-12) flowers, pe- 
duncle 4-10 cm long, 1 .2-2 mm thick, densely brownish 
puberulent, pedicels 3-8 mm long. Flowers 5-7 mm long, 
6-8 mm broad, campanulate, densely puberulent on the 
outside, tepals ca. 3.5 mm long and 3 mm broad, drying 
dark and papillate-puberulent within; outer stamens with 
prominent (0.6 mm) broad filaments and oblong anthers 
ca. 1 mm long and 1 mm broad, inner stamens ca. 2 
mm long with glands 0.7-1 mm thick, staminodia ab- 
sent; pistil 2.3-4 mm long, ovary narrowed at the base, 
style 1.5-2.3 mm long, stigma subcapitate. Fruits borne 
on a flat or shallow (0-3 mm) cup 11-14 mm broad, to 
8 mm long and conical or as little as 3 mm long and 
saucer-like, the cup gradually or abruptly expanded above 
the thickened pedicel, red at maturity; berry 2.2-2.8 cm 
long and 1.5-2 cm in diameter (dry), broadly ellipsoid 
or oblong, becoming black. 

TYPE Costa Rica, San Jose Province, east of 
La Chonta, 1 1 May 1969, Roy W. Lent 1679 (ho- 
lotype, F 1749018; negative, 61218; isotype, CR). 

Trees of wet evergreen montane forest forma- 
tions, from 1500 to 2500 m elevation in areas 
subjected to moist winds from the Caribbean and 
adjacent areas. Flowering material has been col- 
lected in February-September; mature fruits have 
been collected in April-October. This species is 
only known from the slopes of Volcan Irazu and 
eastward through the Cordillera de Talamanca as 
far as the border with Panama. 

Ocotea pseudopalmana is recognized by the 
dense pubescence on the underside of the leaves 
and exterior of flowers, the stiff leaves that usually 
dry dark brown above, the few-flowered inflores- 
cences, the larger flower size, and the cloud forest 
habitat. This species is related to O. gomezii with 
much larger flowers, larger more rounded leaves, 
and lower-altitude (800 to 1400 m) habitat. This 
species resembles O. mollifolia, but that species 
has broader thinner leaves, smaller flowers, and 
fruit cups with persisting perianth lobes. This 
species also resembles members of the O. helic- 
terifolia-O. valeriana complex, but those species 
have smaller glabrescent flowers with staminodes 



and deeply cupulate fruiting receptacles. The ma- 
terial placed here resembles Phoebe amplifolia Mez 
& J. D. Smith, from highland Guatemala; but while 
that species has rather similar flowers, the small 
(0.7 mm) sagittate-capitate staminodes are dis- 
tinctive, and the leaves are larger and often with 
long (3 cm) petioles. Some of the collections placed 
here were earlier identified as Ocotea palmana, 
but that species is actually a Pleurothyrium (q.v.). 
Material placed here includes: Almeda & Nakai 
3834 (CAS, CR, F), Brown 17518 (F, LSU), Davidse 
et al. 28550, 28906 (CR, MO), Holdridge6555, 6575 
(CR, NY), Rossbach 3171 (GH), Standley 38633, 
38848 (us), Wilbur & Almeda 16714 (DUKE). 



Ocotea puberula (Rich.) Nees, Syst. Laurin. 472. 
1836. Laurus puberula Rich., Actes Soc. Hist. 
Nat. Paris 1: 108. 1792. O. pyramidata Blake 
ex Brandegee, Univ. Calif. Publ. Bot. 7: 326. 
1920. 

Trees 7-15(-27) m tall, to 50 cm d.b.h., bark pale 
gray, leafy branchlets 1 .7-7 mm thick, flattened or ribbed 
with longitudinal ridges extending downward from the 
leaf bases, glabrous or sparsely and minutely (0. 1 mm) 
puberulent, young stems drying dark, older stems pale 
grayish. Leaves with petioles 10-26(-30) mm long, 0.7- 
2 mm broad, glabrous distally but with minute (0. 1 mm) 
hairs near the base adaxially, distally sulcate above; leaf 
blades 7-2 1(-25) cm long, 2.5-8(-l 2) cm broad, elliptic- 
oblong to narrowly elliptic or narrowly ovate-elliptic, 
tapering gradually to an acuminate apex (or tapering 
abruptly to a short-acuminate apex in broader leaves), 
acute to obtuse at the base and slightly decurrent on the 
petiole, drying stiffly chartaceous and grayish green to 
olive green, dull or lustrous above, essentially glabrous 
above, glabrous beneath or with minute hairs along the 
major veins, with 5-9 major secondary veins, lacking 
domatia, tertiary veins elevated on the dried surfaces 
above and below and forming a weak reticulum of broad 
(1-3 mm) areolae. Inflorescences axillary, 4-7 cm long, 
peduncles 1-8 mm long, rachis minutely puberulent, with 
a few proximal lateral branches to 15mm long, pedicels 
1-3 mm long. Flowers unisexual and dioecious, glabrous 
distally on the outside; male flowers 3-5 mm long, peri- 
anth lobes 2-2.7 mm long, 1.7-2.2 mm broad; outer 
stamens 2-2.5 mm long, filament flattened, anthers ca. 
1.2 mm long and 0.7 mm broad, valves superposed, 
inner stamens slightly shorter, glands 0.7-1 mm wide, 
staminodes absent; pistillode ca. 1.3 mm long and 0.4 
mm in diameter. Female flowers 2-3 mm long, 2.5-3.5 
mm broad, perianth parts ca. 1.5 mm long, nonfunc- 
tional stamens 1-1.2 mm long and with anthers ca. 0.5 
mm long; pistil ca. 2.2 mm long, ovary ca. 0.9 mm in 
diameter, style ca. 0.8 mm long, stigma flat, disclike, and 
drying dark. Fruits borne on a short (2-4 mm) distally 
flattened receptacle 4-8 mm broad at the top (but more 
cuplike in life), with undulate or 6-lobed margin, pedicels 
4-16 mm long; berry 6-8 mm in diameter (dried), glo- 
bose, green becoming black and lustrous. 



BURGER: FLORA COSTARICENSIS 



93 



A species of lowland evergreen and partly de- 
ciduous forest formations. Flowers have been col- 
lected in December-January in Panama; fruits have 
been collected in January-February. Though little 
collected in Central America, this species ranges 
from Mexico to Argentina. 

Ocotea puberula is recognized by the function- 
ally unisexual flowers on unisexual (dioecious) 
trees, small globose fruits subtended by flattened 
receptacles or weakly developed cups, and larger 
elliptic-oblong Nectandra-like leaves. Fruiting 
material closely resembles a number of our species 
ofNectandra, but the inflorescences of O. puberula 
have fewer branches and the cups are not as well 
developed. This species was first collected in Costa 
Rica near Palmar Norte in 1988, after the manu- 
script was completed; it was not possible to include 
this species in the keys. We thank J. Rohwer for 
the determination. 



Ocotea rivularis Standl. & L. O. Williams, Ceiba 
1: 238.1951. Figure 2. 

Medium-sized trees to 1 5 m tall, leafy branchlets 4 
12 mm thick. Leaves alternate, petioles ca. 1 cm long 
and 5 mm thick, sulcate and winged, glabrous; leaf blade 
30-45 cm long, 15-23 cm broad, obovate, abruptly 
rounded near the apex to a bluntly short-acuminate apex, 
tapering gradually to the attenuate base, margin entire 
and slightly revolute, drying stiffly chartaceous, smooth 
and glabrous above, essentially glabrous and reddish 
brown beneath, with 10-14 major secondary veins on 
each side, central secondaries arising at angles of 50- 
65, tertiary venation subparallel but obscure. Inflores- 
cences axillary to distal leaves, 1 8-30 cm long, panicu- 
late, peduncles 7-10 cm long, reddish and 2-3.5 mm 
thick, pedicels 1-2 mm long, minutely puberulent. Flow- 
ers ca. 3 mm long and 3 mm broad, greenish, tepals to 
2 mm long and glabrous within; outer stamens with fil- 
aments almost equalling the length of the anthers, outer 
anthers ca. 0.6 mm long and 0.5 mm broad, staminodes 
absent; pistil ca. 2 mm long with slender style 0.8 mm 
long. Fruits borne in a small (4 mm long and 6 mm 
broad) cupulate receptacle 2-3 mm deep with persisting 
small broad perianth bases; berry ca. 1 2 mm long and 
ellipsoid. 

Trees of the evergreen rain forest formations of 
the Pacific lowlands; flowering in July-early Au- 
gust. This species is endemic to southern Puntar- 
enas Province in the Golfo Dulce area. 

Ocotea rivularis is easily recognized by its very 
large obovate thin-textured leaves, attenuate lam- 
ina base with revolute margins, large inflores- 
cences with small flowers, and small lobed cups 
with ellipsoid fruits. The leaves are essentially gla- 
brous but may have very thin appressed hairs (0.2 



mm long) on the underside which are difficult to 
see. This species is very closely related to O. in- 
sularis, with smaller thicker leaves. Paul Allen 
(1956, pp. 276-277) stated that the species was 
common around the type locality near Esquinas. 
This species is only known from Allen 5590 (F, 
MO, us, the type) and Grayum et al. 4068 (CR, MO), 
from near Rincon. 



Ocotea skutchii C. K. Allen, J. Arnold Arbor. 26: 
352. 1945. Nectandra producta C. K. Allen, loc. 
cit. 397. 1945. O. williamsii P. A. Allen, The 
Rain Forests of Golfo Dulce, 410, pi. 27, 1956, 
non O. williamsii (O. C. Schmidt) Kosterm., J. 
Sci. Res. (Jakarta) 1: 122. 1952. Figure 11. 

Trees to 30 m tall, often with prop roots near the base, 
leafy branchlets 2.5-5 mm thick, minutely (0.1-0.3) pu- 
berulent with brownish or silvery appressed hairs but 
quickly becoming glabrous, terete. Leaves alternate, pet- 
ioles 0.5-5 cm long but difficult to delimit because of 
the decurrent and revolute lamina base, narrowed por- 
tion of the leaf base to 5 cm long, minutely puberulent 
or glabrescent, flat above; leaf blades 8-13 cm long, 2.5- 
5 cm broad, elliptic-obovate to elliptic-oblanceolate or 
narrowly elliptic-oblong, usually short-acuminate at the 
apex, tapering gradually to a cuneate or acute base and 
decurrent on the petiole, margin strongly revolute near 
the base, drying stiffly chartaceous to subcoriaceous, gla- 
brous except for minute hairs above the midvein near 
the petiole, often lustrous above and glaucous beneath, 
minutely and obscurely puberulent beneath with thin 
appressed hairs ca. 0.3 mm long and parallel with the 
secondary veins, with 5-9 major secondary veins on each 
side, central secondaries arising at angles of 35-55. In- 
florescences axillary to distal leaves, 8-18 cm long, open- 
ly branched panicles, peduncles to 6 cm long, ca. 1.5 
mm thick and sparsely appressed puberulent, pedicels 
l-3(-5) mm long. Flowers 2.5-3.5 mm long, 3.5-5.5 mm 
broad, campanulate, tepals ca. 2 mm long and 1.5 mm 
broad, obtuse, densely puberulent on the outside and 
minutely papillate within; outer stamens 0.6-1.3 mm 
long, filaments 0.1-0.7 mm long, outer anthers 0.5-0.8 
mm long and equally broad or broader, thecae super- 
posed or occasionally the lower somewhat lateral to the 
upper (Nectandra-\ike), inner stamens ca. 1.5 mm long 
and with large glands, staminodes not seen; pistil 1.5- 
2.5 mm long, style slender and equalling the ovary in 
length, stigma slightly discoid. Fruits borne in a broadly 
cupulate receptacle ca. 1 cm long and 10-15 mm broad, 
red; berry 3-4 cm long, 13-18 mm in diameter, ellipsoid- 
oblong. 

Trees of evergreen lower montane forest for- 
mations in the central volcanic highlands and the 
Cordillera de Talamanca, from (30) 600 to 1400 m 
elevation. Flowering material has been collected 
in December (Skutch 3062, GH, NY, the type ofO. 
skutchii) and January (Skutch 3906, AA, NY, the 



94 



FIELDIANA: BOTANY 



type of N. productd). Fruits were collected in April 
(Solas 392, usj). This species is only known from 
the central highlands and the General Valley in 
Costa Rica. 

Ocotea skutchii is recognized by the average- 
sized elliptic-obovate leaves with long-decurrent 
and revolute lamina base, the slender appressed 
hairs on the lower leaf surfaces, and the puberulent 
flowers with variable stamens. The leaves are often 
lustrous above and glaucous beneath. This species 
is part of a group of taxa related to O. glaucoser- 
icea, which often become tall trees with basal prop 
roots and have the same characteristic pubescence 
on the lower leaf surfaces in early stages. There is 
the possibility that the plants placed here may in- 
tergrade with O. glaucosericea at slightly higher 
elevations. A few collections of O. austinii (such 
as Solano 3 CR, F) also appear as if they could 
represent intermediates with O. skutchii. There is 
also the possibility that this species intergrades 
with O. whitei of somewhat higher elevations. 
Whether collections which appear intermediate are 
simply extremes of variation within their species, 
or whether this is actually a single polymorphic 
complex of intergrading populations, are ques- 
tions that cannot be resolved without further study 
and more collections. 

Caroline Allen (1966) placed Paul Allen's Oco- 
tea williamsii (1956) into synonymy under O. 
skutchii, and this may be correct. The long narrow 
fruits (4.8 x 2 cm) on small cupular receptacles 
are distinctive. The flowers of the type (Allen 5983, 
us) are abnormally developed, lacking both sta- 
mens and pistils. The tufts of hairs along the mid- 
vein on the lower leaf surface may also be abnor- 
mal. Regardless, the name O. williamsii is a later 
homonym and cannot be used as a specific epithet. 



Ocotea stenoneura Mez & Pittier, Bull. Herb. Bois- 
sier, ser. 2, 3: 233. 1903, emend. C. K. Allen, 
J. Arnold Arbor. 26: 334-335. 1945. 

Trees to 20 m tall, leafy branchlets 3-7 mm thick, 
densely reddish brown tomentulose, becoming grayish 
and terete. Leaves alternate, petioles difficult to delimit 
because of the strongly revolute lamina margin, nar- 
rowed basal portion of the leaf 1.5-3 cm long and 2.5- 
7 mm broad, flat or broadly sulcate above; leaf blades 
ca. 1 8-20 cm long, 9 cm broad, broadest at or near the 
middle, elliptic-oblong, abruptly narrowed at the acute 
to short-acuminate apex, tapering and slightly rounded 
above the revolute base, drying subcoriaceous, glabrous 
above or with short hairs above the midvein and with 
the suhparallcl tertiary venation often slightly raised 
above, densely ferruginous-pilose (or tomentulose?) be- 



neath with appressed hairs 0.1-0.5 mm long, with 10- 
1 2 major secondary veins on each side, the central sec- 
ondaries arising at angles of 35-60. Inflorescences ax- 
illary to distal leaves, somewhat shorter than the leaves, 
peduncles 4-8 cm long to 4 mm thick and densely pu- 
berulent. Flowers unknown, dried stamens (on the fruit- 
ing cupules) resembling those of O. insularis (fide Roh- 
wer, 1986, p. 145). Fruits borne on flat saucer-like 
receptacles, ca. 1 cm broad and 2-3 mm deep, often with 
triangular perianth lobes persisting on the margin; berry 
(perhaps not mature) globose, 10-13 mm in diameter. 

A poorly known species of evergreen forests, 
from near sea level to 1000 m elevation. Collected 
near Las Vueltas, Tucurrique, Cartage Province; 
fruiting in February (Tonduz 13377, photo of B at 
F, the lectotype; C. K. Allen, 1945). Skutch 3014 
(MO, NY), flowering in December in the General 
Valley, is provisionally placed here. The species 
is endemic to Costa Rica. 

Ocotea stenoneura is distinguished by its larger 
puberulent leaves with short strongly revolute 
lamina base, and the small globose fruits on rel- 
atively flat receptacles. Whether this material rep- 
resents a very rare species or aberrant individuals 
of a population placed under another name in this 
treatment is difficult to say. Some specimens ear- 
lier identified as Ocotea stenoneura are now placed 
under O. dentata, a member of the O. insularis 
alliance, with very different fruits and more ob- 
ovate leaves. 



Ocotea tenera Mez & J. D. Smith ex Mez, Bull. 
Herb. Boissier, ser. 2, 3: 234. 1903. Figure 13. 

Shrubs or small trees 3-8(-12) m tall, to 15(-20) cm 
d.b.h., leafy branchlets 1.5-3.8 mm thick, glabrous from 
the earliest stages, drying terete and dark. Leaves alter- 
nate, usually distichous in a single plane and not crowd- 
ed, petioles 6-12 mm long, ca. 1 mm thick, lateral mar- 
gins forming a sulcus adaxially, glabrous; leaf blades 
(5-)7-16(-20) cm long, (1.5-)3-6(-8) cm broad, elliptic 
to elliptic-oblong or ovate-elliptic, tapering gradually to 
the acute or acuminate apex, acute to obtuse at the base, 
usually drying very dark and thin-chartaceous in texture, 
glabrous above and below, minor veins slightly raised 
above, with 4-7 major secondary veins on each side, 
central secondaries arising at angles of 50-70, often 
loop-connected near the margin distally, often with pit 
domatia. Inflorescences 4-8(-12) cm long, axillary or 
extra-axillary on distal stems, paniculate with relatively 
few widely spaced flowers, glabrous, peduncles to 4 cm 
long, pedicels slender, ca. 3.5 mm long. Flowers greenish 
white to yellowish (but drying black), 1.5-3 mm long, 
glabrous, tepals ca. 1.5 mm long; outer stamens 0.6-1.2 
mm long, with broad short filaments, outer anthers ca. 
0.7 mm long, narrowed apically and with superposed 
thecae, staminodes absent or minute, glabrous or with 
a few hairs around the edge of the floral tube within; 



BURGER: FLORA COSTARICENSIS 



95 



pistil 1.5-2.3 mm long, ovary globose or ovoid, style 
usually equalling the ovary in length, stigma simple. Fruits 
borne on the flattened apex of the thickened pedicel, a 
cuplike receptacle absent or shallow (2-3 mm), 1 cm 
broad and with an undulate margin, usually pendant, 
becoming rose red or bright red, thickened portion of 
the pedicel 1-3 cm long; berry 2-3 cm long and 1-2 cm 
thick, ovoid or oblong, becoming purple or black. 

Trees usually found at forest edges and in sec- 
ondary vegetation at lower elevations and in forest 
understory at higher elevation, in evergreen wet 
forest formations of the Caribbean slope and cen- 
tral highlands, between 50 and 1500 (1700) m 
elevation. Flowering collections have been made 
in August-March, and fruits have been collected 
in August-September. This species is endemic to 
Costa Rica, ranging from the Cordillera de Tilaran 
to the valley of the Rio Reventazon and Puerto 
Limon. 

Ocotea tenera is a distinctive species of Laura- 
ceae with its small stature, limited range, thin el- 
liptic leaves that dry very dark, minute flowers 
which almost become black when dry, ellipsoid 
fruits on a flattened receptacle at the apex of a 
thickened pedicel, and the lack of puberulence. 
The species has been reported to be dioecious (C. 
K. Allen, 1945, p. 357) but this may be an error, 
as most plants appear to have bisexual flowers. 
Ocotea tenera is closely related to O. bernoulliana 
Mez of Guatemala, but that species has a more 
deeply cupulate fruiting receptacle and the leaves 
do not become as dark when dry. Ocotea brenesii, 
with broader leaves, larger flowers, and the floral 
cup hairy within, appears to be closely related to 
O. tenera and may be difficult to separate. Ocotea 
cernua and O. meziana are also similar, but their 
leaves do not become so dark when dried. 



Ocotea valeriana (Standl.) W. Burger, comb. nov. 
Phoebe valeriana Standl., Field Mus. Nat. Hist., 
Hot. Ser. 18: 460. 1937. P. smithii C. K. Allen, 
J. Arnold Arbor. 26: 317-318. 1945. Figure 6. 

Trees 3-12(-20) m tall, leafy branchlets 2.5-8 mm 
thick, densely strigulose to tomentulose with yellowish 
brown to dark brown hairs 0.1-0.5 mm long. Leaves 
alternate, differing greatly in size and shape on different 
trees, petioles 4-22(-28) mm long, 1.7-3 mm thick, sul- 
cate above, tomentulose to strigulose; leaf blades 8-25 
cm long, 3.5-14 cm broad, broadly obovate to oblong 
(usually widest above the middle), abruptly narrowed to 
the short-acuminate, obtuse or rounded apex, obtuse to 
rounded (cordulate) at the base, equal or subequal, drying 
subcoriaceous and yellowish brown to dark brown, mi- 
nutely puberulent on the vein above and sparsely pu- 



berulent to glabrous between the veins, dull or lustrous, 
strigulose on the veins beneath with slender hairs ca. 0.3 
mm long, with (4-)5-10 major secondary veins on each 
side, central secondaries arising at angles of 35-60. In- 
florescences 5-25 cm long, distal and with the subtending 
leaves often caducous, peduncles (l-)3-9 cm long, 1-3 
mm thick, densely strigulose, dark or with yellowish 
brown hairs, lateral branches of the rachis with (l-)3-7 
flowers and not further branched, pedicels slender, 2-10 
mm long. Flowers 4-6 mm long and 6-15 mm broad, 
essentially glabrous outside and within, perianth pellucid 
punctate, tepals ca. 3 mm long; outer stamens with an- 
thers 0.9-1. 5(-2) mm long, 1-1.3(-1.7) mm broad, ob- 
long to slightly broader than long, filaments very short, 
staminodes absent or small or up to 1 mm long; pistil 
ca. 2.5 mm long, ovary ca. 1.5 mm in diameter, narrowed 
at the base, style ca. 1 mm long, stigma discoid. Fruits 
borne in a cup 9-14 mm long, 10-18 mm broad, 5-8 
mm deep, cupulate or hemispheric, margin entire or with 
the persisting perianth bases, red; berry ca. 3 cm long 
and 1.8 cm in diameter, obovoid to ellipsoid, obtusely 
apiculate at the apex, black. 

Trees of evergreen montane forest formations, 
from (1000) 1400 to 2200 m elevation. Flowering 
in January-May; fruits have been collected in De- 
cember-February and May-June. The species 
ranges from the Cordillera de Tilaran, along the 
Caribbean escarpment and continental divide of 
the central volcanic highlands as far east as the 
western slopes of the General Valley in the Cor- 
dillera de Talamanca. Endemic to Costa Rica (but 
see below). 

Ocotea valeriana (formerly Phoebe valeriana) is 
a very variable species characterized by the stiffly 
chartaceous to subcoriaceous laminae that are pu- 
berulent beneath and usually broadest above the 
middle. The glabrous flowers drying black, pel- 
lucid-punctate perianth, distinctive puberulent in- 
florescences with lateral branches usually bearing 
long-pediceled flowers (and not further branched), 
and middle elevation habitat further distinguish 
this species. Staminodes may be present or absent, 
but there are rarely three well-developed stami- 
nodes, as in species of Phoebe. Ocotea valeriana 
may prove to be a high elevation derivative of O. 
helicterifolia, but the fruit cups are larger and deep- 
er than in Costa Rican material of that species. 
There are highland specimens from Guatemala 
that are very similar to O. valeriana, but these are 
placed in O. helicterifolia and seem to intergrade 
with more typical lowland forms of O. helicteri- 
folia in Guatemala and Mexico. There is little evi- 
dence for such intergradation in Costa Rica, but 
it may be present (see the discussion under O. 
helicterifolia). 

Phoebe smithii (based on A. Smith P.C. 367, F) 
has very short petioles, short inflorescences, and 



96 



FIELDIANA: BOTANY 



more oblong laminae slightly rounded at the base. 
In addition, the anthers are more often broad (their 
length equaling their width), and the staminodes 
are smaller or more often absent. Specimens here 
placed under Ocotea valeriana, but more like the 
type of P. smithii appear to predominate in col- 
lections made from the Cordillera de Tilaran. 
However, there are many intermediates through- 
out the range of the species, and populations at 
several localities have both the extreme forms and 
the intermediates. 



Ocotea valerioides W. Burger, sp. nov. Figure 2. 

Arbor 5-1 5 m alta, ramulis foliiferis 5-1 2 mm crassis. 
Folia alterna, saepe aggregata. petiolis 4-14 mm longis, 
3-6 mm crassis; laminis 13-38 cm longis et 10-22 cm 
latis, obovatis vel oblongo-obovatis, subtus tomentosis, 
nervis secondariis 8-12 paribus. Inflorescentiae pani- 
culatae, 9-30 cm longae, pedunculis usque 1 8 cm longis. 
Flores 4-5(-7) mm longi et 8-10 mm lati, extus parce 
puberuli, tepalis intus minute papillatis; stamina ser. I- 
II antheris 1-1.2 mm longis, staminodiis nullis. Fructus 
ellipsoideus, 28-36 mm longus. 

Small trees 5-1 5 m tall, to 40 cm d.b.h., leafy branch- 
lets 5-12 mm thick, densely tomentulose with yellowish 
to dark grayish brown hairs to 0.5 mm long. Leaves 
usually clustered at the ends of branchlets, petioles 4- 
14 mm long, 3-6 mm thick, densely tomentulose, broad- 
ly sulcate above; leaf blades 13-38 cm long, 10-22 cm 
broad, obovate to oblong-obovate or slightly pandurate, 
abruptly rounded and short-acuminate at the apex, ta- 
pering gradually to the cuneate base and slightly rounded 
at the petiole, drying stiffly chartaceous, dark brown above 
and slightly lustrous, minutely puberulent above the ma- 
jor veins, lower surface paler in color and with a dense 
tomentum of short (0.3 mm) thin straight hairs slightly 
rough to the touch, with 8-12 major secondary veins on 
each side, central secondaries arising at angles of 40- 
60, tertiary veins prominent beneath and often subpar- 
allel. Inflorescences axillary to distal leaves or scalelike 
undeveloped leaves, racemiform panicles with short lat- 
eral branches to more openly paniculate with spreading 
side branches, (5-)9-30 cm long, primary peduncles to 
18 cm long, 1-2.5 mm thick, densely yellowish brown 
puberulent, pedicels 3-1 1 mm long. Flowers 8-10 mm 
broad, 4-5(-7) mm long, rotate at anthesis, perianth parts 
ca. 4 mm long and 2mm broad, sparsely puberulent on 
the outside and minutely papillate within; outer stamens 
subsessile and sometimes broader than long, outer an- 
thers 1-1.2 mm long and 0.9-1.2 mm broad, thecae 
usually superposed but the lower occasionally lateral (re- 
sembling Nectandra), inner stamens biglandular, stam- 
inodes not seen, the edge of the floral tube with straight 
erect hairs around the base of the stamens; pistil 1.7- 
2.5 mm long, ovary rounded, style slender, 0.7-1 . 1 mm 
long, stigma discoid. Fruits borne on a broad (12-16 
mm), shallow (ca. 5 mm deep) cup gradually expanded 
from a thickened (3-5 mm) pedicel, longitudinal ridges 
sometimes present and giving the margin of the cup an 



undulate appearance, becoming red; fruits 28-36 mm 
long and 14-22 mm in diameter, ellipsoid or slightly 
obovoid. 

TYPE Costa Rica, Alajuela Province, 1 km S 
of Cariblanco along the Sarapiqui road, elevation 
about 850 m, 15 Sept. 1974, Gary S. Hartshorn 
1530 (holotype, CR 71922; isotypes, F [negative, 
61118], MO). 

Trees of the very wet Caribbean slope and low- 
lands, between 40 and 850 m elevation. Flowers 
have been collected in July and September; fruits 
were collected March-May and July. This species 
has only been collected in central Costa Rica (from 
Cariblanco, Alajuela, to between Siquirres and 
Turrialba, Limon) and in the province of Bocas 
del Toro, Panama. 

Ocotea valerioides is recognized by the large ob- 
ovate puberulent leaves with short thick petioles, 
the larger flowers with subsessile anthers, and ra- 
cemiform panicles with densely yellowish brown 
puberulent rachis. This material had been filed 
under Phoebe valeriana (now Ocotea valeriana) in 
herbaria, but O. valerioides has much larger leaves, 
a lower elevation habitat, more puberulent flow- 
ers, and stamens with the anthers often developing 
a Nectandra-like configuration. In fact, this species 
may represent a tendency fully expressed in Nec- 
tandra belizensis, which appears to be related, as 
does O. helicterifolia. All these species are part of 
a complex that deserves serious study. Within our 
material the lower elevation collections have larg- 
er leaves (Gomez & Herrera, 23619, CR, MO; Gra- 
yum et al. 3519, CR, F, MO; Zamora & Elizondo 
674, CR, F), and those around 800 m have some- 
what smaller leaves (Hartshorn 1530, CR; Hold- 
ridge 6779, CR, NY; Poveda & Castro 3920, CR, F). 
The altitudinal separation of O. valerioides and O. 
valeriana is consistent with patterns seen in many 
genera and families; a clinal connection is unlikely. 
There may be a close relationship with O. lentii, 
and the possibility of intermediates with that 
species should be explored. The leaves of a fruiting 
collection from Panama (Kirkbride & Duke 486, 
MO) differs from Costa Rican collections in vesture 
and prominence of the tertiary veins, while another 
collection from Panama (von Wedel 2264, MO) is 
more typical. 

Ocotea veraguensis (Meissn.) Mez, Jahrb. Konigl. 
Bot. Gart. Berlin 5: 240. 1889. Sassafridium 
veraguense Meissn. in DC., Prodr. 1 5, pt. 1 : 171. 
1864. O. paradoxa Mez, Bot. Jahrb. Syst. 30, 



BURGER: FLORA COSTARICENSIS 



97 



Beibl. 67: 16. 1901. O. bakeri Blake, Contrib. 
Gray Herb. 52: 65. 1917. Figure 14. 

Small trees 4-10 m tall, leafy branchlets 1.2-5 mm 
thick, at first minutely puberulent with grayish appressed 
hairs 0.1-0.3 mm long, surfaces becoming grayish or 
dark and smooth to striate. Leaves separate along the 
stem or somewhat clustered distally, petioles (3-)5- 
10(-1 5) mm long, ca. 1 mm thick, lateral margins form- 
ing an adaxial sulcus and the midvein prominent within 
the sulcus; leaf blades 6-14 cm long, 2-5 cm broad, 
elliptic to elliptic-oblong, elliptic-obovate or narrowly 
ovate-lanceolate, obtuse to bluntly acute at the apex 
(sometimes rounded and rarely acute), acute to obtuse 
at the base, drying stiffly chartaceous to subcoriaceous, 
smooth and glabrous above and below (rarely minutely 
puberulent beneath), often drying grayish and slightly 
lustrous above, with 5-8 major secondary veins on each 
side and the central secondaries arising at angles of 30- 
60. Inflorescences axillary to distal leaves or pseudo- 
terminal, 5-16 cm long, basal branches of the inflores- 
cence longer than the distal and forming a rounded pan- 
icle of 8-50 flowers, peduncle 3-9 cm long, very minutely 
puberulent. Flowers 5-10 mm broad, white turning 
brown, with the odor of jasmine, tepals imbricate, to 3 
mm broad and rounded at the apex, minutely papillate 
puberulent; outer anthers 1.5-2.5 mm long and sessile 
or subsessile, ovate-triangular to lanceolate and flat, pap- 
illate puberulent, staminodes inconspicuous or absent; 
pistil 1.5-2 mm long. Fruits borne in a shallow cup 1- 
1.6 cm broad and 1-5 mm deep, usually with an outer 
undulate ridge and an interior entire margin, the cup 
abruptly expanded above the thickened pedicel, red at 
maturity; berry up to 2.5 cm long and 1.8 cm thick, 
ellipsoid, becoming black. 

A species of deciduous and partly deciduous 
forest formations, from near sea level to about 
1200 (1600) m elevation along the Pacific slope 
and Meseta Central. The principal flowering time 
is in December-March, and fruiting is mostly in 
February-May. This species ranges from central 
Mexico to Panama. 

Ocotea veraguensis is recognized by its stiff gla- 
brous leaves that often dry grayish (or yellowish), 
inflorescences with long peduncles, relatively large 
flowers with flattened papillate puberulent outer 
stamens, and the short broad fruit cup with 2- 
ridged margin. The restriction to deciduous or sea- 
sonally very dry forests along the Pacific slope is 
another distinction. The margins of the fruit cups 
resemble those found in Licaria, and collections 
from moister forests, such as on the Osa peninsula, 
resemble L. cufodontisii in their darker more acu- 
minate leaves. 



Ocotea viridiflora Lundell, Wrightia 5: 36. 1974. 

