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ILLINOIS LIBRARY

,M URBANA-CHAMPAIGN

BIOLOGY

o vu,^

rt.xv,
no. 33

FLORA COSTARICENSIS

t\ iiliam Burger, Kditor

Family #202 Rubiaceae

vVilliam Burger
Charlotte M. Tavlor

Kurcniber 30. 1993
'ublication 1454

>U BLUSHED BY !

TURA

?) This paper meets the requirements of ANS1/NISO Z39. 48-1992 (Permanence of Paper).

FTELDIANA

Botany

NEW SERIES, NO. 33

FLORA COSTARICENSIS

William Burger, Editor

Family #202 Rubiaceae

William Burger

Curator

Department of Botany

Field Museum of Natural History

Chicago, Illinois 60605-2496

Charlotte M. Taylor

Missouri Botanical Garden

St. Louis, Missouri 63166-0299

Accepted April 16, 1993

Published December 30, 1993 BIOLOGY LIBRARY

Publication 1454 101 "

PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY

1993 Field Museum of Natural History
Library of Congress Catalog Card Number: 93-73814

ISSN 00 15-0746
PRINTED IN THE UNITED STATES OF AMERICA

Table of Contents

INTRODUCTION v

ACKNOWLEDGMENTS v

RUBIACEAE 1

KEYS TO THE RUBIACEAE OF COSTA RICA 2

Key 1 : Technical Keys to the Traditional

Tribes and Genera of Rubiaceae 2

Key 2: Artificial Key to Genera and

Illustrations 8

Illustrations of Rubiaceae 15

Description of Genera and Species (Aliber-
tid) 82

Psychotria 220

LITERATURE CITED 324

LIST OF ACCEPTED SPECIES 325

INDEX . 328

List of Illustrations

1 . Twining shrubs (Manettia spp.) and
subshrubs with small stiff leaves
(species of Arcytophyllum, Declieux-

ia, and Diodid) 15

2. Twining herbs: species of Coccocyp-
selum and Geophila 16

3. Herbs with small leaves and slender
stems: species of Didymaea, Galium,
Nertera, and Oldenlandia 17

4. Erect herbs with narrow lanceolate
leaves and capitate or verticillate
flowers: species of Crusea, Mitracar-

pus, Richardia, and Spermacoce 18

5. Erect herbs with narrow lanceolate
leaves: Spermacoce spp 19

6. Erect herbs with narrow lanceolate
leaves: Diodia spp. and two species

of Spermacoce 20

7. Herbs or subshrubs with larger
leaves: species of Amphidasya, Hoff-
mannia, Lasianthus, and Psychotria ... 2 1

8. Herbs or subshrubs with axillary
flowers: unusual species of Hoffman-

nia 22

9. Herbs or subshrubs with axillary
flowers: pubescent species of Hoff-
mannia and H. congesta 23

10. Subshrubs with axillary flowers: spe-
cies of Hoffmannia with larger leaves . . 24

1 1 . Subshrubs with axillary flowers: spe-
cies of Hoffmannia with leaves ta-
pering gradually to the base 25

12. Subshrubs with axillary flowers: spe-
cies of Psychotria 26

13. Subshrubs with axillary flowers: spe-
cies of Psychotria 27

14. Trees with very large or lobed leaves:
three species of Pentagonia 28

15. Flowers with very long corolla tubes:
species of Lindenia, Osa, and Poso-
queria 29

16. Inflorescences with greatly expanded
petal-like calyx lobes: species of Ca-
lycophyllum, Mussaenda, Pogonopus,

and Warszewiczia 30

17. Inflorescences of involucrate heads:
species of Psychotria (formerly Ce-
phaelis spp.) 31

18. Inflorescences of involucrate or con-
spicuously bracteate heads: species of
Psychotria 32

19. Inflorescences of compact heads with
flowers connivent at the base: species
ofAppunia, Morinda, and Schroder a . . 33

20. Inflorescences long and narrow: spe-
cies of Gonzalagunia 34

2 1 . Inflorescences long and narrow: spe-
cies of Gonzalagunia and Rondeletia . . 35

22. Fruits usually terminal and solitary:
species of Randia with small leaves ... 36

23. Fruits usually terminal and solitary:
species of Randia with medium-sized
leaves 37

24. Fruits usually terminal and solitary:
species of Randia with larger leaves ... 38

25. Fruits usually terminal and solitary:
species ofAlibertia, Duroia, Genipa,

and Hippotis 39

26. Fruits usually terminal and solitary:
species of Borojoa and Genipa 40

27. Plants usually epiphytic: species of
Cosmibuena and Hillia with smaller
leaves 41

28. Plants usually epiphytic: species of
Cosmibuena and Hillia with larger
leaves 42

29. Trees with large open inflorescences:
species of Ladenbergia and Conda-
minea corymbosa 43

30. Inflorescences with clusters of long-
tubular flowers: species ofAmaioua,
Guettarda, Isertia, and Tocoyena 44

3 1 . Showy flowers: species of Coutarea,
Crusea, Exostema, Ixora, and Pen-
tas

32. Inflorescences with scorpioid or heli-
coid branches: species of Guettarda ... 46

33. Flowers with narrow corolla tubes:
species of Guettarda and a species of
Chomelia 47

34. Flowers with narrow corolla tubes:
species of Chomelia, Guettarda, and
Hamelia 48

35. Inflorescences mostly axillary: spe-
cies of Sabicea (vines) and Sommera
(trees) 49

36. Inflorescences axillary or terminal:
species of Chiococca and a species of
Chione 50

37. Many small flowers in dense inflores-
cences: species of Chimarrhis, Cin-
chona, Machaonia, and Uncaria 51

38. Flowers in much-branched open in-
florescences: species of Deppea, Rus-

tia, and Simira 52

39. Small flowers in dense or open pani-
cles: species of Elaeagia 53

40. Small flowers and capsular fruits:
species of A Iseis, Exostema, Ferdi-
nandusa, and Macrocnemum 54

4 1 . Rondeletia spp 55

41 A. Rondeletia spp 56

42. Hamelia spp 57

43. Bertiera, Ixora, and Raritebe spp 58

44. Faramea: species with larger leaves ... 59

45. Faramea: species with smaller leaves. . 60

46. Coussarea and Rudgea spp 61

47. Coussarea spp. and two similar Psy-
chotria spp 62

48. Coussarea: species with larger leaves . . 63

49. Palicourea: species with conspicuous
bracts . . .64

50. Palicourea: species of lower eleva-
tions and a species of Isertia 65

45 51. Palicourea: species with larger leaves

and yellow or orange flowers 66

52. Palicourea: species with larger leaves
and blue, lavender, purple, or white
flowers 67

53. Palicourea: species with smaller

leaves 68

54. Psychotria subg. Heteropsychotria:
larger-leaved pubescent species and a
species of Palicourea 69

55. Psychotria subg. Heteropsychotria:
species with smaller leaves 70

56. Psychotria subg. Heteropsychotria:
species with very small inflorescences ..71

57. Psychotria subg. Heteropsychotria:
species with larger open inflorescenc-
es 72

58. Psychotria subg. Heteropsychotria:
species with conspicuous open
inflorescences 73

59. Psychotria subg. Heteropsychotria:
species of deciduous habitats and

some with smaller inflorescences 74

60. Psychotria subg. Psychotria: species
with very small leaves and a com-
plex of epiphytic species 75

6 1 . Psychotria subg. Psychotria: species

with smaller narrow leaves 76

62. Psychotria subg. Psychotria: high-ele-
vation species and those with Ficus-

like stipules 77

63. Psychotria subg. Psychotria: densely
pubescent species 78

64. Psychotria subg. Psychotria: large-
leaved species 79

65. Psychotria subg. Psychotria: decidu-
ous and unusual species 80

66. Psychotria subg. Psychotria: several
unusual species 81

Introduction

This is the eighth issue in the Flora Costaricensis
series. The first dealt with the Piperaceae (Field-
iana, Bot. 35, 1971). The second included families
numbered 42 through 53, Chloranthaceae through
Urticaceae (Fieldiana, Bot. 40, 1977). The third
issue covered the Gramineae and was authored by
Richard Phol (Fieldiana, Bot., new series, No. 4,
1980). The fourth issue included families num-
bered 54 through 70, Podostemaceae through Car-
yophyllaceae (Fieldiana, Bot., new series, No. 13,
1983). The fifth issue covered families 200 and
201, the Acanthaceae authored by L. H. Durkee,
and the Plantaginaceae (Fieldiana, Bot., new se-
ries, No. 18, 1986). The sixth issue included fam-

ilies 80 and 81, Lauraceae and Hernandiaceae
(Fieldiana, Bot., new series, No. 23, 1990). The
seventh issue included families numbered 97
through 103, Krameriaceae through Zygophylla-
ceae (Fieldiana, Bot., new series, No. 28, 1991).

In the figures, leaves and leafy stems are drawn
to the same scale throughout. Enlarged flowers and
fruits are drawn to the same scale on an individual
plate unless otherwise noted. The closed scales
represent centimeters and the open scales repre-
sent millimeters. The figures are somewhat dia-
grammatic and represent the senior author's con-
cept of a common or characteristic morphology.

Acknowledgments

We wish to thank the staff, and especially Pablo
Sanchez, of the Museo Nacional de Costa Rica for
their assistance over many years. A grant from the
Museo Nacional allowed the senior author to work
at the Herbario Nacional for several weeks in No-
vember 1990. Charlotte Taylor received support
for travel from the National Science Foundation
(BSR 83-10702 and BSR 87-00068), the Fondos
Institucionales Para Investigacion of the Univer-
sity of Puerto Rico, and the Dee Scholarship Fund
of Field Museum. Collecting programs by the Mis-
souri Botanical Garden (MO), Institute Nacional
de Biodiversidad and the Museo Nacional (CR),
supported in part by grants from the National Sci-
ence Foundation and the National Geographic So-
ciety, have added significantly to our knowledge
of Costa Rica's Rubiaceae. The recent collections
of Jorge Gomez-Laurito, Michael Grayum, Wil-
liam Haber, Barry Hammel, Gerardo Herrera,
Quirico Jimenez, and Nelson Zamora have been

especially significant. Loans from the U.S. Na-
tional Herbarium (us) and the Duke University
Herbarium (DUKE) were important for our work
on this family.

A number of our colleagues have been especially
helpful in preparing this treatment. The annota-
tions and advice of C. Dennis Adams, John Dwyer,
Barry Hammel, and David Lorence were especial-
ly important. In addition, John Dwyer and David
Lorence have provided descriptions of new spe-
cies, and Roy Gereau corrected all the Latin de-
scriptions. The Flora Mesoamericana project un-
der the leadership of Geritt Davidse (MO) has
provided information and assistance on many oc-
casions. We are also indebted to the Missouri Bo-
tanical Garden for allowing Charlotte Taylor to
contribute her time and effort to this treatment.
Finally, we thank three anonymous reviewers who
made many corrections and suggested useful im-
provements for the text.

FLORA COSTARICENSIS

Family #202 Rubiaceae

RUBIACEAE

By William Burger and Charlotte M. Taylor

Herbs, shrubs, or small- to medium-sized trees (rarely
vines or tall canopy trees), stems glabrous to pubescent
with simple hairs, terete or angular; stipules of opposing
leaves usually united across the stem (interpetiolar),
sometimes united to the petioles and forming a broad
sheath (Spermacoceae), rarely separate and paired at the
leaf base, stipules of the same leaf sometimes also united
above the petioles (intrapetiolar) and forming a short
tube, often with hair-like or tooth-like colleters at the
adaxial base or along the edge, persistent to caducous
and leaving a scar across the stem (stipules transformed
into small leaves in Galium and Sherardia). Leaves op-
posite or sometimes whorled (very rarely alternate), al-
ways simple, petiolate or occasionally sessile, glabrous
or pubescent, nearly always entire and without lobes
(pinnatind in Pentagonia spp., with small lobes in Simira
spp., minutely serrate or crenate in some Spermacoceae),
nearly always pinnately veined (palmately veined in some
weedy Spermacoceae), domatia of pits or tufted hairs
present in the vein axils on the lower surfaces of leaves
in some genera. Inflorescences terminal or axillary, sol-
itary to several at each node, very variable in form (open
paniculate to cymose, racemose, spicate or capitate),
branches of the inflorescence often opposite, bracts and
bracteoles often present, the flowers often borne in distal
cymes or dichasia, sometimes cincinnoid and 1 -sided
(rarely solitary, fasciculate, or united), sessile or pedi-
cellate. Flowers usually bisexual and radially symmet-
rical (rarely unisexual and dioecious), most often 4- or
5-parted, epigynous, the hypanthium narrowly tubular
to subglobosc. calyx tube usually present, calyx lobes
usually present and equal or subequal (rarely with 1 lobe
greatly expanded and colorful); corolla often salverform
with a narrow tube (funnelform to rotate or tubular),
corolla lobes valvate, imbricate or contorted in bud; sta-
mens usually as many as the corolla lobes and alternating
with them, nearly always borne on the corolla tube, fil-
aments long to short, anthers usually narrowly oblong
and basi fixed to dorsifixed, dehiscing longitudinally (with
terminal pores in Rustia); ovary inferior (rarely half-
inferior), usually with a ring-like disc on the upper sur-
face, with 2 (1-8) locules, placentation apical, basal or
from the median septum (parietal in some Gardineae),
style solitary from the center of the apex of the ovary,
stigmas usually 2 or solitary (clavate to capitate). Fruits
capsular, baccate or drupaceous and often with 2 (4-5)
pyrenes (a syncarp in Morinda and Schradera, a samara
in Allenanthus), berries sometimes large with the seeds
imbedded in a fleshy pulp, capsules opening along the
locules (loculicidal) or along the septum (septicidal); seeds
sometimes with wings or tufted hairs.

The Rubiaceae are one of the largest families of
flowering plants, with an estimated 10,700 species
(Mabberley, 1987). The family is best represented
in the evergreen tropics and is often an important
component of the lower strata of such forests.

In most cases the family is easily recognized.
The simple opposite leaves are nearly always en-
tire and pinnately veined. Only a few weedy spe-
cies have subpalmate venation, and a few of our
woody species have lobed leaves (Pentagonia spp.
and Simira maxonii). The trichomes are never
branched or stellate. Some species have domatia
in the form of pits or tufted hairs in the vein axils
on the undersides of leaves. Though variable in
presence, domatia can be helpful in identifying
species. Too small to be useful to ants, these leaf
domatia probably offer shelter for predatory and
fungivorous mites (Pemberton & Turner, 1989).
The nodes are nearly always marked by interpetio-
lar stipules or interpetiolar lines if the stipules have
fallen. The stipules can be important in identifying
species but may be apparent only on young shoots;
they may be greatly enlarged when subtending in-
florescences. Persisting stipules may be distorted
or torn apart as the stem expands. Hair-like or
tooth-like structures between the base of the stip-
ule and the stem are called colleters. These are
usually Anger-like with elongate axial cells and a
palisade epidermis (Lersten, 1974). They are be-
lieved to secrete mucilage, gums, or resins.

Inflorescences vary greatly in some genera. In
some species the bracts subtending the first pair
of opposite branches of the inflorescence may be
replaced by smaller leaves. In this case an inflo-
rescence that is solitary and terminal can be in-
terpreted as being a group of three inflorescences:
a terminal one and two axillary to the distal leaves
(bract homologs). The flowers are often borne in
distal cymes on opposite branches of the inflores-
cences. Many species are distylous with long-styled
(pin) or short-styled (thrum) flowers on different
plants. The corolla is nearly always radially sym-
metrical and with a conspicuous tube. Curvature
of the tube or assymetry of the corolla lobes is
rare. The inferior ovary is usually two-locular, and
the number of ovules per locule has been used as

FIELDIANA: BOTANY, N.S., NO. 33, DECEMBER 30, 1993, PP. 1-333

a primary criterion for distinguishing the subfam-
ilies.

While a very distinctive family, there are a few
genera of other families that can be mistaken for
Rubiaceae. Collections ofCassipourea (Rhizopho-
raceae), Hedyosmum (Chloranthaceae), and Neea
(Nyctaginaceae) are often found among specimens
of Rubiaceae. There are also look-alikes in Acan-
thaceae, Loganiaceae, and Onagraceae.

The Rubiaceae of Central America are relatively
well understood, and their taxonomy is in good
order. This is the result of intensive study by many
workers, past and present. Among these, the work
of Paul Carpenter Standley provided a solid foun-
dation. His publications and many annotations
have been particularly useful in preparing the pres-
ent account. The treatments of the family for the
Flora of Guatemala (Standley & Williams, 1975),
Flora of Panama (Dwyer, 1 980), and Flora of Ven-
ezuela (Steyermark, 1 974) have also been very use-
ful. The recent studies by Dennis Adams, John
Dwyer, Clement Hamilton, Joseph Kirkbride, Da-
vid Lorence, and others have clarified many dif-
ficult species groups and are cited in the text. Many
collectors have contributed substantially to our
knowledge of this family in Costa Rica (an index
to exsiccatae is available on request).

tation, ovule orientation, and characteristics of
seeds and fruits. This key follows those presented
in the Flora of Guatemala (Standley & Williams,
1975) and the Flora of Panama (Dwyer, 1980).
While often very difficult to implement, this key
has wide application and places the genera into
the traditional tribes. More modern keys can be
found in Robbrecht (1988).

As Verdcourt (1976, p. 5) has stated, "[T]he
family Rubiaceae contains so many genera and
species, many of which resemble each other even
when not closely related, that it is impossible to
make a useable key which does not involve looking
at small and difficult characters." To provide an
alternative, we give an additional artificial key that
is much simpler and attempts to make the illus-
trations more readily accessible. Scanning the il-
lustrations with the help of the second key will,
hopefully, allow determination of many species
without having to ascertain all the morphological
details required by the technical key. Commentary
under the genera and species gives characteristics
that can be helpful in distinguishing the taxa; the
detailed descriptions are useful in confirming a
determination. Nevertheless, there is no substitute
for careful comparisons with annotated herbarium
collections to verify a determination.

Keys to the Rubiaceae of Costa Rica

We provide a technical key to the genera that
requires ascertaining corolla aestivation, placen-

Key 1: Technical Keys to the Traditional Tribes and Genera of Rubiaceae
(see Robbrecht, 1988, for a more modern system)

1 a. Ovules more than 1 in each locule of the ovary (subfamily Cinchonoideae) 2

1 b. Ovules solitary in the locules of the ovary (subfamily Rubioideae, except Naucleeae) 9

2a. Fruits fleshy and indehiscent (baccate or berry-like) 3

2b. Fruits dry and dehiscent (capsule-like) 5

3a. Corolla lobes valvate in bud 1 . Isertieae

3b. Corolla lobes imbricate or contorted in bud 4

4a. Corolla lobes imbricate in bud 2. Hamelieae

4b. Corolla lobes contorted in bud 3. Gardenieae

5a. Flowers in compact spherical heads 8. Naucleeae

5b. Flowers not in compact spherical heads 6

6a. Seeds with wings, tufted hairs or appendages, arranged vertically imbricate on the placenta

4. Cinchoneae

6b. Seeds angled but not winged, or if winged then arranged horizontally on the placenta ... 7

7a. Corolla lobes imbricate or contorted in bud 6. Rondeletieae

7b. Corolla lobes valvate in bud 8

8a. Seeds horizontal, usually many (> 25); stipules entire or bifid; trees and shrubs

FIELDIANA: BOTANY

5. Condamineeae

8b. Seeds vertical and imbricate, usually few; stipules usually setose; herbs and subshrubs ....

7. Oldenlandieae

9a. (from Ib) Seeds pendulous, the radicle superior; trees, shrubs, or woody lianas 10

9b. Seeds ascending, the radicle inferior; trees, shrubs, or herbs 12

lOa. Flowers in spherical/globose heads 8. Naucleeae

I Ob. Flowers never in globose heads 11

II a. Stamens usually borne at apex of corolla tube (the rarely encountered Vangueria of the

Vanguerieae will key out here; see text) 9. Guettardeae

1 Ib. Stamens borne at base of the corolla tube or from the disc 10. Chiococceae

12a. Corolla lobes contorted in bud; trees and shrubs 11. Ixoreae

1 2b. Corolla lobes valvate in bud; trees, shrubs, and herbs 13

1 3a. Ovules borne on base of the locule; mostly woody plants 14

1 3b. Ovules borne from the septum in the center of the ovary; herbs, shrubs, or trees 17

1 4a. Ovary with 7-8 locules; inflorescences globose; fruits multiple, of 4-50 united flowers ....

14. Morindeae

14b. Ovary with 1-5 locules; inflorescences various; fruits simple or with 2 united flowers if

multiple 15

15a. Ovary 1- or 2-locular and with a thin partial septum; fruits with 1 seed . . 12. Coussareeae
15b. Ovary 2-locular (5-locular) and with thick well-developed septum; fruits with 2(-5) seeds

1 6a. Stamens usually inserted near the apex of the corolla tube; flowers bisexual

13. Psychotrieae

1 6b. Stamens usually inserted near the base of the corolla tube; flowers often unisexual

15. Anthospermeae

1 7a. (from 1 3b) Stipules not leaf-like nor setose; trees and large shrubs; flowers united near the base;

fruits united or partly united into a syncarp 14. Morindeae

1 7b. Stipules either setose with awn-like appendages or leaf-like (and the small leaves apparently whorled
and lacking stipules); herbs or small shrubs; flowers often congested but not united at the base;

fruits never united into a syncarp 18

18a. Stipules usually bearing 3-30 narrow setae or awns; leaves usually opposite . 16. Spermacoceae
1 8b. Stipules leaf-like; leaves and leaf-like stipules appearing as whorls of 4 or more leaves per node

. 17. Rubieae

1. ISERTIEAE (MUSSAENDEAE)

la. Leaves apparently alternate (a minute opposing leaflet often present, not known from Costa Rica)

Didymochlamys

1 b. Leaves opposite, both leaves of the node developed 2

2a. Leaves with the minor venation parallel (lineolate) 3

2b. Leaves with the minor venation not parallel 5

3a. Leaves large, to over 1 m long, with pinnatifid lobes in some species; rows of glands (colleters)

present on the interior of the calyx cup; plants often monopodial Pentagonia

3b. Leaves up to 35 cm long, never with pinnatifid lobes; calyx cup without glands on the interior;

plants often much-branched 4

4a. Calyx 5-lobed Sommera

4b. Calyx 2-lobed or spathe-like Hippotis

5a. Inflorescences axillary 6

5b. Inflorescences terminal 8

6a. Erect unbranched plants to 50 cm tall, with long closely clustered leaves; ovary 2-locular;

corolla more than 30 mm long Amphidasya

6b. Plants with leaves well spaced along the twining or creeping stems; ovary 2-5-locular; corolla
less than 1 2 mm long 7

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

7a. Plants prostrate, herbaceous; ovary 2-locular; fruit bright blue Coccocypselum

7b. Plants erect or climbing, herbs or subshrubs; ovary 3-5-locular; fruits reddish becoming

purple or bluish black Sabicea

8a. Flowers and inflorescences drying black; inflorescence capitate; leaves and flowers semisucculent

Schradera

8b. Flowers and inflorescences not drying black; inflorescences subcapitate only in Amphidasya; leaves

and flowers not semisucculent 9

9a. Stipules fimbriate distally; herbaceous with erect unbranched stems to 0.8 m tall . . Amphidasya

9b. Stipules not fimbriate distally; woody plants with branched stems to 3 m tall 10

1 Oa. Inflorescences spike-like; shrubs Gonzalagunia

I Ob. Inflorescences cymose to paniculate; shrubs or trees 11

I 1 a. Anthers not transversely locellate (not divided by transverse walls); corollas less than 2 cm long;

ovary 2-locular Raritebe

1 Ib. Anthers transversely locellate; corollas 3 or more cm long; ovary (2-)5-6-locular Isertia

2. HAMELIEAE

la. Inflorescences always axillary; ovary with 2 or 3 locules; stamens with connective, rarely prolonged
distally [corolla lobes imbricate or subvalvate] Hoffmannia

Ib. Inflorescences usually terminal; ovary with 4 or 5 locules; stamens with the connective often pro-
longed distally 2

2a. Corolla lobes imbricate in bud; raphides present in leaves; inflorescences often with few cincinnoid
branches Hamelia

2b. Corolla lobes contorted in bud; raphides absent in the leaves; inflorescences with many branches,
not cincinnoid . . Bertiera

3. GARDENIEAE

la. Ovary with a single locule and intruding parietal placentas (but often difficult to see and the abutting

placentas appearing as a septum); pollen grains in tetrads 2

Ib. Ovary with usually 2 locules (the septum often thin or obliterated as the seeds develop); pollen

separate 3

2a. Plants native and wild, usually armed with spines on branches, frequently with short-shoots