Trees 3-8(-20) m tall, leafy branchlets 1-3 mm thick, 
glabrous, longitudinally ridged in early stages but soon 



becoming terete and dark or grayish. Leaves alternate in 
a spiral, petioles (4-)5-12 mm long, 0.7-1.2 mm thick, 
glabrous, with adaxial ridges forming a deep narrow sul- 
cus above near the stem; leaf blades 3-1 0(-1 2) cm long, 
1-4 cm broad, narrowly elliptic to elliptic-oblong or 
slightly oblanceolate, gradually narrowed to the acumin- 
ate or caudate-acuminate apex, the narrow (2-3 mm) tip 
1-2 cm long on some leaves, acute at the base and slightly 
decurrent on the petiole, drying stiffly chartaceous, gla- 
brous and slightly lustrous above with the venation 
slightly elevated, glabrous beneath, with (3-)4-7 major 
secondary veins on each side, central secondaries arising 
at angles of 30-60, the minor venation raised beneath 
when dry and forming a reticulum of weakly denned 
areolae, distinctive pit domatia often present in the vein 
axils and on the secondary veins beneath. Inflorescences 
axillary to distal leaves, small (2-5 cm) and racemose or 
umbellate (rarely paniculate), peduncles 1-3 cm long and 
ca. 0.5 mm thick, glabrous, pedicels 2.5-7 mm long. 
Flowers 2.5-4 mm long and 3-4 mm broad, glabrous on 
the outside, tepals ca. 2 mm long and glabrous within; 
outer stamens 1.3-2 mm long, outer anthers 0.8-1 mm 
long and 0.6-0.8 mm broad, rectangular with superposed 
thecae, inner stamens ca. 2.2 mm long with large (2.2 
mm) glands and strigulose filaments, staminodes slender 
and small (0.5 mm), slightly strigulose; pistil 2-3 mm 
long, ovary ellipsoid, style half the length of the pistil, 
stigma slightly discoid. Fruits borne in a small deep cup 
ca. 7 mm long and 10 mm broad, perianth parts per- 
sisting or deciduous and the cup entire, red; berry ellip- 
soid to oblong, 12-25 mm long and 6-12 mm in di- 
ameter, green (becoming black?). 



Trees of cloud forest formations at elevations 
of 1400 to 2000 m, along the central Cordilleras 
and in the Chiriqui highlands. Flowers have been 
collected in April-May; fruits have been collected 
in March and May-June. A poorly understood 
species of western Panama, possibly also repre- 
sented in Costa Rica (see below). 

Ocotea viridiflora is recognized by its small nar- 
rowly elliptic glabrous leaves with elevated ve- 
nation (when dry) and distinctive pit domatia, long- 
acuminate leaf tips, small few-flowered glabrous 
inflorescences, and oblong fruits in well-developed 
cups. This species appears to be a close relative of 
the larger-leaved O. meziana, and perhaps O. lae- 
tevirens. The type of O. viridiflora differs from O. 
meziana in having completely glabrous terminal 
buds. Our use of this name for specimens from 
Monteverde may be incorrect: the Panamanian 
populations appear to have lobed fruit cups, while 
the Costa Rican populations appear to have entire 
cups. Nectandra davidsoniana C. K. Allen of the 
Chiriqui highlands is another small-leaved species 
with Ocotea-like stamens, easily confused with this 
species. Nectandra salicina has similarly shaped 
small leaves and lives in similar forests, but lacks 
pit domatia. 



98 



FIELDIANA: BOTANY 



Ocotea wedeliana C. K. Allen, J. Arnold Arbor. 
26: 339. 1945. 

Trees 3-15 m tall with slender (10 cm) trunks, leafy 
branchlets 1.2-4 mm thick, minutely (0.1 mm) ap- 
pressed puberulent and dark but soon becoming glabrous 
and grayish, usually with 5 narrow longitudinal ridges, 
the central part of the stem often with a narrow hollow 
passage. Leaves alternate, petioles 6-12 mm long, 1.4- 
2.5 mm thick, with 2 lateral ridges and usually sulcate 
above, becoming dark and glabrous; leaf blades 1 2-27 
cm long, 4-8(-10) cm broad, narrowly elliptic to nar- 
rowly elliptic-oblong or narrowly elliptic-obovate, ta- 
pering to an obtuse apex with acuminate tip, the slender 
tip 7-20 mm long, tapering gradually to the cuneate or 
acute base, drying thin-chartaceous and often grayish to 
dark brown, glabrous above and the major veins flat or 
slightly impressed, sometimes minutely dark punctate 
above, glabrescent beneath, with 6-9 major secondary 
veins on each side, the central secondaries arising at 
angles of 40-60. Inflorescences axillary to distal leaves, 
10-15 cm long, racemose panicles with short (15 mm) 
few-flowered (3-9) lateral branches, peduncles 7-1 1 cm 
long, ca. 1 mm thick and glabrous, pedicels 2-4.5 mm 
long. Flowers white, 2.5-3.5 mm long, 2.5-3.5 mm broad, 
campanulate, perianth very minutely (0.05-0. 1 mm) pu- 
berulent on the outside, tepals ca. 2 mm long and 2 mm 
broad, broadly ovate and obtuse; outer stamens ca. 1.5 
mm long with puberulent poorly denned filaments, outer 
anthers ca. 1 mm long and 0.8 mm broad, oblong, thecae 
superposed, inner stamens ca. 2 mm long and puberulent 
beneath the anthers, glands sessile and reniform, stam- 
inodes absent; pistil 1.8-2.3 mm long with ovoid ovary 
1 . 1 mm in diameter, style ca. 1 mm long with slightly 
discoid stigma. Fruits borne on an expanded flat or sau- 
cer-like receptacle 1-2 mm long, 5-8 mm broad, and 0- 
2 mm deep, margin undulate, becoming red; berry ob- 
long, 12-20 mm long and 8-11 mm in diameter, be- 
coming bluish green. 



Trees of the evergreen lowland Caribbean (trop- 
ical moist) forest formations. Flowers have been 
collected in May, and fruits have been collected 
in January-March. The species ranges from south- 
ernmost Costa Rica (Grayum & Schatz 5286, CR, 
MO) and adjacent Bocas del Toro Province to cen- 
tral Panama. 

Ocotea wedeliana is recognized by the long thin 
narrowly elliptic-oblong leaves, narrow longitu- 
dinal ribs on distal stems, small oblong fruits, and 
flat or saucer-like cup with undulate margin. It is 
not clear whether the open spaces in the center of 
the twigs are a consistent feature of this species, 
or if ants inhabit these spaces. This species appears 
to be closely related to the larger-leaved O. atir- 
rensis, and O. paulii of higher elevations. The two 
latter species appear to intergrade at about 500 to 
1000 m elevation, and some of those collections 
resemble O. wedeliana quite closely. This species 
is called sigua in Panama. 



Ocotea white! Woodson, Ann. Missouri Bot. Gard. 
24: 188. 1937. Nectandra whitei (Woodson) C. 
K. Allen, J. Arnold Arbor. 26: 398. 1945. O. 
eusericea Lundell, Wrightia 5: 338. 1977. Figure 
11. 

Trees 10-35 m tall, leafy branchlets 2-5 mm thick, 
minutely sericeous at first but soon becoming glabrous 
and dark brown. Leaves alternate, usually clustered at 
the ends of branchlets, petioles poorly differentiated, 4- 
1 8 mm long, sulcate near the base and with lateral mar- 
gins continuous with the lamina margins, narrowed por- 
tion of the leaf to 3 cm long; leaf blades (5-)7-14 cm 
long, (1.5-)2-4.5 cm broad, narrowly obovate to oblan- 
ceolate or elliptic-obovate, short-acuminate to bluntly 
obtuse (rarely rounded in Costa Rica), tapering gradually 
to an attenuate and decurrent base, margin entire and 
conspicuously recurved near the base, drying stiffly char- 
taceous to subcoriaceous, upper surface usually dark and 
dull with minor venation inconspicuous in Costa Rican 
collections (but often lustrous and with the tertiary ve- 
nation slightly raised in Panamanian collections), lower 
surface glabrous or with thin appressed hairs ca. 0.2 mm 
long and usually difficult to see, with 6-12 major sec- 
ondary veins on each side, central secondaries arising at 
angles of 35-50, tertiary venation often slightly ele- 
vated beneath, domatia of tufted hairs or pits very rarely 
present in vein axils beneath. Inflorescences axillary to 
distal leaves, to 1 5 cm long, peduncles to 6 cm long and 
2-3 mm thick, minutely appressed puberulent or gla- 
brous, reddish brown, pedicels 1-2 mm long, puberulent. 
Flowers ca. 3 mm long and 3.5 mm broad, yellowish, 
puberulent on the outside, tepals ca. 2 mm long and 
minutely puberulent within; outer stamens with short 
filaments, outer anthers 0.6-0.9 mm long and 0.7-1 mm 
broad, often slightly longer than broad with thecae su- 
perposed or the lower somewhat lateral (as in Nectan- 
dra), staminodes usually absent (in Costa Rican collec- 
tions); pistil ca. 2 mm long, style slender, ovary ovoid. 
Fruits borne in a cupulate or funnelform receptacle 6- 
12 mm long, 8-12(-14) mm broad, usually shallow (1- 
3 mm) but occasionally deeply cupulate, margin entire 
or undulating, red; berry 15-38 mm long, 10-18 mm 
thick, ellipsoid or oblong, green. 

Trees of wet evergreen cloud forest formations, 
from 1 200 to 2500 m elevation. Flowering collec- 
tions have been made in March-September; fruits 
have been collected in May-September and De- 
cember-January. The species, as here understood, 
ranges from the Cordillera de Tilaran through the 
Cordillera de Talamanca into Panama (but see 
below). 

Ocotea whitei is recognized by the smaller and 
narrow (often oblanceolate) leaves with decurrent 
lamina base and lamina margin usually revolute 
near the petiole, and the ellipsoid fruits borne on 
a small shallow cup. Collections from the area near 
Monteverde placed here have leaves that are often 
long-acuminate and usually dry dark with a flat 
dull upper surface. However, collections near the 



BURGER: FLORA COSTARICENSIS 



99 



border with Panama tend to have lustrous upper 
leaf surfaces and short-acuminate apices. This 
species appears to be very variable in the Chiriqui 
highlands, and the type collection (Seibert 307, AA, 
MO, NY), with leaves drying yellowish brown and 
the slightly raised tertiary venation on the lustrous 
upper surfaces, is very different from Monteverde 
material. Nevertheless, the pattern of variation seen 
in O. whitei in the Chiriqui highlands makes it 
appear reasonable to place the Monteverde col- 
lections under this name; also, the flowers and 
fruits appear to be identical. This species is closely 
related to O. austinii, and specimens which may 
represent intermediates have been collected in the 
Chiriqui highlands. Ocotea skutchii is also closely 
related and there may be intergradation between 
the two around 1000 m elevation. All these species 
are, in turn, related to O. glaucosericea and its 
allies, and they form a complex that is difficult to 
decipher; it may even be a single polymorphic 
entity with local subspecies (but considered as sep- 
arate species here). 



Ocotea sp. A. aff. Ocotea laetevirens Standl. & 
Steyerm. 

Trees 15-25 m tall, leafy branchlets 5-7 mm thick, 
glabrescent, longitudinally ridged but soon becoming ter- 
ete. Leaves alternate, petioles 10-25 mm long, 2-4 mm 
thick, narrowly sulcate above with thin elevated adaxial 
margins (but with rounded or broadly sulcate petioles in 
Standley & Valeria 44651), glabrous; leaf blades 14-30 
cm long, 7-18 cm broad, broadly obovate to broadly 
elliptic-obovate, abruptly narrowed or rounded to the 
bluntly obtuse and slightly decurrent base, margin slight- 
ly revolute near the base, drying stiffly chartaceous or 
subcoriaceous and yellowish brown, glabrous and lus- 
trous above, with the minor venation slightly raised (flat 
in Standley & Valeria 44651), glabrous beneath (but with 
a few hairs in the vein axils in Poveda et al. 3501), with 
4-7 major secondary veins on each side, central second- 
aries arising at angles of 40-60, lower surface with the 
minor venation raised and forming a distinct or some- 
what obscure reticulum. Inflorescences in anthesis not 
seen; infructescences short (6 cm in Standley & Valeria 
44651) to long (22 cm in Allen 1740), the peduncles 
becoming 3-4 mm thick. Fruits borne in pale brown 
gradually expanded conical cups (in Standley & Valeria 
44651) or dark cups abruptly expanded from the base 
(in Allen 1740), 10-15 mm long and 10-20 mm broad 
at the top, 3-7 mm deep, the margins slightly irregular 
with persisting perianth bases; fruits about 10 mm long 
and 10 mm broad (but perhaps not mature) and with 
flat distal surfaces and rugose sides (based on Allen 1740; 
the other collection lacks fruits). 

Trees of evergreen lower montane forest for- 
mations from near Tilaran at about 750 m ele- 



vation (Standley & Valeria 44651, us), near Tur- 
rialba (Poveda et al. 3501, CR, F), and the north 
rim of El Valle de Anton, between 600 and 1000 
m elevation in Code Province, Panama (Allen 
1740, F, MO). Fruiting material was collected in 
January at Tilaran and March in Panama. 

Ocotea sp. A (aff. O. laetevirens) is distinguished 
by its larger glabrous obovate leaves with reticu- 
late minor venation and slightly decurrent lamina 
base, and the deep, slightly warty, fruit cups with 
irregular rims. The fruits of the Allen collection 
differ from all our other Lauraceae with their lus- 
trous flattened tops, but this may be an artifact of 
immaturity (these flattened tops do not exceed the 
rims of the cups). The specimens placed here may 
not be conspecific; only more and better collec- 
tions can help resolve the status and placement of 
these collections. Several aspects of leaf structure 
and the fruit cup suggest a relationship with O. 
laetevirens. A collection with immature fruits from 
Rincon de Osa (Liesner 2091, CR, MO) may also 
belong here. Grayum et al. 7859 (CR, MO) from the 
Atlantic slope of Volcan Barva at 1 200 m and with 
fruits 4.5 x 2.5 cm is provisionally placed here. 



Ocotea sp. B. 

Trees, ca. 15m tall, trunk with broad-based prickles 
(projections) 1-3 cm long, leafy internodes l-2(-4) mm 
thick, minutely (0. 1 mm) puberulent (juvenile) or gla- 
brescent, becoming dark reddish brown. Leaves alter- 
nate, petioles 4-7 mm long; leaf blades 6-10 cm long, 
(1.5-)2-3 cm broad, elliptic-oblanceolate, bluntly short- 
acuminate at the apex, tapering gradually to the decur- 
rent base, drying chartaceous, glabrescent above, sparse- 
ly and minutely puberulent along the midvein beneath, 
with 10-14 obscure secondary veins on each side, the 
distal secondaries loop-connected near the margin, dom- 
atia absent. Inflorescences solitary and axillary, 5-10 cm 
long, paniculate, peduncles 3-6 cm long, subglabrous, 
distal flowers in cymose pairs. Flowers 2-3 mm long, 3 
mm broad, puberulent on the outside, tepals 1 .2-1 .4 mm 
long; outer stamens ca. 0.6 mm long, glabrous, outer 
anthers 0.4 mm long and 0.5 mm broad, thecae super- 
posed, inner stamens ca. 0.8 mm long with glands 0.3 
mm thick, staminodes absent; pistil 1.5 mm long, with 
a short (0.5 mm) style and simple stigma. Fruits un- 
known. 

Ocotea sp. B is distinguished by the Zanthox- 
ylum-\ike projections on the trunk (absent on 
younger individuals). The smaller subglabrous el- 
liptic-oblanceolate leaves with decurrent bases and 
small paired flowers are also unusual. We have 
only two collections from near the Marenco Bio- 
logical Station on the Osa Peninsula: Burger et al. 
12376 (juvenile foliage) and 12377 with a few 



100 



FIELDIANA: BOTANY 



flowers in February 1 988. More material is needed 
for a formal description and to determine possible 
relationships. 



Persea Miller 

REFERENCES L. E. Kopp, A taxonomic revision 
of the genus Persea in the Western Hemisphere 
(Perseae-Lauraceae). Mem. New York Bot. Card. 
1 4( 1 ): 1- 1 20. 1 966. L. O. Williams, The avocados, 
a synopsis of the genus Persea, subg. Persea. Econ. 
Bot. 31: 315-320. 1977. 

Trees or shrubs, bisexual. Leaves alternate or rarely 
verticillate, petiolate, the leaf blades entire and pinnately 
veined. Inflorescences axillary or pseudoterminal, soli- 
tary or in fascicles, paniculate to racemose (rarely cap- 
itate), with persistent or deciduous bracts, puberulent 
and usually with many flowers, pedicels usually present. 
Flowers bisexual, perianth of 6 parts (tepals) in 2 whorls 
(series), campanulate to rotate, the outer tepals usually 
shorter than the inner and glabrous on the inner (adaxial) 
surface as in subgenus Eriodaphne or the outer tepals 
are equal to the inner tepals and puberulent on their 
inner surface in subgenus Persea, tepals often persisting 
in fruits, a floral cup or tube absent or only slightly 
developed; androecium of 9 stamens and 3 staminodes, 
stamens of the outer whorls (series I-II) introrse and with 
long filaments (in our species), outer anthers 4-thecous 
(rarely 2-thecous), the thecae superposed in 2 planes (in 
Central America), inner stamens (series III) with prom- 
inent filaments and each with 2 stipitate or sessile glands 
near the base, with 4-thecous anthers dehiscing laterally, 
lateral-extrorse or the upper thecae lateral and the lower 
extrorse (rarely 2-thecous or the inner series nonfunc- 
tional), staminodes (series IV) usually large and con- 



spicuous with sagittate-acute apices; pistil glabrous or 
puberulent, ovary globose to ellipsoid (occasionally stip- 
itate), the style slender and longer than the ovary, stigma 
small to discoid or triangular-peltate. Fruits borne on an 
enlarged woody or succulent pedicel, the receptacle not 
forming an enlarged cup, perianth parts often persisting 
but not enlarging beneath the fruits; berry globose to 
ellipsoid or pyriform, 1-5 cm in diameter (to 15 cm in 
P. americana). 

Persea is a genus with over 200 named species, 
largely in the Western Hemisphere (ca. 80 spp.), 
with additional species in India, Southeast Asia, 
and outliers in Australia, the Mascarene Islands, 
the Azores, and the Canary Islands. Costa Rican 
members of this genus are recognized by the larger 
flowers (for Lauraceae) with anthers borne on well- 
developed filaments, the large staminodia, the 
perianth often persisting at the base of the fruits, 
and the lack of cupulate or broadly flattened re- 
ceptacles beneath the mature fruits. In addition, 
most Costa Rican species of Persea tend to have 
longer petioles than other Costa Rican species of 
Lauraceae, and the minor venation usually forms 
a visible reticulum (sometimes raised) over the 
surface of the dried lamina. Despite the distinc- 
tiveness of the genus, there may be species that 
intergrade with Phoebe in Central America. It 
should be noted that Kostermans (1957) suggested 
that some species of Phoebe should be placed in 
Persea. Some of our species of Beilschmiedia re- 
semble Persea in fruit and vegetative character- 
istics, but they have 2-thecous anthers. 



Key to Species of Persea 

la. Leaves often subopposite and crowded near the ends of branchlets, stems often with broadly 
thickened nodes where the leaf clusters were borne and from which multiple branches emerge; 

leaves glabrous and elliptic-oblong; tepals subequal 2a 

Ib. Leaves rarely subopposite or crowded at a thickened distal node, stems rarely with larger nodes 

from which multiple branches emerge; leaves glabrous or puberulent 3a 

2a. Flower buds densely whitish tomentulose; fruits said to be broader than long and reniform 

P. rigens 

2b. Flower buds sparsely puberulent; fruits unknown P. silvatica 

3a. Fruits often exceeding 3 cm in length, not subtended by persisting tepals; leaves usually drying 
chartaceous, often broadly ovate or obovate; often puberulent beneath with spreading or erect 

hairs; tepals subequal 4a 

3b. Fruits rarely exceeding 3 cm in length, usually subtended by the persisting tepals; leaves usually 
drying subcoriaceous to coriaceous, with appressed parallel hairs, spreading hairs or glabrous; 

tepals unequal with the outer clearly shorter than the inner (subequal in P. albida) 5a 

4a. Flower pedicels 2-5 mm long; leaves broadly ovate to broadly obovate, abruptly rounded 
and short-acuminate at the apex, acute to obtuse at the base, sparsely puberulent beneath; 
0-2500 m . . .P. americana 



BURGER: FLORA COSTARICENSIS 



101 



4b. Flower pedicels 10-25 mm long; leaves very broadly elliptic to broadly obovate, rounded at 
the apex and without a tip, obtuse to subtruncate at the base, densely puberulent beneath; 

1000-2600 m P. schiedeana 

5a. Leaves 3-9 cm long and 1-3 cm broad, with a dense indumentum of sericeous parallel hairs 
beneath; fruits globose and 7 mm in diameter; rarely collected trees of the Pacific slope, ca. 1 200 

m P. brenesii 

5b. Leaves larger, or if of similar size highland species 6a 

6a. Trees usually found only above 2000 m elevation; leaves often small (less than 10 cm long) and 

very coriaceous; fruits ca. 1 cm in diameter 7a 

6b. Trees rarely found above 2000 m elevation (except in P. veraguasensis); leaves rarely small and 

coriaceous 8a 

7a. Leaves ferrugineous puberulent (at least in early stages), variable in size and form but often 

ovate; branchlets becoming black, 3-9 mm thick; high Cordilleras P. vesticula 

7b. Leaves glabrous or minutely villous; elliptic-oblong to oblong-obovate; branchlets very dark 

reddish brown, 2-5 mm thick; Chiriqui highlands and adjacent Costa Rica . . P. obtusifolia 

8a. Leaves lanceolate to oblong or narrowly ovate, tapering gradually to the acute or bluntly obtuse 

apex, usually drying yellowish or pale brown; fruits 5-12 mm in diameter; 300-2300 m elevation 

9a 

8b. Leaves broadly elliptic to oblong-obovate, rounded to bluntly obtuse or short-acuminate at the 
apex, usually drying dark brown; fruits 12-17 mm in diameter; rarely collected, around 1000 m 

elevation lOa 

9a. Flowers with outer tepals 2-2.5 mm long and inner tepals 3-5 mm long, tips of the inner 
tepals persisting and the fruits subtended by the unequal parts; leaves with 5-10 pairs of 

major secondary veins P. veraguasensis 

9b. Flowers with outer tepals 1-2 mm long and inner tepals 4-6 mm long; tips of the inner tepals 
breaking off and the fruits subtended by subequal parts; leaves with 8-12 pairs of major 

secondary veins P. caerulea 

lOa. Leaves elliptic to elliptic-oblong, often slightly glaucous beneath, flat between the secondaries and 
the tertiaries not raised beneath, with (4-)5-8 pairs of major secondary veins; outer tepals puber- 
ulent on their inner surfaces P. albida 

lOb. Leaves broadly elliptic to oblong-obovate, the tertiary veins raised on the lower surface, never 
glaucous, with 5-13 pairs of major secondary veins; outer tepals glabrous on their inner surfaces 
P. povedae 



Persea albida Kosterm., Reinwardtia 7: 51. 1969, 
nomen novum for P. pallida Mez & Pittier, Bull. 
Herb. Boissier ser 2, 3: 23 1 . 1 903, non P. pallida 
(Nees) Oliver, 1880. Figure 15. 

Trees, ca. 1 5 m tall, leafy branchlets 3-6 mm thick, 
minutely appressed puberulent (sericeous to strigulose), 
becoming (sub)glabrous and dark in color, longitudinally 
striate. Leaves alternate (rarely subopposite), petioles 1.5- 
3.5 cm long, 1-2 mm thick, dark and glabrous, slightly 
sulcatc above; leaf blades 7-18 cm long, (3-)4-8 cm 
broad, elliptic to elliptic-oblong, short acuminate to 
bluntly obtuse or rounded at the apex, obtuse to acute 
at the base, margin becoming revolute when dry, drying 
stiffly chartaceous to subcoriaceous and often dark brown, 
upper surface glabrous, slightly lustrous and with a fine 
(0.2-0.5 mm) reticulum of raised minor venation, sparsely 
and minutely (0.2-0.4 mm) puberulent beneath, often 
somewhat glaucous beneath becoming glabrescent, with 
(4-)5-8 pairs of major secondary veins on each side, 
central secondaries arising at angles of 40-60, a retic- 
ulum of small (0.2-0.5 mm) areolae formed by the raised 
minor venation beneath. Inflorescences 10-15 cm long, 



in axils of distal leaves, primary peduncles 5-9 cm long 
and more than half the length of the inflorescence, sparse- 
ly puberulent. Flowers yellowish, ca. 6-8 mm long and 
4-6 mm in diameter, perianth densely and minutely se- 
riceous on the outer surfaces, both outer and inner whorls 
of tepals puberulent within, the whorls subequal; outer 
stamens with narrow anthers ca. 1.7 mm long, on short 
puberulent filaments, inner stamens 3 mm long with 
sessile glands, staminodes 1.2-1.6 mm long with a large 
(1-1.3 mm) glabrous triangular-sagittate apex; pistil 3- 
5 mm long with a slender style 2-3 mm long, ovary 
glabrous, stigma discoid. Fruits subtended by the per- 
sisting campanulate perianth parts, outer tepals ca. 5 mm 
long and 5 mm broad at the base, inner parts ca. 6 mm 
long and 4-5 mm broad; berry globose, 10-15 mm in 
diameter. 



Trees of lower montane evergreen forest for- 
mations, at about 1000 to 1500 m elevation on 
the Pacific slope of southernmost Costa Rica (but 
see below). Immature flowers were collected in 
February (Pittier 11111, CR, us, the type from Valle 



102 



FIELDIANA: BOTANY 



de Goto), mature flowers were collected in March 
(Burger et al. 12182, CR, F, MO, NY, at Las Alturas 
de Goto Brus), and fruits were collected in August 
(Raven 21891, CR, F, south of San Vito de Goto 
Brus). The species is known from only these col- 
lections and a recent collection from highland (2000 
m) Honduras (Keyser 1397, EAP, F). 

Persea albida is recognized by the slender long 
dark petioles, elliptic-oblong leaves minutely pu- 
berulent and often glaucous beneath, and with a 
fine reticulum of raised minor venation on both 
surfaces. The slightly unequal perianth parts pu- 
berulent on all surfaces, prominent sagittate stam- 
inodes, and smaller globose fruits also help distin- 
guish this species. An additional sterile collection 
(Standley 51268, us) from El Muneco in Cartago 
Province at 1400 m elevation may be this species. 
Persea albida is similar to P. povedae. 



Persea americana Miller, Card. Diet. ed. 8. (with- 
out pagination) 1768. Laurus persea L., Sp. PI. 
370. 1753. P. drymifolia Schlechtend. & Cham., 
Linnaea 6: 365. 1831. Figure 15. 

Small to large trees 5-40 m tall, often with a dense 
rounded crown, leafy branchlets 1 . 5-6 mm thick, densely 
to sparsely puberulent with slender pale brownish hairs 
0.1-0.4 mm long, remaining puberulent or glabrescent. 
Leaves alternate or subopposite, petioles 1-6 cm long, 
1-2.5 mm thick, sulcate above, glabrous or puberulent; 
leaf blades 6-22(-30) cm long, 3-1 4(-l 9) cm broad, nar- 
rowly to broadly ovate or ovate-elliptic, sometimes ob- 
ovate or suborbicular, usually short-acuminate at the 
apex (acute to obtuse), acute to obtuse or rounded and 
often slightly asymmetric at the base, margin entire, drying 
chartaceous, smooth and glabrescent above, sparsely to 
moderately puberulent beneath with short (0. 1-0.4 mm) 
slender hairs, with 5-9 major secondary veins on each 
side, central secondaries arising at angles of 30-50, ter- 
tiary veins often raised on both surfaces when dry. In- 
florescences axillary to distal leaves or in terminal clus- 
ters, 4-12 cm long, compact or loosely branched panicles, 
peduncles 1-7 cm long, slender and puberulent, pedicels 
2-5 mm long, yellowish brown puberulent. Flowers 5- 
8 mm long, campanulate, outer tepals 4-6 mm long and 
1-3 mm broad, ovate-elliptic to oblong, acute at the 
apex, inner whorl 4-7 mm long, equalling or slightly 
longer than the outer, both whorls yellowish brown to- 
mentulose on inner and outer surfaces; outer stamens 
3.5-4.5 mm long, outer anthers 1.2-1.5 mm long and 
0.9 mm broad, the 4 thecae superposed, inner stamens 
with slender filaments and stalked glands, staminodes 
2-3 mm long and broadly or narrowly sagittate at the 
apex; pistil 3-4 mm long, puberulent with a long slender 
style, stigma simple. Fruits borne on a thick (3-5 mm) 
stalk not usually swollen near the apex, perianth decid- 
uous and leaving a rim ca. 6 m broad; berry 5-15 cm 
long, globose to pyriform or obovoid, dark green, seed 
2-5 cm in diameter. 



Trees widely cultivated and growing wild in both 
evergreen and partly deciduous forest formations, 
from near sea level to about 2500 m elevation 
throughout Costa Rica and on Cocos Island. Flow- 
ering collections have been made primarily in Jan- 
uary-March, with a few in August-September. 
Fruiting specimens have been collected in Feb- 
ruary, May, August, and November. This species 
is now widely cultivated throughout the tropics 
and subtropics of the world. 

Persea americana is recognized by its large dark 
green fruits on a terete pedicel (lacking both a fruit 
cup and a persisting perianth), the puberulent yel- 
lowish flowers, and the usually thin ovate leaves 
on long petioles, with the upper leaf surfaces dark 
lustrous green and the underside very pale green. 
The species is best interpreted in a wide sense; 
however, the considerable range of diversity often 
makes the identification of individual sterile col- 
lections difficult. Variation within the species in 
Central America probably reflects a long history 
of cultivation in this region. The fruits of this 
species may be the most nutritious of all the cul- 
tivated fleshy fruits. The high food quality of the 
avocado is probably due to coevolution with birds 
that are fruit-eating specialists and depend on these 
fruits for nearly all their nutrition. Common names 
are abacate, aguacate, aguacatillo, and cura agua- 
cate. English names include alligator pear, avo- 
cado, and butter pear. A large number of names 
in native Amerindian languages attest to the an- 
tiquity of cultivation in Mesoamerica. For a nar- 
rower interpretation of this species see Williams 
(1977). 



Persea americana var. nubigena (L. O. Williams.) 
Kopp, J. Arnold Arbor. 14: 19. 1966. P. nubi- 
gena L. O. Williams, Ceiba 1: 55. 1950. P. gi- 
gantea L. O. Williams, Ceiba 4: 39. 1953. 

These trees of montane cloud forests reach 40 
m in height. Their leaves are more coarsely tex- 
tured with more prominent venation, and they are 
more puberulent than typical P. americana. Also, 
the leaves tend to be broadly ovate or suborbi- 
cular. If it were not for the great variability within 
P. americana, these highland populations might 
be worthy of specific status, but there do appear 
to be intermediates between typical variety amer- 
icana and the more distinctive specimens of va- 
riety nubigena. Unfortunately, some specimens 
placed here seem to be intermediate with P. schie- 
deana, and it is possible that there has been gene 



BURGER: FLORA COSTARICENSIS 



103 



flow between P. schiedeana and P. americana at 
higher elevations. 

Two unusual collections with ovate leaves 
somewhat intermediate between P. americana and 
P. veraguasensis on long (1 5-45 mm) petioles and 
with globose fruits 3 cm in diameter are provi- 
sionally placed here (Davidse et al. 24447, 28401, 
CR, MO, from 1 500 to 1 600 m near the valley of 
the Rio Colon in southern Puntarenas Province). 
They may represent a new species. 



Persea brenesii Standley, Publ. Field Mus. Nat. 
Hist, Bot. Ser. 18: 458. 1937. 

Trees 5-9 m tall, leafy branchlets 3-5 mm thick, densely 
yellowish brown sericeous with straight slender ascend- 
ing hairs 0. 1-0.3 mm long, becoming dark brown in age. 
Leaves alternate and somewhat crowded on short distal 
twigs, petioles 7-16 mm long, ca. 2 mm broad, with 
lateral margins but without a sulcus above; leaf blades 
(3-)5-9 cm long, l-2.4(-3) cm broad, elliptic to elliptic- 
oblong, acute to obtuse at the apex, cuneate at the base, 
margins entire and slightly involute, drying subcoria- 
ceous, sparsely puberulent above, the lower surface 
densely grayish brown or yellowish brown sericeous with 
slender parallel hairs ca. 0.4 mm long, with (3-)4-7 ma- 
jor secondary veins on each side, central secondaries 
arising at angles of 20-40. Inflorescences axillary to 
distal leaves, to 15 cm long, paniculate, peduncles 3-7 
cm long, sericeous, pedicels ca. 2 mm long. Flowers ca. 