Randia

2b. Plants grown in gardens for ornament, usually lacking short-shoots Gardenia

3a. Flowers bisexual 4

3b. Flowers unisexual 6

4a. Inflorescences with 1-few flowers, flowers not in a candelabra-like arrangement; leaves drying

black Genipa

4b. Inflorescences with several to many flowers; flowers usually in a candelabra-like arrangement;

leaves drying black or not 5

5a. Flower buds curved at the apex, corolla white and becoming yellowish in age; seeds with testa

cells more than twice as long as wide Posoqueria

5b. Flower buds straight at the apex, corolla bright yellow at anthesis; seeds with testa cells less

than twice as long as wide Tocoyena

6a. Fruits rounded or globose; terminal stipules not forming a conical cap, triangular and persisting

6b. Fruits oblong; terminal stipules forming a conical cap, caducous 8

7a. Fruits subtended by several whorls of persisting bracts (stipules), fruits more than 5 cm diam.,

pericarp thick, carnose, surfaces rough Borojoa

7b. Fruits not subtended by several persisting bracts; fruits to 3 cm diam., pericarp thin and hard,
smooth . . . Alibertia

FIELDIANA: BOTANY

8a. Female flowers usually solitary; fruits hirsute Duroia

8b. Female flowers capitate or cymose; fruits not densely hirsute .... . Amaioua

4. CINCHONEAE

la. Vining or clambering with slender herbaceous stems (genus placed in the Hedyotideae in modern

systems) Manettia

Ib. Shrubs or trees, stems not slender and clambering 2

2a. Anthers dimorphic or trimorphic Ferdinandusa

2b. Anthers monomorphic (all alike within the flower) -3

3a. Calyx with 1 expanded (2-5 cm) white lobe in 1 or 2 flowers of the inflorescence

Calycophyllum

3b. Calyx lobes equal or subequal, inflorescences without expanded large white calyx lobes 4

4a. Plants epiphytic; leaves semisucculent, lateral veins often obscure when dried 5

4b. Plants terrestrial; leaves not semisucculent, lateral veins clearly evident 6

5a. Seeds winged at either end Cosmibuena

5b. Seeds with tufted hairs at one end Hillia

6a. Inflorescences spike-like; stamens attached at the base of the corolla tube [filaments hirsutulous]

Alseis

6b. Inflorescences not spike-like, various; stamens attached above the base of the corolla tube (except

in Coutarea and Exostema) 7

7a. Corolla lobes imbricate or contorted in bud 8

7b. Corolla lobes valvate in bud 9

8a. Stamens conspicuously exserted; corolla not inflated in bud; fruits not compressed or lenti-
cellate Exostema

8b. Stamens not conspicuously exserted (may become exserted as corolla ages); corolla buds
inflated in late stages (before anthesis); fruits strongly flattened, surfaces often lenticellate

Coutarea

9a. Flowers 4-parted; capsules rounded Bouvardia

9b. Flowers 5-parted; capsules oblong 10

lOa. Capsule splitting from below to the apex [a ring of hairs present within the mouth of the corolla]

Cinchona

lOb. Capsule splitting from above to the base 11

I la. Corolla lobes split at the apex; dehiscing capsules forming 4 coiled valves (not known from Costa

Rica) Joosia

I 1 b. Corolla lobes not split at the apex; capsules valves not becoming coiled 12

1 2a. Capsules opening into the locules Macrocnemum

1 2b. Capsules opening along the septum Ladenbergia

5. CONDAMINEEAE

la. Anthers opening by terminal pores; leaves with pellucid glandular dots Rustia

Ib. Anthers opening by longitudinal slits; leaves without pellucid glandular dots 2

2a. Inflorescences axillary 3

2b. Inflorescences terminal 4

3a. Flowers solitary or few, ca. 25 cm long Osa

3b. Flowers many, ca. 3 mm long Chimarrhis

4a. Calyx lobes equal or subequal, small; stipules large and bifid [leaves large and subsessile]

Condaminea

4b. Calyx with 1 lobe enlarged (2-6 cm) and colored in 1 or a few flowers of each inflorescence; stipules

small, not bifid Pogonopus

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

6. RONDELETIEAE

la. Corolla lobes contorted in bud 2

1 b. Corolla lobes imbricate in bud 5

2a. Corolla tube becoming very long (+ 10 cm), much longer than the corolla lobes; capsule valves

becoming coiled; shrubs of stream sides Lindenia

2b. Corolla never exceeding 5 cm, tube shorter than the corolla lobes; capsule valves not becoming

coiled; if woody not restricted to river edges and wet sites 3

3a. Small herbs of wet sites Sipanaea

3b. Trees and shrubs 4

4a. Corolla 4-parted, yellowish, glabrous within Deppea

4b. Corolla 5-parted, greenish white, villous within Elaeagia

5a. Calyx lobes unequal, often expanded into a broad lobe 6

5b. Calyx lobes equal or subequal, small and unexpanded 7

6a. Expanded calyx lobe bright red Warszewiczia

6b. Expanded calyx lobes whitish (in some spp.) Rondeletia

7a. Capsules ca. 1 cm diam., opening into the septum; seeds winged or flat and enlarged; wood turning
reddish when cut and exposed (in ours) Simira

7b. Fruits to 5 mm diam., opening at the septum or locule; seeds lacking wings, not flattened; wood
not turning reddish when cut and exposed 8

8a. Corolla tubes usually more than 8 mm long, stamens included; capsule valves usually not woody
and bifid at apex; common plants in Central America Rondeletia

8b. Corolla tubes to 5 mm long, stamens usually exserted; capsule valves woody, bifid at the apex;
rarely collected in southern Central America Bathysa

7. OLDENLANDIEAE

la. Plants subshrubs, leaves usually small and stiff (ericoid); seeds plano-convex or carinate; plants of

high montane formations 2800-3400 m elevation Arcytophyllum

Ib. Plants herbs or subshrubs, leaves not stiff and ericoid; seeds angular or winged; 0-2000 m elevation

2a. Garden ornamentals; flowers red, pink, or white Pentas

2b. Weedy plants of wet or moist sites; flowers white 3

3a. Flowers 4-parted, common introduced weeds Oldenlandia

3b. Flowers 5-parted, rare introduced weeds , Pentodon

8. NAUCLEEAE

1 a. Woody vines with recurved spines; inflorescences axillary, pedunculate, each with 2-5 globose heads

(some modern treatments place this genus in Cinchonieae) Uncaria

Ib. Trees planted for wood, without spines; inflorescences terminal, each with 1 globose head

. Neolamarckia

9. GUETTARDEAE

la. Woody lianas with vining branches [leaves with subparallel 3 veins; fruits fleshy; rare in Costa
Rica] Malanea

Ib. Trees or shrubs, not lianas with vining branches (except in some species of Chomelid) 2

2a. Corolla lobes valvate in bud; some species with subparallel (lineolate) minor venation [spines/thorns
sometimes present on stems and twigs] Chomelia

2b. Corolla lobes imbricate in bud; minor venation of the leaves not subparallel nor lineolate 3

FIELDIANA: BOTANY

3a. Fruits dry, separating into 2 indehiscent mericarps (cocci); flowers not secund on inflorescence

branches; spines sometimes present on stems and branches Machaonia

3b. Fruits fleshy; flowers often in secund arrangements; spines absent on stems and branches 4

4a. Fruits covered with a fine dense pubescence Guettarda

4b. Fruits glabrate (formerly Antirhea spp.) Chomelia

10. CHIOCOCCEAE

la. Flowers 4-parted; fruits dry, flat and broadly winged Allenanthus

Ib. Flowers 5-parted; fruits fleshy, not winged 2

2a. Fruits compressed laterally (oblong in cross-section); corolla lobes valvate in bud .... Chiococca

2b. Fruits rounded (terete in cross-section); corolla lobes imbricate in bud Chione

11. IXOREAE

la. Inflorescences axillary; floral bracts connate and calyx-like or involucrate beneath the flowers;

cultivated Coffea

1 b. Inflorescences terminal; floral bracts separate, not calyx-like; wild or cultivated for ornament ....

. Ixora

12. COUSSAREEAE

la. Ovules separate in a 1-locular ovary; seeds horizontal and fruits often broader than long, exocarp
usually leathery; flowers blue or white Faramea

Ib. Ovules connate from a basal column, ovary 1- or 2-locular; seeds longitudinal, fruits longer than
broad, exocarp spongy or carnose; flowers white Coussarea

13. PSYCHOTRIEAE

1 a. Plants with creeping prostrate stems and long internodes; leaves often cordate Geophila

Ib. Plants erect, herbaceous subshrubs to small trees; leaves various (rarely subcordate) 2

2a. Herbaceous subshrubs; fruits laterally compressed Declieuxia

2b. Shrubs, trees, or rarely subshrubs; fruits terete 3

3a. Stipules with a group of small digitate teeth at the apex; seeds with an incurved adaxial (ventral)

surface Rudgea

3b. Stipules without digitate teeth at the apex, simple to bifid; seeds with a flat but sulcate adaxial

surface 4

4a. Corollas usually yellow, orange, reddish, purple, or blue (rarely white), often swollen at the base,

corolla tube often long (+ 1 cm) and slender, with a ring of pubescence below the middle internally;

stipules usually persisting; most often found at higher elevations Palicourea

4b. Corollas usually white or green to yellowish, not gibbous near the base, corolla tubes generally short

(- 1 cm), with a ring of pubescence at or above the middle internally or glabrous within; stipules

persisting or deciduous; widespread with many species Psychotria

14. MORINDEAE

la. Base of hypanthium free, not united to adjacent flowers, not forming a syncarp in fruit; ovary and
fruits with 7-8 locules . . . Lasianthus

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

1 b. Base of hypanthium united to adjacent flowers, forming a syncarp in fruit; ovary and fruits with 1
or 2 locules 2

2a. Flowers and fruits drying black; syncarps fleshy, more than 1 cm diam., calyx not elevated on fruits
Morinda

2b. Flowers and fruits drying dark brown; syncarps dry, less than 1 cm diam., calyx tube prominent on
fruits Appunia

15. ANTHOSPERMEAE

la. Wild plants forming prostrate mats at high elevations; leaves very small and rounded; fruits orange

Nertera

Ib. Cultivated ornamental small (< 1 m) erect shrubs with small narrow acute leaves; fruits brownish

. Serissa

16. SPERMACOCEAE

1 a. Fruits with circumscissile dehiscence Mitracarpus

Ib. Fruits indehiscent or opening by slits, pores or valves 2

2a. Fruits breaking with 3-6 separate indehiscent cocci (note that cocci are borne on the exterior of the

fruiting axis and may resemble seeds; compare Crusea) Richardia

2b. Fruits usually with 2 separate or united cocci, cocci dehiscent or indehiscent 3

3a. Cocci opening distally or longitudinally (sometimes 1 of the 2 cocci not opening in a fruit)

Spermacoce

3b. Cocci not opening or opening only at the base 4

4a. Cocci borne on and separating from a central persisting axis, indehiscent; calyx usually with rounded

lobes Crusea

4b. Cocci not borne on a central axis, indehiscent or dehiscent from the base; calyx usually with acute

lobes . . Diodia

17. RUBIEAE

1 a. Leaves opposite (interpetiolar stipules connate and small) Didymaea

Ib. Leaves whorled (stipules leaf-like) 2

2a. Native herbs; flowers on separate pedicels or solitary and involucrate Galium

2b. Rare introduced procumbent herbs; flowers 4-10 and subsessile in distal involucrate heads

. . Sherardia

Key 2: Artificial Key to Genera and Illustrations

Note that small plants, leafy twigs, and large leaves are all drawn to the same scale throughout the 67
figures. Closed scales represent centimeters; open scales are millimeters. Unless otherwise indicated,
enlarged flowers or fruits are to the same scale on the same figure. The illustrations represent typical or
common morphologies; they cannot show the range of variation.

la. Herbaceous plants or slender few-branched subshrubs, usually less than 1.5 m tall 2

Ib. Trees, shrubs, vines, or lianas 23

2a. Creeping plants with slender flexible stems, often rooting at the nodes [flowers usually axillary]

2b. Erect or prostrate plants, lacking slender consistently creeping stems, stems slightly woody
if vining 8

FIELDIANA: BOTANY

3a. Leaf blades less than 1 cm broad 4

3b. Leaf blades usually more than 1 cm broad 7

4a. Leaves in whorls, often linear or narrow; flowers separate (Galium) or in terminal

heads (Sherardia, not illustrated) Fig. 3

4b. Leaves opposite, linear to ovate 5

5a. Leaves often linear; fruits dry capsules with few seeds; weeds below 1500 m

elevation (Oldenlandid) Fig. 3

5b. Leaves not linear; fruits fleshy, 1-2-seeded, rarely found below 1 500 m elevation

6a. Fruits orange to red, globose (Nertera) Fig. 3

6b. Fruits blue to black, usually bilobed (Didymaed) Fig. 3

7a. Fruits with many seeds, blue (Coccocypselum) Fig. 2

7b. Fruits with 1-2 seeds, red, blue-black, or black (Geophild) Fig. 2

8a. (from 2b) Leaves usually less than 4 cm broad and lanceolate, often subsessile 9

8b. Leaves usually more than 4 cm broad and petiolate 17

9a. Vining plants often climbing over other plants, with axillary flowers; fruits capsular

with many seeds (Manettid) Fig. 1

9b. Plants erect or if vining then close to the ground and not usually climbing over other

plants; flowers various 10

lOa. Leaves usually less than 15 mm long, stiff or coriaceous 11

lOb. Leaves more than 15 mm long, or thin and herbaceous when less than 15 mm long

1 la. Plants of high elevation often in exposed sites; leaves thick and blunt at the apex,

usually closely spaced (Arcytophyllurri) Fig. 1

lib. Plants of mid-elevations; leaves stiff and sharp at the apex (Diodia brasiliensis

and the cultivated Serissa, not illustrated) Fig. 1

1 2a. Ovules more than 2 in each locule; fruits with more than 2 seeds; rarely collected plants

usually found in wet or partly submerged sites (not illustrated) 13

1 2b. Ovules 1 in each locule; fruits with no more than 2(-3) seeds or mericarps; commonly

collected plants in many open weedy habitats, dry or wet 14

13a. Plants with sparse small hairs; corolla pink, tube 5-14 mm long Sipanea

13b. Plants glabrous, slightly succulent; corolla white, tube 1.5-4 mm long

Pentodon

14a. Stipules not clearly united to petioles, awns only 1-2 on each side of the node; stiff

erect plants from a woody base (Declienxia) Fig. 1

14b. Stipules united with the petioles forming a truncated sheath, the sheath usually with
more than 3 thin erect awns on each side; fruits dry and usually 2-seeded (genera of

Spermacoceae; see the technical keys and the figures below) 15

15a. Leaves usually less than 2 cm long Figs. 1, 5-6

1 5b. Leaves usually more than 2 cm long 16

16a. Terminal capitula of flowers consistently present, often subtended by bract-like leaves

(Crusea, Mitracarpus, Richardia, Spermacoce) Figs. 4-5

16b. Terminal capitula rarely present (Diodia, Spermacoce) Figs. 1, 5-6

17a. (from 8b) Inflorescences terminal on short woody stems (compare also Psychotria spp. in

figs. 54-66) Fig. 7

1 7b. Inflorescences axillary on semisucculent or woody stems to 2 m tall 18

1 8a. Locules with 2 or more ovules; fruits with many seeds (Hoffmannia spp.) 19

1 8b. Locules with 1 ovule; fruits usually 2-seeded 22

19a. Leaves 3/node or petioles with vesicles Fig. 8

19b. Leaves 2/node, petioles without vesicles 20

20a. Species conspicuously pubescent Figs. 7, 9

20b. Species mostly glabrescent 21

2 la. Leaves larger and decurrent Figs. 9-10

21b. Leaves various, inflorescences smaller Fig. 11

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

22a. Ovary usually with 8 locules (Lasianthus) Fig. 9

22b. Ovary with 2 locules (Psychotria spp., but note that Psychotria aubletiana with sessile in-

volucrate axillary capitulae is not illustrated) Figs. 12-13

23a. (from Ib) Plants vines or lianas 24

23b. Plants shrubs, trees, or subshrubs 26

24a. Slender-stemmed vines (Manettia and Sabicea) Figs. 1, 35

24b. Woody climbers or lianas 25

25a. Inflorescences pedunculate globose capitula; stems with sharp recurved spines; leaf blades

without parallel or lineolate 3 venation (Uncarid) Fig. 37

25b. Inflorescences never globose capitula, with many small flowers in panicles (and leaves with
parallel 3 venation in Malanea, not illustrated) or with larger (> 2 cm) flowers in few-
flowered inflorescences in species ofHillia and Randia; spines sometimes present in Randia.

26a. Epiphytic shrubs and small trees 27

26b. Terrestrial shrubs or trees 29

27a. Flowers small (< 15 mm), ovary with 4 locules; fruits baccate with 4 pyrenes (Psychotria

spp.) Fig. 60

27b. Flowers large (> 15 mm), ovary with 2-4 locules; fruits elongate capsules with many seeds

or united at the base into a partial syncarp 28

28a. Corolla tube less than 2 cm long; fruits united at the base, fleshy (Schraderd) Fig. 19

28b. Corolla tubes more than 2 cm long; fruits free at the base, elongate capsules (Cosmibuena

with winged seeds and Hillia with a tuft of hairs at 1 end of the seed) Figs. 27-28

29a. Larger leaves usually becoming 40-50 cm long, entire or lobed in a few species; fruits many-seeded

29b. Larger leaves not usually becoming 40-50 cm long, never lobed; fruits 1 -many-seeded 33

30a. Minor venation of the leaves subparallel (lineolate), some species with deeply lobed leaves;

fruits baccate or hard, seeds angular (Pentagonid) Fig. 14

30b. Minor venation reticulate, leaves entire or with small lobes; fruits capsular, seeds mostly flat

3 la. Stipules almost separate, 4/node; inflorescences with few 1 branches and no bracteoles

(Condamined) Fig. 29

31b. Stipules united, 1-2/node; inflorescences with many 1 lateral branches and small bracteoles

32a. Fruits small, ca. 4 mm long (Elaeagid) Fig. 39

32b. Fruits large, ca. 5 cm long (Simird) Fig. 38

33a. Flowers with corolla tubes more than 10 cm long, white (native species not found in gardens) . .

33b. Flowers with corolla tubes less than 10 cm long (or, if close to 10 cm, red and planted for ornament)

34a. Flowers funnelform distally, with a gradually expanded tube, corolla lobes broadly triangular

(Osa) Fig. 15

34b. Flowers salverform distally and with a narrow tube throughout, corolla lobes narrowly ovate

to oblong 35

35a. Leaves narrowly elliptic, to 14 cm long, low shrubs of streamsides (Lindenid) .... Fig. 15
35b. Leaves not narrowly elliptic, usually more than 14 cm long, shrubs and trees of forests

(Posoquerid) Fig. 15

36a. Inflorescences with some calyx lobes greatly expanded to form large leaf-like red or white "petals"

36b. Inflorescences without calyx lobes greatly expanded (slightly expanded in some spp. of Rondeletid)

37a. Inflorescences racemose, to 60 cm long, expanded calyx lobes brilliant red (Warszewiczid)

Fig. 16

37b. Inflorescences not racemose, to 1 5 cm long, expanded calyx lobes white to pinkish red or
dull red . . 38

10 FIELDIANA: BOTANY

38a. Expanded calyx lobes densely pubescent, dull red (white in some forms); cultivated (Mus-

saendd) Fig. 16

38b. Expanded sepals glabrous to glabrescent; calyx lobes white or red; native and also planted

for ornament 39

39a. Corolla tube 3 mm long, white (Calycophylluni) Fig. 16

39b. Corolla tube 25 mm long, red (Pogonopus) Fig. 16

40a. Inflorescences dense heads of closely packed flowers 41

40b. Inflorescences lacking dense heads, if subcapitate the flowers becoming separate in fruit .... 43
4 la. Flowers united or connivent at the base, an involucre of bracts absent (Appunia, Morinda,

Schraderd) Fig. 19

41b. Flowers not united or connivent at the base, an involucre present or absent 42

42a. Inflorescences subtended by an involucre of colorful large bracts; native trees and shrubs

(Psychotria spp.) Figs. 7, 17-18, 56

42b. Inflorescences spherical, without an involucre; introduced trees (Neolamarckia, not illus-
trated).

43a. Inflorescences long and narrow (racemiform to spiciform) 44

43b. Inflorescences not long and narrow 46

44a. Flowers arising separately and sessile, inflorescences spicate; rare in Costa Rica (Alseis sp.)

Fig. 40

44b. Flowers usually in small cymose groups, these often on short secondary peduncles, inflores-
cences racemose 45

45a. Fruits fleshy (Gonzalagunid) Figs. 20.-21

45b. Fruits dry dehiscent capsules (Rondeletia) Fig. 21

46a. Flowers solitary or few at the ends of branches or short shoots, with short inconspicuous peduncles
when few; fruits usually solitary at the tips of branches, usually large (-1-2 cm) and rounded; seeds
many, often imbedded in pulp or horizontal (note: Faramea luteovirens and Rudgea monofructus,
with few-seeded fruits and flowers resembling those in fig. 46, and Serissa, a small ornamental

shrub with short stiff leaves, are not illustrated) 47

46b. Flowers not solitary at the ends of branchlets, on well-developed peduncles when few; fruits rarely

solitary and terminal 48

47a. Ovaries unilocular with parietal placentation (but difficult to see, with placentas often fusing
in the center), seeds variously oriented in white pulp turning black; spines present in some

species (Randia spp.) Figs. 22-24

47b. Ovaries usually 2-8-locular (but the septa often difficult to see), seeds mostly horizontal;
spines absent in all species (other genera of Gardenieae; see the technical key) Figs. 25-26

48a. Corollas 3-10 cm long 49

48b. Corollas less than 3 cm long 58

49a. Fruits elongate and flattened or cigar-like capsules, seeds many and winged 50

49b. Fruits not elongated capsules, seeds not winged 52

50a. Flowers sericeous on the exterior (Ladenbergia and Cinchona) Figs. 29, 37

50b. Flowers glabrous on the exterior 51

51a. Seed with a tuft of hairs (Hillid) Figs. 27-28

5 Ib. Seed without hairs (Cosmibuend) Figs. 27-28

52a. Flowers usually axillary and solitary (Exostema caribaeum) Fig. 31

52b. Flowers neither axillary nor solitary 53

53a. Corollas inflated in bud; capsules flattened and opening on the broad side (Coutared)

Fig. 31

53b. Corollas not inflated in bud; capsules not flattened or with fleshy fruits 54

54a. Flowers red to purplish; garden ornamentals Fig. 31

54b. Flowers white or yellowish; native species 55

55a. Ovule 1 in each locule (Guettarda turrialband) Fig. 30

55b. Ovules many in each locule 56

56a. Corolla densely sericeous (Duroia and Amaioud) Figs. 25, 30

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 1 1

56b. Corolla glabrous on the exterior 57

57a. Corolla lobes convolute; ovary 2-locular; fruits ca. 9 cm diam. (Tocoyena) Fig. 30

57b. Corolla lobes valvate; ovary 2-6-locular; fruits ca. 1 cm diam. (Isertid) Fig. 30

58a. (from 48b) Inflorescences axillary; fruits from axillary peduncles; corollas mostly pubescent on the

exterior (glabrous in Chimarrhis and some species of Hoffmannid) 59

58b. Inflorescences terminal; fruits from terminal peduncles (or from pseudoaxillary peduncles when

lateral shoots continue growth); corollas glabrous or puberulent 69

59a. Ovule 1/locule; seeds 1 /chamber in a bony endocarp, fruits drupaceous; flowers often along
1 side of the inflorescence branches; minor leaf venation parallel in some Chomelia and

Guettarda spp 60

59b. Ovules 3-many/locule; fruits many-seeded berries and capsules; flowers mostly cymose;

minor leaf venation parallel in Sommera 65

60a. Fruits small woody capsules with many seeds 61

60b. Fruits fleshy, drupes or berries with 1-2 pyrenes or with a single stony endocarp . . 62
6 la. Capsules ca. 5 mm long, rounded; corollas 3-5 mm long (Chimarrhis) . . Fig. 37
61b. Capsules 9-20 mm long, elongate; corollas 7-14 mm long (Macrocnemum) . . .