6 mm long and 6 mm broad, the outer tepals ca. 2 mm 
long and rounded at the apex, densely sericeous on the 
outside but glabrous within, inner tepals ca. 4 mm long 
and acute at the apex, puberulent on both surfaces; outer 
stamens ca. 2.8 mm long with narrow oblong anthers ca. 
1 .4 mm long, filaments puberulent, inner stamens ca. 3 
mm long, staminodes ca. 1.5 mm long and puberulent; 
pistil ca. 4 mm long, glabrous, ovary ca. 1.3 mm long, 
stigma discoid. Fruits subtended by the persisting peri- 
anth with the outer series ca. 3 mm long and the inner 
series ca. 5.5 mm long; berry apparently globose and ca. 

7 mm in diameter (dry and perhaps immature). 

Trees of wet evergreen forest formations of the 
northwestern edge of the Meseta Central. The 
species is known only from near La Palma de San 
Ramon, Alajuela, at 1100 to 1200 m elevation. 
Flowers were collected on 27 June 1973 (Primack 
et al. 254, DUKE, F), and immature fruits were 
collected on 29 September 1925 (Brenes 4451, CR, 
F, the type). 

Persea brenesii is recognized by the small stiff 
elliptic leaves densely sericeous beneath, flowers 
with perianth whorls of very different length and 
densely sericeous, and the small globose fruits sub- 
tended by persisting perianth parts. The hairs on 
the lower leaf surfaces are characteristic; they are 
slender, straight, and mostly paralleling the sec- 



ondary veins. There is a possibility that the ma- 
terial placed here is an unusual form of P. vera- 
guasensis, but the small leaves on relatively short 
petioles and the unusual pubescence of lower leaf 
surfaces are distinctive. 



Persea caerulea (Ruiz & Pa von) Mez, Jahrb. Kon- 
igl. Bot. Gart. Berlin 5: 171. 1889. Laurus cae- 
rulea Ruiz & Pavon, Laurografia Peruviana t. 
2, 71802. P. laevigata H.B.K., Nov. gen. sp. 2: 
157. 1817. P. petiolaris H.B.K., loc. cit. 159. 
1817. P. skutchii C. K. Allen, J. Arnold Arbor. 
26: 298. 1945. Figure 9. 

Small to medium-sized trees, 4-25 m tall, leafy 
branchlets 1.2-5 mm thick, at first with yellowish or 
reddish brown hairs 0.1-0.3 mm long, becoming dark 
in color and glabrescent. Leaves alternate, petioles 1.5- 
5(-7) cm long, ca. 1.5 mm thick, dark brown and sparsely 
strigulose; leaf blades 7-15(-24) cm long, 3-7 (-12) cm 
wide, ovate-lanceolate to ovate-elliptic, ovate-oblong or 
elliptic-oblong, tapering gradually to the acute or bluntly 
obtuse apex, acute to obtuse or rounded and subtruncate 
(in larger leaves) at the base, slightly unequal at the base, 
drying stiffly chartaceous and often very pale yellowish 
green in color, glabrous above and very sparsely stri- 
gulose beneath, with 8-12 major secondary veins on each 
side, central secondaries arising at angles of 40-70, ter- 
tiary venation obscure or forming very small (0.3 mm) 
areolae. Inflorescences axillary to distal leaves (occa- 
sionally from older nodes), 5-12 cm long, paniculate, 
peduncle more than half the length, 4-12 cm long and 
ca. 1 .2 mm thick, sparsely to densely pale yellowish brown 
strigulose, pedicels 3-5 mm long. Flowers 6-7 mm long, 
to 15 mm broad, outer tepals 1-2 mm long, inner tepals 
4.5-6.5 mm long, yellowish brown puberulent on the 
outside and glabrous within; outer stamens with anthers 
1.5-2 mm long on filaments 1.5-3(-4) mm long, inner 
stamens with subsessile glands, staminodia sagittate and 
puberulent, 2-3 mm long; pistil ca. 3.5 mm long, style 
longer than the ovary and slender, stigma simple or cap- 
itate. Fruits subtended by persisting perianth parts of 
about equal length (because the distal ends of the longer 
inner tepals break off), ca. 2 mm long; berry 5-9 mm in 
diameter (dried), globose or subglobose, glaucous (gray- 
ish blue) and subtended by a reddish pedicel, glabrous. 

Trees of evergreen and partly deciduous forest 
formations of the Pacific slope in Costa Rica, be- 
tween (300) 500 and 1500 (2000) m elevation. 
Flowering collections have been made mostly in 
February-May and July-August; a collection in 
November was made in Honduras. Fruits have 
been collected in April-August. The species ranges 
from Honduras through Central America to Co- 
lombia and Venezuela and southward along the 
Andes to Bolivia. 

Persea caerulea is recognized in the dried con- 
dition by its pale colored or orange brown gla- 



104 



FIELDIANA: BOTANY 



brescent leaves on long slender petioles. The peri- 
anth, with much longer inner tepals that break off 
near the ends in fruiting stages, makes both the 
flowers and the fruits distinctive. The species was 
common in the seasonally very dry Meseta Cen- 
tral; at Monteverde it is only found in the drier 
forest below 1400 m elevation. The trees often 
grow along streams and are referred to under the 
general name aguacatillo. This species resembles 
P. veraguasensis and may be difficult to separate 
in the absence of flowers or fruits. 



Persea donnell-smithii Mez, Arbeiten Konigl. Bot. 
Gart. Breslau 1: 113. 1892. 

This is a name that has been applied to a few 
collections from Costa Rica (Holdridge 6641, 6874, 
CR) with large thin obovate leaves, glaucous-gray- 
ish beneath, and with long petioles. They are sterile 
and probably unusual individuals of P. americana. 
Persea donnell-smithii is a species ranging from 
southern Mexico to Nicaragua. 



Persea obtusifolia Kopp, Mem. New York Bot. 
Gard. 14: 81. 1966. Figure 5. 

Shrubs or small trees 0.5-3(-6) m tall, leafy branchlets 
2-4 mm thick, sparsely puberulent with thin ascending 
whitish or yellowish hairs ca. 0.5 mm long (rarely gla- 
brous). Leaves alternate or subopposite, usually clustered 
at the ends of twigs, petioles 3-12 mm long, ca. 2 mm 
thick, sulcate above with 2 adaxial ridges, usually pu- 
berulent with minute (0.1 mm) or small (0.5 mm) thin 
hairs (especially at the base), rarely glabrous; leaf blades 
3.5-9.5 cm long, 2-4 cm broad, elliptic-oblong to ob- 
long-obovate or obovate, bluntly obtuse to rounded at 
the apex, rounded to cuneate at the base, margin revolute 
when dry, leaves drying subcoriaceous to coriaceous and 
yellowish green to pale brown, glabrous above and with 
a very fine (0.2-0.5 mm) reticulum of minor venation, 
minutely villous to glabrous beneath, with 4-8 major 
secondary veins on each side, central secondaries arising 
at angles of (30-)40-70, a fine (0.2-0.5 mm) reticulum 
of minor venation also present on the lower surface. 
Inflorescences axillary to distal leaves, 3-7 cm long, pe- 
duncles 2.5-5 cm long, ca. 1.5 mm thick, villous or 
glabrous, flowers few in cymose groups, pedicels 2-4 mm 
long, densely sericeous. Flowers 4-6 mm long, 3-4 mm 
broad, perianth erect and densely sericeous with yellow- 
ish or ferrugineous hairs on the outside, outer tepals to 
4 mm long and glabrous within, inner tepals to 6 mm 
long and 3 mm broad, puberulent within; outer stamens 
to 4 mm long with prominent hirsute filaments ca. 2.5 
mm long, anthers narrow and ca. 1.5 mm long, inner 
stamens ca. 2.8 mm long with subsessile glands on the 
lower part of the filament, staminodes small (0.5 mm) 
and linear; pistil 3-4 mm long and glabrous, with a slen- 
der style ca. 2 mm long, stigma capitate or simple. Fruits 



subtended by the persisting, erect or spreading tepals, 
the outer ca. 3 mm long and the inner ca. 4 mm long; 
berry globose, ca. 1 cm in diameter, slightly apiculate at 
the apex, green becoming black (not glaucous). 

Small trees and shrubs of high montane elfin 
forest formations and in drier sites at high ele- 
vation, from (1800) 2000 to 3000 m elevation. 
Flowers have been collected in July-August; fruits 
have been collected in November and March. This 
species is only known from easternmost Costa Rica 
in the Cordillera de Talamanca, and in the Chi- 
riqui highlands of Panama. 

Persea obtusifolia is recognized by the small stiff 
leaves often rounded at the apex, the short few- 
flowered inflorescences with long peduncles, 
densely sericeous perianth, and the small globose 
fruits subtended by the persisting perianth parts. 
Both surfaces of the dried leaves display a fine 
reticulum of raised minor venation. This species 
resembles small-leaved specimens of P. vesticula, 
but P. obtusifolia differs in leaf shape and fruiting 
perianth; the two species are closely related. A 
collection by L. D. Gomez et al. (21654, CR, MO) 
is placed here and may represent an ecotype or 
perhaps even a hybrid with Ocotea whitei. How- 
ever, its Ocotea-like flowers may not be mature, 
and its narrow, completely glabrous leaves have 
the reticulation characteristic of P. obtusifolia. The 
only other Costa Rican collection is Davidse et al. 
25542 (CR, MO). 

Persea povedae W. Burger, sp. nov. Figure 1 2. 

Arbor usque 1 2 m alta, ramulis foliiferis 4-9 mm eras- 
sis. Folia alterna vel subverticillata, petiolis 1 5-40 mm 
longis; laminis (9-)l 1-23 cm longis et (4-)6-10 cm latis, 
late ellipticis, elliptico-obovatis vel oblongo-obovatis, 
apice obtuse vel rotundato, glabris, nervis secondariis 
(5-)7-l 3 paribus. Inflorescentiae fructiferae usque 1 5 cm 
longae. Flores ignoti per anthesin; tepala externa brevia 
et intus glabra. Fructus globosus, 1.3-1.7 cm in dia- 
metro. 

Trees to 12 m tall, leafy branchlets 4-9 mm thick, 
glabrous or minutely (0.1-0.4 mm) and very sparsely 
puberulent at the apex, orange brown to almost black, 
becoming longitudinally striate with a few lenticels. Leaves 
alternate to closely clustered (subverticillate) distally, 
petioles 1 5-40 mm long, 1 .2-3 mm thick, glabrous (rare- 
ly sparsely and minutely puberulent), narrowly sulcate 
above; leaf blades (9-)l 1-23 cm long, (4-)6-10 cm broad, 
broadly elliptic to elliptic-obovate or oblong-obovate, 
usually broadest at or above the middle, bluntly obtuse 
to rounded at the apex, gradually narrowed to the obtuse 
or acute base, margin entire and slightly revolute when 
dry, the leaves drying subcoriaceous, glabrous above and 
the midvein impressed, the upper surface with a retic- 



BURGER: FLORA COSTARICENSIS 



105 



ulum of small (0.2-0.5 mm) areolae formed by the raised 
minor venation, lower surface glabrous, with (5-) 7-1 3 
major secondary veins on each side, central secondaries 
arising at angles of 40-70, the lower surface usually 
developing a flat or slightly raised reticulum of minor 
venation forming areolae 0.1-0.3 mm broad. Inflores- 
cences from the axils of distal leaves, to 1 5 cm long in 
fruits, peduncles 3-1 1 cm long in fruits. Flowers not seen 
but the perianth partly persisting in fruits, outer tepals 
shorter than the inner and glabrous on the inner surface, 
inner tepals puberulent on both surfaces; persisting sta- 
mens (series II) 4-thecous (in Poveda 740 at MO), straight 
lustrous hairs ca. 0.7 mm long present around the base 
of the ovary (fruits). Fruits borne on a gradually thick- 
ened pedicel 1-2 cm long and 3-6 mm broad at the top, 
usually with large (2-3 mm) brown lenticel-like protu- 
berances, apex of the receptacle flat or slightly (1 mm) 
depressed; berry globose, 13-17 mm in diameter, be- 
coming dark purple. 



TYPE Costa Rica, Alajuela Province, La Paz 
de San Ramon, 30 Oct. 1973, Poveda 740 (holo- 
type, CR 53308; isotype, MO). 

Trees of evergreen forest formations along the 
continental divide on the western edge of the Mes- 
eta Central, at about 1000 m elevation. Fruits have 
been collected in October-November. This species 
has only been collected at La Palma, Piedades, 
and La Paz, all near San Ramon in Alajuela Prov- 
ince. 

Persea povedae is recognized by the stiff gla- 
brous, medium- to larger- sized leaves with a very 
fine reticulum of minor venation usually visible 
on both surfaces. Globose fruits, thick lenticellate 
fruiting pedicels, partly persisting perianth parts, 
and usually obovate leaves clustered at the ends 
of branchlets further distinguish this species. The 
following collections are placed here: Brenes 337 
(178), 4769 (F) and Poveda 740 (CR, MO). Kopp 
annotated Brenes 6342 as Persea near P. albida, 
and the two species are similar. However, P. albida 
has equal tepals, and the outer tepals are puber- 
ulent within, whereas P. povedae has unequal te- 
pals and the outer tepals glabrous within. This 
species closely resembles P. cuneata Meissner of 
Colombia, but that species has smaller (8 mm) 
ellipsoid fruits, stiffer leaves with more numerous 
secondary veins that are more prominent beneath, 
and anthers with only two thecae. Luis Poveda, 
expert collector and authority on the trees of Costa 
Rica, was the first person to collect this species 
since Brenes, over 40 years earlier. 



Persea rigens C. K. Allen, J. Arnold Arbor. 26: 
297. 1945. Figure 15. 



Trees 6-20(-35) m tall, leafy branchlets 2.5-8 mm 
thick, quickly becoming glabrous and reddish brown, 
smooth or slightly striate, terete, often several arising 
together from a swollen node. Leaves alternate, opposite 
or whorled, often closely congested distally, petioles 8- 
27 mm long, 1.5-4 mm thick, glabrescent and drying 
dark in color, usually sulcate above; leaf blades 9-24(-30) 
cm long, 3.5-9(-l 1) cm broad, obovate-elliptic to ob- 
lanceolate or elliptic-oblong, usually short-acuminate at 
the apex (less often obtuse or rounded), acute to obtuse 
at the base, drying stiffly chartaceous to subcoriaceous 
and often grayish brown to orange brown in color, smooth 
and glabrous above with the tertiary venation slightly 
raised to form a reticulum, lower surface essentially gla- 
brous, with 5-9 major secondary veins on each side, the 
central secondaries arising at angles of 40-60. Inflo- 
rescences clustered at the ends of branchlets, often 2 or 
more from the axils of leaves or fallen leaves, 5-12 cm 
long, thyrsiform panicles, peduncles 2-7 cm long, 1-2 
mm thick, densely and minutely puberulent with thin 
crooked hairs ca. 0.3 mm long, pedicels 1-3.5 mm long. 
Flowers 3-5 mm long, outer tepals ca. 3 mm long and 
3 mm broad, inner tepals ca. 3 mm long and 2 mm 
broad, tomentulose on both inner and outer surfaces; 
outer stamens with anthers 0.8-1.3 mm long on short 
puberulent filaments, inner stamens ca. 2 mm long, 
staminodes prominent, 1.4 mm long and with sagittate 
apices; pistil ca. 2.5 mm long, style slender and with a 
subcapitate stigma. Fruits subtended with short (2-4 mm) 
subequal persisting tepals; berry becoming 1.5-2 cm long 
and 2-3 cm broad, subreniform in shape and somewhat 
depressed at the apex (but see below). 

A poorly defined species of evergreen forest for- 
mations based on a collection by Elbert Little (6058, 
F, the type) from about 1 500 m elevation near the 
Rio Chiriqui Viejo in Chiriqui Province, Panama. 
The flowers of the type were collected in March 
but are not fully in anthesis. This species was 
thought to range from Guatemala, Costa Rica, and 
Panama to Colombia, Venezuela, and Ecuador, 
but its correct circumscription is not resolved (see 
below). 

Persea rigens is distinguished by the occasional 
presence of opposite or whorled leaves clustered 
at swollen nodes at the ends of branchlets. These 
swollen nodes can produce several new twigs in 
later stages. The small puberulent flowers (relative 
to other Persea species with subequal perianth 
parts) and the fruits becoming broader than long 
(in South American collections) further distin- 
guish material placed under this name. The gla- 
brous oblanceolate to elliptic leaves with thick dark 
petioles and fine reticulum of tertiary veins on the 
upper surface, help in the determination of sterile 
collections. We are not sure if Persea rigens has 
been collected in Costa Rica because it is difficult 
to separate this species from the closely related P. 
silvatica, and earlier determinations of Costa Ri- 
can collections as P. rigens may, in fact, be P. 



106 



FIELDIANA: BOTANY 



silvatica. See the discussion under P. silvatica. The 
description of fruits as reniform and the placement 
of a number of South American collections under 
P. rigens (Kopp, 1966, p. 24) require further con- 
firmation. 



Persea schiedeana Nees, Syst. laur. 130. 1836. P. 
pitlieri Mez, Bot. Jahrb. Syst. 30, Beibl. 67: 15. 
1901. Figure 6. 

Trees to 30 m tall, branchlets thick and roughened by 
scars of leaves and bud scales, leafy branchlets 4-8(-16) 
mm thick, at first densely ferrugineous-villous, shoot 
apex with several imbricate series of broad (ca. 6 mm) 
distally rounded bud scales. Leaves alternate and often 
closely clustered at the tips of shoots, petioles 1.5-6 cm 
long. 1.5-3 mm thick, densely puberulent and sulcate 
above; leaf blades 8-26(-33) cm long, 4-20 cm broad, 
very broadly elliptic to broadly oblong or obovate, usu- 
ally bluntly rounded at the apex, rounded and truncate 
to obtuse at the base, margins often somewhat undulate, 
drying stiffly chartaceous to subcoriaceous, upper surface 
usually dark and glabrous, lower surface densely orange 
brown tomentulose with slender hairs 0.3-0.5 mm long, 
with 6-10 pairs of major secondary veins on each side, 
central secondaries arising at angles of 30-60. Inflo- 
rescences axillary to distal leaves or subterminal, 4-20 
cm long, racemose to thyrse like, in the axils of caducous 
bracts to 2 cm long, expanding with a new flush of leaves, 
peduncles 2-7 cm long, densely yellowish or reddish 
brown puberulent, pedicels 10-26 mm long, slender and 
densely puberulent. Flowers 6-10 mm long, becoming 
10-20 mm broad, tepals 6-9 mm long and 2.2-3.3 mm 
broad, broadly lanceolate, densely tomentulose on both 
surfaces, outer tepals slightly shorter or slightly longer 
than the inner; outer stamens ca. 5 mm long with fila- 
ments ca. 3.5 mm long, narrow and with hairs abaxially, 
staminodes 1.5-2.5 mm long and linear; pistil with an 
ellipsoid ovary 2-3.5 mm long, style 1.8-3 mm long, 
stigma simple. Fruits subtended by the persisting peri- 
anth in early stages, the thickened infructescence with 
very few fruits; berry globose and to ca. 5 cm in diameter. 

Trees of evergreen montane rain forest forma- 
tions, between 1000 and 2300 (2500) m elevation 
in Costa Rica. Flowering collections have been 
made in January-May in Costa Rica, and fruits 
have been collected in January-February, May, 
and September. This species ranges from southern 
Mexico through the Central American highlands 
to Colombia. 

Persea schiedeana is recognized by its broad 
leaves densely reddish brown puberulent on the 
undersides, the very rough thick branchlets with 
bud scales or bud scale scars, the yellow flowers 
with long narrow stamens, and the cool wet cloud 
forest habitat. This species has been called agua- 
cate de montana and aguacaton in Chiriqui, Pan- 
ama, and yas in Costa Rica. This species may 



intergrade with P. americana; see the discussion 
under P. americana var. nubigena. 

Persea silvatica van der Werff, sp. nov. 

Arbor ad 10m. Ramuli teretes, apice incrassati. Folia 
chartacea, elliptica vel elliptica-oblonga, 15-25 x 7-12 
cm, glabra. subopposi ta vel apice ramulorum fasciculata, 
apice acuta vel breviter acuminata, basi acuta; costa ner- 
vique (6-10) supra immersi, subtus elevati; reticulatio 
elevata; petioli in sicco nigri, glabri, 1-1.6 cm longi. 
Inflorescentiae apice ramulorum confertae, basi bracteis 
vel cicatribus bractearum circumcincti, ad 20 cm longae, 
minute puberulae. Tepala 6, aequalia vel subaequalia, 
late ovata, 2.5-3.0 mm longa; stamina 9, 4-locellata, 6 
exteriores locellis introrsis, 3 interiores locellis extrorsis, 
filamentis pubescentibus. Staminodia 3, apice sagittata. 
Glandulae staminum interiorum magnae, basi sagittatae 
vel cordatae, vix super basim filamentorum affixae. 
Ovarium glabrum, globosum, ca. 1.1 mm diam.; stylus 
ovarium aequans. Fructus non visus. 

Trees to 10 m tall, leafy branchlets 2.5-6 mm thick, 
glabrous, smooth and brown when young, terete, gray- 
corky when old, usually with several twigs originating 
from a large bud and these twigs with bracts or bract 
scars at the very base, ends of the stems often thickened. 
Leaves subopposite or clustered at the swollen ends of 
branchlets, petioles 10-16 mm long, 1.8-2.5 mm thick, 
drying black, glabrous, with slightly elevated lateral mar- 
gins; leaf blades 15-25 cm long, 7-12 cm broad, elliptic 
to elliptic-oblong or slightly obovate, acute or short acu- 
minate at the apex, acute at the base, drying chartaceous 
to very stiffly chartaceous, primary and secondary veins 
slightly immersed above and prominent beneath, a re- 
ticulum of minor venation slightly raised on both sur- 
faces, glabrous above and below, with 6-10 major sec- 
ondary veins on each side, central secondaries arising at 
angles of 40-65. Inflorescences clustered at the tips of 
branchlets (only seen on the type), subtended at the base 
by bracts or bract scars, ca. 20 cm long, paniculate, pe- 
duncles 4.5-8 cm long, 2-3 mm thick, very sparsely and 
minutely puberulent (more densely only at the base), 
pedicels 2-5 mm long, very minutely (0. 1 mm) whitish 
puberulent. Flowers 3-4 mm long and equally broad, 
campanulate, white to yellowish green, tepals erect and 
equal at anthesis, 2.5-3 mm long, ovate, minutely pub- 
erulent; stamens 9 and all 4-thecous, outer 6 stamens 
dehiscing introrsely, ca. 2.4 mm long with puberulent 
filaments ca. 1.4 mm long, inner stamens ca. 2.1 mm 
long, with large glands attached slightly above the base 
of the filament, Staminodia 3, ca. 1.3 mm long, with the 
sagittate tip curved inward over the ovary; pistil ca. 2.2 
mm long, ovary globose and glabrous, ca. 1.1 mm long, 
stigma discoid. Fruits unknown. 

TYPE Costa Rica, Heredia Province, along the 
Rio Guacimo, southeast of the La Selva Biological 
Station, 80 m elevation, 14 March 1984, G. Schatz 
& H. Young 964 (holotype, MO; isotype, CR). 

Trees of the lowland Caribbean rain forest for- 
mation. Flowering material was collected in March 



BURGER: FLORA COSTARICENSIS 



107 



(the type) and August (Hammel 9453, DUKE). The 
species is only known from the area of the La Selva 
research station (at about 1 00 m elevation) in north- 
central Costa Rica. 

Persea silvatica is recognized by its glabrous el- 
liptic leaves clustered or subopposite at the thick- 
ened (knoblike) ends of slender twigs. This unusual 
growth form is also found in P. rigens, but that 
species has much more densely puberulent whitish 
tomentulose flowers, and less prominently raised 
minor venation on the dried leaves. Earlier, spec- 
imens of this species were identified as P. rigens; 
compare material filed under that name. 

Kopp ( 1 966) divided Neotropical Persea species 
into two subgenera in her revision. Subgenus Per- 
sea is characterized by equal or subequal tepals, 
tepals mostly deciduous in fruits, a pubescent 
ovary, and reflexed perianth at anthesis. Subgenus 
Eriodaphne has unequal tepals (the inner three 
much longer than the outer), tepals persistent in 
fruits, a glabrous or pubescent ovary, and spread- 
ing or erect tepals at anthesis. Persea silvatica does 
not fit well in either subgenus, since it shares equal 
tepals with subgenus Persea and a glabrous ovary 
and persistent tepals with subgenus Eriodaphne. 
Some of the characteristics of Persea silvatica and 
P. rigens suggest a relationship with the Asian sub- 
genus Machilus. Moreover, Hammel (1966, p. 233) 
points out that the flowers of this species are very 
similar to some Costa Rican species of Phoebe (but 
those species develop cupules in fruits). Additional 
research, based on better collections, is needed to 
resolve problems with generic boundaries between 
the subgenera of Persea and some Neotropical 
species currently placed in Phoebe. 



lower surface. Inflorescences axillary to distal leaves or 
clustered near the apex from a leafless short shoot, 5- 
1 2(-l 6) cm long, paniculate, peduncles 2-7 cm long, pale 
yellowish strigulose, pedicels 0-2 mm long. Flowers 3- 
5 mm long, ca. 5 mm broad, densely yellowish or grayish 
strigulose to sericeous on the outer (abaxial) surfaces, 
outer tepals 2-2.5 mm long and 2-2.2 mm broad, ovate 
and acute to obtuse at the apex, glabrous within, inner 
perianth parts 3-5 mm long and 2-2.2 mm broad, acute 
at the apex, sericeous on both surfaces; outer stamens 
1.5-3.5 mm long, anthers ca. 1 mm long with 4 super- 
posed thecae, staminodes ca. 2 mm long, sagittate to 
spatulate; pistil glabrous, 3-4 mm long with style 1.5- 
2.2 mm long, stigma discoid. Fruits subtended by the 
persisting perianth of the shorter outer and longer inner 
whorls; berry globose, 8-12 mm in diameter (dry), green 
and said to become glaucous. 

Trees of evergreen forests of the central high- 
lands and the Pacific slope, between 800 and 2300 
m elevation in Costa Rica. Flowers have been col- 
lected in April-September and December, and 
fruits have been collected in January-April. This 
species ranges from the Cordillera de Tilaran in 
western Costa Rica to Veraguas Province in Pan- 
ama. 

Persea veraguasensis is recognized when dry by 
the pale colored or reddish brown ovate-lanceolate 
laminae borne on long slender petioles and the 
minute, often sericeous, puberulence on young 
vegetative parts and flowers. This species is very 
similar to P. caerulea, but the perianth whorls 
(tepals) do not differ as much in length in flowers 
at anthesis. In addition, the tips of the inner tepals 
do not break off in P. veraguasensis, with the result 
that the tepals differ more in length in this species 
in the fruiting condition than they do in P. cae- 
rulea. 



Persea veraguasensis Seem., Bot. voy. Herald 193. 
1854. P. veraguensis Meissn. in DC., Prodr. 15, 
pt. 1:51. 1864, nomen super/I. Figure 9. 

Trees to 12 m tall, leafy branchlets 2-8 mm thick, 
grayish or yellowish brown with minute (0.2 mm) ap- 
pressed slender hairs but becoming (sub)glabrous and 
dark. Leaves alternate, clustered distally or separate, pet- 
ioles 2-5 cm long, 1-2 mm thick, slightly canaliculate 
above, glabrescent; leaf blades 6-14(-16) cm long, 2.5- 
5.5(-6.5) cm broad, lanceolate to ovate or ovate-elliptic, 
tapering gradually to the acute or bluntly obtuse apex, 
acute to rounded and subtruncate at the base, unequal, 
leaves drying stiffly chartaceous to subcoriaceous and 
pale greenish to reddish brown in color, smooth and 
glabrous above, minutely and densely appressed puber- 
ulent beneath in early stages but soon glabrescent, with 
5-10 major secondary veins on each side, central sec- 
ondaries arising at angles of 40-60 C , tertiary veins form- 
ing a reticulum of small (0.2-0.4 mm) areolae on the 



Persea vesticula Standl. & Steyer., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 23: 1 16. 1944. P. chiapensis 
Lundell, Wrightia 1: 150. 1946. P. popenoei L. 
O. Williams, Ceiba 1: 57. 1950. Figure 5. 

Small shrubs to medium-sized trees 2-12(-25) m tall, 
leafy branchlets 2.5-9 mm thick, at first densely brown- 
ish or yellowish puberulent but glabrescent and dark in 
age. Leaves alternate to subopposite, usually clustered 
near the ends of branchlets, petioles 4-38 mm long, 1 .3- 
3 mm thick, puberulent or glabrous, flat or slightly sul- 
cate above; leaf blades (3-)4.5-14(-l 7) cm long, (1 .8-)2.5- 
7 cm broad, elliptic to ovate, acute to bluntly obtuse 
(rounded in very small leaves) at the apex, obtuse to 
rounded and subtruncate (rarely acute) at the base, mar- 
gin becoming revolute, drying subcoriaceous to coria- 
ceous, glabrous to sparsely and minutely puberulent 
above, lower surface glabrous to sparsely puberulent with 
slender hairs 0.5 mm long or with a dense tomentum 
between the veins (in some individuals), with (3-)5-l 1 



108 



FIELDIANA: BOTANY 



major secondary veins on each side, central secondaries 
arising at angles of 30-60. Inflorescences axillary to 
distal leaves, 3-15 cm long, paniculate, peduncles 1.5- 
1 1 cm long, densely brownish tomentulose or glabres- 
cent, pedicels ca. 2 mm long. Flowers 4-6 mm long, outer 
tepals 2.5-4.5 mm long and 2-3 mm broad, tomentulous 
on the outside and usually glabrous within, inner tepals 
4-6.5 mm long, densely puberulent on both surfaces; 
outer stamens 1.5-3.5 mm long, outer anthers 1-2.2 mm 
long, narrowly oblong with 4 superposed thecae, inner 
stamens 2.2-4 mm long, staminodes 1-1.5 mm long; 
pistil 2.2-4 mm long, glabrous, ovary ovoid or ellipsoid, 
style slender and 1.1-3 mm long, stigma discoid. Fruits 
subtended by the persisting perianth flattened in a plane 
ca. 6-8 mm broad, dark brown and puberulent; berry 
8-12 mm in diameter (dry), globose and glaucous. 

Shrubs and trees of high montane cloud forests 
and elfin forest formations, between 2000 and 3200 
m elevation in Costa Rica. Flowering material has 
been collected in September-March in Costa Rica 
and in April-August in Guatemala and Honduras. 
Fruiting material has been collected in November- 
April in Central America. The species ranges from 
Chiapas, Mexico, southward to the Cordillera de 
Talamanca of Costa Rica, and may extend into 
adjacent Panama (see below). 

Persea vesticula is recognized by its high-ele- 
vation habitats, the thick dark twigs with leaves 
usually clustered at the ends, the puberulent flow- 
ers with perianth whorls of differing length, and 
the small globose glaucous fruits subtended by the 
persisting tepals. The specimens placed here span 
a great variety of leaf forms. These range from 
larger elliptic laminae with numerous strongly as- 
cending secondary veins to very short rounded 
leaves with only a few secondary veins. Interme- 
diate collections span this range, and the small 
thick rounded leaves appear to be associated with 
very high altitudes or exposed windy sites. The 
flowers also range considerably in size; anthers of 
Costa Rican material are about 1 mm long, while 
the type from Guatemala has anthers about 2mm 
long. Considering this range of variation it may 
be difficult to separate some collections from the 
closely related P. obtusifolia. The species is called 
asca. 



Phoebe Nees 

Trees or shrubs, bisexual. Leaf blades alternate (rarely 
subopposite), petiolate, leaf blades entire with palmate, 
tripliveined or pinnate venation, domatia sometimes 
present in the vein axils of the lower leaf surfaces. In- 
florescences usually solitary and axillary to distal leaves 
(subterminal or rarely fasciculate), paniculate and with 
spreading lateral branches or with short lateral branches 



and racemose, distal flower groups often cymose, gla- 
brous or puberulent. Flowers bisexual, radially sym- 
metrical, usually campanulate in form, perianth of 6 
parts in 2 whorls, outer and inner whorls equal or sub- 
equal with the outer parts somewhat shorter, glabrous 
or puberulent, floral tube short; androecium with 9 fertile 
4-thecous stamens (in Central America), the thecae su- 
perposed in 2 planes, the 6 outer stamens (series I-II) 
with introrse dehiscence, filaments usually equalling the 
anthers in length or slightly shorter, the 3 inner stamens 
(series III) with well-developed filaments and each with 
2 sessile or subsessile glands, inner anthers 4-thecous 
with the lower having extrorse and the upper lateral or 
lateral-extrorse dehiscence, staminodes (series IV) 3 and 
prominent, with cordate-sagittate apices usually borne 
on a short stipe, usually with hairs at the base; pistil 
glabrous (in Costa Rica), ovary subglobose to ellipsoid, 
style slender and often equalling the ovary in length, 
stigma discoid to obtuse (simple). Fruits borne in a cu- 
pulate or saucer-like receptacle, expanded above the 
thickened pedicel, perianth persisting at the edge of the 
receptacle or more often deciduous (never enlarging); 
berry ovoid to ellipsoid or globose. 