Fig. 40

62a. Fruits usually with 2 pyrenes 63

62b. Fruits with a single hard endocarp with 2-5 locules [fruits not flattened or economically

useful] 64

63a. Fruits usually flattened laterally and with a lustrous white surface [corolla tubes

< 9 mm long; native plants] (Chiococcd) Fig. 36

63b. Fruits rounded, becoming red 64

64a. Corolla tubes > 10 mm long; widely cultivated (Coffea, not illustrated).

64b. Corolla tubes < 1 5 mm long; wild or rarely cultivated (species of Psychotrid)

Figs. 12-13

65a. Corolla lobes valvate or slightly imbricate in bud, often with a distal appendage (Cho-
melia) Figs. 33-34

65b. Corolla lobes broadly imbricate with 1-2 exterior lobes, lacking distal appendages

(Guettarda) Figs. 32-34

66a. (from 59b) Plants generally with few lateral branches, weak subshrubs to 3 m tall (Hoffmannid)

Figs. 7-11

66b. Plants trees, shrubs, or clambering 67

67a. Fruits capsular (Chimarrhis) Fig. 37

67b. Fruits berry-like 68

68a. Stems clambering; locules 3-5 (Sabiced) Fig. 35

68b. Trees and shrubs; locules 2 (Sommera) Fig. 35

69a. (from 58b) Fruits dry and mostly capsules; ovaries with usually more than 1 ovule per locule . .

69b. Fruits fleshy or with arenchymatous tissue; ovaries with 1-many ovules per locule 76

70a. Fruits dry samaras with rounded circumferential wings, red and making a colorful display;
rarely collected trees (Allenanthus, not illustrated).

70b. Fruits not flattened samaras with a winged margin 71

7 la. Flowers small ( 1 cm) and often closely congested in dense inflorescences or in small groups

on open branched inflorescences 72

71b. Flowers usually more than 1 cm long, not usually closely congested in the inflorescence . .

72a. Spines often present; capsules opening from the base (Machaonid) Fig. 37

72b. Spines absent; capsules opening from the top 73

73a. Anthers opening by terminal pores; leaves with pellucid dots (Rustid) Fig. 38

73b. Anthers opening by lateral slits; leaves lacking pellucid dots 74

74a. Corolla yellow; shrubs or small trees of higher elevations (Depped) Fig. 38

74b. Corolla white; medium to large trees of low and high elevations (Elaeagid) . . . Fig. 39

12 FIELDIANA: BOTANY

75a. Capsules usually small (2-5 mm) and rounded; corollas often pubescent (Rondeletia spp.)

Figs. 41-41A

75b. Capsules usually more than 9 mm long and elongated, rounded or flattened; corollas puber-

ulent (Cinchona and Exostemd) or glabrous (Ferdinandusa and Macrocnemum)

Figs. 37, 40

(from 69b) Ovaries with 3-many ovules per locule; fruits usually many-seeded 77

Ovaries with 1 ovule per locule; fruits with 1-2 seeds (usually 5 in Psychotria racemosd) ... 80

77a. Anthers with the thecae divided into small sections [ovaries 2-6-locular] (Isertid)

Figs. 30, 49

77b. Anthers with the thecae not divided into sections 78

78a. Ovaries and fruits 5-locular (Hamelid) Figs. 34, 42

78b. Ovaries and fruits 2-locular 79

79a. Corolla lobes valvate in bud; inflorescences with cymose branches (Raritebe) Fig. 43

79b. Corolla lobes contorted in bud; inflorescences with flowers along 1 side of branches (Bertierd)

Fig. 43

(from 76b) Fruits usually with only 1 pyrene (seed), rounded in cross-section; the ovary 2-locular
in early stages, with a thin septum or with only 1 locule; flowers white to brilliant blue or lavender

(rarely yellowish) 81

Fruits usually with 2 pyrenes or seeds, the pyrene plano-convex in cross-section; the ovary 2-locular
and with a well-developed septum in early stages; flowers white to yellow, orange, red, purple, or

bluish purple (rarely blue) 84

8 la. Flowers brilliant blue, blue-lavender, or white; fruits usually broader than long to globose;

stipules acute to long-awned at the apex 82

8 1 b. Flowers white to yellowish white; fruits usually longer than broad; stipules obtuse to acute,

not awned 83

82a. Larger-leaved species of Faramea Fig. 44

82b. Smaller-leaved species of Faramea Fig. 45

83a. Smaller-leaved species of Coussarea Figs. 46-47

83b. Larger-leaved species of Coussarea Figs. 48

Corolla lobes contorted in bud, corollas white to red; wild plants and cultivated ornamentals

(Ixord) Fig. 43

Corolla lobes valvate in bud, corollas white to red, yellow, or purple; plants not cultivated orna-
mentals 85

Stipules usually rounded distally and with several short indurated tooth-like appendages at the

apex; inflorescences often few-flowered (Rudged) Fig. 46

Stipules not rounded distally and with thickened tooth-like structures at the apex; inflorescences

with few to many flowers 86

Flowers usually brightly colorful, yellow to orange, purple, or bluish purple (rarely white), corolla
tubes often slightly enlarged on 1 side at the base, a ring of hairs present on the interior of the

swollen lower half of the corolla tube (Palicourea spp., see also fig. 54) 87

Flowers usually white or slightly yellowish, corolla tubes not expanded on 1 side at the base, a
ring of hairs not present in the cylindrical lower half of the interior of the corolla tube (Psychotria

spp.) 88

87a. Palicourea spp.: inflorescences with conspicuous bracts and a species with spathaceous calyx

Fig. 49

87b. Palicourea spp.: lowland species Fig. 50

87c. Palicourea spp.: flowers mostly yellow-orange Fig. 51

87d. Palicourea spp.: flowers mostly purple-violet Fig. 52

87e. Palicourea spp.: smaller-leaved species Fig. 53

Fruits becoming blue, purple, or black (orange in P. racemosa with 5 pyrenes, red in P. haema-
tocarpa with very small inflorescences); leaves drying greenish to brown (except when treated with
isopropyl alcohol), domatia usually absent (except P. acuminata); stipules often persisting and not
subtending a ring of reddish colleters (subgenus Heteropsychotria and other species) 89

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 13

88b. Fruits becoming red at maturity; leaves drying grayish, grayish pink, or reddish brown to black,
domatia present in a few species; stipules usually caducous and enclosing a ring of reddish colleters

at their base (subgenus Psychotrid) 90

89a. Heteropsychotria: large-leaved and pubescent Fig. 54

89b. Heteropsychotria: smaller-leaved species Fig. 55

89c. Heteropsychotria: smaller inflorescences Fig. 56

89d. Heteropsychotria: large open inflorescences Fig. 57

89e. Heteropsychotria: conspicuous inflorescences Fig. 58

89f. Heteropsychotria: deciduous and other species Fig. 59

90a. Subg. Psychotria: species with very small leaves Fig. 60

90b. Subg. Psychotria: species with small leaves Fig. 61

90c. Subg. Psychotria: species with Ficus-\ike stipules Fig. 62

90d. Subg. Psychotria: species with dense pubescence Fig. 63

90e. Subg. Psychotria: species with larger leaves Fig. 64

90f. Subg. Psychotria: deciduous and unusual species Fig. 65

90g. Subg. Psychotria: miscellaneous unusual species Fig. 66

14 FIELDIANA: BOTANY

Manettia reclinata

Arcytophyllum muticum

FIG. 1 . Twining shrubs (Manettia spp.) and subshrubs with small stiffleaves (species of Arcytophyllum, Declieuxia,
and Diodia).

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Coccocypselum hirsutum

C. cordifolium

Coccocypselum lanceolatum

Geophila repens

G. cordifola

Geophila macropoda

FIG. 2. Twining herbs: species of Coccocypselum and Geophila.

FIELDIANA: BOTANY

Galium aschenbornii

Oldenlandia

corymbosa

FIG. 3. Herbs with small leaves and slender stems: species of Didymaea, Galium, Nertera, and Oldenlandia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Spermacoce verticillata

S. suaveolens

S. densiflora

Richardia scabra

Crusea parviflora

Mitracarpus hirtus

FIG. 4. Erect herbs with narrow lanceolate leaves and capitate or verticillate flowers: species of Crusea, Mitracarpus,
Richardia, and Spermacoce.

FIELDIANA: BOTANY

Spermacoce exilis

Spermacoce prostrata

10 cm

Spermacoce ovalifolia

S. assurgens

latifolia

confusa

FIG. 5. Erect herbs with narrow lanceolate leaves: Spermacoce spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

D. serrulata

cJi

FIG. 6. Erect herbs with narrow lanceolate leaves: Diodia spp. and two species of Spermacoce.

FIELDIANA: BOTANY

Hoffmannia davidsoniae

Amphidasya ambigua

Psychotria guapilensis

Lasianthus panamensis ^ GJ3

FIG. 7. Herbs or subshrubs with larger leaves: species of Amphidasya, Hoffmannia, Lasianthus, and Psychotria.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Hoffmannia

subauriculata

H. vesiculifera

FIG. 8. Herbs or subshrubs with axillary flowers: unusual species of Hoffmannia.

FIELDIANA: BOTANY

FIG. 9. Herbs or subshrubs with axillary flowers: pubescent species of Hqffmannia and H, congesta.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 23

Hoffmannia
leucocarpa

11111111111 10cm

H. dotae

<J5

H. asclepiadea

FIG. 10. Subshrubs with axillary flowers: species of Hoffmannia with larger leaves.

FIELDIANA: BOTANY

H. hamelioides

H. longipetiolata

H. psychotriifolia

Hoffmannia
pallidiflora

H. inamoena

H. laxa

FIG. 1 1 . Subshrubs with axillary flowers: species of Hoffmannia with leaves tapering gradually to the base.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

P. macrophylla

sycnotria aggregata
Psychotria cartagoensis

Psychotria uliginosa
10 cm

FIG. 12. Subshrubs with axillary flowers: species of Psychotria.

FIELDIANA: BOTANY

FIG. 1 3. Subshrubs with axillary flowers: species of Psychotria.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Pentagonia
donnell-smithii

FIG. 14. Trees with very large or lobed leaves: three species of Pentagonia.

FIELDIANA: BOTANY

Osa
pulchra

Lindenia rivalis

Posoqueria grandiflora

FIG. 15. Flowers with very long corolla tubes: species of Lindenia, Osa, and Posoqueria.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Calycophyllum candidissimum

Pogonopus speciosus

Warszewiczia coccmea

ussaenda erythrophylla

FIG. 1 6. Inflorescences with greatly expanded petal-like calyx lobes: species of Calycophyllum, Mussaenda, Pogono-
pus, and Warszewiczia.

FIELDIANA: BOTANY

Psychotria poeppigiana

FIG. 1 7. Inflorescences of involucrate heads: species of Psychotria (formerly Cephaelis spp.).

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Psychotria glomerulata

c-8

FIG. 18. Inflorescences of involucrate or conspicuously bracteate heads: species of Psychotria.

FIELDIANA: BOTA1

Appunia guatemalensis

Schradera costaricensis

FIG. 19. Inflorescences of compact heads with flowers connivent at the base: species of Appunia, Morinda, and
Schradera.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Gonzalagunia panamensis

ovatifolia

FIG. 20. Inflorescences long and narrow: species of Gonzalagunia.

FIELDIANA: BOTANY

Rondeletia brenesii

Rondeletia buddleoides

Rondeletia urophylla

Gonzalagunia bracteosa

FIG. 2 1 . Inflorescences long and narrow: species of Gonzalagunia and Rondeletia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Randia brenesii

Randia
loniceroides

Randia
thurberi

Randia aculeata

R. armata

FIG. 22. Fruits usually terminal and solitary: species of Randia with small leaves.

FIELDIANA: BOTANY

Randia
altiscandens

FIG. 23. Fruits usually terminal and solitary: species of Randia with medium-sized leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

R. genipoides

FIG. 24. Fruits usually terminal and solitary: species of Randia with larger leaves.

FIELDIANA: BOTANY

FIG. 25. Fruits usually terminal and solitary: species ofAlibertia, Duroia, Genipa, and Hippotis.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Genipa americana

FIG. 26. Fruits usually terminal and solitary: species of Borojoa and Genipa.

FIELDIANA: BOTANY

FIG. 27. Plants usually epiphytic: species of Cosmibuena and Hillia with smaller leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Hillia macrophylla

Cosmibuena macrocarpa

FIG. 28. Plants usually epiphytic: species of Cosmibuena and Hillia with larger leaves.

FIELDIANA: BOTANY

Ladenbergia
brenesii

Condaminea
corymbosa

L. sericophylla

FIG. 29. Trees with large open inflorescences: species of Ladenbergia and Condaminea corymbosa.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Tocoyena
pittieri

FIG. 30. Inflorescences with clusters of long-tubular flowers: species ofAmaioua, Guettarda, Isertia, and Tocoyena.

FIELDIANA: BOTANY

Exostemma
caribaeum

FIG. 3 1 . Showy flowers: species of Coutarea, Crusea, Exostema, Ixora, and Pentas.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Guettarda crispiflora

FIG. 32. Inflorescences with scorpioid or helicoid branches: species of Guettarda.

FIELDIANA: BOTANY

Guettarda foliacea

G. brenesii

FIG. 33. Flowers with narrow corolla tubes: species of Guettarda and a species of Chomelia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Chomelia microloba

Hamelia rovirosae

Chomelia recordn
Chomelia spinosa

FIG. 34. Flowers with narrow corolla tubes: species of Chomelia, Guettarda, and Hamelia.

FIELDIANA: BOTANY

Sabicea panamensis

Sommera donnell-smithii

FIG. 35. Inflorescences mostly axillary: species of Sabicea (vines) and Sommera (trees).

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

FBCX 3*. b

species of Chucocca aad a species of Omome.

FIELDIANA;

Machaonia
martinicensis

Chimarrhis latifolia

FIG. 37. Many small flowers in dense inflorescences: species of Chimarrhis, Cinchona, Machaonia, and Uncaria.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Rustia costaricensis

Simira maxonii / \ Rustia occidentalis

FIG. 38. Flowers in much-branched open inflorescences: species of Deppea. Rustia, and Simira.

FIELDIANA: BOTANY

FIG. 39. Small flowers in dense or open panicles: species of Elaeagia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Alseis sp. aff. hondurensis

Exostemma mexicanum

Ferdinandusa panamensis f^~ ^Macrocnemum glabrescens

FIG. 40. Small flowers and capsular fruits: species of Alseis, Exostema, Ferdinandusa, and Macrocnemum.

FIELDIANA: BOTANY

R. monteverdensis

FIG. 4 1 . Rondeletia spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Rondeletia
povedae

Rondeletia
chaconii

R. hamelifolia

FIG. 41 A. Rondeletia spp.

FIELDIANA: BOTANY

axillaris

FIG. 42. Hamelia spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Ixora nicaraguensis

Raritebe palicoureoides

FIG. 43. Bertiera, Ixora, and Raritebe spp.

FIELDIANA: BOTANY

FIG. 44. Faramea: species with larger leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

parvibractea

6J3

FIG. 45. Faramea: species with smaller leaves.

FIELDIANA: BOTAN\

R. cornifolia

Rudgea reducticalyx

C. impetiolaris

R. trifurcata

FIG. 46. Coussarea and Rudgea spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Psychotria
umbelliformis

FIG. 47. Coussarea spp. and two similar Psychotria spp

FIELDIANA: BOTANY

Coussarea talamancana

FIG. 48. Coussarea: species with larger leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

tilaranensis

FIG. 49. Palicourea: species with conspicuous bracts.

FIELDIANA: BOTANY

Palicourea
guianensis

1 cm

P. crocea

FIG. 50. Palicourea: species of lower elevations and a species of Isertia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Ufl

P. lasiorrhachis

FIG. 5 1 . Palicourea: species with larger leaves and yellow or orange flowers.

FIELDIANA: BOTANY

Palicourea
purpurea

FIG. 52. Palicourea: species with larger leaves and blue, lavender, purple, or white flowers.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

FIG. 53. Palicourea: species with smaller leaves.

FIELDIANA: BOTANY

Palicourea
standleyana

i i i i i i i i i I i i i i i i i i i I

FIG. 54. Psychotria subg. Heteropsychotria: larger-leaved pubescent species, and a species of Palicourea.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Psychotria goldmanii , ^

rt.

P. steyermarkii

FIG. 55. Psychotria subg. Heteropsychotria: species with smaller leaves.

FIELDIANA: BOTANY

Psychotria
haematocarpa

deflexa

FIG. 56. Psychotria subg. Heteropsychotria: species with very small inflorescences.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Psychotria
tapantiensis

FIG. 57. Psychotria subg. Heteropsychotria: species with larger open inflorescences.

FIELDIANA: BOTANY

berteriana

FIG. 58. Psychotria subg. Heteropsychotria: species with conspicuous open inflorescences.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

2 mm

irticinaiis ^v^ ---^ - ^C*TI . .'_ . vi-i*ssss5Sfc. J / T x.'^-^^x" rgcsmosa

FIG. 59. Psychotria subg. Heteropsychotria: species of deciduous habitats and some with smaller inflorescences.

FIELDIANA: BOTANY

Psy. guadalupensis (sensu lato)

FIG. 60. Psychotria subg. Psychotria: species with very small leaves and a complex of epiphytic species.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

marginata

Psychotria
monteverdensis

FIG. 6 1 . Psychotria subg. Psychotria: species with smaller narrow leaves.

FIELDIANA: BOTAN\

Psychotria
mexiae

FIG. 62. Psychotria subg. Psychotria: high-elevation species and those with Ficus-like stipules.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

FIG. 63. Psychotria subg. Psychotria: densely pubescent species.

FIELDIANA: BOTANY

panamensis var.
compressicaulis

FIG. 64. Psychotria subg. Psychotria: large-leaved species.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Psychotria
rosulatifolia

viridis

FIG. 65. Psychotria subg. Psychotria: deciduous and unusual species.

FIELDIANA: BOTAW

Psycnotna lamarinensis

FIG. 66. Psychotria subg. Psychotria: several unusual species.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Alibertia A. Richard

Trees or shrubs, dioecious, lacking spines, glabrous or
sparsely puberulent; stipules interpetiolar (and intrapeti-
olar in a few spp.), acute, usually persisting. Leaves op-
posite, petiolate; leaf blades entire, often with domatia.
Inflorescences terminal, 6 flowers fasciculate or capitate,
9 flowers solitary or 2, flowers of both sexes sessile or
subsessile, subtended by persisting stipule-like bracts.
Flowers unisexual, (3-)4-5(-8)-parted, hypanthium
hemispheric (in 9 flowers) to tubular (in <5 flowers), calyx
tube truncate or dentate; corolla salverform, fleshy or
subcoriaceous, corolla tube cylindrical, glabrous or vil-
lous within, 3-8-lobed, lobes short to long, obtuse to
acute, convolute in bud; stamens 3-8, filaments short or
absent, anthers linear, dorsifixed, included within the

tube; ovary 2-8-locular, ovules 3-many on axile placen-
tas within each locule, few-seriate to multi-seriate, often
imbedded in a pulpy placenta. Fruits terminal and sol-
itary, baccate, fleshy, sessile or subsessile, globose, usu-
ally over 2 cm diam., 2-8-locular with thin septa, peri-
carp firm and coriaceous, placentas becoming pulpy; seeds
usually many, compressed to rounded.

A genus of about 35 species, ranging from Mex-
ico, the West Indies, and Central America into
South America. Alibertia is recognized by its dioe-
cy, sessile or subsessile terminal flowers, and the
large terminal subglobose fruit with a persisting
terminal calyx tube. This genus is similar to
Amaioua and Borojoa.

Key to the Species of Alibertia

la. Stipules 7-20 mm long; flowers 16-34 mm long; leaves elliptic-oblong; usually tapering gradually
to the apex A. edulis

Ib. Stipules to 4 mm long; flowers 7-10 mm long; leaves usually somewhat obovate, abruptly narrowed
to an acuminate or rounded apex A. garapatica

Alibertia edulis (L. C. Rich.) A. Rich, ex DC.,
Prodr. 4: 443. 1830. Genipa edulis L. C. Rich.,
Actes Soc. Hist. Nat. Paris 1: 107. 1792. Gar-
denia edulis (L. C. Rich.) Poir. in Lam., Encyc.
Meth. Bot. Suppl. 2: 708. 1812. Figure 25.

Shrubs or rarely small trees, l-4(-6) m tall, sometimes
forming thickets, bark often peeling in longitudinal strips,
leafy branchlets (1.5-)2-4.5 mm thick, glabrous or
sparsely puberulent, reddish brown and becoming gray-
ish; stipules 7-15(-20) mm long, 2-4(-5) mm broad at
base, triangular to deltoid, acute or acuminate, longi-
tudinally striate, acute or acuminate, brownish and gla-
brous, subcoriaceous, persisting with older leaves. Leaves
with petioles 2-5(-10) mm long, 0.5-1.8 mm thick, gla-
brous; leaf blades 5-14(-20) cm long, 1 .5-5(-8) cm broad,
narrowly elliptic-oblong to broadly ovate-oblong, apex
acute to long-acuminate, base gradually narrowed and
attenuate (in narrow leaves) to rounded and subtruncate
(in broader leaves from Caribbean lowlands), drying stiff-
ly chartaceous to subcoriaceous and often grayish green,
lustrous above in life, glabrous on both surfaces (or pu-
berulent beneath in material from Caribbean lowlands),
2 veins 6-1 2/side, small domatia of pits and tufted hairs
often present in vein axils beneath. Inflorescences of
subsessile <5 or 9 flowers, subtended by stipule-like bracts
7-15 mm long. Flowers 4- (rarely 5-) parted, 16-34 mm
long, hypanthium 4-8 mm long, 3-5 mm diam., calyx
tube truncated except for the short (0.3-1 .7 mm) narrow
(0.3 mm) lobes; corolla white, tube 2-3 cm long, lobes
to 2 cm long and 12 mm broad, triangular, minutely
puberulent; stamens usually 4, subsessile, anthers ca. 1 3
mm long in <5 flowers; staminodes 5 mm long in 9 flowers.
Fruits 2-3 cm diam.. obovoid to subglobose, yellowish
brown, slightly truncated distally, persisting calyx tube

4-6 mm long and 4-6 mm diam.; seeds 3-8 mm long,
oblong, slightly flattened longitudinally, striate.

Shrubs of both deciduous and evergreen forest
formations, from near sea level to 500(-1000) m
elevation. Probably flowering and fruiting
throughout the year, with the main flowering sea-
son April-July. The species ranges from Mexico
to northern South America.

Alibertia edulis is recognized by its solitary ter-
minal rounded fruit with persisting calyx tube,
generally narrow oblong-elliptic leaves, stiff striate
interpetiolar stipules, and generally shrubby habit.
In Central America the species is most common
in deciduous or partially deciduous woodland.
Common names for this species in Central Amer-
ica are lagartillo, trompillo, trompo, trompito, and
"wild guava." The fruit is occasionally eaten by
local people and sporadically cultivated. This spe-
cies may intergrade with material currently placed
under other names in South America; compare A.
acuminata (Benth.) Sandwith and A. latifolia
(Benth.) Schum. Specimens may resemble some
species of Randia.

Material from the Caribbean lowlands placed
under this name differs in having broader leaves
that dry dark and have short straight hairs on their
undersurfaces. This material, while quite different
from that found on the Pacific slope, appears to
intergrade with the more typical forms in Guate-

FIELDIANA: BOTANY

mala and Belize; it is not often collected in Costa
Rica.

Alibertia garapatica K. Schum. in Mart., Fl. Bras.
6(6): 384. 1889.

Shrubs or small trees to 5 m tall, leafy branchlets
slender (1-2 mm thick) with slightly thickened nodes,
terete, brownish, minutely (0. 1 mm) puberulent and gla-
brescent; stipules 2-4 mm long, ca. 2 mm broad at the
base, persisting or deciduous. Leaves with petioles 2-
7(-10) mm long, 0.7-1 .2 mm thick, minutely puberulent;
leaf blades 5- 1 2(- 1 5) cm long, 2-5(-7) cm broad, broad-
ly elliptic-obovate to broadly oblong-obovate or elliptic-
oblong, apex abruptly narrowed and acuminate, caudate-
acuminate or rounded, tip 5-15(-20) mm long, base
obtuse to cuneate, drying stiffly chartaceous and usually
grayish in color, glabrous above and below, 2 veins 5-
7/side, domatia of tufted hairs in the vein axils beneath
(in Colombian material). Inflorescences terminal and
capitate, sessile or subsessile, with 4-8(-16) S flowers,
the 9 flowers solitary or paired, flowers subtended by 2
triangular stipules (bracts). Flowers aromatic, 6-10 mm
long, hypanthium 1-2 mm long, ca. 1.3 mm broad, ob-
conic, distal margin entire, calyx cup and teeth minute
(0.5 mm) or absent; corolla tube 3-7 mm long, greenish,
lobes white, 1.5-3 mm long, ca. 1.5 mm broad near the
base; stamens 4. Fruits solitary, sessile or subsessile, glo-
bose or subglobose to obovoid, ca. 25 mm long and 30
mm diam., drying black.