Phoebe, with over 175 specific epithets, is not 
well defined. Many species that have been de- 
scribed in Central America are better placed in 
Ocotea, and we have made a number of transfers 
in this text. Moreover, Kostermans (1957) defined 
Phoebe so as to exclude all the American species, 
placing them in Cinnamomum. Transfer of Phoebe 
cinnamomifolia and its allies to Cinnamomum is 
probably correct, but should be part of a global 
survey of the species. Kostermans' work (1957) is 
quite superficial regarding Neotropical taxa, and 
we prefer to maintain current usage at this time. 
The three large staminodes with cordate-sagittate 
apices in each flower, the nine fertile 4-thecous 
stamens, and fruits subtended by an expanded re- 
ceptacle, characterize the Costa Rican species 
placed here. They differ from Ocotea primarily in 
the consistent presence of three well-developed 
staminodes and from Persea because of the de- 
velopment of the receptacle in fruits. Tripliveined 
leaves characterize a few species in our area. 

In this treatment, Phoebe helicterifolia, P. mol- 
licella, P. pittieri, and P. valeriana are considered 
as species of Ocotea and have been transferred. 

There is a serious problem regarding the cir- 
cumscription of Phoebe costaricana and P. cin- 
namomifolia. The senior author was at first unable 
to distinguish these two entities, which appear to 
behave as two different species in Costa Rica, and 
most specimens were annotated as P. cinnamom- 
ifolia; see the discussion under P. cinnamomifolia. 

Phoebe amplifolia Mez and J. D. Smith was said 
to occur in Costa Rica by C. K. Allen ( 1 945), based 
on Popenoe 984 from near Rancho Redondo in 



BURGER: FLORA COSTARICENSIS 



109 



the province of San Jose. We have not seen this 
collection and doubt that it is correctly identified. 



Phoebe amplifolia is a larger-leaved species of 
Guatemala, resembling P. hammeliana. 



Key to Species of Phoebe 

la. Leaves tripliveined or subtripli veined, with the basal lateral secondary veins strongly ascending; 

the leaves often drying subcoriaceous and often grayish green to yellowish, glabrous 2a 

Ib. Leaves pinnately veined or if subtripliveined then the laminae drying thin in texture and usually 

puberulent 4a 

2a. Leaves strongly tripliveined with the basal lateral veins reaching the distal part of the lamina, 
mid vein with weakly developed secondary veins; fruiting receptacle becoming 12-14 mm wide; 

southeastern Costa Rica P. neurophylla 

2b. Leaves weakly tripliveined, the basal lateral veins rarely reaching the distal part of the lamina, 

midvein usually with prominent secondary veins (at least in the distal half) 3a 

3a. Leaves usually elliptic-oblong to lanceolate, (6-)9-29 cm long, tapering gradually to an acute 
or acuminate apex; fruit cups 8-12 mm broad; wide-ranging, 0-1900 m . . P. cinnamomifolia 
3b. Leaves ovate to broadly elliptic, usually tapering abruptly to a short-acuminate or caudate- 
acuminate apex; fruit cup 5-8 mm broad; Pacific slope, 600-1200 m P. brenesii 

4a. Leaves glabrous beneath, usually drying pale grayish or yellowish green, coriaceous to subcoriaceous 

5a 

4b. Leaves puberulent beneath, usually drying dark brown to yellowish brown; species now transferred 

to the genus Ocotea 6a 

5a. Leaves 20-50 cm long, petioles 2.5-6 cm long; 0-1000 m elevation P. chavarriana 

5b. Leaves 9-22 cm long, petioles 1 .5-3 cm long and often reddish (in life); 1 500-2500 m elevation 

P. hammeliana 

6a. Leaves 1-4 cm broad and 3-10 cm long, usually narrowly elliptic; flowers often with 3 staminodes 

7a 

6b. Leaves 4-14 cm broad and 8-30 cm long, usually broadly elliptic to obovate; flowers rarely with 3 

well-developed staminodes 8a 

7a. Fruit cups ca. 6 mm broad; pubescence on the lower leaf surfaces grayish and soft to the touch; 

1400-2300 m O. mollicella 

7b. Fruit cups 10-15 mm broad; pubescence on the lower leaf surfaces brownish and slightly rough 

to the touch; 1000-3200 m O. pittieri 

8a. Fruit cups 10-18 mm broad and 5-6 mm deep; trees of montane forests, 1000-2000 m 

O. valeriana 

8b. Fruit cups 7-14 mm broad and 2-3 mm deep; lowland rain forests, 0-500 m . . . . O. helicterifolia 



Phoebe brenesii Standl., Publ. Field Mus. Nat. 
Hist. Hot. Ser. 18: 459. 1937. Figure 1. 

Small or medium-sized trees 5-15(-20) m tall, trunk 
to 75 cm in diameter, leafy branchlets 1 .2-3.5 mm thick, 
very minutely (0. 1 mm) puberulent at first but soon be- 
coming (sub)glabrous, dark in color and longitudinally 
striate. Leaves alternate, often distally clustered, petioles 
4-30 mm long, ca. 1 mm thick, with 2 adaxial ridges 
forming an adaxial sulcus; leaf blades 2.5-10.5 cm long, 
1 .3-4(-5) cm broad, broadly elliptic to elliptic-oblong or 
ovate-elliptic, usually tapering abruptly to the short-acu- 
minate or caudate-acuminate apex, usually obtuse to 
rounded at the base, slightly decurrent on the petiole, 
drying stiffly chartaceous to subcoriaceous and dark or 
pale yellowish gray, smooth and glabrous above and be- 
low, tripliveined with a prominent pair of secondary 



veins arising from the primary vein 2-12 mm above the 
petiole, 1 or 2 additional secondaries arising from the 
central or distal part of the midvein on each side, tufted 
hairs (domatia) usually present in the axils of the basal 
veins beneath, tertiary and smaller veins sometimes 
forming a slightly raised reticulum beneath. Inflores- 
cences axillary, 3-15 cm long with relatively few flowers 
in a distal open panicle, peduncle to 8 cm long and 
usually as long or longer than the flowering rachis, gla- 
brous, pedicels 2-4 mm long. Flowers 2.5-4 mm long, 
outer tepals ca. 2.5 mm long and glabrous on the outside; 
outer anthers narrow and ca. 1 mm long with filaments 
ca. 0.5 mm long, inner stamens ca. 2 mm long, stami- 
nodes 1-1.3 mm long with puberulent stipe and glabrous 
acute apex; pistil 2-2.5 mm long, ovary globose to tur- 
binate, style slightly longer than the ovary, stigma cap- 
itate. Fruits borne on a broadly expanded (5-8 mm) 
shallow (2-3 mm deep) obconical receptacle ca. 4 mm 



110 



FIELDIANA: BOTANY 



long, rim apparently entire but often with 6 slits marking 
the bases of the tepals, pedicel slightly thickened and 4- 
5 mm long; berry 10-13 mm long and 5-9 mm in di- 
ameter (dry), ellipsoid, becoming black on a red or rose- 
colored base. 

Trees of the Pacific slope between 600 and 1 200 
m elevation, but reaching 2000 m in Chiriqui. 
Flowers have been collected in January-April, 
while fruits have been collected in March-May 
and September. This species may be restricted to 
premontane moist forest formations. It is only 
known from the Meseta Central (San Ramon to 
Villa Colon), near Boquete in the Chiriqui high- 
lands and in Veraguas Province of Panama. 

Phoebe brenesii is recognized by the usually small 
stiff and abruptly acuminate leaves with tripli- 
veined venation. In addition, the leaves tend to 
be broad and are often borne on long slender pet- 
ioles. The glabrous flower buds, prominent stam- 
inodes with caudate-acuminate apices, and small 
fruits on shallow receptacles further distinguish 
this species. Compare material of this species with 
the very variable P. cinnamomifolia; P. brenesii 
is probably a derivative of that species. 



Phoebe chavarriana Hammel, J. Arnold Arbor. 
67: 131. 1986. Figure 2. 

Small or medium-sized trees 8-15 m tall, 15-35 cm 
d.b.h., bark reddish brown, leafy branchlets 4-10 mm 
thick, at first with minute (0. 1 mm) appressed ascending 
grayish hairs, glabrescent and usually drying dark in col- 
or, terete. Leaves alternate, petioles 2.5-6 cm long, 3-6 
mm thick, narrowly sulcate above, glabrous; leaf blades 
20-50 cm long, 10-23 cm broad, elliptic to somewhat 
ovate-elliptic, elliptic-oblong or slightly elliptic-obovate, 
acute to obtuse or very short acuminate at the apex, acute 
to obtuse or rounded at the base, drying subcoriaceous 
and grayish green, glabrous above and below, the mid- 
vein impressed above, with 6-10 major secondary veins 
on each side, the central secondaries arising at angles of 
45-65, leaves appearing glaucous beneath. Inflores- 
cences axillary from distal leaves or clustered near the 
apex, 10-23 cm long, paniculate, peduncles 1-3 cm long 
and ca. 1 mm thick, sparsely and minutely puberulent, 
pedicels 1-3 mm long. Flowers 2.5-4 mm long, sparsely 
and minutely puberulent on the outside, perianth whorls 
subequal, tepals ca. 2.5 mm long, greenish white; outer 
stamens 1.5-1.9 mm long, outer anthers 1-1.2 mm long, 
oblong with superposed thecae, inner stamens ca. 2 mm 
long, with sparsely puberulent filaments, stum modes 1.2- 
1.8 mm long and with sagittate-cordate apex; pistil ca. 
2.2 mm long, glabrous, ovary globose, style ca. 1 mm 
long, stigma simple. Fruits borne on a flattened recep- 
tacle ca. 2 mm deep and 1 2 mm broad with persisting 
perianth parts (but only 1-2 mm long), pedicel 1-1.5 cm 
long and 4-6 mm thick below the expanded apex, conical 



and becoming red; berry 15-18 mm long and 10- 12 mm 
in diameter, ovoid, dark green to almost black. 

Trees of the lowland rain forest and escarpment 
on the Caribbean slope, between 50 and 1000 m 
elevation. Flowering occurs in April-May, and 
fruits have been collected in July-August. This 
species is only known from near the confluence of 
the Rio Sarapiqui and Rio Puerto Viejo in Her- 
edia, and from near Moravia de Turrialba in Car- 
tago (but see below). 

Phoebe chavarriana is recognized by its very 
large glabrous leaves, and fruits borne on flattened 
receptacles with very short persisting perianth parts 
above a conical pedicel. The large staminodes are 
characteristic of Phoebe and Persea, but the fruits 
are similar to those of several other species of 
Phoebe. The species was named in honor of Rafael 
Chavarria, who helped make the La Selva research 
station a success. 

An unusual collection (Holdridge 6336, CR, usj) 
with large (50 x 1 8 cm) obovate leaves and large 
(30 x 15 mm) oblong fruits from 200 m in the 
Caribbean lowlands is provisionally placed here. 
Another unusual collection (Hartshorn 1456, CR, 
F) along the Sarapiqui road at 950 m elevation 
may be a small-leaved form, with leaves that dry 
yellowish and similar infructescences (collected in 
April). Phoebe chavarriana is closely related to 
Phoebe hammeliana of higher elevations. 



Phoebe cinnamomifolia (H.B.K.) Nees, Linnaea 2 1 : 
488. 1 848. Persea cinnamomifolia H.B.K., Nov. 
gen. sp. 2: 1 60. 1817. Phoebe mexicana Meissn. 
in DC., Prodr. 15: 31. 1864. Persea mexicana 
(Meissn.) Hemsl., Biol. cent. amer. Bot. 3: 71. 
1882. Phoebe tonduzii Mez, Bot. Jahrb. Syst. 
30, Beibl. 67: 1 5. 1 90 1 . Phoebe costaricana Mez 
& Pittier, Bull. Herb. Boissier 3: 230. 1903. Fig- 
ure 1. 

Trees (rarely shrubs), 5-25 m tall, trunks often 30-40 
cm in diameter, leafy branchlets 1 .5-6 mm thick, at first 
with minute (0. 1-0.2 mm) appressed ascending hairs but 
usually becoming (sub)glabrous and dark brown. Leaves 
alternate, petioles l-3.5(-5)cm long, 1-3 mm thick, with 
a broad or narrow sulcus above, minutely puberulent or 
glabrous; leaf blades (6-)9-29 cm long, (3-)4-9(-14) cm 
broad, elliptic-oblong to oblong, ovate-oblong, lanceo- 
late or falcate, tapering gradually to the acute or acu- 
minate apex, acute to obtuse at the base, drying subcor- 
iaceous to coriaceous and often grayish green or yellowish, 
usually glabrous above and below but often with tufted 
hairs in the vein axils beneath, venation usually tripliv- 
eined (rarely pinnate) with the basal secondaries arising 
from 2-20 mm above the petiole and with (2-)3-6 ad- 



BURGER: FLORA COSTARICENSIS 



111 



ditional prominent secondaries on each side, basal sec- 
ondaries arcuate-ascending and usually reaching the 
middle of the lamina, tufted hairs or pit domatia usually 
present in the axils of the major veins beneath. Inflo- 
rescences (5-) 1 0-25 cm long, solitary or in clusters from 
axillary short shoots, racemose (with individual or small 
groups of flowers arising from the primary rachis) to 
paniculate with lateral flowering branches, peduncles 
slender (ca. 1 mm) and shorter than the flowering rachis, 
puberulent or glabrous, pedicels 3-5 mm long. Flowers 
2-4 mm long, 2.2-3.8 mm broad, glabrous or sometimes 
minutely puberulent, tepals 1.5-2 mm long, erect and 
often subequal with the outer slightly shorter than the 
inner; outer stamens with filaments 0.2-1 mm long, the 
narrow outer anthers 0.6-1 mm long, inner stamens 1.5- 
2 mm long, staminodes 0.8-1.5 mm long with a thick 
acute apex 0.5-0.8 mm; pistil 2-3.5 mm long, ovary 
rounded, style ca. 1 mm long, narrow, stigma capitate 
or simple. Fruits borne in a cupulate campanulate re- 
ceptacle 5-10 mm long, 8-12 mm broad and 2-3 mm 
deep, the perianth parts persisting or deciduous (and then 
the margin with obscure indentations), becoming red; 
berry 1-1.8 cm long, 0.7-1.3 cm thick (dry), ellipsoid, 
dark green. 

Trees of evergreen or partly deciduous forest 
formations on both the Pacific and Caribbean 
slopes, and ranging from near sea level to 1500 
( 1 900) m elevation. This species has not been col- 
lected from below 600 m on the Pacific slope, and 
it appears to be quite rare below 400 m on the 
Caribbean slope. The species is common at Mon- 
te verde, the eastern portion of the Meseta Central 
(but not near the volcanoes), the valley of the Rio 
Reventazon, and from the Cordillera de Tala- 
manca to the Chiriqui highlands. Flowering ma- 
terial has been collected in Costa Rica November- 
May and July. Fruits have been collected in Jan- 
uary-May, July-August, and November. This 
species ranges from southern Mexico through Cen- 
tral America into South America. 



Phoebe cinnamomifolia is recognized by the 
usually larger stiff leaves on long petioles, and the 
tripliveined laminae. The usually glabrous leaves 
(except for the tufted domatia), the often clustered 
racemose inflorescences, small flowers with prom- 
inent staminodes, and fruit cup with persisting 
perianth (or notches in the rim of the cup) further 
distinguish this species. Specimens placed under 
this name include a rather heterogeneous assem- 
blage as regards vegetative morphology (see be- 
low), but the floral characters are quite uniform. 
Phoebe brenesii and P. neurophylla are very closely 
related to this species; see the discussions under 
those species. 

Phoebe cinnamomifolia is used here in a very 
broad sense, and includes a variety of forms. Most 
of the unusual collections are linked by interme- 
diates to the more common forms. The type of 
Phoebe tonduzii (based on Tonduz 11753, us) rep- 
resents a very puberulent form with shorter, more 
ovate, leaves; it is rarely collected. A few collec- 
tions with very small elliptic leaves and very small 
glabrous inflorescences have been collected in the 
Caribbean lowlands and are place here provision- 
ally (see Stork 2314, F). More significant is the 
type of P. costaricana Mez & Pittier (based on 
Pittier 11107, us) This specimen can be used to 
characterize a number of higher-altitude ( 1 200 to 
1800 m) collections which may represent a dis- 
tinctive species; it is especially common in the 
Co to Brus area and adjacent Chiriqui highlands 
of Panama. The following key attempts to separate 
P. costaricana from most collections of P. cinna- 
momifolia in Costa Rica, but there are many col- 
lections that appear to be intermediate and the 
question of recognizing P. costaricana as a species 
or subspecies requires further study. 



Diagnostic Features of Phoebe costaricana and P. cinnamomifolia 

\ a. Inflorescences nearly always solitary and axillary to distal leaves, glabrous and with few-flowered 
lateral branches, pedicels to 4 mm long, the flowers not closely clustered; axillary buds inconspicuous 
and usually less than 2 mm long; the leaves usually strongly tripliveined with only a few prominent 

secondary veins in the distal half of the lamina, leaf blades rarely more than 12 cm long 

P. costaricana 

1 b. Inflorescences solitary or several on leafless short shoots, pseudoterminal or axillary to distal leaves, 
densely grayish puberulent to glabrous, pedicels to 3 mm long and flowers often clustered in cymose 
groups of 7 or more; larger (3-4 mm) axillary buds with whitish hairs and overlapping scales often 
present and conspicuous in leaf axils and at the tips of twigs; leaves tripliveined but usually with 
well-developed secondaries throughout the length of the midvein, leaf blades often more than 1 5 
cm long p. cinnamomifolia 



112 



FIELDIANA: BOTANY 



Phoebe hammeliana W. Burger, sp. nov. Figure 
12. 



Arbor 8-1 5 m alta, ramulis foliiferis 2.5-8 mm crassis. 
Folia alterna, pctiolis 1.4-2.8 cm longis; laminis 9-21 
cm longis et 5-12 cm latis, ovatis, elliptico-ovatis, vel 
oblongo-ovatis, apice obtuso vel breviter acuminate, gla- 
bris, coriaceis, nervis secondariis 4-8 paribus. Inflores- 
centiae usque 20 cm longae, pedunculis usque 10 cm 
longis, rubris. Flores 3-4 mm longi et 5-7 mm lati, extus 
glabri, stamina ser. I-II filamentis longis et antheris 1- 
1 .2 mm longis, staminodiis 1 .5-2 mm longis et capitatis; 
gynoecium 2.2-2.6 mm longo, ovario subgloboso. Fruc- 
tus ellipsoideus, 1 5 mm longus et 1 mm crassus; cupula 
8-10 mm lata, ca. 2 mm profunda. 



Trees 8-15 mm tall, leafy branchlets 2.5-8 mm thick, 
glabrous or with minute (0. 1 mm) grayish hairs, stems 
becoming reddish brown and longitudinally striate. 
Leaves alternate or occasionally subopposite, petioles 
1.4-2.8 cm long, 1.3-4 mm thick, glabrous or with mi- 
nute grayish hairs, narrowly sulcate above; leaf blades 
9-21 cm long, 5-12 cm broad, ovate or elliptic-ovate to 
ovate-oblong, obtuse to short-acuminate at the apex (and 
the end of the lamina often twisted to one side), rounded 
and truncate to obtuse (acute) at the base, often slightly 
decurrent on the petiole, margin entire or somewhat un- 
dulate, drying coriaceous or subcoriaceous and often lus- 
trous above, glabrous on both surfaces, yellow-green in 
life but drying reddish brown to orange-green, midvein 
impressed above, with 4-8 major secondary veins on 
each side, central secondaries arising at angles of 30- 
60 (the basal secondaries rarely strongly ascending), ter- 
tiary venation obscure. Inflorescences axillary to distal 
leaves or clustered near the apex, to 20 cm long, reddish, 
glabrous, peduncle to 10 cm long, pedicels 0.5-3. 5(-4.5) 
mm long. Flowers 3-4 mm long and 5-7 mm broad, 
tepals 2.5-3.5 mm long, 1.5-2 mm broad, glabrous; out- 
er stamens 2-2.5 mm long, anthers 1-1.2 mm long and 
narrowly ovate-oblong, inner stamens ca. 2.2 mm long, 
staminodes 1.5-2 mm long and with a prominent sag- 
ittate apex, a few hairs present within the floral tube; 
pistil 2.2-2.6 mm long, ovary subglobose, style ca. 1.5 
mm long, slender, stigma simple or slightly discoid. Fruits 
borne on a shallow (2 mm) cup 8-10 mm broad, bases 
of the broadened perianth parts persisting or breaking 
off, pedicel to 2 cm long, conical and often warty, be- 
coming red; berry ca. 15 mm long and 10 mm in di- 
ameter (dry), ellipsoid. 



TYPE Costa Rica, San Jose Province, 0.9 km 
above La Chonta and 100 m from the Pan-Amer- 
ican Highway, elevation 2460 m, 11 May 1969, 
Roy W. Lent 1678 (holotype, CR; isotypes, DUKE, 
F, negative, 61 123, NY). 

Trees of wet evergreen montane forest forma- 
tions along the continental divide and the Carib- 
bean side of the Central Volcanic highlands, and 
in the western part of the Cordillera de Talamanca, 



between 1 200 and 2500 m elevation. Flowers have 
been collected in May-July, and fruits have been 
collected in January, March-April, and Novem- 
ber. The species is only known from central Costa 
Rica and the Chiriqui highlands of Panama. 

Phoebe hammeliana is recognized by the thick 
yellowish green leaves usually drying orange- 
brown, the prominent reddish petioles, small gla- 
brous flowers with prominent staminodes, and the 
fruits borne on a flattened cup at the apex of a 
conical pedicel. Growth patterns of distal twigs 
often resemble those of Persea rigens (q.v.). This 
species is closely related to the lowland Phoebe 
chavarriana, with much larger leaves that are more 
elliptic in form. A closely related species, but with 
more prominent basal secondary veins, occurs at 
similar elevations in Colombia and appears to be 
undescribed. This species seems to suffer from gall 
formations, both on the leaves and in the fruiting 
inflorescences. The following Costa Rican collec- 
tions are placed here: Gomez 23977 (CR), Gon- 
zalez 44 (CR, F), Hammel 14091 (DUKE), Hart- 
shorn 1125 (CR, F), Holdridge 5151 (CR, NY), A. 
Jimenez 2552 (CR, F), Oersted 14938 (F), Poveda 
1061 (CR, F), Utley & Utley 3648 (CR, F), Stork 
2404 (F). This species is named for Barry Hammel, 
whose studies and collections at La Selva have 
made a major contribution to our knowledge of 
Costa Rica's Lauraceae. 



Phoebe neurophylla Mez & Pittier, Bull. Herb. 
Boissier 3: 231. 1903. Figure 1. 

Trees 15-18 m tall, leafy branchlets 1.5-4 mm thick, 
glabrous but the shoot apex with appressed sericeous 
hairs. Leaves alternate, petioles 9-16 mm long, 1-2 mm 
thick, sulcate above, glabrous; leaf blades (6-)9-2 1 cm 
long, (2.3-)3-7 cm broad, elliptic-oblong to oblong, ovate- 
elliptic or slightly obovate-oblong, short-acuminate at 
the apex, acute to obtuse and short-decurrent on the 
petiole, drying stiffly chartaceous and grayish green or 
pale brown, glabrous on both surfaces, midvein im- 
pressed above, venation strongly tripliveined with the 
major basal secondaries arising 5-10 mm above the pet- 
iole and parallel with the margin more than half the 
length of the lamina, major distal secondaries beginning 
near the middle of the lamina and 2-3 on each side, 
prominent pit domatia with tufted hairs present in the 
basal vein axils beneath. Inflorescences 3-7 cm long, 1- 
5 arising together from axillary short shoots, usually ra- 
cemose with most of the flowers solitary or in groups 
directly from the central rachis or paniculate with 
branched lateral branches, peduncle 1-3.5 cm long, slen- 
der (0.5 mm) and glabrous, pedicels 4-5 mm long. Flow- 
ers 4-5 mm long and 3-4 mm broad, glabrous on the 
outside, tepals ca. 2.3 mm long, glabrous but minutely 



BURGER: FLORA COSTARICENSIS 



113 



ciliolate along the distal edge; outer stamens with prom- 
inent (0.6-0.9 mm) filaments bearing narrow anthers 
0.7-0.9 mm long, inner stamens with flattened glands, 
staminodes 1.2-2 mm long with thick acute apices 0.8- 
0.9 mm long; pistil ca. 2.5 mm long, ovary about 1 mm 
thick and gradually narrowed into the style, stigma cap- 
itate. Fruits borne on an open receptacle 1 0- 1 4 mm wide 
and 2-5 mm deep, the broadly obconic receptacle ca. 5 
mm long and with the margin entire or obscurely lobed, 
pedicel becoming 12 mm long and expanded to 4 mm 
thick beneath the receptacle, red; berry becoming 20 mm 
long and 1 5 mm in diameter, ellipsoid, green. 

Trees of the western General Valley and the 
Golfo Dulce area of the Pacific slope of southern 
Costa Rica, between 500 and 1000 m elevation 
(to 1 500 m in Nicaragua). Flowering material has 
been collected in January; fruits have been col- 
lected in March (Pittier 12054, us, the type) and 
June-August. The species is found in Nicaragua 
and southern Costa Rica. 

Phoebe neurophylla is recognized by its larger 
elliptic-oblong leaves with strongly triplinerved 
venation with the basal secondaries paralleling the 
margins from near the base to beyond the middle 
of the lamina. The lack of pubescence, large stam- 
inodes, shallow and subentire fruit cup, and re- 
stricted range further distinguish this species. Car- 
oline Allen (1945, p. 315) placed a specimen from 
Zarcero (A. Smith 142, F) under this name, but 
that collection probably belongs under P. cinna- 
momifolia. Likewise, specimens from Mexico 
ascribed to this species by Allen (1945) will prob- 
ably prove to be another species. Phoebe neuro- 
phylla is very closely related to P. cinnamomifolia, 
but the unusual venation, deeper floral tube, slightly 
larger fruits, and deeper cup support the recogni- 
tion of this taxon as a species rather than a sub- 
specific element of the very variable and widely 
ranging species. Also, the floral parts seem to dif- 
fer, but this is based on a single collection and may 
be atypical. This species is similar in some ways 
to the type of P. costaricana; see the discussion 
under P. cinnamomifolia. 



Pleurothyrium Nees 

Medium-sized trees of evergreen forest formations. 
Leaves alternate or occasionally subopposite, petiolate, 
the leaf blades often obovate (elliptic-oblong in ours), 



entire and pinnately veined, often minutely puberulent 
to tomentose beneath. Inflorescences solitary and axil- 
lary or pseudoterminal, paniculate or racemose, the flow- 
ers usually in cymose or umbellate groups on short lateral 
branches of the main inflorescence axis, an involucre 
absent. Flowers bisexual, perianth of 2 equal (or sub- 
equal) whorls of 3 parts (6 tepals), floral tube short or 
equalling the ovary; androecium of 9 fertile 4-thecous 
stamens, the outer 6 anthers (series I-II) with the upper 
thecae usually introrse and the lower thecae lateral-ex- 
trorse, often varying in dehiscence by bending and cur- 
vature of the stamen, interior 3 stamens (series III) usu- 
ally with the upper thecae opening laterally and the lower 
thecae extrorse, the 6 glands of the inner stamens much 
enlarged and extending outward to the periphery of the 
androecium, and closely adjacent to the outer stamens, 
staminodes (series IV) absent or minute; pistil with glo- 
bose to ellipsoid ovary, slender style and discoid stigma. 
Fruits borne in a cupulate receptacle, surface of the cup 
often pustulate or warty; berry usually ellipsoid. 

A genus of perhaps 25 species ranging from 
Guatemala to northern South America and the 
Amazon Basin. The glands are enlarged basally 
and come to lie between the outer stamens; ap- 
pearing to be associated with the outer stamens, 
though they originate at the base of the inner sta- 
mens. We believe that the presence of these en- 
larged glands in the periphery of the androecium 
and the unusual form of the stamens, argues against 
the submersion of Pleurothyrium under Ocotea, 
as advocated by Bernardi (1962) and Kostermans 
(1957). In some species the glands are connivent 
and difficult to interpret; or they may be lobed and 
appear to be more numerous than six. Rohwer 
and Kubitzki (1985) clarified the nature of the 
androecium, and their findings are consistent with 
our observations. 

The unusual bent stamens of Pleurothyrium and 
the unique pattern of anther dehiscence may be 
better understood in the context of Pleurothyrium 
golfodulcense, where the androecial elements are 
tightly compressed into a dome: dehiscence is out- 
ward from the surface of this dome. By becoming 
bent, all the valves seem capable of "apical" (or 
outward) dehiscence from the surface of the dome. 
Thus, small pollinators can work over the surface 
of the dome, rather than in and among the an- 
droecial elements. 



Key to Species of Pleurothyrium 

la. Leaves becoming more than 30 cm long, usually with an arcuate submarginal vein connecting the 
secondary veins distally; small trees of lowland rain forests . 2a 



114 



FIELDIANA: BOTANY 



Ib. Leaves never becoming more than 30 cm long, a distinct submarginal vein not developed . . . 3a 
2a. Leaves drying chartaceous and dark, usually obovate, petioles 4-8 mm long; inflorescences to 

65 cm long P. hexaglandulosum 

2b. Leaves drying subcoriaceous, grayish above and reddish brown beneath, petioles 15-25 mm 
long Pleurothyrium sp. A 

3a. Inflorescences becoming more than 1 cm long, lateral branches of the inflorescences with secondary 
branches; flowers ca. 6 mm broad, androecium not densely congested; hairs on stems usually less 
than 0.3 mm long; from below 400 m elevation Pleurothyrium trianae 

3b. Inflorescences to 10 cm long, the lateral branches often with only 1 flower; flowers ca. 10 mm broad, 
androecium densely congested; hairs on stems and petioles 0.2-0.5 mm long 4a 

4a. Leaves obovate to broadly oblong-obovate, leaves 6-15 cm broad, rounded and short-acuminate 
to caudate-acuminate at the apex; on the Caribbean slope and near the continental divide, 800- 
1 600 m elevation P . palmanum 

4b. Leaves elliptic to narrowly elliptic-oblong, leaves 4-7.5 cm broad, acute to long-acuminate at the 
apex; hills bordering Golfo Dulce below 500 m elevation P. golfodulcense 



Pleurothyrium golfodulcense W. Burger & N. Za- 
mora, sp. nov. Figure 9. 

Arbor ca. 10 m alta, ramulis foliiferis 2-4 mm crassis. 
Folia alterna, petiolis 1 1-28 mm longis, 1.3-2 mm cras- 
sis, hirsutulis; laminis 9-20 cm longis et 4-7.5 cm latis, 
elliptico-oblongis, elliptico-obovatis vel oblongo-obov- 
atis, apice breviter acuminato, subtus puberulis, nervis 
secondariis 5-9 paribus. Inflorescentiae racemosae, 3-9 
cm longae, pedunculis 2-4 cm longis. Flores ca. 5 mm 
longi et 8-12 mm lati, extus puberulenti, tepalis intus 
papillato-puberulis; stamina ca. 0.7 mm longa, antheris 
ca. 0.5 mm longis, congestis, staminodiis nullis, glan- 
dibus magnis; gynoecium angustum, ca. 2 mm longo. 
Fructus ignotus; cupula ca. 22 mm lata, 10 mm profun- 
da. 

Trees ca. 10 m tall, leafy branchlets 2-4 mm thick, 
densely yellowish brown or grayish brown hirsutulous 
with straight or slightly crooked ascending hairs 0.2-0.5 
mm long. Leaves alternate in a spiral or occasionally 
subopposite, petioles 1 1-28 mm long, 1.3-2 mm thick, 
terete, flattened or slightly sulcate above (adaxially), 
densely hirsutulous; leaf blades 9-20 cm long, 4-7.5 cm 
broad, elliptic-oblong to elliptic-obovate or oblong-ob- 
ovate, abruptly narrowed to a short-acuminate or cus- 
pidate apex (rarely obtuse or acute), the tip to 1.5 cm 
long, usually obtuse at the base (less often acute or slight- 
ly rounded) and often asymmetric, margin slightly re- 
volute (dry), drying stiffly chartaceous or subcoriaceous, 
flat dull and glabrescent on the upper surface but mi- 
nutely puberulent above the midvein, puberulent on the 
lower surface and slightly rough to the touch, sparsely 
or obscurely puberulent between the veins beneath, with 
5-9 major secondary veins on each side, central second- 
aries arising at angles of 40-60. Inflorescences solitary 
and axillary to undeveloped leaves near the shoot apex 
or between the older leaves, 3-9 cm long, racemose with 
lateral branches becoming 1-2 cm long and bearing only 
a single flower, peduncles 24 cm long, densely ferru- 
gineous puberulent, lateral branches subtended by per- 
sisting bracts ca. 4 mm long with 2 bracteoles often mid- 



way between the rachis and the flower, pedicels 2-7(-l 2) 
mm long. Flowers 8-12 mm broad, flower buds ca. 5 
mm long, densely puberulent on the outside, perianth 
becoming rotate, tepals ca. 6 mm long and 3-4 mm 
broad, papillate-puberulent within; androecium in the 
form of a dome 2.5 mm broad, stamens and glands 
tightly congested with the outer glands connivent and 
forming a peripheral ring around the androecium, sta- 
mens ca. 0.7 mm long and 0.4 mm broad, anthers ca. 
0.5 mm long, dehiscence essentially apical from the sur- 
face of the dome, glands and stamens closely appressed; 
pistil slender, ca. 2 mm long. Fruits borne in a warty 
cup 2 cm long, ca. 2.2 cm broad and 1 cm deep, margin 
slightly (1 mm) undulate with the persisting bases of the 
perianth parts, becoming pink; berry not seen. 