Trees of evergreen and partly deciduous for-
mations on both the Caribbean and Pacific slopes
in central Panama, from near sea level to 500 m
elevation. In Costa Rica it is known only from
near Punta Mala on the Pacific coast (A. Jimenez
3912, flowering in March) and the Reserva Biol.
Carara (Zuniga 232, fruiting in May), both in
southern Puntarenas Province. The species ranges
from Costa Rica to Colombia and occurs in south-
ern Mexico.

Alibertia garapatica is recognized by its terminal
subsessile flowers and solitary fruit, small inter-
petiolar stipules, relatively broad and slightly ob-
ovate leaves that dry chartaceous, and smaller
flowers. A short tube may be present on some
stipules. This species is poorly known; it may be
mistaken for some species of Randia.

Allenanthus Standley

Small to medium size trees, branchlets glabrous or
sparsely puberulent; stipules interpetiolar, persistent or
deciduous. Leaves opposite, short-petiolate, leaf blades
acuminate, entire, often with minute domatia. Inflores-
cences panicles with opposite branching, broadly cor-
ymbose in form, terminal or axillary to distal leaves,

bracteate, flowers pedicellate. Flowers bisexual, small (3-
6 mm), hypanthium obovoid to urceolate, truncated dis-
tally, laterally compressed, calyx lobes 4, small; corolla
whitish, tubular and with 4 spreading lobes, valvate to
somewhat imbricate; stamens 4, borne in the throat of
the corolla tube, filaments slender, anthers oblong; ovary
2-locular, with 1 ovule in each locule, style distally bifid.
Fruits becoming dry and samara-like, flattened with broad
lateral wings surrounding the 2 central narrow longitu-
dinally parallel seed chambers, material of the wings
slightly spongy, calyx lobes persisting; seeds laterally
compressed, pendulous.

A small genus with two species, ranging from
central Mexico to western Panama. Allenanthus
hondurensis Standley is found in central and
southern Mexico and in Honduras; our species
also appears to have a disjunct distribution in Cos-
ta Rica and Panama. The flattened fruit, resem-
bling that of Ulmus or some Terminalia species,
is unique among Central American Rubiaceae.

Allenanthus erythrocarpa Stand!.. Ann. Missouri
Bot. Gard. 27: 344. 1940. Chimarrhis decurrehs
Steyerm., Ceiba 3: 18. 1952.

Trees, 6-20 m tall, leafy branchlets 1.5-5 mm thick,
internodes 4-8 cm long, usually glabrous, subterete; stip-
ules 3-6 mm long, 2-3 mm broad at the base, apex acute,
puberulent within, deciduous. Leaves with petioles 6-
15 mm long, 1-1.5 mm thick, sulcate with adaxial mar-
gins with punctate (gland-like) projections along the edge;
leaf blades 6-1 1 cm long, 3-5 cm broad, ovate-elliptic
to ovate-oblong, apex tapering gradually and acuminate,
tip to 1 .5 cm long, base obtuse and slightly decurrent on
petiole, drying thin chartaceous and sometimes dark in
color, 2 veins 5-7 /side, glabrous above or puberulent
only along the major veins, with small (0.2 mm) as-
cending hairs on the major veins beneath, usually with
small tufted domatia in slight depressions in the vein
axils beneath (with 2-lipped structures ca. 1 mm long at
the vein axils in Zamora & Poveda 825). Inflorescences
both terminal and sometimes also axillary to distal leaves
and together forming a single conspicuous panicle (thyrse)
to 15 cm long and 10 cm broad, becoming 20 cm long
and 18 cm broad in fruit, primary peduncles 3-7 cm
long, terete, shorter toward apex of the inflorescence,
peduncles and rachis with opposite lateral branching,
with 1 or 2 longitudinal lines of dense short (0.2-0.4 mm
long) ascending hairs, bracts 1-2 mm long, pedicels 1-
2 mm long. Flowers with the hypanthium 1-2 mm long,
somewhat flattened (compressed), calyx lobes 4, 0.5-1
mm long; corolla becoming 4 mm long, tube ca. 3 mm
long, lobes 4, rounded; stamens 4, exserted on slender
filaments ca. 1.5 mm long, anthers ca. 0.8 mm long.
Fruits flat and samara-like, pink to red, 5-7(-8) mm long,
3-4(-6) mm broad, oblong-elliptic in outline, the base
of the fruit decurrent on pedicel, body of the fruit ca.
0.7 mm thick, fruiting pedicels ca. 4 mm long; seeds
forming an oblong area in the center of the fruit ca. 2
mm long and 1 mm broad.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Trees of evergreen or partly deciduous forest
formations of the Pacific slope, at around 500-700
m elevation. The species is known from near Par-
que Nacional Rincon de la Vieja in Guanacaste
Province (Herrera & Rivera 843 CR, MO, Zamora
& Poveda 825 CR, F). Flowers were collected in
June (Panama); fruiting in August-September
(Panama) and October (Costa Rica). The species
is known only from Costa Rica and western Pan-
ama.

Allenanthus erythrocarpa is recognized by its
flattened reddish samara-like fruit with small dis-
tal calyx lobes, conspicuous infructescences, and
unusual lines of hairs along branches of the inflo-
rescences. With their broad terminal inflores-
cences and bright red or pink fruits, these trees are
very conspicuous when fruiting (Zamora & Po-
veda 825 CR, F).

Allenanthus hondurensis Standl. of northern
Central America is a smaller tree found in decid-
uous forest, with the leaves more puberulent be-
neath and smaller (5x3 mm) yellowish fruit.

Alseis Schott

Trees or large shrubs, branchlets terete, glabrous or
more often puberulent; stipules interpetiolar, triangular
to subulate, caducous or persisting. Leaves opposite, of-
ten clustered at the ends of branchlets, petiolate; leaf
blades often narrowly obovate, drying thin-chartaceous,
sometimes with domatia. Inflorescences terminal or ax-
illary, solitary in each axil, usually spike-like or racemose
and cylindrical in form, simple or with lateral branches
(paniculate and racemiform), flowers lacking pedicels or
the pedicels merging gradually into the base of the ovary.
Flowers bisexual, small, white to yellow, protogynous;
hypanthium obconical to subcylindrical, calyx lobes 5,
deciduous; corolla tube cylindrical to campanulate or
urceolate, villous within, corolla lobes 5, valvate (?rarely
open) in bud; stamens 5, filaments attached near the base
of the corolla tube, anthers exserted, oblong and sagittate,
dorsifixed; ovary 2-locular, septum thin, placentas apical
with numerous ovules in each locule, style long, distally
bifid with recurved stigmas. Fruits capsular, cylindrical,
2-locular and bivalvate, dehiscing septicidally from apex
to base; seeds numerous, linear-fusiform, the testa re-
ticulate and prolonged at apex and base.

A genus of about 20 species found in Mexico
and Central America and southward to Peru and
Brazil. The often long (ca. 20 cm) obovate leaves
clustered at the ends of twigs, long (15-30 cm)
spicate or racemiform inflorescences with many
flowers, and narrow capsular fruit splitting into
two parts distinguish members of this genus. Su-
perficially, these plants may resemble some species

of Gonzalagunia and Rondeletia. Alseis blackiana
Hemsl., with leaves to 30 cm long, is found in
central and eastern Panama. Alseis hondurensis
Standl. occurs in northern Honduras, Guatemala,
and Belize, while A. yucatanensis Standl. occurs
in southern Mexico, Belize, and Guatemala.

Alseis sp. aff. A. hondurensis Standl., Trop. Woods
16:48. 1928. Figure 40.

Small trees, ca. 15m tall, trunk ca. 30 cm dbh with
soft bark, leafy branchlets 1.5-7 mm thick, glabrous, pale
brownish, smooth; stipules 3-8 mm long, 1-2 mm broad
at the base, triangular-subulate, apex acute, caducous.
Leaves clustered at the ends of twigs, petioles 10-55 mm
long, 0.8-2 m thick, glabrous and drying dark; leaf blades
(7-)9-19 cm long, (2-)3-8 cm broad, elliptic-obovate to
obovate or ovate-elliptic, apex short-acuminate, tip 4
7 mm long, base acute (obtuse) and occasionally slightly
decurrent on petiole, glabrous above, glabrous below ex-
cept for some thin hairs ca. 0.7 mm long near the vein
axils (domatia?), 2 veins 6-10/side. Inflorescences ter-
minal or axillary, solitary or 3, 1 1-18 cm long, ca. 2 cm
diam., spicate with flowers sessile on the rachis or with
opposite basal spicate lateral branches, peduncles to 4
cm long, ca. 1 mm thick, minutely grayish puberulent,
bracts and pedicels not apparent. Flowers with minute
(0. 1-0.2 mm) tomentulous grayish hairs, hypanthium ca.
1.5 mm long and 0.6 mm thick, cylindric to obconic,
calyx lobes ca. 0.5 mm high and 0.6 mm broad at the
base, triangular, brownish and mostly glabrous; corolla
white, short-tubular campanulate, 2-3 m long, corolla
lobes little differentiated; stamens exserted, anthers ca.
0.8 mm long, style branches recurved. Fruit apparently
narrowly obovoid and splitting into 2 valves, each valve
ca. 8 mm long and 2.5 mm broad, with a notch 1 mm
deep at apex, yellowish and smooth-lustrous within.

This species is presently known from only two
collections. Flowering material was collected in
February 1989 east of Bahia de Drake on the Osa
Peninsula (Q. Jimenez et al. 670 CR, F, MO). Old
fruit were collected in July (Hammel et al. 17120
CR, F, MO) at the Reserva Forestal El Cangrejo (near
the road from Puriscal to Quepos) at ca. 400 m
elevation in San Jose Province.

Alseis sp. aff. A. hondurensis is distinguished by
its sessile flowers on spicate inflorescences (rarely
paniculate with one or two lateral spicate branches
near the base), longer petioles drying dark, and
narrowly obovoid capsule splitting into two sep-
arate valves. The type of A. hondurensis differs in
the shorter petioles, minute puberulence on the
lower leaf surfaces, the leaf blades more often ob-
lanceolate with a gradually tapering base, and much
larger minutely puberulent inflorescences. In ad-
dition, A. hondurensis is a species of the Caribbean
lowlands, whereas our species is found on the Pa-

FIELDIANA: BOTANY

cific slope. Alseis blackiana Hemsl. of Panama dif-
fers in the much larger leaves with more secondary
veins. Both those species have clearly pedicellate
flowers, while the Costa Rican collections have
sessile flowers. However, species of Alseis appear
to be very variable, and it is possible that the Costa
Rican material will prove to be conspecific with
one of those other species.

Amaioua Aublet

Trees or shrubs, dioecious, branchlets usually puber-
ulent; stipules united, both interpetiolar and intrapetio-
lar, forming a conic cap over the shoot-apex and tearing
irregularly, caducous. Leaves opposite (rarely 3/node),
often crowded at the distal ends of stems, petiolate; leaf
blades entire, often with minute domatia in vein axils
beneath. Inflorescences terminal on the main stem or on
short lateral branches, usually fasciculate, with or with-
out primary peduncles, often 3-branched, flowers in ul-

timate cymose or capitate groups or solitary. Flowers
unisexual, hypanthium hemispheric to cupulate or tu-
bular, calyx tube truncate distally or dentate, calyx lobes
6 (5) or none; corolla usually salverform, corolla tube
terete, sericeous externally and minutely puberulent
within, corolla lobes 6 (rarely 5), spreading, oblong, con-
torted in bud. cream white to greenish; stamens 6 (5),
borne on the middle or lower part of the corolla tube,
filaments very short, anthers narrow, dorsifixed, includ-
ed; ovary 2-locular, placentas borne on the septa, ovules
many and biseriate in 2 horizontal rows in each locule,
style short with coherent(?) style branches. Fruits bac-
cate, oblong, areolate at apex (calyx scar), 2-locular, seeds
many, imbedded in a pulp, horizontal, suborbicular and
laterally compressed.

A small genus of about seven species, mostly in
South America; two species reach our area. The
compact terminal inflorescences with unisexual
sericeous flowers and many-seeded baccate fruit
help to distinguish this genus.

Key to the Species of Amaioua

1 a. Fruit in clusters on short peduncles, sessile or subsessile; ring of colleters or hairs above the new
stipule scars ca. 0.5 mm long and usually obscure A. corymbosa

Ib. Fruit usually borne individually on long pedicels in an umbel-like group at apex of stems; ring of

colleters above the new stipule scars ca. 1 mm long, visible and drying dark reddish

A. pedicellata

Amaioua corymbosa H.B.K., Nov. gen. sp. 3: 419,
pi. 294. 1820.

Shrubs or small trees, l-8(-l 5) m tall, leafy branchlets
2-5 mm thick, at first angular but becoming terete, ap-
pressed sericeous and glabrescent, with conspicuous leaf
scars, a very short (0.5 mm) ring of colleters present just
above the stipule scar on young stems; stipules 8-20 mm
long and 5-8(-10) mm broad at the base, sericeous ex-
ternally. Leaves with petioles 3-18(-30) mm long, to 3
mm thick, with stiff ascending sericeous hairs; leaf blades
5-14(-23) cm long, 3-8(-13) cm broad, elliptic-ovate,
ovate-oblong, broadly obovate, or broadly oblong-ellip-
tic, apex abruptly rounded or obtuse and short-acumi-
nate, base obtuse to acute and slightly decurrent on pet-
iole, drying stiffly chartaceous or subcoriaceous, glabrous
above, glabrous to sparsely appressed puberulent on the
veins beneath, 2 veins 5-8(-10)/side, some of the 3
veins subparallel and at right angles to the secondaries,
usually with small tufts of hairs in the vein axils beneath.
Inflorescences of $ flowers to 10 cm long, corymbose,
primary peduncles 0.5-5 cm long, simple or with 3 pri-
mary branches and the flowers in cymose groupings,
pedicels 1-8 mm long, sericeous; 2 inflorescences to 6
cm long, subtrichotomous to capitate, secondary branch-
es 0-3 mm long, pedicels usually absent. Male flowers
with hypanthium 3-5(-6) mm long and 3-4 mm diam.,
sericeous, calyx teeth 0.5-1.5 mm long, linear, corolla

10-18 mm long, white or grayish green, corolla tube 5-
7(-9) mm long, 1.5-3.5 mm diam., retrorse sericeous,
corolla lobes usually 5 or 6, 5-7(-9) mm long, lanceolate,
papillate-puberulent on the exterior; stamens 6 (5), an-
thers 4-6 mm long, filaments inserted in the middle of
the tube. Female flowers with hypanthium 3-5 mm long,
1-2 mm diam., calyx tube 2-4 mm long, 2.2-3 mm
diam., densely ascending sericeous, calyx teeth 0.5-1
mm long, subulate; corolla 8-1 2 mm long, tube 6-7 mm
long, 2-3 mm diam., densely retrorse sericeous exter-
nally, lobes 6 (5), 4-6 mm long, 1.5-2 mm broad, lan-
ceolate, papillate-puberulent within. Fruits 10-15(-17)
mm long, 4-9(-l 1) mm thick (dried), usually in dense
clusters of 3-10, red or reddish purple becoming black,
drying dark with a pale annular ring distally (scar of the
deciduous calyx tube); seeds irregular, 3-5 mm long to
4 mm broad, flattened, testa striate.

Trees of partly deciduous drier forests of the
Pacific slope but also found in evergreen forest
formations, from near sea level to ca. 300 m ele-
vation. Flowering in July and fruiting in Septem-
ber. This species ranges from southern Mexico
through Central America and southward to Co-
lombia, Venezuela, the Guianas, and Bolivia.

Amaioua corymbosa is recognized by its sub-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

sessile clusters of fruit often on three terminal
branches, densely sericeous flowers with lustrous
retrorse hairs on the corolla, and leaves often with
minute domatia. The ring of colleters just above
the encircling stipule scar are often hidden by the
pubescence. Though often collected in central Pan-
ama and in Nicaragua, we have seen only a few
collections of this species from northern Costa Rica:
Q. Jimenez 376 CR from near Liberia and Zamora
& Chacon 1355 CR from Refugio Cano Negro.

pedicellate fruit in terminal umbel-like groups,
flowers coming directly from the apex of the shoot
on usually unbranched stalks (pedicels), and
broadly elliptic leaves with long thin hairs on the
upper surface in early stages. The unusual glan-
dular teeth (colleters) above the stipule scar near
the apex of the stem are also distinctive. At pres-
ent, this species appears to be limited to a rather
narrow altitudinal range on the Caribbean slope
in Costa Rica.

Amaioua pedicellata Dwyer, Ann. Missouri Bot.
Gard. 67: 30. 1980. Figure 30.

Trees 5-10(-15) m tall, trunks to 22 cm dbh, leafy
branchlets 1.6-6 mm thick, with appressed-ascending
sericeous hairs 0.5-1 mm long, glabrescent, with a ring
of linear colleters ca. 1 mm long encircling the node just
above the stipule scar but breaking off early; stipules 8-
16(-30) mm long, cap-like and caducous, with dense
ascending lustrous sericeous hairs. Leaves with petioles,
8-18 mm long, 1-1.8 mm thick, appressed puberulent;
leaf blades 6-1 3(-l 9) cm long, 3-6(-l 0) cm broad, broadly
elliptic to broadly elliptic-oblong or elliptic-obovate, apex
usually short-acuminate, tip 5-10 mm long, base obtuse
(occasionally acute) and somewhat decurrent on petiole,
drying stiffly chartaceous and dark brown above, upper
surface of the young leaves with scattered slender whitish
appressed hairs to 2 mm long but these quickly falling
and the mature upper surfaces glabrous, lower surfaces
with thin ascending hairs 0.3-0.5 mm long on the major
and minor veins, 2 veins 7-1 I/side, occasionally with
domatia in vein axils beneath. Inflorescences terminal
fascicles of 6-12 pedicellate flowers, the flowers usually
on unbranched pedicels (rarely on peduncles bearing 2-
3 pedicellate flowers), later forming a sessile or umbellate
cluster of long-pedicellate fruit, pedicels 3-8 mm long,
with dense lustrous ascending sericeous hairs. Flowers
with hypanthium and calyx tube ca. 4 mm long and 3
mm diam.. calyx lobes 3-5, 0.5-1.5 mm long, subulate
or linear; corolla rose with pale greenish tube 7-9 mm
long, 1-3 mm diam., densely whitish sericeous, lobes 7-
9 mm long, 3 mm broad at base, narrowly triangular.
Fruits subglobose to oblong, 1 2- 1 7 mm long, 10-14 mm
diam., red to dark reddish purple (but drying black),
sparsely and minutely puberulent near the distal end,
annular calyx scar 34 mm diam., fruiting pedicels 20-
35 m long, 1-1.5 mm thick.

Trees of wet evergreen forest formations of the
Caribbean slope in Costa Rica, and both the Ca-
ribbean and Pacific slopes in Panama, from 600
to 900 m elevation. Flowering in June-July; fruit-
ing in February, September, and December (in
Panama). The species is known only from central
and southern Costa Rica and Code and Veraguas
provinces in Panama.

Amaioua pedicellata is recognized by its long-

Amphidasya Standley

Small shrubs or herbaceous subshrubs, woody at the
base, stems unbranched; stipules connate/interpetiolar,
large, lobed distally or deeply laciniate with filiform seg-
ments, persisting. Leaves often closely clustered near the
ends of stems, often long-petiolate; leaf blades large,
margins entire, domatia absent. Inflorescences terminal
or axillary, cymose to capitate, short, flowers usually
closely crowded, pedicels short. Flowers bisexual, hy-
panthium oblong to turbinate, calyx lobes 4-6, often
unequal, persisting; corolla tubular-salverform, corolla
lobes 4-6, valvate in bud; stamens 4-6, borne on the
middle or upper part of the corolla tube, filaments short,
anthers linear, dorsifixed; ovary 2-locular, with axile bi-
lobed placentas, many ovules in each locule. Fruits fleshy,
indehiscent, crowned by the persistent calyx lobes; seeds
many, angular, testa reticulate.

Amphidasya is a genus of about seven species,
ranging from Costa Rica through Panama to Co-
lombia, Venezuela, and northern Brazil. Our rep-
resentative is distinguished among Costa Rican
Rubiaceae by the short herbaceous habit, large and
long-petiolate leaves, densely clustered flowers with
long calyx lobes, and long corolla tube.

Amphidasya ambigua (Standl.) Standl., Field Mus.
Nat. Hist., Bot. Ser. 11:181.1931. Sabicea am-
bigua Standl., Publ. Field Columb. Mus., Bot.
Ser. 7: 49. 1930. Figure 7.

Herbs, 10-40(-90) cm tall, erect or decumbent, stems
woody at the base, leafy stems 3-7 mm thick, terete,
minutely puberulent, glabrescent, brownish; stipules 1 2-
20(-40) mm long, ca. 4 mm broad at the base, united
basal sheath 3-5 mm long, with long linear acute lobes,
minutely and inconspicuously puberulent. Leaves clus-
tered at the distal part of the stem, opposite or subop-
posite, often rosette-like, petioles 1.5-5 cm long, 1.9-2.8
mm thick, with few minute appressed hairs or glabres-
cent; leaf blades 12-28 cm long, 5-10 cm broad, nar-
rowly elliptic-obovate to narrowly oblong-obovate or
oblanceolate, apex abruptly narrowed and short-acu-
minate, base gradually narrowed and cuneate-attenuate,

FIELDIANA: BOTANY

long-decurrent on petiole, drying stiffly chartaceous, es-
sentially glabrous above, sparsely puberulent with mi-
nute (0. 1-0.2 mm) ascending hairs on the veins beneath
or glabrous, 2 veins 15-25/side. Inflorescences densely
crowded in the axils of distal leaves, to 5 cm long, base
of the inflorescence not usually visible, with 4-20 closely
crowded sessile or subsessile flowers, floral bracts 1-5
mm long, acute. Flowers 5- or 6-parted, the hypanthium
5-8 mm long, glabrous or minutely and sparsely papil-
late-puberulent in later stages, calyx lobes 8-18 mm long,
ca. 1.5 mm broad at the base, often unequal, glabrous
on the surfaces and with minute (0. 1 mm) hairs along
the edge; corolla narrowly salverform, white, puberulent
on the exterior, corolla tube 32-50 mm long, 1-2.5 mm
broad (dried), corolla lobes 6-18 mm long, triangular,
acute. Fruits 8-12 mm long, 4-6 mm diam., cylindrical-
oblong, with persisting calyx lobes, surface of the dried
fruit bullate from pressure of the seeds within; seeds 0.3-
0.4 mm diam., foveolate.

Plants of steep slopes in the shade of evergreen
rain forests on both the Caribbean and Pacific low-
lands, collected at elevations of 10-800 m (to 1 500
m in Panama). Flowering in August and Novem-

ber-December; fruiting in March-July, Septem-
ber, and November. The species ranges from Cos-
ta Rica to Colombia.

Amphidasya ambigua is recognized by its short
stature, distally fimbriate stipules, crowded long-
petiolate leaves, crowded flowers with relatively
long calyx lobes, and relatively long corolla tube.
These plants resemble species of Paradrymonia in
the Gesneriaceae (but the latter have superior ova-
ries). Costa Rican material was earlier thought .to
be a separate species, distinguished in the follow-
ing key. However, recent collections from Panama
and Colombia have produced many intermediate
variants and resulted in a broader interpretation
of A. ambigua. Note, however, that the preceding
description is based on Costa Rican material and
does not represent all the variation found within
the more widely defined taxon. The following key
outlines the differences between the western and
eastern collections.

la. Leaves often elliptic-oblong, petioles 2-7 cm long and densely appressed-puberulent, major sec-
ondary veins 12-18 on each side; calyx lobes densely puberulent, corolla tube 2-3 cm long; central
Panama to Colombia.

Ib. Leaves usually slightly obovate, petioles 1.5-5 cm long and glabrescent, major secondary veins 1 5-
25 on each side; broad surfaces of the calyx lobes glabrous, corolla tube ca. 4 cm long; Costa Rica
and western Panama.

Appunia Hooker f.