TYPE Costa Rica, Puntarenas Province, Alto 
de las Mogas, camino a Rincon de Osa, 14 Feb- 
ruary 1985, P. E. Sanchez, N. Zamora & M. Brenes 
1228 (holotype, CR; isotypes, F, usj; negative 61185, 
F). 

Trees of lowland rain forest formations around 
Golfo Dulce in southernmost Costa Rica, between 
100 and 900 m elevation. Flowering collections 
were made in late January and middle February; 
the January collection had a mature fruit cup. This 
taxon is known from only three collections: Allen 
5885 (us) from above Palmar Norte, Burger & 
Malta 4690 (CR, F) from the hills above Golfito, 
and the type. 

Pleurothyrium golfodulcense is recognized by its 
brownish puberulent stems and leaves, racemose 
few-flowered inflorescences, rotate flowers with 
tightly congested androecium, and deep warty fruit 
cup. The generally elliptic-oblong leaves are sim- 
ilar to many other Lauraceae, but the compacted 



BURGER: FLORA COSTARICENSIS 



115 



dome like androecium is unique among our species. 
The androecium is so compact that it is difficult 
to dissect, and may at first appear to be an ab- 
normality. This species is closely related to P. pal- 
manum. 



nent in this species. The two collections placed 
here differ in a number of ways; emphasis on the 
Costa Rican material accounts for the differences 
in the above description and the original descrip- 
tion. 



Pleurothyrium hexaglandulosum H. van der Werff, 
Ann. Missouri Hot. Gard. 75: 417. 1988. 

Small trees 5-6 m tall, leafy branches 2.5-6 mm thick, 
minutely appressed puberulent but quickly becoming 
(sub)glabrous, dark reddish brown, terete. Leaves alter- 
nate, petioles 4-8 mm long, 3-4 mm thick, appressed 
puberulent at first; leaf blades 27-46 cm long, 8-15 cm 
broad, narrowly elliptic-obovate to oblanceolate or nar- 
rowly elliptic, narrowed to the short acuminate apex, 
gradually narrowed to the base and abruptly rounded 
(subcordate) at the petiole, drying chartaceous or stiffly 
chartaceous, glabrous above with the major venation flat 
or immersed, glabrescent beneath or with minute ap- 
pressed hairs along the major veins, with 9-12 major 
secondary veins on each side, central secondaries arising 
at angles of 35-60, secondaries loop-connected near the 
margin (in the type collection) or only weakly loop-con- 
nected (in the Costa Rican collection). Inflorescences 
solitary and axillary to distal leaves, (20-)30-65 cm long, 
paniculate with lateral branches up to 25 cm long near 
the base and shorter distally, peduncle brownish puber- 
ulent or glabrous and dark brown, pedicels 8-20 mm 
long, minutely grayish puberulent. Flowers 7-9 mm 
broad, becoming rotate with reflexed tepals, greenish to 
yellowish, tepals equal and 3-4 mm long, minutely pu- 
berulent on the outside and minutely papillate-puberu- 
lent within (with lines of hairs often defined by pressure 
of glands and stamens in bud); glands large (1.5 mm 
broad) and forming a peripheral ring around the an- 
droecium, tightly appressed but not fused, the 9 stamens 
0.8-1.5 mm long with anthers raised above the glands, 
filaments with a few hairs along the back, anthers 0.5- 
0.7 mm long, 4-thecous with the valves on opposite sides 
and lateral dehiscence; pistil 1 .5 mm long, ovary broadly 
ovoid, 1-1.2 mm thick, style 0.2-0.5 mm long, stigma 
discoid. Fruits unknown. 

Understory trees of wet evergreen lowland rain 
forests, and known from only two collections. The 
Costa Rican collection was made on 3 March 1985, 
at 150 to 260 m elevation southwest of Rincon de 
Osa in southern Puntarenas Province (Croat & 
Grayum 59792, CR, F, MO). The type was collected 
near Portobelo, along the Rio Guanche at 50 m 
elevation in the Province of Colon, Panama 
(Hammel & Trainer 14781, MO). 

Pleurothyrium hexaglandulosum is distin- 
guished by its long, narrow, often oblanceolate, 
thin-textured leaves on very short petioles and 
drying dark brown. The large glands at the pe- 
riphery of the androecium are especially promi- 



Pleurothyrium palmanum (Mez & J. D. Smith) 
Rohwer, Mitt. Inst. Allg. Bot. Hamburg 20: 41. 
1986. Ocotea palmana Mez & J. D. Smith, Bot. 
Gaz. 33: 258. 1902. Figure 6. 

Trees to 1 5 m tall and 40 cm d.b.h., leafy branchlets 
3-7 mm thick, densely tomentulose with crooked (often 
appressed) brownish or yellowish brown hairs 0.2-0.5 
mm long. Leaves alternate, petioles 1 5-25 mm long, 1 .8- 
5 mm thick, densely ferruginous tomentulose; leaf blades 
(ll-)15-28 cm long, (6-)7-15 cm broad, obovate to 
broadly oblong-obovate or less often oblong, abruptly 
rounded at the apex and bluntly obtuse to short-acu- 
minate (occasionally caudate-acuminate as in Barbour 
1012), gradually narrowed to the cuneate or obtuse and 
equal or unequal base, drying stiffly chartaceous (sub- 
coriaceous), upper surface drying dark brown and ap- 
pressed puberulent above the major veins, lower surface 
usually drying pale grayish brown or yellowish brown 
and usually densely appressed puberulent over the entire 
surface, with 5-8 major secondary veins on each side, 
the central secondaries arising at angles of 45-60. In- 
florescences axillary to distal leaves or subterminal, 4- 
10 cm long, paniculate or racemose, few (<20) flowered, 
peduncles 1-4 cm long, 0.8-2.3 mm thick, densely fer- 
ruginous puberulent, pedicels 1-10 mm long, bracteo- 
late. Flowers 5-6 mm long and 10-12 mm broad, peri- 
anth densely appressed puberulent on the outside and 
minutely puberulent on the inside, outer tepals to 4.5 
mm long; stamens ca. 1 mm long, closely appressed and 
difficult to interpret, anthers usually bent and with out- 
ward dehiscence, filaments very short, glands of the inner 
stamens enlarged and forming part of the periphery of 
the androecium (between the outer stamens), staminodes 
not seen; pistil papillose (in the type) or puberulent (in 
Barbour 1012). Fruits unknown. 

Trees of evergreen lower montane rain forest 
formations on the Caribbean slope and along the 
continental divide in the central highlands, from 
about 800 to 1 600 m elevation. Only two flowering 
collections have been made: Tonduz 12652 (us, 
the type) collected near La Palma in September, 
and Barbour 1012 (F, NY) from east of Turrialba 
in May. This species is only known from central 
Costa Rica. 

Pleurothyrium palmanum is recognized by the 
larger obovate leaves densely appressed puberu- 
lent beneath and usually cuneate at the base, the 
large puberulent flowers with glands visible at the 
outer periphery of the androecium, and its re- 



116 



FIELDIANA: BOTANY 



stricted ecological range. This species can be con- 
fused with Ocotea valeriana and especially O. 
pseudopalmana, but they tend to grow at higher 
elevation, have darker brown puberulence, and 
have very different flowers. Additional sterile col- 
lections belonging to this species are Holdridge 
6690 (CR, NY) from Tres Marias (Barba) and W. 
& H. Rowlee 210& 233 (F, us) from near La Palma. 
Floral characteristics, pubescence, and geography 
indicate a close relationship with P. golfodulcense. 



Pleurothyrium trianae (Mez) Rohwer, Mitt. Inst. 
Allg. Bot. Hamburg 20: 43. 1986. Nectandra 
trianae Mez, Konigl. Bot. Gart. Berlin 5: 439. 
1889. Figure 9. 

Trees, probably small or medium-sized, leafy branch- 
lets 2-4 mm thick, at first minutely (0.1-0.2 mm) pu- 
berulent with thin brownish ascending hairs, becoming 
(sub)glabrous and dark, terete. Leaves alternate, petioles 
12-18 mm long, 1.2-2.1 mm broad, flat or slightly sul- 
cate above with 2 adaxial margins, minutely puberulent 
but becoming (sub)glabrous; leaf blades 10-16 cm long, 
3-6 cm broad, oblong to elliptic-oblong, acute or short- 
acuminate at the apex, acute to obtuse at the base, drying 
stiffly chartaceous (almost subcoriaceous) and grayish 
brown above, flat and essentially glabrous above, mi- 
nutely and obscurely brownish puberulent beneath and 
soft to the touch with slender hairs 0.1-0.3 mm long, 
with 5-9 major secondary veins on each side, central 
secondaries arising at angles of 35-60. Inflorescences 
apparently axillary to undeveloped leaves among fully 
developed leaves along the distal stems, to 1 3 cm long, 
paniculate with short (2-3 cm) lateral branches, flowers 
in cymose groups on the lateral branches or secondary 
branches, flowers in cymose groups on the lateral branch- 
es or secondary branches, peduncles 3-6 cm long, 1-1.7 
mm thick, pedicels 3-5 mm long, slender and minutely 
(0.2 mm) puberulent. Flowers ca. 4 mm long and 6 mm 
broad, the floral tube ca. 0.7 mm deep, tepals ca. 3 mm 
long and oblong, becoming rotate, puberulent on the 
outside and papillate-puberulent within; outer anthers 
ca. 0.6 mm long and 0.6 mm broad with dehiscence of 
the upper thecae either introrse or lateral and the lower 
thecae extrorse, glands ca. 0.5 mm thick; pistil 2 mm 
long, the slender style 0.7 mm long and with a discoid 
stigma. Fruits and fruiting receptacle not seen. 

Trees of the evergreen lowland Caribbean rain 
forest formations, flowering in late March. The 
only Central American collection was made among 
fallen vegetation overhanging the Rio San Juan, 
between San Juan del Norte (Greytown) and Delta 
de San Juan (Bunting & Licht 872, F, NY). This 
locality is between and 50 m elevation, along 
the border of Costa Rica and Nicaragua. The 
species ranges to northern South America. 

Pleurothyrium trianae is recognized by its ob- 



long grayish brown leaves slightly puberulent and 
soft to the touch beneath, the panicles with short 
lateral branches, the puberulent flowers with con- 
spicuous glands around the periphery of the an- 
droecium, and the unusual modes of dehiscence. 
It may be that the androecium is more tightly 
compacted in life and dehiscence is, in effect, api- 
cal from the outer surface of the domelike an- 
droecium. This species appears similar to Pleu- 
rothyrium zulianense Lasser and P. amapaense C. 
K. Allen of South America. 



Pleurothyrium sp. A. 

Small trees, ca. 8 m tall and 8 cm d.b.h., leafy branch- 
lets appressed puberulent, 4-6 mm thick, dark brown, 
terete. Leaves alternate, petioles 15-25 mm long, 2.5- 
4.5 mm thick, with 2 adaxial ridges and flattened above; 
leaf blades 17-47 cm long, 8-16 cm broad, oblong to 
elliptic-oblong, tapering to an acuminate apex, the tip 
1 5-30(?) mm long, rounded to obtuse at the base, drying 
subcoriaceous, smooth, glabrous and grayish above with 
the midvein raised and the secondaries impressed, gla- 
brescent beneath and reddish brown but minutely (0. 1- 
0.2 mm) puberulent on the midvein, with 10-17 major 
secondary veins on each side, a well-defined arcuate sub- 
marginal vein beginning in the lower fourth of the lam- 
ina, 2-6 mm from the lamina-margin and uniting the 
distal ends of the secondaries, central secondaries arising 
at angles of 55-70. Inflorescences and flowers un- 
known. Fruits borne in a small hemispheric cup 6-8 mm 
long and ca. 1 2 mm broad, drying dark with conspicuous 
(1 mm) lenticels on the outside, margin narrow and en- 
tire, pedicel 12-15 mm long and 2-3 mm thick, also 
lenticellate; berry ca. 2 cm long and apparently ellipsoid, 
dark purple when ripe. 

This species is only known from G. Proctor Coo- 
per 539, near Almirante, Bocas del Toro, Panama. 
This fruiting specimen is labeled Jan.-March 1 928. 
The specimen differs from all our other Costa Ri- 
can Lauraceae by the large leaves with clearly de- 
veloped, arcuate, submarginal veins. Except for 
Pleurothyrium, such venation is rare in Lauraceae 
(but compare our material assigned to Endlichena 
sp., with much smaller, thinner leaves). The len- 
ticellate cups strongly suggest that Pleurothyrium 
is the correct genus for this collection (Bernardi in 
herb.; Rohwer, pers. comm.). 



Povedadaphne Burger 

Medium-sized trees, bisexual, sparsely puberulent, ev- 
ergreen. Leaves alternate, pinnately veined and with pit 
domatia. Inflorescences lacking an involucre of bracts in 
early stages, solitary and axillary to distal leaves, pani- 



BURGER: FLORA COSTARICENSIS 



117 



culatc. Flowers bisexual, perianth of 6 equal parts in 2 
whorls of 3; androecium of 9 fertile stamens, the outer 
6 stamens (series I-II) appearing as a single whorl, the 
3 inner stamens (series III) similar to the outer in size 
and form, stamens thick and the filament not clearly 
differentiated, puberulent, with a flattened glabrous dis- 
tal surface and 4 minute valves dehiscing apically from 
this distal surface. 6 glands present between the inner 
and outer stamens, staminodes absent, a shallow floral 
cup present; pistil slightly narrowed at the base, ovary 
ellipsoid, style slender and equaling or exceeding the 
ovary in length, stigma simple. Fruits borne on a thick- 
ened pedicel, the receptacle only slightly expanded be- 
neath the base of the fruits; berry becoming globose or 
pyriform, seed large. 

The single species is very similar to some of our 
species of Ocotea (such as O. whitei) in the form 
of its flowers, panicles, and foliage. However, the 
nine stamens dehiscing apically by four minute 
distal pores is a unique condition in Lauraceae, 
and the poorly developed cup beneath the large 
fruits is also unusual. This unique species and ge- 
nus is only known from Costa Rica. (Compare 
Williamodendron, which has only three stamens 
of rather similar form.) The genus is named in 
honor of Luis Poveda, who has added much to 
our understanding of the Lauraceae in Costa Rica 
and whose knowledge of Costa Rican trees is un- 
equaled. 



Povedadaphne quadriporata W. Burger, Brittonia, 
40: 277. 1988. Figure 22. 

Trees 6-20 m tall, trunks to 45 cm thick, sometimes 
with sprout-shoots from the base, leafy branchlets 1.5- 
4 mm thick, glabrous or minutely appressed puberulent 
with thin ascending hairs, at first with narrow longitu- 
dinal ridges but becoming terete, dark grayish. Leaves 
alternate, petioles 4-10 mm long but poorly defined be- 
cause of the decurrent lamina-base, 1.2-2.3 mm broad, 
usually glabrous, with thin lateral margins and sulcate 
near the base; leaf blades 4-12.5 cm long, 2-4(-5.5) cm 
broad, elliptic-obovate to elliptic-oblong or oblong-ob- 
ovate, short acuminate at the apex or occasionally ob- 
tuse, gradually narrowed to the cuneate or attenuate base 
and usually decurrent on the petiole, drying subcoria- 
ceous (stiffly chartaceous) and often olive green or brown, 
glabrous above with the major veins slightly elevated, 
glabrous or very sparsely puberulent beneath, axils of 
larger secondary veins usually with conspicuous pit dom- 
atia ca. 1 mm broad and filled with minute whitish hairs, 
with 3-6 major secondary veins on each side, central 
secondaries arising at angles of 30-45, tertiary veins 
slightly raised beneath. Inflorescences solitary in axils of 
distal leaves. 9-1 3 cm long, paniculate with short (3 cm) 
lateral branches, peduncles 4-7 cm long, minutely ap- 
pressed puberulent, flowers often in 3-flowered distal 
cymules, pedicels ca. 2 mm long. Flowers 2.5-3 mm long 
and equally broad, apparently urceolate (campanulate in 



appearance when pressed), minutely sericeous on the 
outside, tepals broadly obtuse and glabrous within; fer- 
tile stamens 9, thick and hairy (a slender filament absent), 
all stamens dehiscing by 4 small pores at the top, stamens 
1.2-1.4 mm long and 0.6 mm broad at the top, the 6 
short-stipitate glands arising exterior to the inner sta- 
mens, ca. 0.6 mm high and equally broad, staminodes 
absent; pistil ca. 2 mm long, ovary ellipsoid and 0.7 mm 
thick, style narrow and 0.9 mm long, stigma simple. 
Fruits borne on a thickened (8 mm) pedicel with the 
receptacle slightly expanded (10-17 mm broad) and 1- 
3 mm deep below the mature fruits; berry oblong in 
development (3 x 2 cm in Chacon 201) with an unusual 
warty surface but becoming globose to pyriform, to 8 cm 
long and 6 cm in diameter, seed reddish purple in cross 
section. 

Trees of the very wet premontane rain forest 
formations of the Caribbean slope, between 200 
and 1000 m elevation. Flowers were collected in 
June (Poveda & Castro 3561, CR, usj, the types), 
immature flowers in July (Chacon et al. 1980, CR). 
Small fruits were collected in January (Chacon 201, 
CR), large fruits in February (Poveda & Holdridge 
2360, CR, NY) and August (D. Smith et al. 1202, 
DUKE, F). The species has been collected in the 
Reserva Forestal de San Ramon, near San Carlos, 
and near Ciudad Quesada in Alajuela Province, 
and just east of the Rio Sarapiqui in Heredia Prov- 
ince. It is not known outside of this restricted area 
in Costa Rica. 

Povedadaphne quadriporata is recognized by the 
small stiff slightly lustrous brownish or olive green 
leaves (dry), with conspicuous small pit domatia 
(filled with white hairs) in the vein axils of the 
lower surfaces, and the decurrent leaf base. The 
large fruits with poorly developed receptacle, and 
flowers with nine hairy stamens dehiscing by four 
little pores at the top, make this species unique 
among known Lauraceae. The foliage resembles 
Ocotea bijuga, O. oblonga, and when dried, O. 
whitei. 



Williamodendron Kubitzki & Richter 

REFERENCE K. Kubitzki & H. G. Richter, Wil- 
liamodendron Kubitzki & Richter, a new genus of 
Neotropical Lauraceae. Bot. Jahrb. Syst. 109: 49- 
58. 1987. 

Moderate-sized to tall trees, branchlets at first puber- 
ulent, becoming grayish. Leaves alternate or closely con- 
gested (whorled), usually clustered at the ends of branch- 
lets, petioles rounded or slightly canaliculate above; leaf 
blades usually large and obovate, pinnately veined. In- 
florescences solitary and axillary, paniculate with short 
lateral branches (thyrso-paniculate), bracts present, brac- 



118 



FIELDIANA: BOTANY 



teoles subtending the pedicels. Flowers small and subgl- 
obose, bisexual, perianth of 6 parts in 2 whorls, tepals 
unequal, the outer tepals broadly triangular and the inner 
more obtuse; androecium of 3 fertile stamens (series III), 
stamens sessile (a narrow filament absent), each anther 
dehiscing distally and extrorsely with 4 small rounded 
valves, glabrous on the exterior and pilose within, stam- 
inodia 3 (series IV), small and lanceolate, floral cup 
subglabrous; pistil glabrous, style absent or short, stigma 
minute. Fruits and fruiting receptacle unknown. 

A genus of two species, one found in Colombia 
and the Amazon basin, and the other in Costa 
Rica. The genus is distinguished by its androecium 
of three functional stamens, each dehiscing by four 
small distal valves. The relatively large obovate 
leaves clustered at the ends of branchlets is also 
distinctive. Williamodendron is related to Mezi- 
laurus but differs in having 4-valved stamens and 
different bark structure, and in lacking the silica 
grains in seriated ray cells unique to the wood of 
Mezilaurus. While flower structure may resemble 
Licaria, that genus has very different wood (see 
reference cited above). The genus was named in 
honor of William Rodriguez for his achievements 
in Amazonian forest botany. 



Williamodendron glaucophyllum (van der Werff) 
Kubitzki & Richter, Bot. Jahrb. Syst. 109: 58. 
1987. Mezilaurus glaucophylla van der Werff, 
Ann. Missouri Bot. Card. 74: 164. 1987. Figure 
19. 

Trees 8-25 m tall, bark smooth and grayish, leafy 
branchlets 5-8 mm in diameter, becoming glabrous and 
grayish, petiole scars 3-4 mm broad and conspicuous. 
Leaves alternate to subopposite (whorled) and often 
closely clustered at the ends of stems, petioles 2-8 cm 
long, 1.5-3 mm thick, grayish with minute (0. 1-0.2 mm) 
appressed hairs, flat or slightly rounded above, the lateral 
(or adaxial) margins weakly developed, thickened near 
the base; leaf blades 14-29 cm long, 6.5-15 cm broad, 
obovate to oblong-obovate, abruptly narrowed or round- 
ed at the obtuse to acute (short-acuminate) apex, grad- 
ually tapering to the acute and often unequal base, drying 
chartaceous, minutely puberulent on the veins above but 
glabrous on the other surfaces above, very minutely (0. 1 
mm) puberulent on the lower surface (especially on the 
veins), conspicuously grayish glaucous on the lower sur- 
face, with 9-14 major veins on each side, central sec- 
ondaries arising at angles of 50-80, tertiary veins ob- 
scure beneath. Inflorescences axillary to leaves, 12-20 
cm long, paniculate with short lateral branches to 2.5 
cm long, the flowers often borne in small cymules of 1- 
3 flowers, peduncle to 3 cm long, ca. 1 mm thick, mi- 
nutely grayish puberulent, pedicels 1-2 mm long, slen- 
der, slightly puberulent. Flowers ca. 1.5 mm long and 
1.5-2 mm broad, white, perianth essentially glabrous, 
outer perianth whorl shorter than the inner whorl, outer 



tepals broadly ovoid or deltoid; fertile stamens 3 (but 1 
sometimes poorly developed), ca. 1 mm long and 0.8 
mm thick, puberulent, free, 4-thecous but the thecae 
sometimes difficult to see on the apex of the anther, 
opening by 4 small circular valves, 3 staminodia present 
with sagittate apices and ca. 0.8 mm long, puberulent; 
pistil ca. 1 .2 mm long, ovary 0.5-0.8 mm long, glabrous, 
stigma simple and acute. Fruits and fruiting receptacles 
unknown. 

Trees of evergreen rain forests on the Pacific 
slope of Costa Rica, from about 50 to 500 m el- 
evation. The first collection was made at Masatal 
de Puriscal (Zamora & Poveda 1014, CR, F, MO, 
the type) with flowers in July. Two additional col- 
lections from the Osa Peninsula are Hammel et 
al. 15214 (CR, MO) and Burger et al. 12348 (CR, F, 
MO, NY). The species is only known from these 
three collections in south-central and southern- 
most Costa Rica, in the provinces of San Jose and 
Puntarenas. 

Williamodendron glaucophyllum is recognized 
by its large thin obovate leaves, glaucous on the 
undersurfaces and clustered near the ends of 
branchlets, long petioles, and the very small flow- 
ers on longer racemose panicles. The small flowers 
with only 3 thick stamens are like those of Licaria, 
but differ in having 4-valved anthers. The small- 
flowered inflorescences in among the clustered 
leaves on tall trees may be why this species had 
not been collected before 1985. These new dis- 
coveries are being made by skilled botanists using 
binoculars and special equipment to collect the 
tall trees of the rain forest. 



A Species of Uncertain Generic Position 

Trees to over 1 m tall, leafy branchlets 2-4 mm thick, 
densely pale grayish tomentulous with curved and mat- 
ted hairs ca. 0.2 mm long, remaining puberulent for some 
time, terete. Leaves alternate, distant, petioles 1 8-30 mm 
long, 1.9-2.7 mm thick, densely grayish tomentulous; 
leaf blades 10-19 cm long, 5-7.5 cm broad, narrowly 
ovate to ovate-elliptic or ovate-oblong, tapering to a 
short-acuminate apex, obtuse to rounded at the base, the 
sides of the lamina unequal at the base with the sides 2- 
6 mm distant on the petiole, drying stiffly chartaceous 
or subcoriaceous, the upper surface becoming glabrous 
and lustrous but with hairs above the slightly impressed 
major veins, tertiary venation slightly elevated, lower 
surface densely yellowish gray or whitish gray tomen- 
tulous, the hairs minute (0.1-0.3 mm) and curved, with 
3-5 major secondary veins on each side, the basal sec- 
ondaries often strongly ascending, central secondaries 
arising at angles of 35-50. Inflorescences not seen at 
maturity (only 5 cm long), solitary and axillary to distal 
leaves or undeveloped leaves near the shoot tip, pani- 
culate with short lateral branches subtended by con- 



BURGER: FLORA COSTARICENSIS 



119 



spicuous (4-7 mm) oblong bracts, peduncle, rachis and Caryodaphnopsis burgeri Zamora & Poveda .... 

bracts densely brownish gray tomentulous. Flowers not CAR burg 

seen at anthesis, flower buds ca. 3 mm long, the outer Cassytha fiiiformis L. . CAS fili 
tepals only sparsely puberulent distally, apparently with 
a normal floral configuration of 6 tepals and 9 fertile 

stamens; a dissection by C. K. Allen showed stamens Endlichena sp.? ?END sp. 

with Nectandra-\ike arrangement of the thecae with a 

prolonged connective and 3 staminodes, another dissec- Licaria brenesii W. Burger LIC bren 

tion by Burger failed to confirm these observations be- cufodontisii Kosterm LIC cufo 

cause the flower was immature. Fruits borne in a well- 

developed cup about 12 mm broad at the top; berry Licaria excelsa Kosterm LIC exce 

ellipsoid-oblong (measurements not available). Licaria multinervis Kurz LIC mult 

Licaria pergamentacea W. Burger LIC perg 

Trees of montane evergreen forest formations Licaria sarapiquensis Hammel LIC sara 

from 1600 to 2300 m elevation, presently known Licaria triandra (Sw.) Kosterm LIC tria 

only from the Pacific slope of the Cordillera de Licaria sp. A LIC sp. A 

Talamanca and adjacent Chiriqui highlands. Im- Litsea glaucescens H.B.K LIT glau 

mature flowers were collected in July (Stern et al. 

7722, NY), and fruits were collected in March Nectandra belizensis (Lundell) C. K. Allen 

(Comacho 7/3/85, seen at CR). The species ranges NEC beli 

from near Division, in the west, to Boquete, Pan- Nectandra cissiflora Nees NEC ciss 

ama. Nectandra cufodontisii (Schmidt) C. K. Allen . . . 

This species is distinguished by the dense gray- NEC cufo 

ish tomentum on the underside of the leaves; no Nectandra globosa (Aubl.) Mez NEC glob 

other species of Lauraceae in our area has such an Nectandra hypoleuca Hammel NEC hypo 

indumentum. The long petioles and relatively few Nectandra kunthiana (Nees) Kosterm 

major secondary veins also make the leaves dis- NEC kunt 

tinctive. If C. K. Allen's dissection is correct, this Nectandra latifolia (H.B.K.) Mez NEC lati 

species would be placed in Nectandra, but we need Nectandra longipetiolata van der Werff 

better material to be sure. NEC long 

Nectandra martinicensis Mez NEC mart 

Nectandra membranacea (Sw.) Griseb 

NOTE ADDED IN PROOF: Gerardo Herrera has NEC memb 

recently collected a very unusual, apparently mon- XTCr , 

T. / 1 -.10 Nectandra mtida Mez NEC niti 

oecious, species in northern Costa Rica (122s, CR, XT . c ,, xrcr- ~, 

Nectandra ramonensis Standl NEC ramo 

F, MO). The male flowers have a slender puberulent XT . . t ,_ . p , . x , 

... Nectandra reticulata (Ruiz & Pavon) Mez 

stammal column with 3 valves on the 3-sided apex; NEC t' 

a pistillode and staminodes are absent. '.' -V \- Mtrr> \f 

Nectandra sahcifoha (H.B.K.) Nees . . . NEC salf 

Nectandra salicina C. K. Allen NEC sali 

List of Accepted Species Nectandra sinuata Mez NEC sinu 

Nectandra turbacensis (H.B.K.) Nees 

Newly described species are in boldface. NEC turb 

Aiouea costaricensis (Mez) Kosterm.. . AIO cost Ocotea atirrensis Mez & J. D. Smith. . OCO atir 

Aiouea obscura van der Werff AIO obsc Ocotea aurantiodora (Meissner) Mez 

Aiouea talamancensis W. Burger AIO tala OCO aura 

Aiouea ?sp AIO ?sp. Ocotea austinii C. K. Allen OCO aust 

Aniba venezuelana Mez ANI vene Ocotea babosa C. K. Allen OCO babo 

Ocotea sp. aff. bijuga (Rottboel) Bernardi 

Beilschmiedia anay (Blake) Kosterm. . BEI anay OCO biju 

Beilschmiedia brenesii C. K. Allen . . . BEI bren Ocotea brenesii Standl OCO bren 

Beilschmiedia ovalis (Blake) C. K. Allen Ocotea calophylla Mez OCO calo 

BEI oval Ocotea sp. aff. caracasana (Nees) Mez 

Beilschmiedia pendula (Sw.) Hemsley OCO cara 

BEI pend Ocotea cernua (Nees) Mez OCO cern 

Beilschmiedia sulcata (Ruiz & Pavon) Kosterm. Ocotea dendrodaphne Mez OCO dend 

. BEI sulc Ocotea dentata van der Werff . . . OCO dent 



120 



FIELDIANA: BOTANY 



Ocotea endresiana Mez OCO endr Phoebe hammeliana W. Burger .... PHO hamm 

Ocotea floribunda (Sw.) Mez OCO flor Phoebe neurophylla Mez & Pittier . . . PHO neur 

Ocotea glaucosericea Rohwer OCO glau Pleurothyrium golfodulcense W. Burger & N. 

Ocotea gomezii W. Burger OCO gome Zamora PLE golf 

Ocotea hartshorniana Hammel OCO hart Pleurothyrium hexaglandulosum van der Werff 

Ocotea helicterifolia (Meissn.) Hemsl PLE hexa 

OCO heli Pleurothyrium palmanum (Mez & J. D. Smith) 

Ocotea holdridgeiana W. Burger OCO hold Rohwer PLE palm 

Ocotea insularis (Meissn.) Mez OCO insu Pleurothyrium trianae (Mez) Rohwer . . PLE tria 

Ocotea laetevirens Standl. & Steyerm Pleurothyrium sp. A PLE sp. A 

OCO laet Povedadaphne quadriporata W. Burger 

Ocotea lentii W. Burger OCO lent POV quad 

Ocotea leucoxylon (Sw.) Laness OCO leuc 

Ocotea meziana C. K. Allen OCO mezi Williamodendron glaucophyllum (van der Werff) 

Ocotea mollicella (Blake) van der Werff Kubitzki & Richter WIL glau 

OCO mole 

Ocotea mollifolia Mez & Pittier OCO molf A species of uncertain generic position 

Ocotea monteverdensis W. Burger . . OCO mont SP. ?gen 

Ocotea nicaraguensis Mez OCO nica 

Ocotea oblonga (Meissn.) Mez OCO oblo 

Ocotea paulii C. K. Allen OCO paul Index to Exsiccatae 

Ocotea pittieri (Mez) van der Werff. . . OCO pitt 

Ocotea pseudopalmana W. Burger OCO pseu Lauraceae of Costa Rica and Adjacent Panama 

Ocotea puberula (Rich.) Nees OCO pubr 

Ocotea rivularis Standl. & L. O. Williams Acronyms refer to genera and species (see p. 120). 

OCO rivu 

Ocotea skutchii C. K. Allen OCO skut Allen, P. H.: 1001 PER vera; 1015 PER caer; 1211. 

Ocotea stenoneura Mez & Pittier OCO sten OCO paul; 1486 OCO whit; 1570 OCO insu; 

Ocotea tenera Mez & J. D. Smith ex Mez 1572 OCO glau; 1611 OCO vera; 1740 OCO 

OCO tene sp. A; 3439 OCOatir; 3482 LICexce; 4672 NEC 

Ocotea valeriana (Standl.) W. Burger cufo; 4846 OCO glau; 4856 PER vera; 4883 

OCO vala PERobtu; 5038 PER vera; 5251 NECsalf; 5503 

Ocotea valerioides W. Burger OCO valo OCO nica; 5552 PER amer; 5590 OCO rivu; 

Ocotea veraguensis (Meissner) Mez . . OCO vera 5629 NEC salf; 5645 OCO insu; 565 1 , 5658 LIC 

Ocotea viridiflora Lundell OCO viri cufo; 5824 OCO molf?; 5861 OCO cern; 5885 

Ocotea wedeliana C. K. Allen OCO wede PLE golf; 5950 LIC perg; 6654 LIC cufo. 

Ocotea whitei Woodson OCO whit Almeda, F., et al.: 3 146 NEC salf; 3762 OCO pitt; 

Ocotea sp. A aff. laetevirens OCO sp. A 3776 OCO aust; 4024 OCO dend; 4138 NEC 

Ocotea sp. B OCO sp. B memb; 45 1 7 PER schi. 

Persea albida Kosterm PER albi Baker, R. & W. Burger: 1 1 7 OCO atir. 

Persea americana Miller PER amer Barbour, W. R.: 1011, 1012 PLE palm; 1019 BEI 

Persea brenesii Standl PER bren pend?; 1032 LIC mult; 1033 OCO sten. 

Persea caerulea (Ruiz & Pavon) Mez. . PER caer Bawa, K. S.: 127 OCO vera; 2055 NEC glob. 

Persea obtusifolia Kopp PER obtu Bernardi, L.: 10498, 10512 NEC memb; 10626 

Persea povedae W. Burger PER pove PHO pitt; 10631 NEC memb. 

Persea rigens C. K. Allen PER rige Biolley, P.: 108 1 , 1 282 OCO vera; 2226 NEC sinu. 

Persea schiedeana Nees PER schi Blum, K. E.: 336 NEC sali; 1409 NEC turb. 

Persea silvatica van der Werff PER silv Brenes, A. M.: s.n. (1902) NEC glob; 337 (178) 

Persea veraguasensis Seem PER vera PER pove; 353 (478) NEC ramo; 362 OCO vera; 

Persea vesticula Standl. & Steyerm. . . . PER vest 371 (515) NEC sali; 377 (538) PHO bren; 3617 

Phoebe brenesii Standl PHO bren OCO tene; 3850 NEC memb; 4059 OCO insu; 

Phoebe chavarriana Hammel PHO chav 4061 NEC memb; 4160 OCO vera; 4185 OCO 

Phoebe cinnamomifolia (H.B.K.) Nees vala; 4200 OCO tene; 4206, 4262, 4272 NEC 

PHO cinn sali; 4344 NEC memb; 4533 OCO tene; 4626 



BURGER: FLORA COSTARICENSIS 



121 



OCO insu; 4439 OCO tene: 4628 OCO insu; 
466 1 (446) NEC sali; 4672 NEC reti; 4762 PHO 
bren; 4769 PER pove; 4780 NEC memb; 4793 
PER amer; 4794 NEC memb; 4877, 4889 LIC 
tria; 4896 PHO bren: 4954 NEC sinu; 496 1 LIC 
tria; 50 1 7 NEC sinu; 50 1 8 NEC reti; 50 1 9 NEC 
memb; 5023, 5066 LIC tria; 5067 NEC sinu; 
5068 NEC memb; 5075 PER amer; 5206 OCO 
insu; 5317 PER amer; 5334, 5384 OCO pitt; 
5389 PER amer; 5393 NEC sali; 5405, 5416 
OCO pitt; 5443 NEC memb; 5455 NEC sinu; 
5489 LIC tria; 55 18 PHO bren; 5535 OCO bren; 
5590 OCO paul; 5714 NEC memb; 5719 PHO 
cinn; 5733 NEC memb; 5826 NEC mart; 5831 
NEC memb; 5846 OCO mezi; 5927 PER rige; 
6099 OCO vala; 62 1 4 BEI pend; 6314 PER pove; 
6317 OCO endr; 6342 PER pove; 6346 OCO 
mezi; 6578, 6586 NEC reti; 6603 OCO vera; 
6605 BEI pend; 6612 NEC ramo; 6650 NEC 
sali; 6660 NEC ramo; 6675 PHO bren; 6704 
NEC sali; 6714 NEC memb; 6729 OCO insu; 
6770 PER amer; 6810 PHO bren; 6817 NEC 
sinu; 6825 NEC sali (or N. smithii); 6834, 9287 
NEC ramo; 9713 NEC memb; 12163 NEC salf 
(or N. niti?); 1 2262 LIC cufo; 1 2278 NEC glob?; 
12314 NEC niti; 12628 OCO vera; 13506 PER 
amer; 13523 LIC bren (the type); 13653 OCO 
bren: 1 3664 NEC glob?; 14288 OCO vera; 14296 
NEC mart; 14403 LIC tria; 14988 NEC ramo; 
1 5090 NEC sinu; 1 5534 OCO vera; 15617 NEC 
mart; 15658 OCO insu; 1 6202 PER amer; 16224, 
16227 OCO endr; 16824 NEC mart; 16833 NEC 
memb; 1 6943 PHO bren; 1 7005 NEC reti; 1 70 1 8 
NEC ramo; 1 7048 PHO bren; 17172 OCO endr; 
17173 PER amer; 1 7457 OCO vera; 1 7722 NEC 
reti; 18942 NEC sali; 19203 NEC mart; 20332 
OCO whit?; 20499 OCO insu; 20539 OCO atir; 
20560 PHO cinn; 2 1 644 OCO atir; 2 1 990 NEC 
ramo; 22563 OCO insu; 22644 OCO atir. 

Brown, C. A.: 138 NEC turb; 17393 OCO hold. 

Bunting, G., & L. Licht: 838, 866 OCO cern; 872 
PLE tria; 1083 OCO aura (Nicaragua). 

Burch, D.: 4589 OCO dend. 

Burger, W., et al.: 3890 NEC memb; 4069, 4101 
OCO vera; 4199 OCO atir; 4236 OCO tene; 
4448 LIC exce; 4690 PLE golf; 4792 PER amer; 
5394 NEC memb: 585 1 OCO dend; 59 1 2 OCO 
dend; 6102 OCO pitt: 6955 NEC memb.; 7045 
OCO vala; 7311 OCO nica; 7405 PER vest; 
7698, 7720 NEC cufo; 7904 OCO cufo; 8602, 
8611 OCO insu; 8665 OCO viri; 8726, 8776 
OCO insu; 8871 OCO cern; 8881 OCO heli; 
9091 PHO cinn; 9282 OCO dend; 9669 OCO 
insu; 9684 PHO cinn; 9852. 10368 OCO atir; 



10501, 10645 PER amer; 10688 NEC sali; 
10718, 11170 LIC bren; 11181, 11248 OCO 
dend; 1 1 690 OCO atir; 1 1 730 OCO molf; 1 1 966 
NEC sali; 11979 PER amer; 12026 ?END sp.; 
1 2028 OCO dend; 1 2032 OCO cern; 1 2043 OCO 
dend; 12045 OCO cern; 1 2060 OCO pitt; 12063 
OCO aust; 12085 PER vera; 12086 NEC sinu; 
1 2088 BEI pend; 1 2096 OCO whit; 1 2097 OCO 
vala; 12098, 12099 OCO glau; 12101, 12103 
OCO laet; 12104 OCO endr; 12106 OCO bren; 
12110, 121 18OCO atir; 12121 OCO paul; 12128 
OCO insu; 1 2 1 44 OCO gome; 12150 OCO pseu?; 
12 164 OCO endr; 12172 PER vest; 121 73 PER 
schi?; 12174 OCO dent; 12176 PHO cinn; 12177 
OCO skut?; 12178 BEI oval; 12180 NEC cufo; 
12181 OCO insu; 12182 PER albi; 12 183 NEC 
cufo; 12184 BEI oval; 12185 PHO cinn; 12186 
NEC cufo; 12187 NEC ciss; 12189 BEI oval; 
1 2 1 90 PHO cinn?; 12191 OCO insu; 12193 PHO 
cinn: 12197 ?GENUS?; 1 2 1 99 OCO whit; 1 2200 
OCO vera; 12201 NEC ramo; 12203, 12206 
NEC salf; 12207 LIC exce; 12221 OCO atir; 
12256 OCO leuc; 12334, 12336 LIC cufo; 12337 
OCO pubr; 12341 NEC glob; 12348 WIL glau; 
12356 PHO neur; 12366 LIC perg; 12376, 12377 
OCO sp. B; 12416 OCO mezi; 12418 LIC sara; 
12430 POV quad; 12451, 12467 OCO pseu?. 

Carlson, M.: 3228, 3384, 3392 PER amer; 3584, 

3623 OCO pitt. 
Carvajal, A.: 50 OCO insu; 94 OCO mezi; 103 

OCO insu; 208 NEC sali; 210 PHO cinn. 
Chacon, I.: 426 OCO dend; 600 NEC memb; 704 

OCO hart; 724 OCO dend; 1101 NEC hypo; 

1113 OCO dend; 1 1 29 NEC memb; 1 242 OCO 

nica; 1295 OCO molf; 1327 OCO tene; 1348 

OCO dend; 1418 OCO cern; 1557 OCO vala; 

1574 AIO tala; 1587 AIO cost; 1622 OCO atir; 

1634 OCO valo; 1680 OCO dend; 1722 AIO 

?sp.; 1798 OCO whit; 1804 OCO glau; 1812 

OCO whit; 1894 OCO nica; 1980 POV quad; 

2051 NEC sali; 2086 OCO dend; 2122 PER 

amer; 2279 OCO whit. 
Cook, O. F., & C. R. Doyle: 3 PHO cinn; 647 

OCO vera. 
Cooper, G. P., et al.: 262 OCO cern; 458 PER rige; 

488 NEC memb; 498 NEC kunt; 512 NEC reti; 

532 OCO insu; 539 PLE sp. A; 551 NEC salf?; 

603 OCO atir; 612 OCO nica. 
Cooper, J. J.: 10220 OCO nica; 10286 OCO cern. 
Cordoba, J.: 109 OCO dend; 357 OCO cern; 369 

NEC glob; 821 PER amer; 1008 NEC memb; 

5107 PHO hamm; 5108 NEC cufo. 
Croat, T.: 10430 OCO glau; 13573 NEC memb; 



122 



FIELDIANA: BOTANY 



13677 OCO whit; 22388 PHO hamm; 22484 
NEC mart; 26550, 26560 OCO insu; 26555 NEC 
memb; 26775 PER rige; 26778 PHOcinn; 26934 
OCO whit; 27032 PHO cost; 35242 NEC salf; 
35526 PLE palm; 43304 OCO nica; 4336 1 OCO 
atir, 44480 NEC memb; 469 1 3 OCO vala; 59707 
OCO nica. 
Cufodontis, G.: 187 LIC cufo; 315 NEC cufo. 

D'Arcy, W. G., & J. J. D'Arcy: 6293 OCO vera. 

Daubenmire, R.: 135, 459 OCO vera; 486 NEC 
glob; 515 PER amer; 562 NEC glob; 677 OCO 
vera. 

Davidse, G., et al.: 1510 OCO viri; 23310 NEC 
sali; 23356 NEC sali?; 23390 OCO tene; 24206 
NEC memb; 24222 BEI oval?; 24246 NEC cufo?; 
24287 OCO insu; 24359 OCO flor; 24447 PER 
amer?; 24457 OCO whit; 24482 OCO endr; 
24518 NEC cufo; 24551 PER vera; 24564 OCO 
whit; 24579 PHO cinn?; 24628 PER vera; 25210 
NEC cufo?; 25259 OCO pitt; 25529 AIO cost; 
25542 PER obtu; 25575 PHO cinn?; 25586 OCO 
insu; 25600 NEC kunt; 26063, 26 109 PER vest; 
28226 BEI oval; 28401 PER amer?; 28505 PHO 
cinn?; 28520 OCO whit; 28547 OCO pitt; 2855 1 
OCO pseu; 28555 OCO mole; 28809 OCO pitt; 
29106 OCO hold; 29178 AIO tala; 29213 BEI 
pend; 29381 PER vest. 

Davidson, C: 6726, 6758, 6779, 6887 OCO atir. 

Davidson, M. E.: 268 OCO whit; 361 LIC exce; 
427 PER schi; 435 OCO insu; 516 PER vera; 
531 OCO glau; 566 NEC ramo; 576 PER vera; 
583, 641 PHO cinn; 753 PER vera. 

Dayton, W. A.: 3093 OCO vera; 3123 BEI pend; 
3124 OCO nica?; 3126 BEI oval. 

Dodge, C. W., et al: 6269 OCO tene; 6312, 6361, 
6471 OCO vera; 9761 PER amer. 

Dryer, V. J.: 390, 425, 470 OCO insu; 509 OCO 
leuc; 734 OCO insu; 862 OCO endr; 939 BEI 
pend; 943 PER amer; 954 NEC sali; 992 OCO 
mezi; 1011 OCO tene; 1032 PER amer; 1050 
OCO insu; 1052 OCO mezi; 1076 PHO cinn; 
1081 BEI pend; 1 154 PER schi; 1 179 BEI pend; 
1200 OCO mezi; 1201 PER schi; 1203 OCO 
insu; 1241, 1300 PER amer; 1315 NEC sali; 
1 332 OCO mont; 1 333 NEC sali; 1 334 BEI pend; 
1335 PHO cinn; 1336 OCO vala; 1362 OCO 
insu; 1422 OCO mezi; 1466 PHO cinn; 1591 
OCO mont; 1599 OCO flor; 1609 NEC memb; 
1612 OCO pseu?; 1613 OCO tene. 

Echeverria, J. A.: s.n. AIO cost; 526 NEC memb; 

1197 AIO cost. 
Englesing, F.: 65, 123 NEC reti; 162 PER amer. 



Folsom, J.: 8776, 8974, 9062 OCO atir; 91 12 OCO 
biju; 9283 OCO atir; 9288 OCO dend; 9512 
OCO mezi; 9529 OCO atir; 9864, 9904 OCO 
cern; 9930 OCO hart. 

Foster, R.: 730 NEC glob; 4114 OCO insu (Cocos 
Is.). 

Fournier O., L. A.: 389, 390 LIC tria; 400 NEC 
ramo; 649 OCO mole; 82 1 OCO vera; 868 PER 
amer; 873 PHO cinn; 905, 906 NEC ramo; 925 
OCO vera; 929 PER caer; 935 PHO bren; 941- 
943 PHO cinn; 945, 946 PER caer; 949 NEC 
ramo; 957 PER caer; 958, 960, 961 PHO cinn; 
966 PER caer; 967, 968, 1004 PHO cinn; 1010, 
1011, 1040 PER caer; 1053 PHO cinn; 1067 
PER schi; 1071 OCO glau; 1102 OCO cufo; 
1104 AIO cost; 1156 PHO bren; 1179 PHO 
cinn; 1190 PER amer; 1191 PER schi; 1233 
PHO cinn; 1235 PHO bren; 1245 OCO mole; 
1263 PHO bren; 1291 PER caer; 1330 OCO 
pitt; 1356 OCO vera; 1409-1419 PHO bren; 
1499 OCO aust; 1519 NEC memb; 1649 OCO 
dend; 1668 NEC mart; 1684, 1685 BEI pend; 
1728 OCO atir; 1771 OCO cern; 1796 OCO 
hold; 1797 NEC cufo; 1799 AIO cost; 1800 NEC 
cufo. 

Frankie, G. W.: 34c OCO dend; 52c NEC hypo; 
58c OCO atir; 412 OCO atir. 

Gentry, A.: 1 1 1 8 OCO glau; 2024 OCO nica; 59 1 8 
OCO whit; 5991 OCO glau; 6035 OCO paul; 
48524, 48567 NEC hypo; 48711 BEI pend; 
48736 OCO mont; 48762 OCO insu; 4877 1 PER 
amer; 488 1 1 OCO endr. 

Godfrey, R. K.: 66480, 66486 PHO bren; 67049 
NEC glob; 67078 OCO vera; 67200 PER caer. 

Gomez, L. D., et al.: 20768 OCO dend; 20851 
OCO nica; 2095 1 OCO atir; 20957 OCO bren?; 
20977, 2 1 007 OCO atir; 2 1 1 02 OCO nica; 2 1 1 30 
PER schi; 21143 LIC sara; 21175 NEC sinu; 
21496 OCO tala; 21654 OCO obtu; 22697, 
22705 NEC ramo; 22722 OCO dend; 22864 
NEC cufo; 23064 NEC kunt; 23208 OCO tene; 
23216 PHO hamm: 23227 BEI oval; 23304 NEC 
reti; 23344 NEC reti; 23445 OCO atir; 23619 
OCO valo; 23653 OCO dent; 23875 OCO whit; 
23977 PHO hamm; 24101 NEC mart. 

Gomez-Laurito, J.: 79 ANI vene; 82 OCO flor; 87 
OCO molf; 57 1 OCO nica; 588 OCO dend; 1 207 
PER vest; 1558 PHO pitt; 27 1 1 LIC bren?; 3066 
OCO pseu; 3113 NEC memb; 3610 NEC glob; 
4580 OCO vera; 4590 OCO pseud; 4743 PER 
vest; 4945 PLE palm; 5178 OCO insu; 5262 
PHO cinn; 5287 OCO endr; 6307 PER amer; 



BURGER: FLORA COSTARICENSIS 



123 






6372 OCO atir; 6609 OCO dend; 6613 OCO 
atir; 6938 OCO insu; 6946 OCO insu (Cocos 
Is.); 7632 OCO atir?; 7645 OCO molf; 7669 
OCO heli; 7836 OCO dend; 7957 OCO insu; 
8113 PER amer; 8141 OCO mezi; 8812, 8864 
OCO insu; 9304 OCO atir?; 9619 OCO 
insu; 97 1 6 OCO atir; 9785 OCO endr; 9800 BEI 
oval?; 9849 OCO bren?; 9897 OCO pseu; 9900 
ANI vene; 9906 NEC ramo; 99 1 1 NEC memb; 
9989 OCO cern; 10216 NEC salf; 10217 NEC 
memb; 10255 OCO atir; 1 03 1 6 NEC sali; 10459 
OCO cern; 1061 1 OCO paul; 1 1086 OCO endr; 
11098 OCO leuc; 11262 NEC memb; 11237 
OCO laet; 1 1272 NEC memb; 1 1329 NEC sali; 
1 1 385 OCO pitt; 1 1 390 OCO bren; 1 1400 OCO 
paul; 11447 OCO pseu; 11450 OCO gome; 
1 1453 OCO endr; 1 1464 AIO cost; 1 1467 PHO 
hamm. 

Gonzalez Meza, R.: 23 OCO aust; 24 NEC cufo; 
34 PER vest. 

Grayum, M., et al.: 1007 OCO atir; 1290 OCO 
dend; 1398 OCO molf; 1480 OCO tene; 1482 
NEC hypo; 1818 OCO dend; 2201 OCO mont; 
2388 OCO leuc; 2767 PHO chav; 3030 NEC 
memb; 3416 OCO atir; 3845 OCO pitt; 3875 
OCO tene; 3969 NEC niti; 3995 OCO atir/nica; 
4069 OCO rivu; 4091 OCO atir; 4374 OCO 
cern; 4525 NEC reti; 4702, 4709 LIC cufo; 4749 
OCO nica; 4775 NEC niti; 4894, 4947 OCO 
vera; 5 1 04 PHO cinn; 5 1 23 PER schi; 5151 OCO 
laet; 5266 OCO cern; 5286 OCO wede; 6101 
OCO nica; 6260 NEC sali; 6263 OCO tene; 6355 
OCO nica; 6876, 6888 OCO atir; 6923 LIC sara; 
6995 OCO dend; 7169 OCO hold; 7589 OCO 
whit; 7605 LIC cufo; 7970 PHO cinn; 8060 NEC 
memb; 8064 CAS fili. 

Haber, W. H.: 161 OCO mont; 191 OCO insu; 
198 OCO flor; 222 NEC sali; 238, 249 OCO 
mezi; 250, 252 PHO cinn; 259 OCO mezi; 262 
NEC glob; 268 NEC sali; 276 PER amer; 289 
NEC sali; 314 OCO vala; 318 NEC sali; 321 
PHO cinn; 324 OCO whit?; 346 OCO pitt?; 385 
OCO whit; 390 PER vera; 439 NEC sinu; 494 
BEI pend; 525, 526 OCO whit; 579 PER vera. 

Hammel, B., et al.: 1218 OCO hart; 5623 PER 
obtu; 7013 OCO pitt; 7823, 7844 OCO atir; 
7950 OCO dend; 7957 OCO mezi; 8034A OCO 
dend; 8075, 8201 OCO mezi; 8131, 8223 OCO 
atir, 8299 OCO dend; 8390 OCO cern; 8663 
LIC sara; 8864 NEC hypo; 89 1 4 OCO flor; 9111 
NEC hypo; 9192 OCO dend; 9705 OCO flor; 
9807 OCO atir; 10073 NEC hypo; 10146 OCO 
atir; 10229 OCO biju; 10348 NEC hypo; 10479 
NEC reti; 10491 OCO dend; 10501 OCO atir; 



10503 OCO dend; 10532 LIC sara; 10538 OCO 
insu; 10561 OCO mezi; 1 06 1 8 OCO atir; 10630 
OCO hart; 10657 NEC reti; 10670 LIC sp. A; 
10693 NEC kunt; 10736 OCO molf; 10871 OCO 
babo; 1 09 1 2 OCO leuc; 1 1 034 OCO insu; 11036 
LIC tria; 1 1058 OCO leuc; 1 1095, 1 1 164 OCO 
mezi; 1 1 168 NEC ciss; 1 1 170 OCO flor; 11212 
OCO atir; 1 1221 OCO mezi; 1 1226 NEC kunt; 
1 1 252 OCO atir; 1 1 522 OCO molf; 1 1 530 OCO 
babo; 1 1663 OCO biju; 1 1665 NEC ciss; 1 1679 
PHO chav; 1 1 762 OCO dend; 1 1 928 OCO leuc; 
11932 OCO hart; 11981 OCO leuc; 11995, 
12018, 12080 OCO dend; 12163 OCO atir; 
12198 OCO dend; 1 2235 LIC sara; 1 2332 OCO 
dend; 12397 OCO atir; 12409 NEC lati; 12738 
NEC lati; 12762 OCO atir; 12786 NEC memb; 
12840, 12848 NEC hypo; 12893 NEC memb; 
12909 OCO dend; 12922 PHO chav; 12957 
OCO leuc; 12961, 13029 NEC memb; 13036 
OCO mezi; 1 3074 NEC hypo; 13111 ANI vene; 
13123 OCO leuc; 1 3 1 44 OCO tene; 1 3249 NEC 
memb; 13366 ANI vene; 13771 NEC niti; 13797 
OCO whit; 1 4086 OCO endr; 1 4088, 1 4090 NEC 
cufo; 14091 PHO hamm; 14111 LIC exce?; 
15044 OCO pseu?; 15214 MEZ glau. 
Hartshorn, G.: 926, 949 OCO cern; 950 NEC hypo; 
970 NEC reti; 1005 OCO hart; 1043 NEC memb; 
1063 OCO tene; 1080 OCO dend; 1082 NEC 
kunt; 1103 OCO molf; 1121 AIO cost; 1122 
PER schi; 1125 PHO hamm; 1127 BEI oval; 
1140 OCO hart; 1156 PER caer; 1169 PHO 
cinn; 1179 PHO chav; 1209 PHO cinn; 1218 
OCO hart; 1229 PHO insu; 1234 NEC lati; 1252 
NEC memb; 1256 NEC kunt; 1258 OCO tene; 
1276 NEC hypo; 1284 PHO cinn; 1289 PHO 
chav; 1300 OCO dend; 1333 OCO atir; 1347 
OCO insu; 1351 OCO flor; 1388 NEC hypo; 
1414 OCO mezi; 1426 OCO atir; 1441 OCO 
nica; 1445 OCO cern; 1456 PHO hamm; 1463 
OCO vala; 1465 BEI pend; 1469 NEC sali; 1472 
OCO mezi; 1514 NEC memb; 1518 OCO dent; 
1 5 1 9 BEI sulc; 1 530 OCO valo; 1 543 OCO dent; 
1550 OCO mezi; 1563 OCO flor; 1567 OCO 
molf; 1576 OCO insu; 1585 OCO biju; 1588 
LIC sara; 1 589 OCO mezi; 1 590 LIC tria; 1 595 
OCO atir; 1 600 NEC reti; 1 628 OCO nica; 1 635 
OCO tene; 1636 OCO insu; 1638 OCO hart, 
1751 NEC hypo; 1753 NEC memb; 1819 PER 
schi; 1865 NEC salf; 1874 PHO neur?; 1885 
OCO vera; 1900 OCO mont; 2115 OCO insu; 
2117, 2123 OCO tene; 2126 OCO insu; 2130 
PHO vala; 2159 BEI pend; 2161 OCO skut; 
2 1 64 NEC ramo; 2 1 65 OCO flor; 2 1 66 BEI oval; 
2167 NEC memb; 2182 OCO vera. 



124 



FIELDIANA: BOTANY 



Hatheway, W. H.: 1423 AIO cost; 1427 PER caer; 
1463 NEC cufo; 1466AIOcost; 1478 OCO mole; 
1678 NEC cufo. 

Hazlett, D.: 2781 OCO cern; 5242 OCO whit. 

Herrera, G.: 504 OCO gome. 

Hill, S. R.: 11958OCOatir. 

Holdridge, L. R.: 5126 OCO tene; 5128 NEC reti; 
5138 OCO vera; 5145 OCO atir; 5150 NEC 
cufo; 5151 PHO hamm; 5 1 53 OCO dend; 5 1 55 
NEC glob; 5 1 56 NEC reti; 5 1 57 OCO oblo; 5 1 59 
NEC cufo; 5164 OCO dend; 5168 NEC ramo; 
5172 NEC sinu; 5173, 5175 OCO aust; 5179 
NEC sinu; 5217, 5227 OCO vera; 6328 OCO 
cern; 6336 OCO dend; 6339 NEC niti; 6544 
PER caer; 6545 NEC sinu; 6546 PHO cinn; 6547 
NEC mart; 6548 PHO bren; 6549 OCO vera; 
6550 NEC reti; 6551 NEC memb; 6552 PER 
schi; 6553 PER amer; 6554 NEC cufo; 6555 
OCO pseu; 6558 PHO hamm; 6559 AIO cost; 
6560 PHO hamm; 6562 PER vera; 6564 OCO 
leuc?; 6565 NEC cufo; 6569 OCO endr; 6570 
OCO hold; 6571 OCO laet; 6572 OCO endr; 
6574 NEC turb; 6575 OCO pseu; 6576 OCO 
vala; 6578 OCO leuc?; 6580 OCO endr; 6581 
OCO insu; 6583 OCO nica?; 6584 NEC memb?; 
6586 OCO oblo; 6587 NEC kunt; 6588 OCO 
dend; 6590 ?END sp.; 6591 OCO leuc; 6594 
OCO cern; 6595 BEI oval; 6639 OCO oblo; 6646 
OCO insu; 6649 OCO cern; 6650 OCO leuc; 
6652 OCO atir; 6654 OCO leuc; 6655 NEC salf; 
6663 OCO vala; 6671 OCO pitt; 6687 NEC 
cufo; 6688 NEC sali; 6690 PLE palm; 6691 OCO 
insu; 6696 OCO dend; 6700 OCO hart?; 6701 
NEC sali; 6702 OCO atir; 6704, 6706 OCO hart; 
6713 PHO cinn; 6714 NEC memb; 6715 OCO 
oblo; 6719 OCO valo; 6724 OCO endr; 6739 
OCO flor; 6758 PHO vale; 6767 NEC sali; 6779 
OCO oblo; 6783 OCO pseu?; 6808, 6813 OCO 
flor; 68 1 4 OCO oblo; 68 1 5 NEC cufo; 68 1 9 PER 
rige; 6820 OCO oblo; 682 1 LIC mult; 6825 PER 
rige; 6826 OCO aust; 6827, 6829 OCO pseu?; 
6831 NEC sali; 6868 NEC turb?; 6872 NEC 
glob; 6873 OCO atir. 

Holm. R., & H. Iltis: 212 OCO dend. 

Jacobs, B.: 2540 NEC lati; 2876 OCO atir; 2966 
NEC memb; 2967 OCO mezi. 

Jimenez, A.: 63 OCO atir; 124 PHO cinn; 302 
NEC glob; 303, 304, 355 OCO vera; 371 NEC 
glob; 379 OCO nica; 404 PHO vala; 479 AIO 
cost; 487, 491 OCO vera; 822 LIC tria; 1166 
OCO insu; 1190 OCO tene; 1192 OCO insu; 
1242 OCO mezi; 1333 OCO atir; 1461 OCO 
laet; 1506 NEC glob; 1568 OCO vera; 1671 NEC 



reti; 1722 NEC kunt; 1723 OCO dent; 1744 
NEC memb; 1864 OCO atir; 1867 NEC memb; 
1971 OCO pitt; 2001 LIC tria; 2142 NEC mart; 
2334 OCO atir; 2544 PER schi; 2552 PHO 
hamm; 2720 NEC glob; 2746 OCO atir; 2752 
OCO leuc; 2905 OCO dend; 3006 OCO cern; 
3117, 3125 OCO vera; 3183 OCO insu; 3219 
OCO glau; 3440 OCO endr; 3540 OCO vera; 
3548 PER schi; 3567 OCO endr; 3673 OCO 
vera; 3682 NEC sinu; 3801 PHO cinn; 3835 
PHO neur; 3868 PHO bren; 3942 PER caer; 
3977 OCO dend; 3989 OCO pitt; 4116 AIO 
cost; 4125 NEC memb. 

Jimenez, Oton: 379 NEC glob; 822 LIC tria; 1298 
PER amer. 

Jimenez-Saa, H.: 88 OCO insu; 98 OCO hart. 

Kirkbride, J., & J. Duke: 997 PER caer. 

Lankester, C.: 260 OCO tene; 1335 OCO aura; 
1340 NEC glob. 

Lao, E. A.: 326 PER schi; 337 PER vera; 342 OCO 
whit; 350 PER amer; 351 NEC cufo; 395 BEI 
oval; 599 PER caer. 

Lawton, R.: 1043 OCO tene; 1 197 PHO cinn; 1225 
OCO pitt; 1265 BEI pend. 

Lellinger, D., & J. White: 1641 OCO atir. 

Lems, K.: s.n. & 5067 PER amer; 5164 BEI pend; 
5168 PER amer; 5174 OCO nica?; 5345 OCO 
endr; 5352 OCO calo; 5552 PER amer. 

Lent, R.: 727 OCO endr; 794 OCO lent; 1677 
OCO hold; 1678 PHO hamm; 1679 OCO pseu; 
1735 OCO insu; 1737 PER caer; 2070 OCO 
lent; 235 1 LIC tria?; 2467 NEC salf; 2705 OCO 
flor; 2878 OCO atir; 3116 OCO insu; 3181 OCO 
atir; 3297 NEC sali; 3386 OCO atir; 3490 NEC 
sinu; 3660 AIO cost; 3884 OCO insu. 

Leon, J.: 481 OCO insu; 528 NEC sinu; 543 PER 
schi; 578 NEC sinu; 868 LIC tria?; 895 OCO 
vera; 1047 OCO dend; 1104 LIC mult; 1168 
OCO laet; 1237 OCO vera; 1599, 1644 NEC 
glob; 1679 NEC mart; 1720 NEC memb; 1762 
OCO endr; 1861 OCO dend; 2066 OCO tene; 
2166 OCO calo; 2391 OCO pitt; 2452 NEC 
memb; 2477 OCO dend; 2509 OCO cern; 25 1 1 
OCO dend; 2744 OCO cern; 3119 OCO vera; 
3203 LIC mult; 33 1 9 NEC reti; 3378 OCO dend; 
3429 OCO cern; 4262 PER amer?; 4266 OCO 
vera; 4344 OCO glau; 4533 OCO cern. 

Liesner, R.: 1789 PHO heli; 1982 OCO nica; 2151 
OCO cern; 2 1 62 NEC reti; 3034, 3048 NEC salf; 
3166 OCO nica; 4494, 4775, 4941, 4943, 4972 
OCO vera; 502 1 NEC reti; 5022 NEC glob; 5 106 
NEC sali; 5123 OCO atir; 5189 OCO dend; 



BURGER: FLORA COSTARICENSIS 



125 



14310, 15052 OCO atir; 15319, 15394 OCO 
dend; 15580 OCO insu. 

Little, E. L.: 6007 OCO aust; 60 1 2 AIO cost; 60 1 3 
NEC cufo; 6023 AIO cost; 6047, 6056 OCO 
whit; 6057, 6058 PER rige; 6059, 6062, 6069 
OCO whit; 6078 LIC exce?; 20013 OCO mezi; 
20059 NEC memb; 20060 BEI sulc?; 20088 NEC 
memb; 20093 OCO aust; 20125 PHO cinn; 
20174 NEC reti; 20276 NEC sinu; 20336 PHO 
cinn; 20412 PHO hamm; 20415 NEC cufo. 