Shrubs or small trees, glabrous or puberulent; stipules
interpetiolar and united at the base with the petioles,
subulate-acuminate, persisting. Leaves opposite, short
petiolate, leaf blades often lanceolate and acuminate,
entire, mostly drying thin-chartaceous, domatia absent.
Inflorescences terminal or axillary, capitate with a few
flowers congested at apex of a short to long peduncle,
flowers free and subtended by bracteoles but without
pedicels. Flowers bisexual, small, white; hypanthium
hemispheric or oblong, calyx tube short-cylindrical, usu-
ally truncate, lobes absent or minute (5); corolla fun-
nelform to urceolate, corolla lobes usually 5, valvate in
bud; stamens 5, filaments short, anthers dorsiflxed and
included; ovary 4-locular, each locule with 1 ascending
ovule, style slender, stigma capitate. Fruits fleshy and
baccate, sessile and loosely aggregated on apex of the
peduncle, each fruit with 4 (or fewer) nutlets, each nutlet
with 2 unequal locules (a seed-bearing locule and an
empty locule).

A genus of about 1 species in Central and South
America. Most authors have placed this genus into
synonymy under Morinda, but that genus has the

basally united flowers developing into a fleshy syn-
carp and two stigmas.

Appunia guatemalensis J. D. Smith, Hot. Gaz. 48:
294. 1909. Morinda guatemalensis (J. D. Smith)
Steyerm., Mem. New York Bot. Gard. 23: 385.
1972. Figure 19.

Low or slender-branched shrubs, (0.5-)l-3(-4) m tall,
leafy stems 1.2-4 mm thick, terete, glabrous or rarely
minutely (0.0 1 mm) puberulent; stipules 1.5-3 mm long,
2-4 mm broad at the base, with a narrowed simple or
bifid tip ca. 0.5 mm long, glabrous. leaves opposite,
petioles 2-5 mm long, 0.8-1.6 mm thick, glabrous or
sparsely and minutely (0.05 mm) puberulent; leaf blades
7-1 6 cm long, 3-7 cm broad, elliptic-oblong to narrowly
obovate, apex acuminate to acute, base acute, drying
grayish green to dark olive green and often lustrous above,
glabrous above and below, 2 veins 4-6/side. Inflores-
cences solitary in leaf axils (2/node), borne on glabrous
peduncles 3-27 mm long and 0.5-1 mm thick (dried),
capitula with 3-12 sessile and congested flowers sub-
tended by triangular bracts ca. 1.5 mm long. Flowers

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

glabrous, hypanthium and calyx tube ca. 2 mm long, 1.7
mm diam. distally, calyx lobes not developed; corolla
ca. 1 5 mm long, white or greenish, corolla lobes 5-6 mm
long, 1.5 mm broad at the base, becoming recurved,
greenish within. Fruits 6-8 mm long, 4-6 mm diam.,
subglobose, sessile, purple to brownish or black.

Plants of low elevation in open grassy sites or
thickets, 0-300 m elevation. Flowering in Janu-
ary-September in northern Central America. The
species ranges from central Mexico along the Ca-
ribbean coast to southern Nicaragua and has been
only rarely collected in the Pacific lowlands of
northern Costa Rica.

Appunia guatemalensis is distinguished by its
small capitate inflorescences on slender peduncles
in the axils of leaves, sessile flowers and fruits, and
usual lack of pubescence. The inflorescences are
at first borne on very short peduncles, but these
elongate during anthesis and fruiting. This species
resembles Morinda royoc (flowers fused at the base),
Psychotria erecta (blue fruits), and Alibertia gar-
apatica (terminal sessile inflorescences). Appunia
seibertii Standley of Panama has cuneate-decur-
rent leaf bases. It is possible that the few collections
from near Liberia represent disjunct individuals
and not well-established populations.

Arcytophyllum Willdenow ex Schultes

REFERENCE P. Mena V., Revision of the genus

Arcytophyllum (Rubiaceae, Hedyotideae). Mem.
New York Hot. Gard. 60: 1-26. 1990.

Shrubs or small subshrubs, stems woody, erect or
prostrate, usually with short internodes and congested
leaves, nodes thickened; stipules united and interpetio-
lar, entire to bifid or setose distally, persisting. Leaves
opposite, small, often closely crowded and imbricate,
sessile or subsessile; leaf blades entire, thick-coriaceous,
glabrous, venation often obscure, domatia absent. Inflo-
rescences terminal (sometimes apparently axillary to dis-
tal leaf-like bracts), with cymose or clustered flowers on
short peduncles, or of solitary flowers, pedicels short.
Flowers bisexual, glabrous externally; hypanthium hemi-
spheric to obovoid, calyx lobes 4(-5), often with glands
between the lobes; corolla campanulate to funnelform,
corolla lobes 4, often papillate-puberulent within, val-
vate in bud; stamens 4, free portion of the filament
emerging from between the corolla lobes; anthers dor-
sifixed, exserted or partly included; ovary 2-locular, pla-
centas borne on the septum, ovules 4-12/locule, style
slender, stigmas 2. Fruits capsular, turbinate to subglo-
bose, usually dehiscing septicidally and basipetally,
2-locular; seeds few, oblong and plano-convex to con-
cave-convex, punctate.

A genus of 1 5 species, ranging from Costa Rica
through Panama into the Andes as far south as
Bolivia. These plants are distinguished by their
small stiff ericoid leaves, short internodes, and
small stature in paramo or similar open high-al-
titude vegetation types. Standley (1938, p. 1273)
suggested that the genus might be congeneric with
Houstonia.

Key to the Species of Arcytophyllum

la. Plants shrub-like, rooting only at the base, with many erect branching stems to 70 cm tall; leaves
4-8 mm long; flowers in cymose groups on short peduncles A. lavarum

Ib. Plants prostrate with main stems rooting at the nodes, short erect stems less than 20 cm tall; leaves
3-6 mm long; flowers solitary on short leafy stems A. muticum

Arcytophyllum lavarum K. Schum. ex Standl.,
Contr. U.S. Natl. Herb. 18: 127. 1916. Mallos-
toma lavarum (K. Schum.) J. D. Smith, Enum.
PI. Guatem. 5: 36. 1899, nom. nud. (based on
A. lavarum K. Schum. in herb.). A. chirropoense
Suesseng., Bot. Jahrb. Syst. 72: 285. 1942. Fig-
ure 1.

Subshrubs with creeping and erect woody stems, 10-
40(-70) cm tall, sometimes forming mats, with many
erect branches, nodes thickened with the bases of per-
sisting stipules and leaf bases, internodes 2-7(-18) mm
long, leafy branchlets 0.5-1.5 mm thick, glabrous, with
4 longitudinal ridges, becoming silvery gray to black;

stipules 1-2.5 mm long, distal margin entire to erose or
spiny, thickened at the base. Leaves sessile or with pet-
ioles ca. 1 mm long, glabrous throughout, articulate at
the base; leaf blades 4-8 mm long, 24 mm broad, ovate-
elliptic to ovate-oblong or oblong, apex obtuse or round-
ed, base obtuse to subtruncate, drying thick and coria-
ceous, darker and lustrous above, with a rim of lustrous
tissue along the edge beneath, midvein impressed above,
other veins not visible above or below. Inflorescences
to 2 cm long, usually with peduncles to 1.5 cm long,
branches of the inflorescence sometimes subtended by
leaf-like bracts, glabrous, flowers usually in cymose
groupings (fasciculate), pedicels 0.5-2 mm long. Flowers
ca. 7 mm long, hypanthium 1-1.5 mm long, obconic
(turbinate) to hemispheric, calyx lobes 4, 1-1.5 mm long,
ovate-oblong to triangular and persistent, often with 1-

FIELDIANA: BOTANY

3 setae between each pair; corolla tinged with blue, pur-
ple, or pink in bud, campanulate, corolla tube 2-3 mm
long, corolla lobes 2-3 mm long, white and minutely
puberulent on the inner surfaces; stamens 4, filaments
ca. 1.5 mm long, attached near the mouth of the tube,
anthers 0.8-1 mm long, purple; ovary with ovules borne
together on a stipe from the base of the septum, style ca.

4 mm long, stigmas 2 and often connate. Fruits short-
pedicellate, 1.5-2 mm long, subglobose, with a ring of
tissue and the persistent sepals distally; seeds 4-8/locule,
ca. 1 mm diam.

Small shrubby or mat-forming plants of open
or partly shaded sites in Paramo formations and
open high elevation sites, from (1800-)2500 to
3500 m elevation. They have also been found as
pioneers on volcanic substrates at 800-900 m el-
evation in the Cordillera de Guanacaste. Flower-
ing throughout the year (mostly in January-March
and July-August). The species ranges eastward
from Volcan Rincon de la Vieja to the Chiriqui
highlands of Panama.

Arcytophyllum lavarum is distinguished by its
short shrubby habit, short internodes with thick-
ened nodes, small stiffopposite ericoid leaves, and
four-parted campanulate flowers with corolla lobes
bluish or purple on the outer (abaxial) surfaces and
white on the inner (adaxial) surfaces. This species
and its congener differ from all our other Rubi-
aceae in habit and appearance with their small
thick leaves, miniature shrubby form, and exposed
high -elevation habitat. These plants often grow
among similar-looking species of Hypericum
(Guttiferae, yellow flowers with many stamens),
Ugni myricoides (H.B.K.) Berg (Myrtaceae, lack-
ing interpetiolar stipules), and Ericaceae (alternate
leaves). References to a published description by
Schumann are incorrect.

Arcytophyllum muticum (Wedd.) Standl., J. Wash.
Acad.Sci. 18: 163. 1 928. Hedyotis mutica Wedd.,
Chloris Andina 2: 43, pi. 50. 1857. A. recur-
vatum Suesseng., Bot. Jahrb. Syst. 72: 286. 1942.
Figure 1.

Small prostrate subshrubs, 3-10(-20) cm tall, often
forming short dense mats 5-10 cm thick, usually rooting
from the nodes on thicker horizontal stems, much-
branched, the erect leafy flowering stems without roots,
internodes 0.2-6 mm long; stipules ca. 0.5 mm long,
glabrous or with a few hairs distally, near the base and
on lines beneath the stipule. Leaves sessile, usually close-
ly spaced, glabrous throughout; leaf blades 3-5(-6) mm
long, 0.5-2 mm broad, lanceolate to linear-lanceolate or
narrowly oblong, apex acute to obtuse, base cuneate,
drying thick-coriaceous and with similar color above and
below, venation obscure. Inflorescences of solitary flow-

ers terminal on short leafy branchlets, borne on slender
peduncles (pedicels) 2-4 mm long or sessile. Flowers to
12 mm long and 7 mm broad, hypanthium ca. 1 mm
long, calyx lobes 1-2 mm long, narrow, corolla campan-
ulate-funnelform, 5-8 mm long, white with purple or
lilac on the outer surfaces, corolla tube 3-4.5 mm long,
corolla lobes 2-4 mm long and 1-2 mm broad, papillate-
puberulent on the lower half within (adaxially); anthers
borne just beneath the sinuses of the corolla lobes. 0.7-
0.8 mm long. Fruits 1-1.5 mm diam., subglobose, with
4-6 seeds per locule.

Small moss-like plants of paramo vegetation and
in bogs and along open slopes in high montane
formations, from 2700 to 3400 m elevation. Flow-
ering in January, March, and July-August in Costa
Rica. The species is found in the Cordillera de
Talamanca of Costa Rica and adjacent highlands
of Panama, to Colombia, Ecuador, and Venezuela.

Arcytophyllum muticum is distinguished by its
short moss-like habit, very small stiff narrow op-
posite leaves, and woody stems with short inter-
nodes and interpetiolar stipules. The four-parted
flowers and inferior ovary help distinguish these
plants from similar species of Ericaceae and Hy-
pericum. We have only seen six collections from
Costa Rica. The diminutive size may cause many
collectors to overlook this species.

Bathysa Presl

Trees or shrubs, often puberulent; stipules interpetio-
lar, entire, acute to bifid at apex, deciduous or persisting.
Leaves opposite, petiolate; leaf blades entire and pin-
nately veined, domatia absent. Inflorescences terminal
and solitary, paniculate with opposite branching, often
much-branched with many small flowers. Flowers bi-
sexual, often small, calyx cupular and truncated distally
or with 4-5 calyx lobes; corolla funnelform to subrotate,
corolla lobes 4-5; stamens 4-5, inserted on the throat of
the tube, anthers dorsifixed and exserted; ovary 2-locu-
lar. ovules many in each locule. Fruits capsular, 2-locular
with septicidal dehiscence, splitting from apex into 2
valves; seeds horizontal, compressed or angular, with or
without marginal wings.

A genus of about 1 2 species, nearly all from
eastern Brazil or Amazonia. The lack of intrapeti-
olar stipules distinguishes these plants from Elae-
agia, while the short corolla tubes and slightly ex-
serted stamens separate it from Rondeletia.

Bathysa veraguensis Dwyer, Ann. Missouri Bot.
Gard. 67: 40. 1980.

Small trees to 5 m tall, leafy branchlets 2.5-6 mm
thick, minutely appressed-puberulent with yellowish hairs

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

0.2-0.4 mm long, terete; stipules 22-32 mm long, 2-6
mm broad, narrowly oblong to falcate, densely sericeous
with lustrous ascending yellowish hairs. Leaves with pet-
ioles 4-16 mm long, 2-2.8 mm thick, densely puberu-
lent; leaf blades 12-36 cm long, 9-18 cm broad, obovate
to broadly oblanceolate or oblong, apex short- or long-
acuminate, tip to 1 8 mm long, base gradually narrowed
to obtuse but often abruptly rounded at the petiole, dry-
ing chartaceous and brown or reddish brown, minutely
(0. 1-0.3 mm) puberulent above and below, 2 veins 1 2-
20/side. Inflorescences 1 5-30 cm long, 1 2-22(-30) cm
broad, open paniculate with a larger pair of lateral
branches and much smaller distal branching, peduncles
5-9 cm long, 2-3.5 mm thick, densely sericeous with
ascending hairs, pedicels 6-12 mm long, usually with
bracteoles 3-4 m long in the middle, flowers 1-3 in distal
cymules. Flowers with hypanthium ca. 3 mm long and
3 mm diam. distally, conical, densely sericeous together
with the calyx, calyx lobes 5 (4), 3-5 mm long, 3-4 mm
broad at the base; corolla white, glabrous on the exterior,
tube 3-5 mm long, to 5 mm diam.; anthers 5, 3-4 mm
long. Fruits 8-15 mm long to 8 mm broad (including
the large persisting calyx lobes), ellipsoid-cupulate from
a narrow (0.7 mm) pedicel, densely sericeous.

Plants of the evergreen Pacific lowlands of the
Osa Peninsula, collected at 400 m elevation. Flow-
ering material was collected in February in Pan-
ama; old fruits were collected in June in Costa
Rica (Hammel et al. 17029 CR, MO). This species
is known only from southern Costa Rica and Code
and Veraguas provinces in Panama.

Bathysa veraguensis is recognized by its often
larger puberulent leaves with many secondary
veins, large open terminal panicles with frequent
distal dichotomous branching, larger distant flow-
ers, white corollas glabrous on the exterior, and
sericeous capsules with broad persisting calyx lobes.
Leaf shape and pubescence appear to vary consid-
erably, making it likely that the single Costa Rican

collection (cited above) and the Panamanian type
(Lao & Gentry 531 MO) are conspecific.

Bertiera Aublet

Shrubs or small trees, branchlets terete, glabrous or
puberulent; stipules connate both interpetiolar and in-
trapetiolar and forming a short sheath above the node
(often difficult to see or interpret), interpetiolar portion
triangular and acute, persistent. Leaves opposite, disti-
chous, petiolate or rarely sessile; leaf blades entire, dry-
ing chartaceous, domatia present or absent. Inflores-
cences solitary and terminal, pedunculate panicles with
a prominent central rachis and opposite or alternate lat-
eral branches bearing flowers in cymose or helicoid (cin-
cinus-like) arrangements, bracts narrow, flowers often
sessile. Flowers bisexual, small, white or greenish white,
hypanthium turbinate to subglobose, entire distally or
with 5-6 small persisting calyx lobes; corolla funnelform,
corolla tube narrow, usually strigillose externally, gla-
brous or puberulent on the throat within, corolla lobes
5 (4, 6), short, convolute in bud; stamens 5 (4, 6), inserted
on the distal part of the corolla tube, filaments very short,
anthers dorsifixed, often with the connective slightly pro-
longed, included in the throat; an ovarian disc or annular
ring present; ovary 2-locular, placentas borne on the sep-
tum, with many ovules in each locule, style slender and
glabrous, stigma simple or 2-lobed. Fruits berries, glo-
bose to ellipsoid, purple or black; seeds many, small,
angular, foveolate or granular.

A genus of perhaps 30 species, found in the
American tropics and in Africa. The genus is dis-
tinguished by its unusual stipules, thyrse-like in-
florescences, and many-seeded fleshy fruit. These
plants resemble some species of Psychotria (but
those have two-seeded fruit) and some species of
Gonzalagunia and Rondeletia with cymose-heli-
coid branching.

Key to the Species of Bertiera

la. Leaves with petioles 1-4 mm long, with 5-8 strongly ascending major secondary veins on each side;

stipules 7-14 mm long; Cocos Island and Panama B. angustifolia

Ib. Leaves with petioles 3-9 mm long, with 4-6 major secondary veins on each side; stipules 5-8 mm

long; wide ranging continental B. guianensis

Bertiera angustifolia Benth., Bot. voy. Sulph. 103.
1845. Figure 43.

Shrubs or small trees, 3-6 m tall, leafy branchlets 1 .2-
4 mm thick, with appressed-ascending sericeous hairs
0.7-1.8 mm long, internodes often uniform (ca. 2-3 cm)
in length; stipules 7-20 mm long, 1.5-2.7 mm broad at

the base, with a narrow tip, persisting or deciduous. Leaves
with petioles 1-3.5 mm long, sericeous with appressed-
ascending hairs; leaf blades 9-17 cm long, 2-3.5(-5) cm
broad, lanceolate to very narrowly ovate-elliptic, apex
gradually narrowed and acute or acuminate, base acute
to obtuse or slightly rounded, drying dark, glabrous on
the upper surface except for the midvein, sericeous on
the veins beneath, 2 veins 5-7/side and strongly as-

FIELDIANA: BOTANY

cending, with minute tufted domatia in the leaf axils
beneath. Inflorescences 10-18 cm long, 3-6 cm broad,
peduncles 4-10 cm long and often pendulous, lateral
branches 1.5-3 cm long and alternate, with straight as-
cending hairs ca. 0.6 mm long, bracts 5-13 mm long,
linear, distal bracteoles ca. 1 mm long, flowers usually
sessile. Flowers 6-7 mm long, hypanthium 1-1.5 mm
long, sericeous, calyx lobes 4 or 5, 0.2-0.5 mm long,
acute; corolla white, sparsely pubescent, tube 2-3 mm
long, 0.7 mm diam., lobes 5 (rarely 4), 1.3-2 mm long;
stamens 5, anthers 1-1.5 mm long. Fruits ca. 10 mm
diam., mostly sessile, drying black and with 10 longi-
tudinal ribs (not always apparent at maturity), glabres-
cent.

Plants of moist evergreen lowland forest for-
mations, from near sea level to 500 m elevation.
Flowering in February and April on Cocos Island;
fruiting in February. This species is known only
from Cocos Island and Panama.

Bertiera angustifolia is recognized by its narrow
leaves, terminal panicles with mostly sessile flow-
ers on helicoid lateral branches, and unusual stip-
ules. This species may be no more than a variant
of B. guianensis, but the narrower leaves with more
strongly ascending veins do give the Cocos Island
plants a rather distinctive appearance.

Bertiera guianensis Aubl., Hist. pi. Guiane 1: 180,
pi. 69. 1775. Figure 43.

Shrubs or small trees, l-6(-10?) m tall, leafy branch-
lets 0.9-4.5 mm thick, with appressed-ascending hairs
ca. 0.4 mm long, stems becoming glabrescent, internodes
often quite uniform (4-5 cm) in length; stipules 5-15
mm long, 3-4 mm wide at the base, basal sheath 3-4(-6)
mm long (above the node), acuminate (rarely slightly
bifid). Leaves often distichous, petioles 3-10 mm long,
0.8-1.8 mm thick, strigulose; leaf blades 8-18(-21) cm
long, 2-6(-8) cm broad, narrowly oblong to narrowly
elliptic-oblong or lanceolate, apex gradually narrowed
and acute or acuminate, base gradually cuneate to ob-
tuse, drying chartaceous and dark olive green to grayish,
glabrous above or with a few hairs on the midvein,
sparsely strigillose with hairs 0.4-0.8 mm long beneath
(the hairs on the veins longer), 2 veins 4-6 (3-8)/side,
arcuate-ascending. Inflorescences 8-24 cm long, often
pendant, lateral branches 1-5 cm long, lower branches
longer and with more secondary branching (pyramidal),
peduncles 2-10 cm long, 0.7-1 .5 mm thick, densely strig-
ulose with stiff whitish ascending hairs 0.5-1 mm long,
bracts 3-9(-15) mm long, triangular to linear, flowers
sessile or subsessile. Flowers ca. 8 mm long, hypanthium
0.7-1.8 mm long, pubescent, calyx lobes 5-6, 0.3-1 mm
long; corolla white, tube 3-5 mm long, 1-2 mm wide
with short stiff ascending hairs or glabrescent, corolla
lobes 5, 1.5-3 mm long, ovate-oblong and acute, pu-
berulent within; stamens 5-6, anthers 0.8-1.8 mm long,
the connective prolonged 0.3-0.6 mm long, sagittate at
the base; ovary with resinous dots, style ca. 2.5 mm long,

stigmas bifid and oblong, ca. 2 mm long. Fruits sessile,
subglobose, 3-8 mm diam., with 6-10 longitudinal ribs,
blue drying black; seeds 1-2 mm long, muricate.

Shrubs and small trees of wet evergreen lowland
forest formations, from near sea level to ca. 1000
m elevation. Probably flowering and fruiting
throughout the year (most flowering collections
from January to August). The species ranges from
Mexico, Central America, and the western Greater
Antilles to Bolivia and the Guianas.

Bertiera guianensis is characterized by its nar-
row leaves, unusual stipules, characteristic pubes-
cence, thyrse-like inflorescences with sessile flow-
ers often on helicoid distal branches, and 1 0-ribbed
immature fruit. This species is usually found on
ridges in primary forest at La Selva.

Borojoa Cuatrecasas

REFERENCE J. Cuatrecasas, Borojoa, Nuevo
genero Rubiacea. Revista Acad. Colomb. Ci. Ex-
act. 7: 474-477. 1950.

Small trees, dioecious, glabrous; stipules interpetiolar
and sometimes intrapetiolar with a short sheath above
the node and with 2 large free interpetiolar lobes pro-
duced above the basal sheath, usually persisting. Leaves
opposite and decussate, often large, petiolate; leaf blades
entire, domatia sometimes present. Inflorescences soli-
tary and terminal, subtended by 1-3 pairs of bracts re-
sembling the stipules, <5 flowers cymose or sessile in a
congested head of few to many flowers, 9 flowers usually
solitary. Flowers unisexual and differing in form, $ flow-
ers 4- or 5- (to 8-) parted, corolla usually funnelform,
puberulent on both inner and outer surfaces, corolla lobes
convolute in bud, stamens 5, anthers linear; 9 flowers 6-
8-parted, ovary 6-8-locular, placentation axile, ovules
many in each locule, stigmas 6-8. Fruits berry-like, large,
subglobose, pericarp usually thick-walled and fleshy, in-
dehiscent; seeds imbedded in a mucilaginous pulp, at-
tached horizontally in longitudinal rows, flattened.

Borojoa is a genus of about 10 species occurring
in Costa Rica, Panama, Colombia, and Venezuela.
The genus is distinguished by the solitary and ter-
minal female flowers and fruit, and the male flow-
ers terminal and sessile or in solitary heads. The
larger leaves, unusual stipules, larger than average
flowers, and fruits with thick pericarp are also dis-
tinctive. These rarely collected trees of evergreen
lowland rain forests are not well understood. It is
not clear at this time whether our species are pe-
ripheral elements of other species or distinct spe-
cies deserving recognition (see below). The fruits
are used in Choco, Colombia, to make a refreshing
drink.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Key to Two Putative Species of Borojoa

la. Leaves essentially glabrous, drying chartaceous to subcoriaceous, often elliptic-ovate, major sec-
ondary veins 8-12 pairs; fruits 6-10 cm diam., glabrous B. panamensis

Ib. Leaves glabrous to pubescent beneath, drying thin-chartaceous, usually broadly elliptic, major sec-
ondary veins 6-9 pairs; fruit 3-6 cm diam., densely velutinous or glabrescent on the outer surface

. B. atlantica

Borojoa atlantica Dwyer, Ann. Missouri Bot. Card.
67:46. 1980. Figure 26.

Trees to 10 m tall, leafy stems 3-5, thick, glabrescent
or densely pubescent with soft erect hairs ca. 0.5 mm
long, terete; stipules ca. 10 mm long, 5 mm diam., with
a basal sheath 2-5 mm long and a free distal portion
triangular with acuminate apex, persisting with the leaves.
Leaves with petioles 10-26 mm long, 1.5-2.5 mm thick,
densely pubescent to glabrescent; leaf blades 1 2-26 cm
long, 7-16 cm wide, broadly elliptic to broadly elliptic-
obovate, apex short-acuminate, base obtuse, drying thin-
chartaceous to chartaceous and usually dark brown or
dark greenish brown, glabrous to sparsely pubescent
above, minutely puberulent to velutinous on the veins
beneath with hairs ca. 0.5 mm long, 2 veins 7-1 I/side,
with tufts of hairs in the vein axils. Inflorescences not
seen. Fruits 27-60 mm long, globose to slightly obovoid,
minutely velutinous, subtended by bracts ca. 5 mm long
and 6 mm broad.

Plants of the wet Caribbean lowlands, 0-300 m
elevation. The Costa Rican material was collected
in fruit in June. The species is known from Costa
Rica and Panama, but its circumscription is not
yet certain. The broad leaves velutinous on the
veins beneath (in our material) are distinctive, but
the original description states that these plants may
be almost glabrous.

Borojoa panamensis Dwyer, Phytologia 17: 446.
1968. Figure 26.

Trees 4-13 m tall, trunks to 25 cm dbh, leafy inter-
nodes 3-8 mm thick, essentially glabrous, drying brown;
stipules 12-28 mm long, 5-16 mm broad, united above
the node for 2-8 mm, stiff and longitudinally striate,
acuminate. Leaves with petioles 13-30 mm long, 2-4
mm thick, with 2 lateral adaxial ridges, glabrous; leaf
blades 13-27(-38)cm long, 7-14(-17)cm broad, elliptic-
oblong, to elliptic-obovate or broadly elliptic, apex usu-
ally acuminate, base obtuse to acute (sometimes slightly
decurrent on petiole), drying stiffly chartaceous to sub-
coriaceous and grayish green, glabrous above and below
but with small tufted domatia in vein axils beneath, 2
veins (5-)7-12/side, 3 veins weakly subparallel. Inflo-
rescences of 2-9 terminal sessile $ flowers (9 flowers prob-
ably solitary), subtended by a pair of stipules ca. 10 mm
long. Flowers with hypanthium and calyx tube not dif-
ferentiated, ca. 8 mm long and 6 mm diam. at apex.

subglabrous and drying dark, calyx lobes 0.4-1 mm long;
corolla white, sericeous with downward-pointing lus-
trous hairs, corolla tube ca. 12 mm long, 4 mm diam.
near apex, corolla lobes 5-6, ca. 10 mm long, triangular
and acute. Fruits 5-1 1 cm long, 6-10 cm diam., subglo-
bose, the surface smooth, glabrous and yellowish brown,
umbonate at apex, persisting calyx tube ca. 4 mm high,
outer wall 8-15 mm thick; seeds 4-8 mm broad, 2-3
mm thick, angular or rounded.

Trees of evergreen forest formation, from near
sea level to 600(-1500) m elevation. Flowering in
March and May; fruiting in January-August and
November. The species ranges from northern Cos-
ta Rica (in the Caribbean lowlands) to Panama.

Borojoa panamensis is distinguished by its gla-
brous (except for the domatia) stiff leaves, dis-
tinctive stipules, sessile terminal flowers with gla-
brescent calyx, sericeous corolla, and the large
solitary terminal globose fruit. A specimen from
1 500 m on Cerro Turrubares (Q. Jimenez 836 CR)
is disjunct as regards both elevation and coming
from the Pacific slope. New collections are pro-
viding a better overview of variation within this
species but more material is needed. Herbarium
specimens can be very similar to Genipa ameri-
cana, but the latter have pedunculate inflores-
cences and short corolla tubes and the stipules lack
prominent parallel venation.

Borreria G. F. W. Meyer

Borreria G. F. W. Meyer is here considered part
of Spermacoce.

Bouvardia Salisbury

REFERENCE W. H. Blackwell, Jr., Revision of
Bouvardia. Ann. Missouri Bot. Gard. 55: 1-30.
1968.

Shrubs, subshrubs or perennial herbs; stipules inter-
petiolar, with a very short sheath united to the petioles,
entire or with 1 -several slender teeth or awns. Leaves
opposite or in whorls of 3-4(-6), usually short-petiolate

FIELDIANA: BOTANY

and puberulent, entire, domatia absent. Inflorescences
terminal, usually solitary, cymose to corymbose or sub-
capitate (rarely of solitary flowers). Flowers bisexual, di-
morphic, glabrous or puberulent on the exterior, calyx
lobes 4(-5), usually lanceolate, persisting; corolla long-
tubular to salverform, usually more than 20 mm long,
white to yellow, red, pink, or purple, corolla lobes 4,
valvate in bud; stamens 4, borne above the middle of
the corolla tube, anthers linear or oblong, sessile and
included in pin flowers, with filaments and exserted in
thrum flowers; ovary 2-locular, ovules many on a peltate
placenta borne from the lower part of the septum, style
1 , slender, exserted in pin flowers and included in thrum
flowers. Fruits capsular, globose or obovate, didymous-
globose, dehiscing at first loculicidally, then septicidally;
seeds many and vertically imbricate, with entire wings.

Bouvardia contains about 35 species, primarily
Mexican and Guatemalan but ranging to Nicara-
gua. It seems probable that the few specimens col-
lected in Costa Rica and Panama over the last 100
years represent escaped cultivated material.

Bouvardia glabra Polak., Linnaea 41: 565. 1877.

Ornamental shrubs, usually 1-1.5 m tall, leafy stems
0.6-3 mm thick, terete, pubescent or glabrescent; stipules
2-4 mm long, with a short (0.5 mm) base and slender
awn, minutely puberulent. Leaves opposite, petioles 1-
5 mm long; leaf blades 2.5-5(-10) cm long, 0.7-1. 8(-3)
cm broad, narrowly ovate-elliptic to lanceolate, apex ta-
pering gradually and acute, base obtuse, drying dark
brown above and much paler beneath, minutely pubes-
cent beneath, 2 veins 4-6/side, ascending. Inflorescence
4-8 cm long, terminal or axillary to distal leaves, with
(l-)3-9(-18) flowers, pedicels 3-7 mm long. Flowers with
hypanthium ca. 1.5 mm long, calyx with unequal lobes
3-8 mm long, 0.4-2 mm broad, glabrous or sparsely
puberulent; corolla glabrous, white, tube 1.5-3 cm long,
1.5-3 mm diam., lobes 4, ca. 5 mm long.

Ornamental plants not known to grow wild in
Costa Rica. The type (Polakowsky 337 photo B &
fragment F) was collected in Costa Rica. Blackwell
recognized B. glabra, but Williams (Standley &
Williams, 1975, p. 26) considered it to be a syn-
onym of B. longiflora (Cav.) H.B.K. We have seen
only two collections, both from gardens: Brenes
24418 (16) CR and M. Valeria 33 F. Note: The
latter has more than 18 flowers in the inflores-
cence. Common names arejazmin andjazmin de
la virgen.

Calycophyllum DeCandolle

Trees, often attaining a large size, branchlets terete;
stipules united and interpetiolar, caducous. Leaves op-
posite, petiolate; leaf blades entire, pinnately veined. In-

florescences terminal (lateral branches apparently axil-
lary when subtended by distal leaves), corymbiform
panicles, often many-flowered, pedunculate, flowers ses-
sile or short-pedicellate, at first completely enclosed within
close-fitting membranous (perianth-like) bracts. Flowers
bisexual, radially symmetrical except when the calyx de-
velops a single large petaloid structure; hypanthium ob-
long to obconic, terete, calyx lobes minute, absent, or 1
developed into a large petiolate and petal-like blade;
corolla short funnelform to campanulate, radially sym-
metrical, corolla tube short, villose within the upper part,
corolla lobes 4-8, broad, imbricate or contorted in bud,
with 1 lobe exterior; stamens 4-8, borne on the corolla
tube, filaments slender, anthers oblong, versatile, ex-
serted; ovary 2-locular, placentas borne on the septum,
with few or many ovules in each locule, ovules imbricate
and ascending, style slender and glabrous, stigmas 2,
linear-oblong. Fruits a capsule, oblong-cylindrical, trun-
cated apically , septicidally 2-valved, coriaceous or slight-
ly woody; seeds few to many, the testa expanded and
wing-like at both ends.

A genus of seven or eight species in the West
Indies and northern South America, with one spe-
cies ranging through Central America to Mexico.
The development of a large whitish petal-like
structure from the distal edge of an otherwise trun-
cated calyx in some flowers distinguishes this ge-
nus, but not all flowers have this structure. The
hard wood, height of the trees, and bi valvate cap-
sule are also distinctive.

Calycophyllum candidissimurn (Vahl) DC., Prodr.
4: 367. 1830. Macrocnemum candidissimurn
Vahl, Symb. 2: 38, pi. 30. 1791. Figure 16.

Trees (rarely shrubs), (2-)5-18(-28) m tall, bark red-
dish brown and often stripping off in longitudinal strips,
leafy branchlets 1-4 mm thick, glabrous or puberulent;
stipules 5-10 mm long, 2-3 mm broad, ovate-lanceolate,
caducous and exposing a ring of stiff colleters ca. 1 mm
long at the node. Leaves with petioles (4-)8-22(-30) mm
long, glabrous or puberulent; leaf blades 4-10(-13) cm
long, 1 .5-7(-8) cm broad, broadly elliptic-ovate to broadly
elliptic or broadly obovate, apex abruptly narrowed and
short-acuminate (obtuse), base cuneate and decurrent on
petiole, drying chartaceous and brown, glabrous above
and glabrous between the major veins beneath, 2 veins
4-7/side, often with minute pits and tufts of hairs (doma-
tia) in vein axils beneath. Inflorescences corymbose to
broadly cymose in form, often flat to broadly rounded
distally. (2-)5-12(-20) cm long, often with 3 major pe-
duncles from the end of the stems, the lateral peduncles
sometimes subtended by smaller leaves (and appearing
to be axillary), minutely puberulent or glabrescent, flow-
ers in small compound dichasia with a central sessile
flower, young flowers enclosed in glabrous calyptrate
bracts 5-10 mm long. Flowers 5-9 mm long, hypanthi-
um obconical, 2-3.5 mm long, 1.5 mm diam., glabrous
or puberulent, often with glandular dots, calyx lobes usu-
ally absent, some flowers with a petiolate (clawed) petal-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

like blade 2-4 cm long and 1.5-3.5 cm broad, the blade
suborbicular or reniform to broadly obovate, rounded
distally, obtuse to truncate or subcordate at the base,
white or pale greenish white, palmately veined, the pet-
iole-like base 1-2.5 cm long; corolla 5-7 mm long, white,
campanulate to funnelform, corolla tube 2-3.5 mm long,
ca. 1.5 mm broad at the base and 3 mm broad distally,
often densely villose at apex of the throat with erect hairs
ca. 1 mm long, lobes 4, 3-4.5 mm long, 2-2.5 mm broad,
becoming reflexed; stamens 4, filaments 1.5-2.5(-3.5)
mm long, anthers 1.2-1.5 mm long; style 3-5 mm long,
stigmas 1.3-2 mm long. Fruits (6-)8-12 mm long, 3-4
mm diam., oblong-cylindrical, sessile or subsessile, gla-
brous or sparsely puberulent, with 8 longitudinal ribs;
seeds 3-5 mm long, fusiform with wings at 2 ends, body
of the seed ellipsoid, 1-1.5 mm long.

Conspicuous trees of deciduous and partly de-
ciduous forest formations in the Pacific lowlands,
from near sea level to about 450 m elevation (to
700 m elsewhere). Flowering in November-Feb-
ruary and May; fruiting in January-August. The
species ranges from central Mexico, Belize, and
Guatemala, along the Pacific slope of central and
southern Central America to Colombia and Ven-
ezuela; it also occurs in the West Indies.

Calycophyllwn candidissimum is recognized by
the bright whitish petaloid structures developed
from the calyx of some flowers. The trees bear
many inflorescences over their crowns, and the
bright petal-like sepal lobes make a striking visual
display when in full flower. The large size of these
trees in deciduous woodland also contributes to
the effect. The species has been called madrono,
salamo, and surra in Costa Rica. The wood is hard
and highly elastic and fine textured and finishes
smoothly; it has been used for tool handles, ar-
chery bows, and many other purposes (Standley,
1938).

Cephaelis Swartz

A poorly defined genus of about 100 species in
the American tropics and southern Asia. The ge-

nus was distinguished by the involucrate heads of
flowers, two-locular ovary with solitary basal
ovules, and drupaceous fruits with two nutlets.
Most authors now agree that the species of Cepha-
elis are polyphyletic and cannot be clearly segre-
gated from Psychotria (Taylor et al., 1 99 1 , p. 1 39).
See the treatment of Psychotria (key 3) for species
formerly placed in Cephaelis, and Figures 1 7 and
18.

Chimarrhis Jacquin

Trees, often growing to large size and with buttressed
trunks; stipules interpetiolar and intrapetiolar, leaving a
scar encircling the stem above the node (and above the
petiole attachment), caudate to acuminate, persistent or
caducous. Leaves opposite, often clustered at the ends
of twigs, short-petiolate; leaf blades large- to medium-
sized, sometimes with domatia (absent in our spp.). In-
florescences solitary or paired in leaf axils (rarely pseu-
doterminal), paniculate and often corymb-like in form,
flowers in open cymose groupings, bracts present. Flow-
ers bisexual and radially symmetrical, monomorphic,
protogynous in Costa Rica, hypanthium cupulate to tu-
bular, truncated to dentate distally, calyx lobes 5 (4) and
very small or none; corolla funnelform, white, corolla
tube short and broad, villous within, corolla lobes 5 (4),
valvate in bud; stamens 5 (4), borne on the throat of the
corolla tube between the corolla lobes, filaments slender
and villous at the base, anthers dorsifixed, often exserted;
ovary 2-locular, placentation peltate on the septum, ovules
many in each locule, style short, stigmas 2, obtuse. Fruits
capsular, small and woody, oblong, dehiscing septici-
dally from apex and 2-valved; seeds many, compressed
or angulate, horizontal, testa reticulate.

A genus of about 14 species ranging from Costa
Rica into South America and in the West Indies.
Chimarrhis is recognized by the axillary inflores-
cences, small flowers with poorly developed calyx
lobes, corolla tube villous within, and small
rounded bivalved capsules with many horizontal
seeds. The buttressed trunks, great height of some
individuals, and the stipular scar encircling the
stem above the nodes are also distinctive.

Key to the Species of Chimarrhis

la. Fruit 45 mm long; leaf blades 1 1-24 cm long and 6-1 1 cm broad; evergreen forests of the Pacific

lowlands C. latifolia

Ib. Fruit 1.5-2.5 mm long; leaf blades 5-15 cm long and 3-7.5 cm broad; Caribbean lowlands

C. parviflora

FIELDIANA: BOTANY

( himarrhis latifolia Standl., Publ. Field Columb.
Mus., Bot. Ser. 4: 265. 1929. Figure 37.

Trees to 30 m tall, with high buttresses and yellow
wood, leafy branchlets 4-9 mm thick, glabrous, leaf scars
prominent (ca. 5 mm broad); stipules 2-3(-7) cm long,
4-10 mm broad at the base, acute, glabrous and reddish
brown, subcoriaceous and caducous, stipular scars often
turning dark. Leaves with petioles 18-45 mm long, 1.2-
1.8 mm thick, glabrous; leaf blades 1 1-24 cm long, 6-
1 1 cm broad, broadly elliptic to elliptic-oblong, apex
obtuse or rounded with a bluntly triangular tip 4-8 mm
long (or short-acuminate), base obtuse to cuneate, drying
chartaceous to stiffly chartaceous, dark brown above and
much paler beneath, glabrous above and below except
for small groups of hairs (domatia) in the vein axils
beneath, 2 veins 7-10/side, 3 veins often subperpen-
dicular to the 2. Inflorescences axillary to distal leaves
(2/node), 8-1 6 cm long, 8-1 2 cm broad, corymbose with
a broadly rounded distal aggregation of many flowers,
primary peduncle 3-8 cm long, 2-3 mm thick, reddish
brown and glabrous, branches of the inflorescence op-
posite or subopposite, distal flowers in cymose groups
of 3, flowers sessile or short-pedicellate, pedicels and
distal branches of the inflorescences minutely puberu-
lent. Flowers ca. 8 mm long, protogynous, hypanthium
1.5-3 mm long, turbinate, glabrous and reddish brown
when dry, calyx tube very short (ca. 0.5 mm), entire or
with 5 broad 0-2 mm long lobes; corolla 4-5 mm long,
white, glabrous externally, tube 1-2 mm long, 1.5 mm
broad, lobes rounded; stamens 5, filaments to 4 mm long,
with whitish hairs on the lower half, anthers 0.8-1 mm
long; pistil with a style to 3.5 mm long, stigmas 2, thick,
ca. 0.7 mm long. Fruits 4-5 mm long, 3 mm broad,
obovoid-oblong with truncated apex, glabrous on the
sides, minutely puberulent on the distal (apical) surface;
seeds ca. 1 mm long.

Trees of evergreen lowland rain forest forma-
tions of the Pacific slope of southern Costa Rica,
below 400 m elevation. Flowering in July-August
(Cooper & Slater 260 F, us the type) and October-
December; fruiting in December-January. The
species is known only from the Pacific slope of
southern Costa Rica (Reserva Biologica Carara to
Golfo Dulce) and adjacent Panama.

Chimarrhis latifolia is recognized by the taller
height of the trees, the generally glabrous parts,
large leaves, corymbose inflorescences, closely
clustered small flowers with short corolla tubes,
and small woody bivalved fruit. Yema de huevo
and jagua amarillo are common names reported
for this species. A sterile specimen collected and
determined by Paul Allen (56 1 3) with large (to 44
cm) leaves, short (1-2 cm) petioles, and minute
puberulence on the lower leaf surfaces and on the
long (7 cm) stipules is tentatively placed here. It
may represent a juvenile shoot, though said to

come from a tree 27 m tall. Allen (1956, pp. 170-
1 72) stated that it is an important timber tree, and
he provided an illustration. Note: This species may
be synonymous with C. cymosa Jacq.

Chimarrhis parviflora Standl., Trop. Woods 11:
26. 1927. Figure 37.

Shrubs or trees to 25 m tall, to 60 cm dbh, with soft
bark and low buttresses, wood yellow, leafy branchlets
1.5-4 mm thick, minutely (0.1-0.3 mm) appressed-pu-
berulent and quickly glabrescent, internodes often short
(1-2 cm); stipules 5-18(-30) mm long, 2-4 mm broad
at the base, narrowly triangular to lanceolate, puberulent
at the base and on the outer surface, caducous. leaves
with petioles 1 1-22 mm long, 1-2 mm thick, minutely
appressed-puberulent and glabrescent; leaf blades 5-
15(-18) cm long, 3-7.5 cm broad, elliptic to elliptic-
oblong or elliptic-obovate, apex tapering abruptly and
short-acuminate, gradually narrowed to the cuneate-at-
tenuate base and slightly decurrent on petiole, drying
chartaceous to stiffly chartaceous, usually dark above,
glabrous above, minutely (0.1-0.4 mm) puberulent be-
neath, often densely puberulent on the major veins be-
neath, 2 veins 5-1 0/side. Inflorescences axillary or pseu-
doterminal, 2-4 at a node, 5-12(-15) cm long, 3.5-8 cm
broad, densely many-flowered, peduncles 2-5(-9) cm
long, minutely puberulent, branches opposite or subop-
posite, pedicels 0-2 mm long, bracts absent or minute
(0.5 mm). Flowers 4-6 mm long, with sweet odor, gla-
brous externally, hypanthium 1-1.5 mm long, turbinate,
calyx tube very short, calyx lobes 4-5, ca. 0.3 mm long,
obtuse and ciliate distally; corolla 2-4 mm long, white,
tubular-funnelform, corolla tube 1.5-2 mm long, villous
within near apex, corolla lobes 4, 1-2 mm long, bluntly
rounded; stamens 4, anthers 0.6-0.7 mm long, exserted;
style 2.5 mm long, stigmas 2, broader than long. Fruits
numerous and tightly grouped at the ends of the infruc-
tescence, 1.5-2.5 mm long, obovoid or turbinate, exo-
carp yellow-brown and woody, with longitudinal ribs,
glabrous on the disc-like apex; seeds 0.8-1.2 mm long.

Trees of evergreen rain forest formations of the
Caribbean slope often found in swampy areas and
along stream edges, from 30 to 900 m elevation.
Flowering in March-June; fruiting in May and
July-September. This species is known only from
Costa Rica and Panama.

Chimarrhis parviflora is recognized by its axil-
lary corymbose inflorescences often four at a node
with many small flowers congested distally, and
the small woody bivalvate capsules. The ability to
grow to considerable height and buttressed trunks
are additional distinctions. Galls are sometimes
present in the infructescences and may be mistak-
en for young capsules.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Chiococca P. Browne

Shrubs, woody climbers or small trees, the branches
often pendant or clambering, branchlets terete, glabrous
or puberulent; stipules interpetiolar and slightly intra-
petiolar (to form a very short tube or cup), usually cus-
pidate, persistent. Leaves opposite, petiolate, entire,
membranaceous to coriaceous, pinnately veined, with-
out domatia. Inflorescences axillary or less often ter-
minal, racemose or paniculate, flowers opposite or along
only 1 side of the rachis, pedicels present or absent.
Flowers bisexual and radially symmetrical, usually
5-parted, hypanthium ovoid to turbinate, calyx lobes 4-
6, short and persisting, corolla campanulate to funnel-
form, white to yellow, lavender or purple, corolla tube
cylindrical to urceolate, often with longitudinal ribs in
line with the sinuses between the lobes, glabrous within
at the mouth, corolla lobes 4-5, valvate in bud, spreading
or reflexed; stamens 4-5, inserted near the base of the
tube, filaments pilose at the base, anthers linear, exserted
or included; ovary 2-locular, with 1 ovule pendulous
from apex of each locule, stigmas 1 or 2. Fruits drupa-
ceous, fleshy to leathery, usually white at maturity, lat-
erally compressed and rounded in outline (in Central
America) or oblong-cylindrical when dried, with 2 py-
renes; seeds pendulous and laterally compressed.

A genus of about 20 species, ranging from the
southern United States through Mexico, Central
America, and the West Indies to southern South
America.

Chiococca is recognized by its often pendant
clambering branches, the very short stipules slight-
ly united above the petioles (and usually with an
awn), and the unusual white fruit flattened on op-
posite sides and rounded in outline (in Central
American species). The ribbed and valvate corolla,
the filaments free to the base of the corolla tube,
the two-locular ovary with solitary pendulous
ovules, and the white drupaceous fruits are also
important distinguishing characters.

All our species are wide-ranging and quite vari-
able; this may make them difficult to separate in
the absence of flowers, since the fruit differ little
among the species. In fact, the patterns of variation
are so broad as to suggest that there may be hy-
bridization between the species.

Key to the Species of Chiococca

la. Stamens usually well exserted at anthesis (with the filaments sometimes visible); corolla often
campanulate-urceolate in Costa Rica; inflorescences usually with opposite branching, flowers sessile
or with pedicels to 3 mm long; secondary veins obscure on the undersides of the leaves, petioles
10-30 mm long; (7700-) 1 600-2200 m elevation C. phaenostemon

Ib. Stamens included within the corolla tube or only the tips exserted; corolla usually funnelform;
inflorescences with few alternate or opposite lateral branches, pedicels 1-6 mm long; secondary
veins visible on the lower leaf surfaces, petioles 3-17 mm long; 0-1200(-1500) m elevation ... 2

2a. Hypanthium/ovary with thin erect hairs 0.1-0.3 mm long (rarely glabrous); leaves densely to sparsely
puberulent beneath with thin straight hairs 0.2-0.3 mm long; inflorescences 24 cm long [petioles
1-4 mm long] C. semipilosa

2b. Hypanthium/ovary glabrous or with a minute (0.05 mm) papillate-puberulence; leaves glabrous or
sparsely papillate-puberulent beneath; inflorescences (2-)4-10 cm long 3

3a. Leaves usually ovate-elliptic and drying grayish or greenish, rarely more than 4 cm broad, petioles
3-8 mm long; corolla 4-8 mm broad distally when open, calyx lobes narrow; fruit strongly com-
pressed C. alba

3b. Leaves usually oblong-elliptic and drying dark brown, often more than 5 cm broad, petioles 5-17
mm long; corolla 7-10 mm broad distally when open, calyx lobes broadly rounded or obscure; fruit
only slightly compressed laterally C. pachyphylla

Chiococca alba (L.) Hitchcock, Ann. Kept. Mis-
souri Bot. Gard. 4: 94. 1893. Lonicra alba L.,
Sp. PI. 175. 1753. Figure 36.