Luteyn, J., & R. Wilbur: 4548 OCO pitt. 

Madriz V., A.: 2 OCO pitt; 3 PER amer; 7 OCO 

aust; 1 2 OCO calo; 20 OCO insu; 34 OCO hold; 

42 PER vest; 44 PHO hamm; 45 PER schi; 65 

OCO mezi?; 69 AIO cost. 
Maxon, W., & A. D. Harvey: 8196 PER schi. 
McDowell, T.: 159 OCO dend; 316 OCO tene; 

401 OCO dend; 509 OCO atir; 582 NEC reti; 

601 NEC hypo; 667 OCO atir; 854 OCO babo; 

1050, 1052 OCO atir; 1 102 NEC memb. 
Menzies, A.: 1 794 OCO insu (type, Cocos Island). 
Molina, A., et al.: 17436 OCO nica; 17524 PER 

amer; 17526, 17734 NEC memb; 17893 OCO 

pitt; 18069 OCO atir; 18140 NEC reti; 18272 

NEC sinu; 18359 AIO tala; 20606 PER caer. 
Mora V., C: 1 3 PER amer; 1 5 OCO endr; 1 7 OCO 

pseu; 19 PER schi. 
Mori, S., et al.: 5625, 5678 OCO whit; 5787 OCO 

viri; 7829 OCO flor. 

Neill: 5030 OCO mont; 5030B OCO insu; 5122 
OCO hart. 

Oersted, A.: Laur. No. 10 LIT glau; 270 PER caer; 

14938 PHO hamm? 
Opler, P.: 88, 1 14, 346, 610 OCO atir; 623 OCO 

cern; 648 OCO vera; 1 592 OCO tene; 1 929 PER 

amer. 
Orozco, J. M.: 38 OCO vera; 341 PER vera; 467 

OCO vera. 

Pittier, H.: 409 OCO vera; 1739 OCO endr; 2040 
BEI oval; 2259 OCO pitt; 2288 PHO vala; 2998 
PHO cinn; 3132 PER schi; 3146 PHO cinn; 
3201 OCO whit; 3633 OCO tene; 3644 OCO 
vera; 3984 NEC reti; 5 145 NEC glob; 5278 OCO 
vera; 5395 OCO paul; 9179 OCO atir; 9184 
OCO tene; 9663 NEC mart; 10351 OCO atir; 
10605 NEC reti; 1 1 102 NEC ciss?; 1 1 107 PHO 
cinn?: 11111 PER albi; 1 1 25 1 , 1 20 1 6 OCO cern; 
1 1282 NEC niti; 1 1283 NEC turb; 1 1489 OCO 
insu?; 1 1 490 NEC mart; 1 20 1 8 NEC reti; 1 2348 
NEC sinu; 1 3396 OCO tene; 16030, 16031 OCO 



molf; 16140 BEI sulc; 16257 OCO insu; 16428 
OCO vera. 

Popenoe, W.: 987, 989 PER schi; 993 NEC mart; 
996 PER schi; 1001 NEC sinu; 1003 PER vera. 

Poveda, L.: 104 OCO valo?; 1 1 1 OCO flor; 112 
OCO leuc; 1 16 NEC reti; 158 OCO oblo; 170 
OCO atir; 391 OCO endr; 392 OCO pseu; 429 
OCO insu; 455 OCO mole; 500 LIC cufo; 514 
OCO calo; 527 NEC memb; 589 POV quad; 
63 1 OCO long; 669 OCO insu; 740 PER pove; 
753 OCO oblo?; 764 OCO molf; 783 OCO tene; 
845 PHO bren; 868 OCO pitt; 926 PER schi; 
931 LIC bren; 933 OCO heli; 947 OCO pseu?; 
956 PHO chav; 998 NEC salf; 1021 ANI vene; 
1061 PHO hamm; 1065 PER vera; 1068 BEI 
pend; 1076 PHO cinn; 1079 OCO laet; 1086 
OCO glau; 1091 OCO insu; 1 108 OCO mont; 
1 130 NEC sali; 1 158 PER rige; 1202 PER vest; 
1299 NEC mart; 1519 OCO pseu?; 1524 BEI 
oval?; 1606 OCO pitt; 1635 OCO dend; 1712 
NEC salf; 1771 OCO insu; 2291 OCO aura; 
2360 POV quad; 2999 OCO pitt; 3000 OCO 
mole; 3019 BEI pend; 3020 PHO cinn; 3021 
OCO endr; 3049 LIC cufo; 3398 NEC sinu; 3399 
PHO bren; 3487 NEC memb; 3488 OCO oblo; 
3489 LIC sara; 349 1 OCO nica; 3492 NEC sinu; 
3493 OCO mezi; 3493A OCO dend; 3494 NEC 
kunt; 3495 OCO leuc; 3496 LIC bren; 3497 PHO 
cinn; 3498 ?END sp.; 3499 OCO skut; 3500 LIC 
mult; 3501 OCO ?sp. A; 3554 OCO nica; 3561 
POV quad; 3566 OCO mole; 3577 LIC exce?; 
3637 LIC exce; 3687 NEC memb; 3688 NEC 
reti; 3795 ?END sp.; 3887 PHO cinn; 3908 CAR 
burg; 39 1 3 OCO vala; 39 1 6 AIO cost; 3920 OCO 
valo; 3994 OCO mole; 3996 NEC sinu; 4042 
CAR burg. 

Primack, R., et al.: 254 PER bren; 332 OCO vala; 
438 NEC cufo; 457 NEC sali. 

Procter, G., et al.: 27421 NEC niti; 32355 OCO 
aust. 

Procter-Cooper, G.: 399, 449 OCO wede; 539 PLE 
sp. A; 603 OCO wede. 

Raven, P.: 21003 OCO atir; 21630 OCO heli; 

2 1 89 1 PER albi; 21916 NEC memb; 21954 PER 

albi?. 

Rodriguez, R. L.: 345 AIO cost. 
Rossbach, G. B.: 3 1 7 1 OCO pseu; 37 14 OCO tene. 
Rowlee, W. W., & H. E. Rowlee: 210, 233 PLE 

palm. 

Salas, S.: 54 LIC tria; 116 OCO hold; 138 NEC 
mart; 219 NEC memb; 240 AIO cost; 293 NEC 
glob; 3 1 9 NEC memb; 343 OCO vera; 462 AIO 



126 



FIELDIANA: BOTANY 



cost; 478, 628 OCO pseu; 1 1 89 NEC sinu; 1 377 
LICtria; 1382 NEC cufo. 

Sanchez, P., et al.: 23 OCO insu; 113 NEC salf; 
377 OCO mezi; 428 OCO mezi; 430 OCO dend; 
1 228 PLE golf. 

Schatz, G., et al.: 613 OCO dend; 970 NEC ciss; 
11 02 OCO dend. 

Schubert, B., et al.: s.n. OCO cern; 627, 627a,b 
NEC niti; 801 OCO atir; 874 OCO cern; 1004 
PER caer; 1031 OCO vera; 1091 NEC glob; 
1301 OCO atir; 1 141, 1306 OCO dend. 

Seibert, R. J.: 307 OCO whit; 308 NEC cufo; 1 603 
OCO dend. 

Shank, P. J., & A. Molina: 4275 OCO atir. 

Shimek, B., & C. L. Smith: s.n. NEC glob; 176 
OCO vera. 

Skutch, Alexander: 2007 OCO dend; 2490 NEC 
reti; 2605 NEC ciss; 2634 NEC mart; 2668 NEC 
memb; 2813 OCO nica; 2821 OCO skut; 3014 
OCO sten; 3117 PER caer; 3606 PER schi; 3745 
OCO mezi; 3755 OCO endr; 3836 NEC glob; 
3844 PHO neur; 4042 PHO cinn; 4172 NEC 
salf; 4182 NEC memb; 4270 OCO cern; 4303 
OCO insu; 4328 NEC salf; 4329 PHO neur; 
4375 NEC mart; 4389 BEI pend; 4591 OCO 
atir; 4687 NEC sinu; 4738 OCO cern; 4757 NEC 
kunt; 4770 LIC cufo; 48 1 2 PER caer; 4905 NEC 
salf; 5116 PHO neur; 5117 NEC mart; 5199 
NEC memb; 5500 OCO insu. 

Smith, Austin: A 125 OCO aust; A 140 PER amer; 
1 42 PHO cinn; 1 59 OCO tene; 1 64 NEC memb; 
181 PER amer; 183 OCO mole; A228 OCO 
tene; A240 OCO vala; A243 NEC memb; H268 
OCO vala; H269 PER amer; 309, 359 OCO 
mezi; 365 OCO laet; 367 OCO vala; A388 PER 
amer; H442 AIO cost; 443 OCO mezi; H469 
OCO bren; 492 OCO endr; 504 OCO aust; H5 1 6 
OCO bren; H522 OCO pitt?; 523 OCO vala; 
H541 NEC sali/smith; A556 OCO tene; 571 
OCO bren; H577 OCO paul; H592 BEI pend; 
593 OCO mezi; 598 OCO vala; H633 OCO paul; 
675 BEI oval; 679 OCO pitt; 68 1 AIO cost; 732 
OCO mezi; 907 OCO paul; NY964 PER vera; 
H969, 972 NEC cufo; P 1 1 50 BEI pend; NY 1 1 44 
OCO insu; 1181 OCO aust; 1323 OCO paul; 
1687 OCO bren; HI 737 OCO atir; 1765 OCO 
dend; 1771 OCO dend; 1774 OCO mezi; 1849 
OCO dend; 2036 NEC cufo; 2309 OCO mole; 
2316 AIO cost; P2409 NEC sinu; 2440 PHO 
cinn; 2464 OCO vera; P2465 NEC ramo; 2479 
PER caer; 2563 OCO atir; 2596 OCO mezi; 
P2615 OCO pitt; 2620, 2710 OCO bren; 2717 
BEI pend; 2722 AIO cost; 2772 OCO tene; 2809 
NEC memb; 2825 NEC cufo; 2842 OCO paul; 



4 1 02 OCO bren; 4 1 68 BEI oval; 4 1 69 AIO cost; 
4171 OCO bren; 4178 OCO mole; 4182 OCO 
laet; 4202 OCO hold; 10007 OCO mole; 10028 
PER amer; 10045 OCO flor; 10066 NEC reti; 
10068 OCO mole; 10072 OCO vala. 

Smith, Damon: s.n. OCO dend; 45 OCO atir; 99 
OCO dend; 245 OCO hart; 300 OCO dend; 406 
NEC reti; 425 OCO mezi; 474 OCO insu; 484 
OCO leuc; 507 NEC reti; 1 202 POV quad; 1210 
LIC sara. 

Smith, John Donnell: 4931 NEC reti; 4932 NEC 
salf; 5114 PER caer; 6751 OCO dend; 6752 
OCO cern; 6754 NEC sinu; 6755 NEC ramo; 
6756 OCO dend; 6757 NEC memb; 7352 LIT 
glau; 7374 PLE palm. (Note: Some of these are 
distribution numbers and not collection num- 
bers.) 

Smith, Lyman B.: 15313 OCO long. 

Snow, B.: 10 OCO insu; 12 OCO whit; 13 PHO 
vala. 

Solano J., I.: 1 PER vera; 3 OCO aust; 4 PER vest. 

Soto, R.: 2379, 2676, 2971 OCO insu. 

Sperry, J.: 695 OCO atir; 797 OCO dend; 946 
OCO dend. 

Standley, P., et al.: 25880 PER caer; 32255, 32705 
PHO cinn; 33679 OCO vala; 34274 PER schi; 
34579 AIO cost; 34824 PHO cinn; 35777 PER 
caer; 36507 OCO atir; 36541 OCO aust?; 37039, 
37188, 37240 OCO atir; 37640 OCO mezi; 
37949 OCO atir; 38940 PHO cinn; 39278 OCO 
pitt; 39289 PER schi; 39433 OCO mezi; 39989, 
40029, 40036 OCO vera; 40238 LIC tria; 40243 
PHO cinn; 42426 OCO mole; 42458 OCO vala; 
42525 LIT glau; 42565 BEI pend; 42585 PER 
vest; 42935 OCO vala; 43050 OCO pitt; 43 1 78, 
43344 OCO vala; 43421 OCO mole; 43960 BEI 
oval; 44 1 1 3 OCO vera; 4465 1 OCO ?sp.; 44907 
OCO vera; 45238 NEC reti; 45242 OCO atir; 
45283 NEC reti; 45340 OCO atir; 45469 NEC 
turb; 4576 1 , 4580 1 OCO tene; 45887 OCO atir; 
45906, 45909 NEC turb; 46121 OCO tene; 
46128 NEC sali; 46167 OCO atir; 46192 NEC 
turb; 46253, 46368 NEC sali; 46524 OCO nica; 
46749 OCO atir; 46773 NEC reti; 46924 OCO 
dend; 47 1 85 OCO atir; 47344 PHO cinn; 47364 
NEC niti; 47475 PER schi; 48403, 48467 OCO 
atir; 48784 LIC sara; 48789 OCO mezi; 50908 
NEC memb; 50962 OCO mezi; 5 1 1 80 OCO 
tene; 5 1 20 1 NEC glob?; 5 1 268 PER albi?; 51271, 
5 1 280, 5 1 285 BEI pend; 5 1 387 OCO pitt; 52072 
AIO cost; 64785, 89348 OCO vera. 

Stern, W. L., & K. L. Chambers: 50, 5 1 OCO whit; 
56 PER rige; 62 PHO cinn?; 95 LIC exce; 111, 
1 1 1A NEC reti; 1 13 NEC memb. 



BURGER: FLORA COSTARICENSIS 



127 



Stern, W. L., et al.: 1026, 1096, 1121 OCO whit; 

1 1 22 SP. ?gen; 1143 NEC cufo; 1 988 PHO cinn. 
Stevens, W. D.: 14131 OCO insu; 23757 OCO 

leuc; 23802 OCO insu; 23838 OCO atir; 24100 

NEC salf. 
Stone, D. E.: 1957 NEC sali; 3195 OCO palm; 

3204 NEC cufo; 3224 NEC mart; 3296 OCO 

insu; 3394 OCO vala. 
Stork, H. E.: 1 1 1 PER caer; 1 399 AIO cost; 1 570 

PER caer; 1 7 1 3 BEI pend; 1 735 AIO cost; 2328 

PER vera; 2332 LIC tria; 2404 PHO hamm; 

2405 OCO vala; 2416 PER caer; 2807 OCO 

babo; 2809 OCO whit?; 3059 NEC mart; 3121 

BEI pend; 3177 OCO mole; 3377, 3577 NEC 

cufo; 4102,4171 OCO bren. 

Taylor, C. M.: 82 AIO cost; 3117 PHO vala. 

Taylor, R. John: 4460 OCO pitt. 

Todzia, C: 813 OCO atir; 1854, 1907 OCO pseu; 
1961 OCO atir; 2022 OCO mezi. 

Tonduz, Adolfo: 1794 AIO cost; 1873 NEC reti; 
1 883 OCO laet; 2226 NEC sinu; 2234 PER caer; 
3598 OCO pitt; 3934, 4876 NEC reti; 6637 OCO 
vera; 6680 PER caer; 7 1 53 NEC glob; 727 1 NEC 
niti; 7374 OCO palm; 7648 OCO vera; 7770 
NEC sinu; 7796 LIT glau; 8555 OCO nica; 8772 
NEC mart; 9005 OCO oblo; 9020 OCO dend; 
9179 OCO atir; 10047 NEC glob; 10104 NEC 
niti; 10999 PER amer; 1 1489 OCO whit?; 11612 
LIC tria; 11638 LIT glau; 11651 NEC sinu; 
1 1676 OCO mole; 1 1 7 1 3 BEI pend; 1 1735 PHO 
cinn; 1 1746, 1 1755 OCO vala; 1 1893 OCO pitt; 
1 1 896 PER amer?; 1 1 939 NEC cufo; 1 2652 PLE 
palm; 12772 OCO atir; 12875, 12878 OCO oblo; 
12913 NEC reti; 12953 OCO sten; 12978 OCO 
atir; 13366 OCO cern; 1 3377 OCO sten; 13794 
PER amer; 13806 NEC glob; 13809, 13845, 
1 3863, 1 49 1 4 OCO vera; 1 6 1 08 PER caer; 17689 
LIC tria; 17692 NEC reti. 

Torres, R.: 7 NEC memb. 

Tyson, E. L.: 7221 NEC ramo; 7410 PER vera. 

Utley, J., & K. Utley: 2779 OCO endr; 2782 OCO 
insu; 3012 NEC cufo; 3015 OCO aust; 3040 
AIO cost; 3238 OCO nica; 3333 NEC turb; 3338 
OCO cern; 3648 PHO hamm; 3699 NEC mart; 
3787 OCO endr; 3853 OCO laet; 4046 NEC 
beli; 4 1 80 OCO laet; 4653 BEI pend; 4750 OCO 
glau; 4760 OCO vera; 4889 NEC salf; 5 140 OCO 
atir; 5220 AIO cost; 5356, 5397 OCO atir; 5407 
OCO mont; 5577 NEC cufo; 5782 OCO vala; 
5788 OCO aust; 5795 OCO pitt; 6087 OCO 
vera. 



Valeric, Juvenal: 21 NEC sali; 23 OCO atir; 33 
AIO cost; 34 OCO vala; 46 OCO atir; 101, 105 
OCO vera; 1 14 NEC memb; 397 OCO cern; 495 
NEC glob; 675 OCO atir. 

Valeric, Manuel: 101 OCO vera; 1 14 NEC memb; 
675 OCO atir; 9 1 4 OCO vera; 1 326 NEC memb; 
1327 NEC mart; 1451 NEC cufo; 1655 OCO 
atir; 1745 OCO vera. 

Von Hagen, C. & W.: 2031 OCO pitt; 2046 OCO 
viri; 2070 OCO pitt; 2 1 20 NEC cufo; 2 1 28 OCO 
leuc; 2140 NEC cufo; 2178 OCO viri?. 

Walker, J.: 188 OCO atir; 414 OCO laet. 

Webster, G., et al.: 12395 OCO aura; 12455 NEC 
sali. 

Wedel, H. von: 388 OCO wede?; 1399 OCO atir; 
2264 OCO vale; 2431 NEC salf; 2889, 2967 
NEC memb. 

Wheelwright, N.: 0/F OCO whit; 1 OCO mont; 3 
OCO flor; 4B PHO cinn; 8 NEC memb; 12B 
OCO tene; 13A PHO cinn; 13C PER caer; 13D 
PER amer; 14B, 15 PHO cinn; 16B PER vera; 
19C OCO whit; 20 NEC sali?; 23 PER amer; 25 
OCO viri; 27 A, 30A PHO cinn; 30B OCO mont; 
3 IE, 3 IF PER caer; 33 OCO mont; 34 OCO 
whit; 36 A PHO cinn; 44 OCO tene; 52 BEI 
pend; 53A OCO whit; 55 BEI pend; 56 OCO 
insu; 58 OCO whit; 59 PHO cinn; 70 BEI pend; 
75 OCO mont; 79 OCO whit; 80 PER amer; 
80C NEC sali; 83 OCO insu; 85, 91 OCO mezi; 
97 PHO cinn; 1 12 OCO whit; 115 OCO insu; 
125 A OCO mezi; 125B PER amer; 126B BEI 
pend; 126D PHO cinn; 133 OCO insu; 136A 
OCO mezi; 140 A PHO cinn; 141 A NEC sali; 
143 A, 144B OCO insu; 162 PER amer; 165 
OCO endr; 167 OCO insu; 169 OCO mezi; 176 
OCO insu; 188 A BEI pend; 201 OCO flor; 206 
??; 207 NEC sinu; 209 OCO vala; 214 PER schi; 
214B OCO insu; 219 BEI pend; 220 ?LIC; 223 
PHO pitt; 224 NEC sinu; 230 NEC sali?; 250 
PHO cinn; 25 1 OCO atir. 

White, P.: 2 1 3 PER caer; 225 OCO insu; 334 PER 
vera. 

Wilbur, R., et al.: 9760 OCO vera; 9945 NEC glob; 
10701 NEC sali; 10894 OCO glau; 167 14 OCO 
palm; 16 762 OCO pitt; 19751 OCO laet; 20133 
NEC salf; 20836 AIO cost; 21726 OCO endr; 
21801 AIO cost; 22076 NEC salf; 22982 NEC 
sali; 24556, 24724 OCO pitt; 24852 OCO endr; 
25407 NEC memb; 30053, 30220 OCO dend; 
30373 OCO atir; 3 1 563B AIO cost; 33575 OCO 
atir. 

Williams, L. O., et al.: 13888 OCO pitt; 13943 



128 



FIELDIANA: BOTANY 



PERcaer; 16064, 16147 PHOcinn; 16 155 PER 
caer; 16160 PHOcinn; 16168 PER schi; 16169 
PER vera; 16235 PER schi; 16236 PER caer; 
1 6238 AIO cost; 16239, 1 6240 PER amer; 16344 
OCOinsu; 16419, 16421 AIO cost; 16443 PER 
caer; 16462 PHOcinn; 1 6469 OCO vala; 16470 
PER schi; 16476 OCO pitt; 16556 PHO cinn; 
16582, 16587 PER caer; 16588 AIO cost; 16598, 
16612 NECcufo; 18916 PERcaer; 20146 PER 
schi; 20 1 47 PER amer; 24 1 1 OCO vala; 24 1 44 
PER vest; 24275 OCO insu; 24307 LIC mult; 
24487 OCO vera; 24488 NEC glob; 24540 OCO 
vera; 24584 OCO nica; 24593 NEC glob; 265 12 
OCO vera; 26527 OCO flor; 26600 NEC glob; 
26612 NEC sali; 28570 PHO cinn; 28623 OCO 
vala; 28706 OCO oblo; 28732 OCO aura; 289 1 
PER schi; 2891 1 PER amer; 28996 OCO vala; 
29044 OCO gome. 

Williams, R. S.: 319 NEC glob. 

Woodson, R. E., et al.: 1022 OCO glau; 1099 PHO 
cinn. 

Woodworth, R. H., & P. A. Vestal: 47 1 OCO cern. 

Zamora, N., et al.: 367 NEC niti; 394 NEC reti; 
399 LIC sp. A; 410 OCO tene; 442 OCO atir; 
674 OCO valo; 719 OCO cern; 729 PHO cinn; 
735 PER schi?; 770 NEC memb; 824 NEC sali; 
879 OCO hold; 880 OCO pitt; 915 OCO vera; 
1014 WIL glau; 11 57 NEC sali; 11 77 NEC glob; 
1 208 CAR burg; 1 2 1 5 BEI oval; 1 287 OCO heli; 
1300 OCO leuc. 



HERNANDIACEAE 

By William Burger 

REFERENCE K. Kubitzki, Monographic der 
Hernandiaceen. Bot. Jahrb. Syst. 69: 78-209. 1969. 

Small to medium-sized trees, less often shrubs or lian- 
as, usually monoecious (bisexual), rarely dioecious, ev- 
ergreen or deciduous, nodes unilacunar, oil cells often 



present; stipules absent. Leaves alternate in a spiral, sim- 
ple (in Central America) or palmately compound with 
3-5 leaflets, petiolate, leaf blades entire or 3-5-lobed, 
venation palmate or pinnate. Inflorescences axillary to 
distal leaves or rarely terminal, dichasial or thyrselike, 
often with a well-developed primary peduncle and much 
branched distally, with or without bracts and bracteoles. 
Flowers usually very small, unisexual or bisexual, radi- 
ally symmetrical; tepals (sepals) 4-10 in 1 or 2 whorls, 
free or united at the base; stamens 3-6 in 1 whorl (often 
opposite the outer whorl of tepals, filaments free, often 
with 2 basal and lateral or abaxial "glands" (usually 
narrowed at the base and often cordate), anthers 2-the- 
cous, opening by large flaps from the bottom upward 
(often remaining attached near the apex of the anther), 
staminodes absent or glandlike (relationship to glands 
and interpretation problematical), pollen inaperturate; 
pistil 1 (carpel apparently 1), ovary inferior and l-lo- 
cular, ovule solitary and pendulous from near the apex 
of the locule, style slender with capitate or broad stigma. 
Fruits 1 -seeded nuts or drupes, with distal wings (in 
Gyrocarpus), lateral wings (in Illigera), or without wings 
(enclosed in a cupule of similar texture as the fruits in 
Hernandia); embryo large, straight, the cotyledons fold- 
ed or rolled up. 

A family of four genera and about 65 species. 
They are pantropical but with some very unusual, 
apparently relict, distributions. Many species oc- 
cur on oceanic island-groups, but the species are 
not adapted to seaside or littoral vegetation (see 
Kubitzki, 1969). Costa Rican species are recog- 
nized by their unusual fruits (very different in the 
three different genera), variable and often long pet- 
ioles, palmately veined leaves (in most) that often 
vary greatly in size, small or very small flowers 
with anthers opening by two flaps, and narrow 
inferior ovary with single locule and ovule. The 
family appears to be closely related to the Lau- 
raceae with which it shares features of androecial 
morphology; it differs in the inferior ovary. These 
plants also resemble some Menispermaceae, both 
morphologically and in the rarity with which they 
are collected. While Kubitzki's monograph is ex- 
cellent, the paucity of collections may require reas- 
sessment of species, if and when additional ma- 
terial becomes available. 



Key to Species of Hernandiaceae 

1 a. Leaves deciduous, often 3- or 5-lobed (in Central America); trees of seasonally very dry and deciduous 
forest formations; fruits with 2 long spatulate or oblanceolate distal wings; flowers 2-4 mm long 
and unisexual Gyrocarpus 

1 b. Leaves evergreen or shortly deciduous; without lobes; trees, shrubs and lianas of evergreen forest 
formations below 1 500 m elevation; fruits without wings; flowers unisexual or bisexual 2a 



BURGER: FLORA COSTARICENSIS 



129 




FIG. 23. Hemandiaceae: A, Sparattanthelium amazonum; B, S. septentrionale, C, Gyrocarpus jatrophifolius; D, 
Hernandia stenura with distal flower-pair; E, H. didymantha. 



130 



FIELDIANA: BOTANY 



2a. Flowers ca. 5 mm long, 1 female and 1 -3 male flowers borne together and subtended by a whorl of 
usually 4 sepaloid bracts; fruits partly enclosed within a cupule Hernandia 

2b. Flowers ca. 2 mm long, apparently bisexual and not subtended by bracts or bracteoles; fruits 

developing at the ends of a few inflorescence branches and often becoming silvery white 

Sparattanthelium 



Gyrocarpus Jacquin 

Trees, rarely shrubs, deciduous, bisexual, branches 
often thick, puberulent or glabrous. Leaves simple, usu- 
ally clustered at the ends of branches, long-petiolate, leaf 
blades often broader than long, entire and lobed or un- 
lobed, venation usually palmate, glabrous or puberulent. 
Inflorescences developing before the leaves or with the 
leaves present, clustered at the ends of branchlets, flowers 
usually in small clusters in dichasial, thyrselike or umbel- 
like arrangements, with or without bracts and bracteoles. 
Flowers bisexual (perfect) or male, very small, parts of 
the flower often early caducous, perianth usually of 7 (6, 
8) parts, 2 parts (lobes) usually much larger and with 2 
smaller lobes attached to each large lobe; stamens 4 or 
5(-7), filaments free, puberulent or glabrous, anthers 
opening by 2 valves (flaps) from the bottom upward, 2- 
thecous; staminodes alternating with the stamens or sol- 
itary and opposing the style, clavate or thickened and 
flattened at the apex; ovary inferior and tomentulose, 
style and stigma 1. Male flowers with reduced ovary, 
more numerous than perfect flowers, wings not devel- 
oping. Fruits drupes with 2 long apical oblanceolate wings 
with rounded tips (narrowly spatulate), developing from 
primordia within the perianth whorl, basal part (ovary) 
globose to ovoid or ellipsoid, puberulent to glabrous; 
seed with leafy cotyledons spirally twisted. 

A genus with only three species: one in eastern 
Africa; G. jatrophifolius from Mexico to central 
Costa Rica; and G. americanus, with a very un- 
usual pantropical distribution. The genus is dis- 
tinguished by the unusual 2-winged fruits, inferior 
ovary, anthers opening by flaps, and restriction to 
seasonally dry areas. The wings do not develop 
from the perianth lobes, but rather from a pair of 
primordia arising interior to the perianth; see dis- 
cussion in the monograph by Kubitzki (1969). 



Gyrocarpus jatrophifolius Domin, Biblioth. Hot. 
22, heft 89: 682. 1925 (1926). Figure 23. 

Small to medium-sized (5-22 m) trees, occasionally a 
shrub, bark corrugated with corky ridges, leafy inter- 
nodes (2-)4-15 mm thick, longitudinally grooved (dry), 
becoming glabrous and grayish. Leaves usually borne 
near the ends of stems, petioles 4-28 cm long, 1-5 mm 
thick, glabrescent and canaliculate above; leaf blades 6- 
20(-30) cm long, 5-25MO) cm broad, 3- or 5-lobed, the 
3 distal lobes with 2 deep sulci (more than Vi the length 
of the lobe) and acuminate apices, cordate to truncate at 
the base with a portion of the major lateral veins forming 



part of the lamina base, margins entire, drying thin- to 
thick-chartaceous, glabrous or minutely (0.1-0.4 mm) 
puberulent on the veins above and below with thin yel- 
lowish hairs, venation palmate with 3 major veins, mid- 
vein with 4-8 pairs of major secondary veins. Inflores- 
cences 4-14 cm long and growing longer in fruits, primary 
peduncles often '/2 the length of the inflorescence, densely 
yellowish puberulent, dichasial or thyrselike, often aris- 
ing from thickened and rounded areas on the stem (these 
with circular peduncle scars after the infructescences fall 
off). Flowers small and green, 2-4 mm long; stamens 
usually 5 (4-7), filaments glabrous or puberulent near 
the base, 1-3 mm long, anthers ca. 0.6 mm long, de- 
hiscing flaps remaining attached at the apex. Fruits with 
2 apical stiffly chartaceous wings, (4.5-)7-l 1(-13.5) cm 
long and 7-14 mm broad, broadest above the middle, 
parallel or spreading and V-shaped, minutely puberulent 
or glabrescent, base of the fruits (ovary) 16-29 mm long 
and 6-10 mm thick, ellipsoid to narrowly ovoid, gla- 
brescent or densely tomentulous with yellowish or gray 
hairs. 

Trees of seasonally very dry deciduous forest 
and woodland formations from near sea level to 
900 (1400) m elevation (rarely higher in Guate- 
mala). Flowers have been collected in September- 
December and March, and fruits have been col- 
lected in December-March and May in Central 
America. The species ranges from Sinaloa, Mex- 
ico, southward along the Pacific slope to central 
Costa Rica. 

Gyrocarpus jatrophifolius is recognized by its 
distinctive fruits with two long narrowly oblan- 
ceolate wings above an inferior ovary, the leaves 
with three prominent distal lobes and two smaller 
lateral lobes (all with acuminate apices), minute 
flowers with anthers dehiscing by two flaps, and 
the restriction to deciduous forest formations. This 
species is known from only a few collections in 
Costa Rica made in lowland Guanacaste Province 
and the western side of the Meseta Central. Ku- 
bitzki (1969) distinguished this species from G. 
americanus Jacq., which has more rounded fruits, 
leaves that are usually unlobed (in Central Amer- 
ica), flowers with usually four stamens, and which 
reaches its southern limit in northern Nicaragua. 
Most authors previously have considered the two 
as parts of a single rather polymorphic species; the 
two species (as delimited by Kubitzki) are sym- 
patric in northern and central Central America. 
The name gallito is commonly used for these plants 



BURGER: FLORA COSTARICENSIS 



131 



in El Salvador and Nicaragua; other names are 
volador, volatin, and caballitos. 



Hernandia Linnaeus 

Trees, rarely shrubs, monoecious (bisexual), usually 
evergreen, stems often thick and with soft wood; stipules 
absent. Leaves simple and alternate, petiolate; leaf blades 
usually with entire margins (rarely 3-5-lobed), venation 
pinnate or palmate. Inflorescences usually axillary to dis- 
tal leaves, corymbiform panicles in Central America, 
usually with an elongate primary peduncle, inflorescence 
and bracts usually minutely tomentulous, the distal flow- 
er clusters often flower-like because they are subtended 
by a whorl of 4 sepaloid bracts, the distal flower clusters 
bisexual with a central female flower (rarely 2 or 3) and 
1 -4 outer short-pedicellate male flowers, the female flow- 
ers subtended by and arising from within a deep cup 
formed by bracteoles (the cup sessile and with an entire 
distal margin in our species), bracteoles of the male flow- 
ers small and basal or absent. Flowers unisexual, outer 
perianth parts imbricate, the inner whorl imbricate to 
valvate, free; male flowers with 6-8 perianth parts (in 
whorls of 3-6 parts), stamens 3-5(-6), filaments free or 
united for '/2 their length, usually each filament with 2 
small appendages (glands) near the base; female flowers 
4-6-parted, staminodia absent, 4 or 5(-10, 12) free or 
connate glands present at the base of the style, often 
opposing the outer perianth segments, ovary inferior and 



slightly compressed, style well developed but not ex- 
ceeding the perianth segments in length, stigma discoid- 
peltate. Fruits ovoid, ellipsoid, or subglobose drupes, or 
becoming hard and nutlike, often subtended and en- 
closed in the expanded and inflated cupule, the cupule 
often with 8 longitudinal ridges, usually minutely to- 
mentulous (both cups and fruits); cotyledons folded. 