Woody climbers, shrubs or less often small trees to 8
m tall and 10 cm trunk diam., distal branches often
pendulous or clambering, distal twigs often opposite and
held perpendicular to the main stems, leafy branchlets
0.7-4 mm thick, glabrous and terete, dark when dried;

stipules 1-5 mm long, the broad basal part 0.5-2 mm
long and slightly (0.5 mm) united above the petioles
(intrapetiolar), with an acuminate or caudate tip 0.5-3
mm long. Leaves distant along the stems, petioles 3-8
mm long, 0.5-1 mm broad, glabrous; leaf blades (2.5-)3-
9(-13)cm long, (l-)1.5-3.8(-6)cm broad, ovate-elliptic,
narrowly ovate, oblong or lanceolate, apex long-acu-
minate (sometimes bluntly acute to short-acuminate),
base obtuse to rounded and slightly decurrent on petiole,
leaves drying stiffly chartaceous to membranaceous, gla-

FIELDIANA: BOTANY

brous above and below or with a few thin hairs ca. 0.2
mm long beneath, 2 veins 3-5/side and weakly loop-
connected distally. Inflorescences mostly axillary, (2-)4-
1 1 cm long, unbranched and racemiform or with few
lateral branches and paniculate, peduncles (0.5-) l-3.5(-7)
cm long, ca. 0.5 mm thick, usually glabrous, bracts 1-
1.5 mm long, narrow, the flowers usually borne along 1
side of the rachis, solitary and separate or in groups of
3, pedicels 1-8 mm long, slender, usually glabrous. Flow-
ers with hypanthium 1-1.8 mm long, flattened laterally
on opposing sides, ellipsoid in outline, glabrous or very
minutely (0.05 mm) papillate-puberulent, calyx tube 0.5-
1 mm long, calyx lobes 0.2-0.6 mm long; corolla fun-
nelform, white to yellowish or rose, usually glabrous ex-
ternally, tube 3-8 mm long, 2-5 mm wide at apex, lobes
5 (4), 3-4 mm long, triangular; stamens 5, included or
slightly exserted, anthers ca. 3 mm long; styles 5-8 mm
long, exserted. Fruits 4-7 mm long, 4-7 mm broad,
rounded-oblong (abruptly rounded at top and bottom)
in outline and flattened laterally on the 2 opposite sur-
faces, white at maturity, persisting calyx ca. 1 mm long
and 1.5 mm diam.

Common clambering shrubby plants along open
forest edges and disturbed areas, in both evergreen
rain forest areas and in seasonally deciduous for-
ests, from sea level to 1300(-1500) m elevation.
Flowering in March-October (mostly June-Au-
gust); fruiting in June-March. The species ranges
from the southernmost United States (Texas and
Florida), through Mexico, Central America, and
the West Indies into tropical South America.

Chiococca alba is recognized by the clambering
stems, the smaller often ovate to lanceolate leaves,
the usually few-branched axillary inflorescences,
funnelform usually yellowish white flowers, and
white flattened fruit with round outline and per-
sisting calyx. Most collections are glabrous, but a
few have minute puberulence on the young stems,
inflorescence, and hypanthium. The disc-like
whitish seeds may have inspired two names used
in Central America for the species: Idgrimas de
Maria and Idgrimas de San Pedro.

Chiococca pachyphylla Wernham, J. Bot. 5 1 : 323.
1913. Figure 36.

Lianas and woody climbers (rarely shrubs?), l-5(-7)
m tall, leafy branchlets 1-5 mm thick, glabrous, terete
and drying dark or grayish; stipules 1-3 mm long, sub-
acuminate to caudate at apex, slightly (0.2-0.5 mm) unit-
ed above the petioles. Leaves well spaced along the stem,
petioles 5-17 mm long, 0.6-1.2 mm broad, glabrous;
leaf blades 6-12 cm long, 3-6 cm broad, oblong-elliptic
to elliptic or ovate-elliptic, apex bluntly acute to short-
acuminate, base acute to obtuse and slightly decurrent
on petiole, drying stiffly chartaceous to subcoriaceous,
glabrous above and below, 2 veins 3-5/side, the sec-
ondaries usually darker than the lower surface and easily

seen. Inflorescences 4-8 cm long, mostly axillary and
with few lateral branches, the distal axes racemose, bracts
1-2 mm long, linear, pedicels 1 .5-4.5 mm long, glabrous.
Flowers glabrous externally, hypanthium 1.5-2 mm long,
calyx tube 0.5-1 mm long, calyx lobes 0. 1-0.4 mm long,
rounded or bluntly triangular, glabrous; corolla funnel-
form, yellowish, corolla tube 5-7 mm long, gradually
expanded to apex and 1-3 mm broad, lobes 2-3.5 mm
long, 1.5-2 mm broad at the base, bluntly acute; stamens
included. Fruits 6-8 mm long, 6-8 mm broad, broadly
ellipsoid-circular to circular in outline, ca. 2 mm thick,
green becoming white, glabrous, persisting calyx ca. 1
mm high and 1.5 mm diam.

Shrubs and climbers of evergreen and deciduous
forest formations, from near sea level to 1 500 m
elevation. Flowering primarily in May-Septem-
ber; fruiting in August-December. The species
ranges from northeastern Mexico to Costa Rica.

Chiococca pachyphylla is recognized by its more
consistently vining habit, stiff usually oblong-el-
liptic leaves, lack of pubescence, racemose inflo-
rescence branches, and flattened white fruit. The
secondary veins on the lower leaf surfaces are much
easier to see than in C. phaenostemon. and the
flowers and fruit appear to be a bit larger than
those of C. alba. In addition, C. pachyphylla has
a number of characteristics that appear to be in-
termediate between C. alba and C. phaenostemon.
Considerable variation in inflorescence and flower
morphology adds to the difficulty.

Chiococca phaenostemonend Schlcctcnd., Linnaca
9: 594. 1834. Figure 36.

Shrubs or small trees (lianas), 3-14 m tall, often with
separate trunks from the base, leafy stems 1.5-6 mm
thick, glabrous, slightly quadrangular at first but becom-
ing terete, older nodes conspicuously thicker than the
internodes; stipules 2-4 mm long, the broad base 1-2
mm long, united around the stem for ca. 0.5 mm, with
a narrow awn 0-2 mm long, the awn often breaking off
to leave a shallow persisting cup at the older node. Leaves
somewhat clustered at the ends of stems, petioles 7-30
mm long, 0.7-1 .5 mm broad, glabrous; leaf blades (4-)6-
13 cm long, (1. 2-) 1.5-4. 8 cm broad, elliptic to elliptic-
oblong, narrowly oblong (rarely elliptic-obovate), apex
gradually tapering and cuneate or acuminate, base ta-
pering gradually and obtuse or acute, decurrent on pet-
iole, leaves drying stiffly chartaceous to subcoriaceous,
glabrous above and below, 2 veins 5-8/side and usually
obscure on the lower surface, weakly loop-connected dis-
tally. Inflorescences axillary or terminal, 3-12 cm long,
paniculate with 3-4 primary branches (and 2-3 of these
with secondary branches) peduncles to 4 cm long, ca. 1
mm thick and sparsely papillate-puberulent, bracts 1-2
mm long, flowers often in cymules, pedicels 0.3-3 mm
long. Flowers with hypanthium 1.2-2 mm long, 0.7-1.3
mm wide, glabrous, calyx tube ca. 0.5 mm long, calyx
lobes 0.5 mm long; corolla usually campanula to to ur-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

ceolate in Costa Rica (less often funnelform), white to
yellowish, glabrous, tube 3-5(-6) mm long, 4-6 mm diam.
at the mouth, lobes 5, 2-4 mm long, 1.5-3 mm broad
at the base; anthers ca. 3 mm long, half to fully exserted.
Fruits 5-6 mm long, 5-6 mm broad, broadly oblong or
broadly obovate in outline, flattened on 2 sides (said to
be thicker and rounded in northern Central America),
persistent calyx 1-1.5 mm long and 1.5 mm diam., ped-
icels to 3 mm long.

Trees of evergreen montane forest formations,
from (?700-)1600 to 2100 elevation (to 2500 m
in Guatemala). Flowering in July-September;
fruiting in July-August and January-February. The
species ranges from northeastern Mexico to the
Chiriqui Highlands of Panama.

Chiococca phaenostemon is characterized by its
highland habitat, larger and campanulate corollas
(in Costa Rica), and the often exserted anthers.
The glabrous often long-petiolate leaves with de-
current base and the secondary veins usually ob-
scure beneath also help to distinguish this species.
While a very distinctive plant in the wild, some
specimens of this species may be difficult to sep-
arate from C. pachyphylla and C. alba. The com-
mon name is chiraquilla.

Chiococca semipilosa Standl. & Steyerm., Publ.
Field Mus. Nat. Hist., Dot. Ser. 22: 279. 1940.
Figure 36.

Shrubs, l-3(-4) m tall, leafy branchlets 1-3 mm thick,
minutely puberulent with thin erect whitish hairs 0.1-
0.2 mm long, soon glabrescent, terete; stipules 2-4 mm
long, broad basal part 0.5-1 mm long, little (0.2-0.5 mm)
united above the petiole, the narrow awn 1-3 mm long,
minutely puberulent. Leaves with petioles 1-4 mm long,
0.5-1 mm broad, with lateral margins continuous with
the lamina margins, minutely puberulent; leaf blades
(l-)3-7.5(-12) cm long, (0.5-)l-2.5(-3) cm broad, nar-
rowly ovate, ovate-lanceolate or ovate-elliptic, tapering
gradually to the acute or acuminate tip, base acute to
obtuse and decurrent on petiole, leaves drying stiffly
chartaceous, glabrous to sparsely puberulent above with
thin whitish hairs 0.2-0.3 mm long, sparsely to densely
soft pubescent beneath with hairs 0.1-0.4 mm long, 2
veins 2-4/side, weakly loop-connected near the distal
margin. Inflorescences axillary, 2-4 cm long, cymose or
racemose with 3-9 flowers (rarely umbellate), peduncles
4-10 mm long, pedicels (0-)l-3(-5) mm long, puberu-
lent. Flowers 5-parted, hypanthium 1-1.5 mm long, 0.7-
1 mm broad, little differentiated from the pedicel, dense-
ly puberulent, calyx tube obscure, calyx lobes 0.5-1 mm
long, acute and drying with little puberulence distally;
corolla cream white to yellowish, broadly funnelform,
usually minutely puberulent externally, tube 4-5 mm
long, 1.5 mm diam. at the base to 3 mm near apex, lobes
1.5-2.8 mm long, 1.5-2 mm broad at the base; stamens
included. Fruits 5-6 mm long, 4-5 mm broad, rounded
in outline and flattened longitudinally, with thin erect

hairs ca. 0.2 mm long, persisting calyx 0.7-1.5 mm long,
1.8 mm broad, drying dark in contrast to the pale fruit.

Shrubs of evergreen and deciduous forest for-
mations, from 200 to 1 600 m elevation. Flowering
in June-July; fruiting in September-January. The
species ranges from Belize and Guatemala to
northwestern Costa Rica.

Chiococca semipilosa is distinguished by the
short thin hairs on the hypanthium/ovary, the pu-
bescence on the lower leaf surfaces, the short few-
flowered inflorescences, and the prominent calyx
lobes that often dry dark. There is the possibility
that material placed here is no more than an un-
usual form of C. alba. The figure is based on the
Guatemalan holotype (Steyermark 31406 F).

Chione DeCandolle

Trees or shrubs, glabrous or glabrescent; stipules unit-
ed (interpetiolar and intrapetiolar) and forming a cap
over the shoot apex, leaving a scar across the stem and
on the adaxial base of the petioles, small, caducous. Leaves
petiolate; leaf blades often coriaceous, entire and pin-
nately veined, domatia present or absent. Inflorescences
terminal, solitary or 3 at a distal node, paniculate with
opposite branching and cymose or corymbose in form,
pedunculate, bracteolate, flowers pedicellate. Flowers bi-
sexual and radially symmetrical, apparently monomor-
phic, hypanthium turbinate, calyx lobes 5 or 6 or un-
developed and the distal margin undulate; corolla
funnelform, white or yellowish, corolla tube short, gla-
brous within, corolla lobes 5(-6), broadly imbricate in
bud with 2 exterior; stamens 5(-6), inserted above the
base of the tube, filaments thick, anthers dorsifixed, ex-
serted; ovary 2-locular, ovules solitary in each locule,
pendulous from apex, style stout, stigmas 2, oblong, ex-
serted. Fruits drupaceous, ovoid to ellipsoid, pyrene sol-
itary and 2-locular; seeds elongate, the testa membra-
nous.

A genus of about 1 5 species; fewer than 6 species
are found in southern Mexico and Central Amer-
ica; the others occur in the West Indies. The genus
is distinguished by its glabrous parts, cap-like stip-
ules (in some species), terminal inflorescences, short
corolla tubes with broadly imbricate corolla lobes
(in bud), and the fleshy fruits with two-locular
pyrene (stone). The genus Oregandra is a syn-
onym; Standley misinterpreted the ovules when
he described that genus.

Chione sylvicola (Standl.) W. Burger, Selbyana 1 2:
138. 1991. Chomelia sylvicola Standl., J. Wash.
Acad. Sci. 18: 182. 1928. Oregandra panamen-
sis Standl., Publ. Field Columb. Mus., Bot. Ser.

FIELDIANA: BOTANY

4: 265. 1929. Anisomeris sylvicola (Standl.)
Standl., N. Amer. Fl. 32: 225. 1934. Chione
costaricensis Stand\., Publ. Field Mus. Nat. Hist.,
Bot. Ser. 22: 111. 1940. Chione panamensis
Steyerm., Ceiba 3: 19. 1952. Chione allenii L.
O. Williams, Phytologia 25: 462. 1973. Figure
36.

Shrubs or trees, (2-)6-l 5(-23) m tall, leafy branchlets
1-5 mm thick, glabrous, drying reddish brown to gray;
stipules 3-8 mm long, obtuse, glabrous, drying dark,
stipule scar crossing the stem between the leaf bases and
1-2 mm high on the adaxial side of the petioles, cadu-
cous. Leaves with petioles (5-)9-24 mm long, 0.7-1.8
mm thick, glabrous, usually sulcate above; leaf blades
6-17(-21) cm long, 2-7(-10) cm broad, elliptic-oblong,
to ovate-elliptic or ovate-oblong, apex bluntly obtuse to
acuminate, tip to 1 cm long, base abruptly rounded to
obtuse or acute, often decurrent on petiole when acute,
drying stiffly chartaceous and pale yellowish green to
dark brown, glabrous above and below or with slender
hairs or pits (domatia) in the vein axils beneath, 2 veins
(3-)4-9/side. Inflorescences, solitary and terminal (but
sometimes the lower branches subtended by smaller
leaves and appearing to be axillary), 5-12 cm long, 3-8
cm broad, glabrous, peduncles 1.5^4 cm long, bracts ca.

I mm long, subulate, pedicels 2-10 mm long and not
clearly distinguished from the hypanthium. Flowers 10-
1 2 mm long, glabrous externally, aromatic, hypanthium
3-4 mm broad at apex, calyx lobes 0-0.5 mm high, ca.
1.5 mm broad, carnose, broadly rounded; corolla white,
glabrous, somewhat fleshy, tube 3-6 mm long, 2-4 mm
diam., lobes 5, 2-3 mm long, 3-5 mm broad, broadly
imbricate in bud, rounded distally; anthers 3-4 mm long
and 0.7 mm thick, exserted. Fruits 14-22 mm long, 7-

I 1 mm diam., ellipsoid or curved, often narrowed below
the persisting calyx, red to purple (black) at maturity,
persisting calyx 1-2 mm long.

Trees, less often shrubs, of evergreen rain forest
formations on both the Caribbean and Pacific
slopes, from near sea level to 2000 m elevation.
Probably flowering and fruiting throughout the year
(flowering mostly in February-June). The species
ranges from southeastern Nicaragua to central
Panama (but see below).

Chione sylvicola is recognized by the lack of
pubescence on both vegetative and reproductive
parts (except for the pubescent domatia in vein
axils), unusual glabrous stipules, terminal inflo-
rescences, versatile exserted stamens and narrowly
ellipsoid fleshy red to black fruits. Specimens of
this species are occasionally mistaken for species
of Neea (Nyctaginaceae). The species concept
adopted here is a broad one. The type of C. cos-
taricensis (A. Smith 1778?) came from 825 m on
the Caribbean slope and has leaves intermediate
between the smaller-leaved collections from the
highlands and the larger-leaved lowland collec-

tions. The specimens of Oregandra panamensis
(Cooper & Slater 144 the type, F, and 149 F) have
unusually large leaves, and they may represent the
same individual. The type of C. allenii (Allen 5321
F) has the larger leaves with greater number of
secondary veins and pubescent domatia charac-
teristic of other collections from the Golfo Dulce
area. The type of C. panamensis (Hagen & Hagen
2137 F) is from 2000 m elevation in the Province
of Chiriqui, and the leaves have minute pit doma-
tia beneath. It may be that the species should be
divided into subspecific elements or that we are
mistaken in placing all this material under a single
name (see Dwyer, 1980, p. 92). The type ofCho-
melia sylvicola (Standley & Valeria 49196 us) is a
smaller-leaved high-elevation (2000 m) specimen
with only a few leaves and fruits.

Chomelia Jacquin

Shrubs or small trees, axillary spines present in some
species, branchlets terete; stipules interpetiolar, acumi-
nate, persistent or deciduous. Leaves opposite, petiolate,
entire, venation pinnate, domatia present in some spe-
cies. Inflorescences solitary, axillary, pedunculate or sub-
sessile, with few to many flowers, cymose or congested
and subcapitate, bracts present, bracteoles free or united.
Flowers bisexual and radially symmetrical, white or yel-
lowish white, hypanthium turbinate to oblong, calyx lobes
4(-5?), narrow and elongate, equal or unequal; corolla
salverform to funnelform, with a narrow elongate tube,
usually sericeous externally, usually glabrous within, co-
rolla lobes 4(-5?), valvate or imbricate in bud, lobes with
or without appendages at apex externally (abaxially); sta-
mens 4, sessile on the throat of the corolla tube, anthers
linear to sagittate, dorsifixed, included or slightly ex-
serted, basal lobes acute to obtuse; ovary 2(-3)-locular,
with 1 ovule pendulous from apex of each locule, style
filiform with 2(-3) short stigmas. Fruits drupaceous, small,
ellipsoid, the pyrene bony, solitary and 2-locular, with
persisting calyx lobes; seeds usually 2. cylindrical, pen-
dulous.

A genus of ca. 50 species in Central and South
America, and with more than 300 species in the
Old World tropics (but these are sometimes placed
under Tarennd). Species without appendages on
the corolla lobes and obtuse basal anther lobes
formerly placed in the genus Anisomeris are here
considered as part of Chomelia, following pre-
vailing opinion. Some species ofGuettarda (with-
out spines), Rondeletia (capsular fruits), and Sa-
bicea (vines with baccate fruits) resemble our
species of Chomelia; the axillary flower and inflo-
rescences, long slender sericeous corolla tubes, and
narrow corolla lobes help to distinguish Chomelia.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Key to the Species of Chomelia

la. Leaf blades with the smallest (4) veins subparallel within areoles denned by the tertiary (3) veins,

or the 3 and 4 veins parallel between the secondaries 2

1 b. Leaf blades with the smallest (4) veins not parallel within areoles denned by the tertiary (3) veins,

3 and 4 veins usually reticulate 4

2a. Leaf blades with both the 3 and 4 veins subparallel and at right angles to the secondary veins;
corolla tubes 6-8 mm long; spines absent; plants of the wet evergreen Caribbean slopes, 600-

900 m elevation C. venulosa

2b. Leaf blades with the 4 veins subparallel within areoles denned by the 3 veins; corolla tubes

12-40 mm long; spines often present; 0-1200 m elevation 3

3a. Flowers borne in pedunculate cymose inflorescences; trees of deciduous and semideciduous

forest formations C. spinosa

3b. Flowers solitary or several in leaf axils, sessile or pedicellate, never cymose; trees of evergreen

forest formations 0-900 m elevation C. recordii

4a. Flowers subsessile in leaf axils and at the apex of short shoots; petioles 2-5 mm long, leaf blades

to 9 cm long C. recordii

4b. Flowers borne on pedunculate inflorescences in the axils of leaves; petioles 2-10 mm long, leaf
blades to 1 5 cm long C. microloba

Chomelia microloba J. D. Smith, Bot. Gaz. 31:
114. 1901. Anisomeris microloba (J. D. Smith)
Standl., Publ. Field Columb. Mus., Bot. Ser. 4:
293. 1929. Antirhea panamensis Standl., N.
Amer. Fl. 32: 264. 1934. Chomelia panamensis
(Standl.) Dwyer, Ann. Missouri Bot. Gard. 67:
100. 1980. Chione chambersii Dwyer & Hay-
den, Ann. Missouri Bot. Gard. 54: 138. 1967.
Chomelia peninsularis Dwyer, Ann. Missouri
Bot. Gard. 67: 101. 1980. Figures 33-34.

Shrubs or small trees, 1.8-15 m tall, leafy branchlets
1.3-3 mm thick, with thin straight ascending hairs 0.5-
2 mm long, glabrescent, spines absent; stipules 2-8 mm
long, ca. 2 mm broad at the base, triangular-cuspidate,
with thin straight hairs, persisting with the leaves. Leaves
often closely clustered distally, petioles 2-18(-50?) mm
long, 0.5-1.3 mm thick, with curved hairs along the
adaxial margins and glabrescent; leaf blades 4-12(-18)
cm long, 2-6(-8) cm broad, narrowly to broadly ovate-
elliptic, to elliptic, oblong or slightly obovate, apex usu-
ally acuminate or with a bluntly triangular tip ca. 1 cm
long, base acute to obtuse, drying stiffly chartaceous, dark
brown or dark green above, glabrous or sparsely pubes-
cent above and below, usually with small (0.5-1 mm)
pubescent domatia in the vein axils beneath, 2 veins 4-
7/side, tissue between the secondary veins smooth when
dry (the minor venation not prominent). Inflorescences
3-10 cm long, to 7 cm broad, with 12-70 flowers, pe-
duncles 2-6 cm long, 0.3-1 mm thick, appressed-pu-
bescent or glabrous, with a terminal flower and 2 lateral
branches or with 3 1 branches and dichotomous 2
branches, pedicels 0-1 mm long. Flowers appressed-pu-
bescent or occasionally glabrous externally, 8-12 mm
long, hypanthium 1.2-2 mm long, 0.5-0.9 mm diam.,
cylindric or turbinate, glabrous, calyx lobes 0.2-0.5 mm
long, obtuse; corolla 7-10 mm long, white, tubular-fun-
nelform, sparsely to densely appressed-pubescent exter-

nally, tube 7-10 mm long, 0.3-1 mm diam. in the mid-
dle, lobes ca. 3 mm long, narrowly ovate and obtuse;
anthers 1 .2-2 mm long; stigma ca. 1 mm long. Fruit 10-
1 7 mm long, 4-8 mm diam., oblong to oblong-obovoid,
fleshy, glabrous and drying black, rounded or truncated
at apex, calyx deciduous, longitudinal ribs absent or
weakly developed.

An uncommon species in evergreen lowland rain
forest formations of the Pacific lowlands in Costa
Rica, to ca. 500 m elevation. Flowering in Feb-
ruary-May; fruiting in July-August. The species
ranges from southwestern Costa Rica to Colom-
bia.

Chomelia microloba is recognized by the small
axillary inflorescences with dichotomous branch-
ing and subsessile flowers along one side, the mi-
nute calyx lobes, narrow corolla tube, and leaves
with pubescent domatia. The leaves of the type
(Tonduz 9874 F, from Sto. Domingo de Golfo
Dulce) are quite small, and they appear to be atyp-
ical for the material placed here. The much longer
(to 1 5 cm) and broader (to 8 cm) leaves of the type
of C. peninsularis (Croat 22440 F, MO, from the
Burica peninsula) are probably more characteristic
of the species. The pubescence can differ greatly
in different collections, with some flowers being
quite glabrous and others appressed sericeous.