A pantropical genus of 24 species best repre- 
sented in the southwestern Pacific and West Indies 
with a few species in Africa, Madagascar, Austra- 
lia, and Central and northernmost South America; 
see the treatment by Kubitzki (1969). Our species 
are recognized by the unusual partial inflores- 
cences with two male and one female flower sub- 
tended by a whorl of four sepaloid bracts, the fe- 
male flower arising from within a cup, the anthers 
opening by two flaps, the inferior ovary and flat- 
tened stigma, and the drupelike fruits included 
within the enlarged cupule. Inflorescence, flowers, 
and fruits are usually covered by fine minute to- 
mentulose (velutinous) hairs. Soft wood, thick 
stems, great variation in leaf size and petiole length, 
and restriction to wet evergreen forests further dis- 
tinguish our species. These plants are poorly rep- 
resented in herbaria. 



Key to Species of Hernandia 

la. Leaves with the basal pair of secondary veins strongly developed and the venation often subpalmate, 
leaf blades usually ovate and often long-acuminate, larger leaves often slightly peltate; partial (distal) 
inflorescences usually 3-flowered; uncommon H. stenura 

Ib. Leaves with the basal pair of secondary veins not strongly developed, the venation pinnate, leaf 
blades often oblong, short-acuminate to acute, larger leaves not known to be peltate; partial inflo- 
rescences usually 2-flowered; rarely collected H. didymantha 



Hernandia didymantha J. D. Smith, Bot. Gaz. 31: 
120. 1901. Figure 23. 

Trees to over 1 5 m high, lower branches descending. 
Leaves simple and unlobed, petioles (1.5-)3-7(-9) cm 
long, canaliculate above (adaxially); leaf blades 9-1 8(-2 1 ) 
cm long, 3.5-7 cm broad, oblong to ovate-oblong or 
slightly obovate. bluntly acute to short-acuminate at apex, 
obtuse or rounded at the base, margins entire, drying 
subcoriaceous, glabrous above and below, venation pin- 
nate with (4-)6-8 pairs of major secondary veins, the 
basal secondaries not more prominent than the distal 
secondaries. Inflorescences terminal or axillary to distal 
leaves, paniculate, distal flower clusters usually with only 
2 flowers, bracts ca. 8 mm long and 3.5 mm broad, 
rounded at the apex and becoming reflexed. Male flower 
usually solitary on a long (5 mm) pedicel, parts in whorls 
of 3; female flower solitary and very short pedicellate, 
4-parted, style with 4 stipitate glands at the base. Fruits 



1.8-2.5 cm in diameter, subglobose, conical at the apex, 
enclosed within a coriaceous cup 2-3 cm in diameter. 



This species is only known from lowland ev- 
ergreen rain forest formations. It has been col- 
lected from Punta Mona (Pittier 12682, us, the 
type), La Selva (Hammel 11656, DUKE; Hartshorn 
802, 1026, CR, F, McDowell 606, DUKE), the Golfo 
Dulce region (Allen 598, F, Q. Jimenez et al. 644, 
CR, F), and near Almirante, Bocas del Toro, Pan- 
ama (Cooper 6 18, MO). These flowering collections 
were made in January-April and November. A 
collection with fruits (Hartshorn 1281, CR, F) from 
La Virgen de Sarapiqui was made in August. This 
collection has the fruits almost entirely enclosed 
by a cupulate receptacle whose distal aperture is 



132 



FIELDIANA: BOTANY 



only 3-6 mm wide; see also the photograph in P. 
E. Sanchez, Florula del Parque Nacional Cahuita, 
p. 141, 1983, where the fruiting season is said to 
be September-November. 

Hernandia didymantha is recognized by the ob- 
long leaves usually lacking strongly developed bas- 
al secondary veins, short acuminate apices, very 
variable petiole lengths, and characters of the ge- 
nus. Additional collections may show that this 
species intergrades with material here treated as 
H. stenura. If that happens, H. didymantha, the 
earlier name, would have to be circumscribed more 
broadly to cover all our material of Hernandia. 
These trees appear to produce leaves in "flushes" 
of new growth and it may be that conditions at 
the time of the leaf flush may determine the size 
and form of the leaves. If this is the case, it may 
mean that H. hammelii D'Arcy of Panama is only 
a small-leaved (6-8 x 2. 5-4 cm) growth form (Ann. 
Missouri Dot. Gard. 68: 224, 1981). Another re- 
cent small-leaved collection is from the Dept. of 
Yoro, Honduras (Hazlett 2714, F). All this ma- 
terial shows considerable variation in leaf size on 
the same branch, and all appear to live in the same 
kinds of moist forests. 



Hernandia stenura Standl., Publ. Field Mus. Nat. 
Hist. Bot. Ser. 18: 1553. 1938. Figure 23. 

Trees to over 25 m tall, leafy internodes 1.8-8 mm 
thick, glabrous or very minutely (0.1 mm) scurfy yel- 
lowish puberulent. Leaves simple and unlobed, juvenile 
and sapling leaves often large and slightly peltate, peti- 
oles 5-18 cm long; leaf blades 9-27(-40) cm long, 7- 
1 8(-25) cm broad, broadly ovate to ovate-elliptic or ovate- 
oblong, abruptly acuminate at the apex with a tip 1-3.5 
cm long (rarely acute), rounded to subcordate at the base, 
the largest leaves sometimes peltate with the petiole at- 
tachment up to 1 cm distant from the lamina margin, 
drying thick-chartaceous or slightly subcoriaceous, mar- 
gin entire or somewhat undulate, glabrous above, gla- 
brous or minutely (0. 1 mm) grayish puberulent on the 
veins beneath, venation pinnate or subpalmate with the 
lowermost secondaries often quite prominent, usually 
with 3-5 pairs of major secondary veins. Inflorescences 
8-30 cm long with prominent primary peduncles 5-20 
cm long, paniculate, sweet-scented, the ultimate parts 
flower-like with usually 4 sepal-like bracts (ca. 10x5 
mm) subtending 1 subsessile female flower (and its cu- 
pule) and 1-4 pedicellate male flowers, all parts covered 
with a fine (0. 1 mm) dense tomentum of soft hairs white 
in life but becoming grayish. Male flowers ca. 6 mm long 
(in bud) on pedicles 4 mm long, parts in whorls of 3, 
filaments with 2 minute glands; female flowers arising 
from within cupules ca. 3 mm long, on a short (3-4 mm) 
pedicel, flower buds ellipsoid, ca. 4-5 mm long, perianth 
4-parted. Fruits drupes ca. 2 cm broad and included 
within the subglobose cupule narrowed at the round dis- 



tal aperture, fruiting cupule 1.3-2.8 cm long and 1.5-2.8 
cm broad (when dry), minutely grayish velutinous/pu- 
berulent on all surfaces, distal aperture 1-2 cm broad. 

Trees of wet evergreen forest formations from 
near sea level to 1000 (1400) m altitude; flowers 
have been collected in November-January and 
March-May; fruits have been collected in March, 
June, and September. The species ranges from 
Guatemala to the province of Code in Panama; 
Costa Rican material has been collected along the 
Caribbean slope and coastal plain and in the Gen- 
eral Valley. 

Hernandia stenura is recognized by the long pet- 
iolate and broadly ovate leaves with subpalmate 
venation (sometimes peltate), thick stems with soft 
wood, distal flower clusters that resemble individ- 
ual flowers, anthers opening by a flap, and the 
inferior ovary developing into a drupe that is large- 
ly surrounded within a slightly succulent cupule. 
The cupule is quite distinctive as its distal aperture 
is entire and somewhat smaller than the enclosed 
fruits. The dense minute grayish tomentum on all 
flowering parts is distinctive and reminiscent of 
that seen in some Lauraceae. These are infre- 
quently collected trees, but they exhibit consid- 
erable variation, both between collections and 
within collections. It is possible that there is in- 
tergradation with H. didymantha; see the discus- 
sion under that species. Material of this species 
was mistakenly assigned to H. sonora L. by Stan- 
dley in "Flora of Guatemala" (Fieldiana, Bot. 24(4): 
346. 1946). 



Sparattanthelium Martins 

Shrubs, small trees, or lianas, evergreen, climbing with 
recurved hooks (modified short shoots) in some species. 
Leaves simple and entire, petiolate, leaf blades often 
narrowly ovate to lanceolate (ovate to obovate), venation 
palmately tripliveined (rarely pinnate), glabrous or pu- 
berulent. Inflorescences axillary from distal leaves, sol- 
itary at a node, much branched dichasia with many small 
distal flowers, bracts and bracteoles absent. Flowers very 
small, radially symmetrical, apparently bisexual, many 
are early caducous after anthesis (perhaps functionally 
male); perianth 4-8-parted, valvate in bud (in 4-5-parted 
flowers) or imbricate, free to near the base, persisting; 
stamens 4, 5(-7), alternating with the calyx lobes, fila- 
ments glabrous and short, anthers elongate, apiculate at 
the apex and opening laterally, flaps attached at the apex, 
both staminodes and glands absent; style straight, stigma 
discoid or lobed. Fruits developing only on the most 
distal branches of the infructescence (with very few fruits 
developing on an infructescence), branches of the in- 
fructescence with thickened nodes and somewhat zigzag 
(fracti-flexuous) in form, both the fruits and the stems 



BURGER: FLORA COSTARICENSIS 



133 



becoming silvery white or yellowish, drupes ovoid to 
elongate ellipsoid, often with longitudinal ribs; testa 
chartaceous, embryo, with inrolled cotyledons. 

An American genus of 1 3 species ranging from 
southern Mexico to Peru, Bolivia, and southern 
Brazil. The genus is recognized by the usually tri- 
pliveined leaves on long slender petioles, the nu- 
merous very small flowers on complex inflores- 
cences with apparently dichotomous (but 



somewhat unequal) branching, the bisexual flow- 
ers with anthers opening by flaps, and the fruits 
apparently developing only from distal nodose in- 
florescence branches. Fruits and inflorescence be- 
come silvery white in some species. Specimens of 
this genus have been collected fewer than five times 
in Central America, but it seems likely that two 
species occur in southern Central America. 



Key to Species of Sparattanthelium 

la. Leaves ovate to oblong or elliptic-oblong, abruptly short-acuminate; flowers usually with 5 perianth 
parts S. septentrionale 

1 b. Leaves narrowly elliptic and tapering gradually to the acuminate apex; flowers mostly with 4 perianth 
parts 5". amazonum 



Sparattanthelium amazonum Martins. Flora 24, 
Beibl 2: 42. 1841, Denkschr. Bayer. Ges. Re- 
gensburg 3: 303, t. 11, f. 2. 1841. S. guatema- 
lenseStondl, Proc. Biol. Soc. Wash. 37:51.1 924. 
S. amazonum ssp. guatemalense (Standl.) Ku- 
bitzki, Bot. Jahrb. Syst. 89: 202. 1969. Figure 
23. 

Shrubs to 6 m tall, scandent or becoming lianas, leafy 
internodes 0.5-5 cm long, 1.5-4 mm thick, glabrous to 
densely puberulent with thin grayish straight hairs to 0.6 
mm long. Leaves with petioles 1 .2-6 cm long, 0.5-1 mm 
thick, glabrous or puberulent; leaf blades 6-16 cm long, 
2.5-5.5 cm broad, lanceolate to narrowly elliptic or el- 
liptic-oblong (rarely ovate), tapering gradually to the acu- 
minate apex, the tip 0.4-1.5 cm long, rounded to obtuse 
at the base, margin entire or minutely undulate, drying 
thin-chartaceous, glabrous or puberulent along the major 
veins above and below, the hairs slender white and straight 
to 0.7 mm long, venation trinerved from slightly above 
the base of the lamina (lateral veins form the lamina 
margin at the very base), midvein with only 1 or 2 pairs 
of major secondary veins. Inflorescences to ca. 1 cm 
long, axillary to distal leaves, branches slender and gray- 
ish puberulent. Flowers small, ca. 2 mm long, perianth 
4 (rarely 5) parted, 1 .2-1 .5 mm long, 0.4-0.8 mm broad, 
acute to obtuse at the apex, puberulent; stamens 4 (5), 
anthers subsessile; ovary (inferior) ca. 0.6 mm long, style 
slender 1-1 .4 mm long. Fruits elongate ellipsoid drupes, 
ca. 1 .7 cm long and 0.8 cm thick, fruits and inflorescences 
becoming white. 

Lianas or shrubs in lowland rain forest forma- 
tions. Only three collections of this species are 
known from Central America: Standley 25066 
(type of 5. guatemalensis) from Puerto Barrios; 
Contreras 8957 from Puerto Mendes, both in the 
Dept. of I/abal. Guatemala; and Herrera 1210, 



CR, F, MO, from near the Nicaraguan border in 
Alajuela Province, Costa Rica. Flowers were col- 
lected in June and August. Immature fruits were 
collected by Herrera in November. The other col- 
lections of this species are from Brazil and Bolivia. 
Sparattanthelium amazonum is recognized by 
the narrow elliptic acuminate leaves on long slen- 
der petioles, the trinerved venation, minute flow- 
ers with four perianth parts on thin-branched little 
inflorescences, and the anthers opening by flaps. 

Sparattanthelium septentrionale Sandwith, Kew 
Bull. 1932: 226. 1932. Figure 23. 

Shrubs (perhaps scandent), branchlets conspicuously 
and densely puberulent in early stages, the hairs slender 
0. 1-0.5 mm long, leafy internodes 0.8-3 mm thick. Leaves 
often very variable on the same stem, petioles 1.3-4.2 
cm long, slender, puberulent; leaf blades 6-15 cm long, 
2-8 cm broad, ovate to elliptic-oblong, abruptly acu- 
minate at the apex with a tip 5-15 mm long, obtuse to 
rounded at the base, margins entire or slightly undulate, 
drying chartaceous, puberulent above and below in early 
stages with slender whitish hairs 0.1-0.8 mm long but 
the longer hairs falling off, trinerved from the base, the 
midvein with 2 or 3 pairs of distal secondary veins. 
Inflorescences said to be 5 cm long (probably becoming 
larger), branches of the inflorescence slender and grayish 
puberulent, pedicels to 4 mm long. Flowers about 1 mm 
in diameter in bud, perianth 5-(6-)lobed, perianth lobes 
1.5-2 mm long and 1 mm wide, the lobes elliptic and 
obtuse at the apex; stamens 5 (4), filaments 0.5 mm long, 
anthers ca. 1 mm long; style 1.5-2 mm long. Fruits un- 
known. 

Probably confined to lowland evergreen forest 
formations and known from only three collections, 



134 



FIELDIANA: BOTANY 



ranging from Tabasco/Yucatan in Mexico to Costa 
Rica. 

Sparattanthelium septentrionale is distin- 
guished by the broader, more ovate or oblong leaves 
and the usually 5-parted perianth. 



Literature Cited 

ALLEN, C. K. 1945. Studies in the Lauraceae, VI. Pre- 
liminary survey of the Mexican and Central American 
species. J. Arnold Arbor., 26: 280-434. 

. 1966. Notes on tropical American Lauraceae. 
II. Costa Rica. Phytologia, 13(3): 232. 

ALLEN, P. H. 1956. The Rain Forests of Golfo Dulce. 
University of Florida Press, Gainesville, Fla., 417 pp. 

BERNARDI, L. 1 962. Lauraceas. Universidad de los An- 
des, Facultad de Ciencias Forestales, Merida, Vene- 
zuela, 355 pp. 

. 1 967. Emendationes laureae imprimis de Nec- 

tandra. Candollea, 22: 49-67, 69-84, 91-101. 

BLAKE, S. F. 1919. The anay, a new edible-fruited rel- 
ative of the avocado. J. Wash. Acad. Sci., 9: 457-462. 

BURGER, W. C. 1988. A new genus of Lauraceae from 
Costa Rica, with comments on problems of generic 
and specific delimitation within the family. Brittonia, 
40: 275-282. 

CROAT, T. B. 1978. Flora of Barro Colorado Island. 
Stanford University Press, Stanford, Calif., 943 pp. 

HAMMEL, B. E. 1986. The vascular flora of La Selva 
Biological Station, Costa Rica. Lauraceae. Selbyana, 
9:219-233. 



HOWARD, R. A. 1981. Nomenclatural notes on the Lau- 
raceae of the Lesser Antilles. J. Arnold Arbor., 62: 45- 
61. 

KOPP, L. E. 1966. A taxonomic revision of the genus 
Persea in the Western Hemisphere (Perseae-Laura- 
ceae). Mem. New York Bot. Gard., 14(1): 1-120. 

KOSTERMANS, A. J. G. H. 1938. A monograph of the 
genera: Anaueria, Beilschmiedia (American species) 
and Aniba. Recueil Trav. Bot. Neerl., 35: 835-928. 

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

. 1964. Bibliographia Lauracearum. Ministry of 

National Research, Bogor, Indonesia. 

KUBITZKI, K. 1 969. Monographic der Hernandiaceen. 
Bot. Jahrb. Syst., 69: 78-209. 

KURZ, H. 1983. Fortpflanzungsbiologie einiger Gat- 
tungen neotropischer Lauraceen und Revision der 
Gattung Licaria (Lauraceae). Diss., Universitat Ham- 
burg, Hamburg, F.R.G. 

MEZ, C. 1 889. Lauraceae Americanae. Jahrb. Konigl. 
Bot. Gart. Berlin, 5: 1-556. 

PEMBERTON, R. W., AND C. E. TURNER. 1989. Occur- 
rence of predatory and fungivorous mites in leaf dom- 
atia. Amer. J. Bot., 76: 105-1 12. 

ROHWER, J. G. 1986. Prodromus einer Monographic 
der Gattung Ocotea Aubl. (Lauraceae), sensu lato. Mitt. 
Inst. Allg. Bot. Hamburg, 20: 1-278. 

ROHWER, J. G., AND K. KUBITZKI. 1985. Entwicklungs- 
linien im Ocotea-Komplex (Lauraceae). Bot. Jahrb. 
Syst., 107: 129-135. 

WILLIAMS, L. O. 1 977. The avocados, a synopsis of the 
genus Persea, subg. Persea. Econ. Bot., 31: 315-320. 



BURGER: FLORA COSTARICENSIS 



135 



Index 

The index includes all accepted names (in Roman type), synonyms (italics), common English names 
(Roman), and vernacular Spanish names (italics). New species are in boldface, and the page numbers of 
illustrations are in boldface. Hyphenated words and multiple words are alphabetized by letter. 



Abacate 103 
Acrodiclidium 47 

excelsum 49 

kunthianum 59 

triandrum 5 1 
Aguacate 103 
Aguacate de montana 107 
Aguacatillo 103, 105 
Aguacatdn 107 
Aiouea 36 

costaricensis 23, 36, 84 

lundelliana 37, 84 

obscura 14, 37 

parvissima 38 

talamancensis 17, 37 

vexatrix 72 
Alligator pear 103 
Aniba 38 

intermedia 39 

venezuelana 26, 39 
Ant plants 16, 78 
Ascd 109 
Avocado 103 



Beilschmiedia 39 

alloiophylla 41 

ana y 20, 40 

austin-smithii 4 1 

brenesii 42 

costaricensis 42 

mexicana 43 

ovalis 18,41 

pendula 26, 42 

rigida 40 

sulcata 28, 42 
Bellota costaricensis 36 
Boldus costaricensis 36 
Butter pear 103 



Caballitos 132 
Caryodaphnopsis 43 

burgeri 14, 43 
Cassytha 44 

filiformis 44 

paradoxa 45 
Chanco bianco 42 
Chanekia 47 
Cinnamomum 45 

camphora 45 

cassia 45 

verum 45 

zeylanicum 45 



Coevolution 

with ants, see species keyed in di- 
chotomy 43a, fig. 3 7, 14 

with mites 2 

with birds 103 
Collectors, important 2 

index to 121 
Come negro 42 
Cryptocarya 44 

kostermansiana 42 
Cura aguacate 103 
Cuscata 44 



Domatia 2 



Endlicheria 45 

formosa 45 

multiflora 46 

sp.? 22, 46 

sprucei 46 

verticillata 46 
Exiccatae, index for Lauraceae 1 2 1 



Gallito 131 
Gyrocarpus 131 

americanus 131 

jatrophifolius 130, 131 



Hernandia 132 

didymantha 130, 132 

hammelii 133 

sonora 133 

stenura 130, 133 
Hernandiaceae 129 
Hufelandia anay 40 

costaricensis 42 

ovalis 4 1 



Index to collections of Lauraceae 1 2 1 
Ira amarillo 92 
Ira rosa 74 
Ira zoncho 76 



Key, artificial 4 
diagnostic 3 



to illustrations (comparative fig- 
ures) 13 



Lauraceae 1 

Laurografia Peruviana 73 

Laurus aurantiodora 72-73 

caerulea 104 

floribunda 79 

globosa 58 

leucoxylon 87 

membranacea 62 

pendula 42 

persea 103 

puberula 93 

purpurea 60 

reticulata 64 

sulcata 41, 42 

triandra 5 1 
Lentisco 53 
Licaria 46 

alata 49 

brenesii 16, 47 

cervantesii 52 

coriacea 52 

cufodontisii 29, 48 

excelsa 29, 49 

guatemalensis 51 

limbosa 52 

misantlae 49 

multinervis 29, 49 

pergamentacea 29, 50 

pittieri 51, 52 

reclinata 52 

sarapiquensis 29, 5 1 

sp. A 52 

tikalana 52 

triandra 29, 5 1 
Litsea 53 

acuminatissima 53 

flavescens 53 

glaucescens 17, 53 

glaucescens var. flavescens 53, 54 

guatemalensis 53 

neesiana 53 



Mespilodaphne aurantiodora 73 

oblonga 90 

Mezilaurus glaucophylla 1 1 9 
Misanteca 47 

costaricensis 5 1 

excelsa 49 

pittieri 5 1 

triandra 5 1 



136 



FIELDIANA: BOTANY 



Nectandra 54 
austinii 56 
belizensis 21, 56 
bijuga 74 
brenesii 75 
caucana 58 
cissiflora 24, 57 
concinna 66 
coriacea 65 

cufodontisii 25, 57, 60 
davidsoniana 58, 98 
gentlei 62 
glabrescens 58 
globosa 31, 58 
heydeana 75 
hypoglauca 80 
hypoleuca 31, 59 
kunthiana 20, 59 
latifolia 30, 60 
longipetiolata 26, 60 
lundellii 60 
martinicensis 30, 6 1 
membranacea 30, 62 
mollis 64 
nervosa 66 
nitida 30, 62 
panamensis 67 
paulii 57 
perdubia 62 
producta 94 
purpurascens 60 
purpurea 60 
ramonensis 31, 63 
reticulata 20, 64 
salicifolia 30, 64 
salicina 17, 65 
savannarum 65 
schippii 56 
sinuata 20, 66 
skutchii 62 
smithii 65 
standleyi 62 
trianae 1 1 7 
turbacensis 31, 66 
whitei 99 
woodsoniana 6 1 



Ocotea 67 
atirrensis 16, 72 
aurantiodora 72 
austinii 18, 73 
babosa 21, 74 
bakeri 98 
bernoulliana 96 
brenesii 26, 75 
calophylla 18, 76 
cernua 27, 77 
cooper i 59 
cufodontisii 57 
cuneata 84 
cuneifolia 9 1 
dendrodaphne 16, 77 
dentata 21, 33, 78 
endresiana 23, 79 



eucuneata 9 1 
eusericea 99 
floribunda 27, 79 
florulenta 75, 91 
fulvescens 76 
glaucosericea 25, 80 
gomezii 19, 8 1 
grandifolia 73 
guianensis 76 
hartshorn iu nu 21, 8 1 
helicterifolia 21, 82 
heydeana 75 
holdridgeiana 25, 83 
insularis 23, 37, 84 
in 84, 85 
irazuensis 73 
laetevirens 25, 85 
latifolia 60 
lenticellata 87 
lentil 15, 86 
leucoxylon 27, 87 
longifolia 73 
meziana 27, 87 
micans 76 
mollicella 17, 88 
mollifolia 20, 88 
mollis 64 

monteverdensis 17, 89 
neesiana 9 1 
nicaraguensis 16, 90 
oblonga 24, 90 
opifera 73 
ovandensis 77 
palmana 93, 116 
paradoxa 97 
paulii 16, 91 
pedalifolia 72 
pentagona 90 
pittieri 17, 92 
portoricensis 90 
pseudopalmana 19, 92 
puberula 93 
pyramidata 93 
quisara 77 
rivularis 15, 94 
salicifolia 64 
seibertii 57 
sericea 76 
skutchii 24, 94 
sp. A aff. laetevirens 100 
sp. aff. bijuga 24, 74 
sp. aff. caracasana 23, 76 
sp. B 100 
standleyi 86 
stenoneura 95 
subsericea 87 
tabascensis 62 
tenera 26, 95 
tonduzii 84, 85 
turbacensis 66 
valeriana 19, 96 
valerioides 15, 97 
velutina 76 

veraguensis 27, 97-98 
verapazensis 86 
viridiflora 98 



wachenheimii 79 
wedeliana 99 
whitei 24, 99 
williamsii 94, 95 
Oreodaphne caracasana 76 
cernua 77 
helicterifolia 82 
mexicana 82 



Persea 101, 108 

albida 28, 102 

americana 28, 103 

americana var. nubigena 103 

austin-smithii 4 1 

brenesii 104 

caerulea 22, 104 

chiapensis 108 

cinnamomifolia 1 1 1 

cuneata 106 

donnell-smithii 105 

drymifolia 103 

gigantea 103 

laevigata 104 

mexicana \ 1 1 

nubigena 103 

obtusifolia 18, 105 

pallida 102 

petiolaris 104 

pittieri 107 

popenoei 108 

povedae 28, 105 

rigens 28, 106 

schiedeana 19, 107 

silvatica34, 107 

skutchii 104 

veraguasensis 22, 108 

veraguensis 108 

vesticula 18, 108 
Persea, subgenus 

Eriodaphne 101, 108 

Machilus 108 

Persea 101, 108 
Phoebe 109 

amplifolia 93, 109, 110 

belizensis 56 

betazensis 82 

bourgeauviana 92 

brenesii 14, 1 10 

chavarriana 15, 111 

cinnamomifolia 14, 109, 111 

costaricana 109, 111, 112 

hammeliana 25, 1 1 3 

helicterifolia 82 

insularis 84 

macrophylla 86 

mayana 90 

mexicana 1 1 1 

mollicella 88 

neurophylla 14, 113 

psychotrioides 92 

neurophylla 14, 113 

pittieri 92 

smithii 96 

tonduzii 111, 112 

valeriana 96 



BURGER: FLORA COSTARICENSIS 



137 



Pleurothyrium 114 
golfodulcense 22, 115 
hexaglandulosum 1 1 6 
palmanum 19, 116 
sp. A 117 
trianae 22, 117 
velutinum 76 

Povedadaphne 1 1 7 
quadriporata 35, 118 



Quina 50 
Quizarrd 58 
Quizarrd amarillo 88 
Quizarrd negra 60 
Quizarrd torita 5 1 



Rhodostemonodaphne 60 

kunthianum 59 
Ruiz & Pavon, publication 73 



Sassafridium veraguense 97 
Sigua 99 
Sparattanthelium 133 

amazonum 130, 134 

amazonum ssp. guatemalense 1 34 

guatemalense 134 

septentrionale 130, 134 
Species of uncertain generic posi- 
tion 119 
Synandrodaphne 60 



Tetranthera glaucescens 53 
Tiguissaro 42 



Volador42, 132 
Volatin 132 



Williamodendron 1 1 8 
glaucophyllum 32, 119 



Yas 107 

Ye ma de huevo 58 

Yema huevo 76 



Zanthoxylum-like projections on 
trunk 100 



138 



FIELDIANA: BOTANY 



Families of seed plants known or expected to occur in Costa Rica and adjacent areas numbered 
according to the sequence of Engler's Syllabus a. . dition 1 1, reworked by L. Diels 



2 Taxaceae 

3 Podocarp 

4 Araucariaceae 

5 Pinaceae 

ipressaceae 

7 Gnetaceae 

8 Typhaceae 

9 Potamogetonaceae 

1 Najadaceae 

1 1 Alismataceae 

12 Butomaceae 

1 lydrocharitaceae 

14 Triuridaccae 
iramineae 

1 6 Cyperaceae 

17 Palmae 

. .lanthaceae 

1 9 Araceae 

20 Lemnaceae 

21 Mayacaceae 

22 Xyridaceae 

23 Eriocaulaceae 

24 Bromeliaceae 

25 Commelinaceae 

26 Pontederiaceae 

27 Juncaceae 

28 Liliaceae 

29 Haemodoraceae 
\maryllidaceae 

3 1 Velloziaceae 

32 Dioscoreaceae 

33 Iridaceae 

34 Musaceae 

35 Zingiberaceae 

36 Cannaceae 

37 Marantaceae 

38 Burmanniaceae 

39 Orchidaceae 

40 Casuarinaceae 

41 Piperaceae 

42 Chloranthaceae 

43 Lacistemaceae 

44 Salicaceae 

45 Garryaceae 

46 Myricaceae 

47 Juglandaceae 

48 Batidaceae 

49 Betulaceae 

50 Fagaceae 

51 Ulmat 

52 Moraceae 

53 Urticaceae 

54 Podostemonaceae 

55 Proteaceae 

56 Olacaceae 

57 Opiliaceae 

58 Loranthaceae 

59 Aristolochiaceae 
60a Hydnoraceae 
60b Rafflesiaceae 

6 ! Balanophoraceae 

62 Polygonaceae 

63 Chenopodiaceae 

64 Amaranthaceae 

65 Nyctaginaceae 

66 Phytolaccaceae 

67 Aizoaceae 

68 Portulacaceae 

69 Basellaceae 

70 Caryophyllaceae 



73 Ranu 

74 Berberidaceae 

76 Magnoh 

79 M 

80 ! 

82 Pi 

incl. Fumariaceae 

84 

85 Tovariaceae 

86 Resedaceae 

87 Moring; 

88 Droseraceae 

89 Crassulaceae 

90 Saxifragaceae 

91 Brunei!: 

92 Cunoniaceae 

93 Hamamelidaceae 

94 Rosai. ' 

onnaraceae 

96 Leguminosae 

97 Krameriaceae 

98 Oxalidaceae 

99 Gerani; 

100 Tropae'i 

101 Linat 

incl. Humiriaceae 
' : rythroxy!;i 
Zygophylku 
Rutaceae 

105 Simarubaceae 

1 06 Burseraceae 

107 Meliaceae 

108 Malpighiaceae 

109 Trigoniaceae 

1 10 Vochysiaceae 

1 1 1 Polygalaceae 

1 1 2 Dichapetalaceae 

1 1 3 Euphorbiaceae 

1 1 4 Callitrichaceae 

1 15 Buxaceae 

116 Coriariaceae 

1 1 7 Anacardiaceae 

118 < 

120 Celastr 

1 2 1 Hippocrateaceae 

123 Icacin;. 

124 Hippocastan > 

125 Sapind 

1 26 Sabiaceae 
Balsaminaceae 

128 Rhamn 

i.arpaceae 
Tiliaceae 

132 

133 Bomba*. 

134 S; 

135 Dillcni 

1 36 Actinidiaceae 

137 Ochna: 

1 38 Caryocaraceae 
Marcgraviaceae 



140 (.. 

142 ( 

inc: 

143 Elatinaceae 
144 

145 Bixac, 
146 

147 . 

1 48 Flacouniaceae 

1 49 Turneraceae 

.ssifloraceae 
151 ' 

153 Begoni 

cae 

? ythraceae 

158 Punica 

159 Lecythidaceae 

160 Rhizophon: 
161 

162 Myrtaceae 

163 Mclastomai 

1 64 ( 

165 Halorrhagat 

166 Araliaceae 

168 Corn a 

169 Clethraceae 

170 Monotropa* 
1 7 1 Pyrola 

172 Ericai 

173 Theo" 

1 74 Myrsinaceae 

175 Primulaceac 
Plumbaginaceae 
iapotaceae 

178 Ebenaceae 

179 Symplocaceae 

1 80 Styracaceae 

1 8 1 Oleaceae 

182 Loganiaceae 

183 Gentianaceae 

184 Apocyrv 

185 Asclepiadaceae 

186 Convolvulaceae 
Polemoniaceae 

188 Hydrophyllaceae 

189 Boraginaceae 

1 90 Verbenaceae 

1 9 1 Labiatae 
Malanaceae 
Scrophulariaceae 

ignoniaceas 
'. 'cdaliaceae 
196 N 1 

200 Acanth 

20 1 Plantaginat 

202 Rubia 

204 V.