Chomelia recordii Standl., Trop. Woods 7: 9. 1926.
C. englesingii Standl., Trop. Woods 16: 45. 1928.
Anisomeris recordii (Standl.) Standl., N. Amer.
Fl. 32: 227. 1934. A. englesingii (Standl.) Standl.,
N. Amer. Fl. 32: 227. 1934. Figure 34.

100

FIELDIANA: BOTANY

Shrubs or trees to 10 m tall, leafy branchlets 0.7-3
mm thick, with straight or crooked strigulose hairs 0.2-
0.5 mm long, becoming gray and glabrescent, spines
present or absent, 7-27 mm long; stipules 3-5 mm long,
triangular and acute, pubescent, usually persisting. Leaves
with petioles 2-5 mm long, 0.5-1 mm thick, with straight
or crooked ascending hairs ca. 0.5 mm long; leaf blades
(2-)3-9 cm long, 2-4.5 cm broad, ovate to ovate-elliptic
or ovate-orbicular, apex acute to obtuse or slightly acu-
minate, base obtuse to rounded and subtruncate, drying
thin-chartaceous, dark brown above, glabrous above or
with hairs above the midvein, puberulent beneath with
straight thin hairs 0.4-0.8 mm long, 2 veins 4-7/side,
vein axils with dense clusters of hairs (domatia) beneath.
Inflorescences of subsessile flowers in the leaf axils or
terminal on short lateral shoots, usually 2 flowers per
node (1-6), with stipule-like bracts and narrow villose
bracteoles. Flowers ca. 30 mm long, white, hypanthium
1-2 mm long, densely villous with straight yellowish
white hairs, calyx lobes 3-6 mm long, narrowly acute;
corolla cream white to greenish white, tube 1 5-20(-24)
mm long and 0.5-1 mm diam., with thin whitish as-
cending hairs 1-1.5 mm long, lobes 5-7(-15) mm long
and 1-1.5 mm broad; anthers sessile, ca. 2.8 mm long,
attached ca. 2 mm below apex of the tube; style linear,
ca. 17 mm long, stigmas ca. 1.5 mm long, narrowly
oblong. Fruits to 14 mm long (including calyx), 3-4 mm
diam., oblong or oblong-obovoid, body of the fruit 8-9
mm long, dull red and often turning blue-black, with
thin whitish ascending hairs, the persistent calyx 4-5 mm
long.

Trees and shrubs of evergreen rain forest for-
mations on both the Caribbean and Pacific slopes,
from near sea level to 1 200 m elevation. Flowering
in April-June and August; fruiting in February,
April, and November. Collections have been made
in the Caribbean slope and lowlands, the General
valley, and the Golfo Dulce area in Costa Rica.
The species ranges from Guatemala to Colombia.

Chomelia recordii is recognized by the few ses-
sile flowers and fruit, the narrow calyx lobes, the
long slender corolla tube, and the relatively short
broad leaves. The 4 veins are usually parallel
within the areolae defined by the 3 veins, but this
cannot be seen in some specimens.

Chomelia spinosa Jacq., Enum. PI. Carib. 1 2. 1 760.
Ixora spinosa (Jacq.) Lam., Encyc. Meth. Bot.
3: 344. 1 789. C.filipes Benth. in Oerst., Vidensk.
Meddel. Dansk. Naturhist. Foren. Kjobenhavn
1852: 41. 1852. Guettarda costaricensis K.
Schum. ex Tonduz, Bull. Herb. Boissier 2: 7.
1895 (nomen). Figure 34.

Shrubs or small trees, 1.5-6(-9) m tall, straight woody
spines often present on older stems, axillary and 1-4 cm
long, ca. 2.5 mm wide near the base, terete, leafy branch-
lets 1-4 mm thick, densely puberulent with thin whitish

hairs 0.4-1 mm long, terete, becoming gray; stipules 4-
8 mm long, ca. 2 mm broad at the base, triangular-
subulate, puberulent, scarious, usually persisting. Leaves
often crowded at the ends of branchlets, petioles 5-20
mm long, ca. 0.5 mm thick, densely puberulent; leaf
blades 3.7-9 cm long, 2-5 cm broad, ovate-elliptic, to
broadly elliptic-oblong, ovate-orbicular or slightly ob-
ovate, apex acute to short-acuminate, base acute to ob-
tuse or slightly rounded and subtruncate, often decurrent
on petiole, drying chartaceous and dark brown above,
sparsely puberulent above with hairs ca. 0.4-0.6 mm
long, sericeous beneath with thin whitish hairs (especial-
ly dense on the major veins), 2 veins (3-)4-7(-8)/side,
strongly ascending, the minor (4) veins subparallel with-
in areolae denned by the 3 veins. Inflorescences ( 1 .2-)3-
7 cm long, axillary, peduncles 12-45 mm long, ca. 0.5
mm thick, densely puberulent, flowers in small distal
clusters of 3-7(-15) near apex of the peduncle. Flowers
sweetly aromatic, hypanthium 2-3 mm long, ca. 1 mm
diam., densely sericeous with whitish ascending hairs,
calyx lobes 0.5-1 mm long, slightly unequal; corolla yel-
lowish white, sericeous externally, tube 12-18(-24) mm
long, 0.5-1 mm diam., lobes 4-7 mm long, 1-3 mm
broad, glabrous along the edges and within (adaxially).
with a short appendage near apex; filaments very short,
anthers ca. 3.5 mm long, sagittate at the base, disc about
0.6 mm long; styles 14-17 mm long, stigmas ca. 1 mm
long, exserted. Fruits sessile, 6-9(-l 2) mm long, 3-6 mm
diam., sparsely puberulent or glabrous, becoming black.

Common shrubs and trees of deciduous and
partly deciduous forest formations on the Pacific
slope of Costa Rica, from near sea level to about
500 m elevation. Flowering in late June-August
in Costa Rica; fruiting in June-December in Cen-
tral America, with a peak of fruiting in November
in Costa Rica. The species ranges from southern
Mexico and Guatemala, along the Pacific coast of
Central America to Colombia, Venezuela, and
northern Brazil.

Chomelia spinosa is recognized by its restriction
to deciduous and partly deciduous vegetation, the
straight woody spines (not usually present on distal
flowering branches), small cymose inflorescences
on slender axillary peduncles, flowers with long
slender corolla tube, and short flowering season.
The minor venation is quite distinctive with the
4 veins parallel only within small areoles defined
by the 3 veins, with the result that the 4 veins of
adjacent areoles often are not parallel with each
other. The species has been called limoncillo, ma-
lacaguite, and malacahuite.

Chomelia venulosa W. Burger & C. M. Taylor, sp.
nov. Figure 34.

Arbores 10-25 m altae, ramulis juvenibus sericeis;
stipulis 11-18 mm longis. Foliae lamina elliptica vel

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

101

oblongo-elliptica, 6-15 cm longa, 3-7 cm lata, venis lat-
eralibus 5-7, venulis parallelis. Inflorescentiae axillares,
ad 9 cm longae, pedunculo ca. 22 mm longo, ramis saepe
dichotomis, floribus secundis. Flores puberuli, hypan-
thio 1-2 mm longo; corolla alba vel cremea, tubo 6-9
mm longo, lobis 4. Fructus 13-15 mm longi.

TYPUS E. Bella 414 (holotypus CR, isotypi F, MO),
from Reserva Biologica Monteverde, Alajuela, Costa
Rica.

Trees, 10-25 m tall, to ca. 35 cm dbh, leafy stems 1-
4 mm thick, sericeous or strigulose with ascending pale
grayish hairs ca. 0.4 mm long; stipules 1 1-18 mm long,
to 4 mm broad, lanceolate with an acute apex, sericeous
at the base and along the midrib. Leaves with petioles
(6-) 1 2-23 mm long. 0.8-2 mm thick, sericeous with pale
grayish hairs; leaf blades 5-15 cm long, 3-7 cm broad,
elliptic to elliptic-oblong, apex short-acuminate with tip
5-8 mm long, base obtuse to acute, drying stiffly char-
taceous, dark brown above, much paler beneath, gla-
brous or very sparsely pubescent above with thin whitish
hairs to 1 mm long, appressed-pubescent beneath with
thin hairs ca. 0.3 mm long and densely sericeous along
the major veins, 2 veins 4-7/side, both the 3 and 4
veins at right angles to the secondaries, depressions
(domatia?) sometimes present in the vein axils. Inflo-
rescences solitary and axillary (2/node), ca. 5 cm long,
to 9 cm in fruit, cymose with 2 lateral simple or bifid
secund branches, peduncles 22-45 mm long and 1 mm
thick, ascending sericeous, distal flowers along 1 side of
the branch, bracts absent, flowers sessile. Flowers pu-
bescent externally, hypanthium 1-2 mm long, 1 mm
diam., calyx tube minute, 1.4-2 mm diam., calyx lobes
0.2-0.8 mm high; corolla tubular, yellow or yellowish
white, fluted distally, tube 6-9 mm long, 0.8-1.5 mm
diam., densely sericeous, lobes 4, 1-2 mm long, 1-2 mm
broad; stamens 4, anthers ca. 3.5 mm long; style ca. 5
mm long, stigmas 1.2 mm long. Fruits 13-15 mm long,
8-9 mm thick, obovoid-oblong, with 6-8 longitudinal
ribs, becoming dark brown, glabrous, persistent calyx ca.
1 mm long, pyrenes ca. 14 x 8 mm; seeds ca. 10 x l
mm.

Plants of the wet evergreen forests of the Carib-
bean slope of Costa Rica, at 600-900 m elevation.
Flowers were collected in May; fruits were col-
lected in October-November. The species is known
from below Monteverde, Alajuela (8443'W), and
the southern Cordillera de Talamanca, Limon
(8259'W). Collections in addition to the type are
Bello 17 2 & 872, Bella & Cruz 457, Herrera 3310,
and Poveda 24.

Chomelia venulosa is recognized by its parallel
minor venation, sericeous flowers on short inflo-
rescences with two simple or bifid lateral branches,
and restricted altitudinal range on the Caribbean
slope. The two-celled deeply ridged pyrenes, the
subimbricate to valvate corolla aestivation, and
the parallel minor venation suggest that this spe-
cies is best placed in Chomelia, as opposed to
Guettarda. It may be allied to the " Anisomeris"
group of Chomelia species with appendages on the

corollas. The overall appearance of the leaves is
similar to that of Chomelia panamensis, but the
minor venation is quite unusual. The 3 and 4
veins are little differentiated, parallel to each other,
and usually perpendicular to the 2 veins.

Cinchona Linnaeus

Trees of medium height or occasionally shrubs,
branchlets terete or tetragonal; stipules interpetiolar, tri-
angular, often large, colleters present within at the base,
caducous or rarely persisting. Leaves opposite, often large,
petiolate; leaf blades with entire margins, coriaceous to
chartaceous, domatia absent. Inflorescences terminal and
solitary (apparently axillary when lateral branches are
subtended by reduced leaves), open-paniculate, usually
large and many-flowered, branching mostly opposite.
Flowers bisexual, small in most species, aromatic, often
puberulent externally, hypanthium turbinate, calyx tube
with small calyx lobes (rarely with the calyx tube entire
distally); corolla salverform to fimnelform, white to pink
or purplish, corolla tube terete or slightly 5-angled, gla-
brous or pilose in the throat; corolla lobes 5 (4, 6), spread-
ing, valvate in bud; stamens 5, inserted in the corolla
tube, filaments short or long, anthers linear dorsifixed,
included or their apices exserted; ovary 2-locular, pla-
centas attached to the septum and spongy, ovules many
in each locule, peltately attached and imbricated, style
narrow, stigmas short and obtuse, included or slightly
exserted. Fruits woody capsules, 2-locular, subcylindri-
cal to ovoid or oblong, dehiscing septicidally from bot-
tom to top; seeds many, peltate, thin and flat, testa with
a broad thin peripheral wing.

A genus of 20 40 poorly defined species, ranging
from Costa Rica southward to Bolivia, mostly along
the Andes mountains. The capsules opening up-
ward from the base helps separate Cinchona from
closely related genera, such as Ladenbergia and
Joosia. This genus has played an important role
in the history of medicine as the source of the
antimalarial drug quinine. The major commercial
sources of quinine are cultivars of Cinchona cal-
isaya Wedd. grown in Indonesia, which originated
from the eastern slopes of the Andes. Species of
the genus were introduced and have become nat-
uralized in Guatemala (see the discussion in
Standley & Williams, 1975, p. 38).

Cinchona pubescens Vahl, Skr. Naturhist. Selsk.
Kjobenhavn 1: 19. 1790. Figure 37.

Small or medium-sized trees, (3-)5-20 m tall, trunks
8-30 cm dbh, bark pale brown and roughened, leafy
branchlets 3-8 mm thick, distinctly 4-angled, minutely
puberulent or glabrous; stipules 4-12(-20) mm long, 3-
8(-12) mm broad at the base, glabrous or minutely ap-

102

FIELDIANA: BOTANY

pressed-puberulent, deciduous. Leaves with petioles
(9_) 1 2-35(-60) mm long, 1 .3-2.5 mm thick, sparsely and
minutely (0.2 mm) puberulent; leaf blades 10-32(-40)
cm long, 7-17(-26) cm wide, broadly ovate to broadly
elliptic-oblong or suborbicular, apex broadly obtuse to
subacuminate, base abruptly cuneate to rounded and
subtruncate, usually slightly decurrent on petiole, drying
stiffly chartaceous or chartaceous, glabrous or very
sparsely puberulent above, sparsely pubescent beneath
with slender hairs ca. 0.3 mm long, 2 veins 7-12/side.
Inflorescences terminal or axillary to distal leaves, 9-40
cm long, 8-24 cm broad, open paniculate with few op-
posite widely spaced (3-1 1 cm) branches, distal branches
minutely tomentulose, the flowers in congested distal
clusters, pedicels 0.5-3 mm long. Flowers white, cream,
or pale pink, densely tomentulose externally, hypanthi-
um 2-3 mm long, 1.3-2 mm diam., calyx tube l-2(-3)
mm long and 2-3 mm diam., calyx lobes 0.2-1 mm long,
acute; corolla 15-16 mm long and funnelform, tube 10-
1 3 mm long, 1.4-2 mm diam.. glabrous within, lobes 5,
lanceolate to oblong, 3-5 mm long and 1.5-2.5 mm wide,
tomentulose externally and villose on the interior mar-
gins; stamens 5, filaments ca. 2 mm long, anthers 2.5-3
mm long; style 6-13 mm long, glabrous. Fruits subcy-
lindrical to narrowly oblong, 12-35(-50) mm long, 6-9
mm diam., glabrescent or minutely (0. 1 mm) puberulent,
brown, lustrous yellowish brown within; seeds 5-1 2 mm
long, 1.5-3 mm broad, flat and oblong-elliptic in outline,
with a thin membranaceous winged margin, surfaces re-
ticulate and the margin erase, dark center of the seed
1.8-3 mm long.

Trees of evergreen forest formation on both the
Caribbean and Pacific slopes of Costa Rica, rang-
ing from (500-)800 to 1700 m elevation. Flow-
ering mostly in June-September, with solitary col-
lections in November, February, and March;
fruiting in February and November-December in
Costa Rica. The species is apparently rarely en-
countered in southern Central America. Our col-
lections come mostly from the Caribbean slopes
of the Cordillera de Tilaran and the Central High-
lands in the Provinces of Alajuela and Heredia. In
Panama the species is known only from the Chi-
riqui Highlands. This species ranges southward to
Venezuela, Peru, and Bolivia.

Cinchona pubescens is recognized by the larger
often broadly rounded leaves, large terminal in-

florescences with small puberulent flowers in distal
clusters, the narrow woody two-locular capsules,
and the seed with a thin elongated membranous
wing. The flowers are said to have the aroma of
Gardenia or Cananga odorata (Annonaceae). The
rarity of this species in southern Central America
suggests that it is not native, and collections may
represent relicts of native pre-Columbian intro-
duction.

Coccocypselum P. Browne,
nomen conservandum

Herbs, annual or perennial, prostrate and creeping to
erect-ascending, usually pubescent with multicellular
hairs; stipules interpetiolar, sometimes reduced to a very
short (0.2 mm) rim, small and simple with a single su-
bulate lobe (2/node) and with or 2-8 smaller lateral
awns, persisting. Leaves petiolate, entire, domatia ab-
sent. Inflorescences terminal or pseudoaxillary, usually
solitary, capitate with (l-)3-20 flowers, sessile or pe-
dunculate, bracts and bracteoles small, flowers sessile.
Flowers bisexual, monomorphic or distylous, small, ca-
lyx lobes 4, narrow and persistent; corolla blue to purple
or white, funnelform, corolla lobes 4, valvate in bud;
stamens 4, filaments borne on the tube of the corolla,
anthers dorsifixed below the middle or near the base;
ovary 2-locular, the placentas borne on the center of the
septum, ovules many and horizontal, style with 2 short
branches. Fruits baccate and arenchymatous, often mealy
and hollow, globose to obovbid, bright blue; seeds many
and small, angled or flattened.

A Neotropical genus of 10-20 species, ranging
from Mexico and the West Indies into South
America. The genus is recognized by its herba-
ceous habit, usually broad puberulent leaves, small
capitula with few flowers, four-parted flowers, two-
locular ovaries, and blue fruit with many small
seeds. These plants are often confused with species
of Geophila. Coccocypselum lanceolatum is our
most distinctive species; the others may be difficult
to distinguish.

Key to the Species of Coccocypselum

la. Leaf blades with 8-13 pairs of secondary veins, narrower than long, ovate-triangular to lanceolate
and usually acute at the apex; with 8-10 flowers in each inflorescence [1000-2000 m elevation]

C. lanceolatum

1 b. Leaf blades with 5-6 pairs of secondary veins, usually about as broad as long, ovate to ovate-
lanceolate, acute to rounded at the apex; with 2-6 flowers in each inflorescence
2a. Inflorescences sessile in the axils of leaves; plants of evergreen lowlands, 0-1000 m elevation

. C. herbaceum

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

103

2b. Inflorescences pedunculate; plants of evergreen lowland and highlands, 0-2000 m elevation ... 3
3a. Leaf blades rounded and cordate or subcordate at the base, often wider than long [ 1 400-2000 m

elevation] C. cordifolium

3b. Leaf blades acute to obtuse or truncated at the base, usually as wide as long and ovate to broadly

elliptic 4

4a. Stems and leaves with hairs 0.5-1 mm long (or occasionally glabrous); calyx lobes 3-4 mm long;

stipules ca. 5 mm long; common in Central America C. hirsutum

4b. Stems and leaves with hairs 0. 1-0.3 mm long; calyx lobes 1.5-2.7 mm long; stipules 2-4 mm long;

not known from Costa Rica (key based on Steyermark, 1972) C. guianensis

Coccocypselum cordifolium Nees & Mart., Nova
Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat.
Cur. 12: 14. 1824. Geophila pleuropoda J. D.
Smith, Bot. Gaz. 52: 50. 1911. Geocardia pleu-
ropoda (J. D. Smith) Standl., Contr. U.S. Natl.
Herb. 17: 445. 1914. Tontanea pleuropoda (J.
D. Smith) Standl., N. Amer. Fl. 32: 148. 1921.
C. pleuropodum (J. D. Smith) Standl., Publ. Field
Columb. Mus., Bot. Ser. 4: 281. 1929. C. roth-
schuhii Loessner, Bot. Jahrb. Syst. 60: 370. 1926.
Figure 2.

Herbs, prostrate or creeping, leafy stems 0.4-1.6 mm
thick, hirsute or villous with thin straight or crooked
hairs 0.5-1.5 mm long; stipules 1.5-3 mm long, united
only at the base (ca. 0.2 mm) and with 2 narrowly linear
awns on each side (4/node). Leaves with slender petioles
4-28(-55) mm long, villous or pilose with thin hairs; leaf
blades 1 1-35 mm long, 1 2-42 mm broad, ovate to ovate-
orbicular or ovate-reniform, apex rounded and bluntly
obtuse (and usually minutely apiculate), base rounded
at the cordate to subtruncate base, drying membrana-
ceous or thin-chartaceous, sparsely to densely pubescent,
the hairs 1-1.7 mm long on the upper surface and ca.
0.7 mm long beneath, 2 veins 3-5/side. Inflorescences
axillary, usually only I/node, 1.4-5 cm long, the capit-
ulum less than 1 cm long and with 3 (2, 4) flowers,
peduncles 4-38 mm long, villous, bracts ca. 4 mm long,
linear, flowers sessile. Flowers with hypanthium ca. 1
mm long, with thin whitish hairs, calyx lobes 1.5-2.5
mm long, linear-lanceolate, sparsely villous; corolla lav-
ender, lilac, pale blue, or whitish, tube 4-6 mm long, 2-
3 mm diam. at apex, sparsely puberulent externally, lobes
3.3-5 mm long, 1.2-2 mm broad at the base, narrowly
triangular to oblong. Fruits 5-6 mm long, 4-5 mm diam.,
ovoid, blue, densely villous and with the persisting calyx
2-3 mm long; seeds 0.2-0.5 mm diam.

Plants of evergreen lower montane rain forest
formations, from 1000 to 1600 m elevation (to
2000 m in Guatemala). Rarely collected in Costa
Rica's major highlands. Probably flowering and
fruiting throughout the year. The species ranges
from Mexico to Panama, and it is also found in
eastern and southern Brazil.

Coccocypselum cordifolium is recognized by the
usually subcordate or truncated leaf bases, the small

pedunculate heads with only two to four blue flow-
ers, and the long hairs on the upper leaf surface.
This species appears to be rare or overlooked in
Costa Rica and Panama. This species is easy to
confuse with Geophila cordifolia, which has one-
or two-seeded red fruits.

Coccocypselum guianense (Aubl.) K. Schum. in
Mart., Fl. Bras. 6(6): 315. 1889. Tontanea gui-
anensis Aubl., Hist. pi. Guiane 1: 108, pi. 42.
1775.

According to Steyermark (1972), this species is
distinguished from similar species by its short (1.5
2.7 mm) calyx lobes, short (0.2-0.5 mm) dense
indumentum on stems and inflorescences, smaller
(34 mm) stipules, and slightly longer (5.5-9 mm)
corolla tubes. The species is said to range from
Florida and the West Indies to Venezuela and the
Guianas (Steyermark, 1972; Hortus Third, 1976).
These plants have been used in ornamental hor-
ticulture as a ground cover and in hanging baskets.
Because of these uses, it is likely that escaped pop-
ulations have become established in some areas.
We have not seen material that can be definitively
ascribed to C. guianense from Costa Rica.

Coccocypselum herbaceum P. Browne, Civ. Nat.
Hist. Jam. 144, pi. 6. 1756; also cited as Aublet,
Hist. pi. Guiane 1: 68. 1775 (fide Adams, 1972);
Lam., Encycl. 2: 56. 1786 (fide Standley & Wil-
liams, 1975). C. repensSv/., Prodr. 31. 1788 (not
C. repens H.B.K. 1819, not Condalia repens Ruiz
& Pav. 1798). Tontanea herbacea (P. Browne)
Standl., N. Amer. Fl. 32: 147. 1921. T. hispidula
Standl., loc. cit. 147. 1921. C. hispidulum
(Standl.) Standl., Publ. Field Columb. Mus., Bot.
Ser. 4: 281. 1929.

Herbs, procumbent or trailing, leafy stems 0.7-2 mm
thick, sparsely to densely pilosulous with erect or ap-

104

FIELDIANA: BOTANY

pressed hairs 0.2-1.2 mm long; stipules with linear awns
3-4 mm long, ca. 0.5 mm broad at the base (2 larger
awns per node), lateral lobes none or 2-4 and 0.5-1 mm
long, puberulent. Leaves with petioles 6-27 mm long,
0.4-1 mm thick, sparsely to densely pubescent; leaf blades
2-5.5 cm long, 1-3.5 cm broad, ovate to ovate-oblong
or ovate-triangular, apex obtuse and sometimes with a
small (0.5 mm) apiculate tip, drying membranaceous to
thin-chartaceous, with short (0.3 mm) or long (1-2 mm)
hairs on the upper surface, glabrous or pilose beneath
with hairs to 1 .2 mm long, 2 veins 5-8/side. Inflores-
cences sessile or subsessile in the leaf axils (rarely with
peduncles to 6 mm long), to 1 cm long, usually with
3(-6) sessile or subsessile flowers, bracts 2-3 mm long,
linear. Flowers monomorphic, hypanthium densely hir-
tellous, calyx lobes 2.5-4.5 mm long, linear-lanceolate,
sparsely pubescent; corolla dark blue to purple or with
a white tube, tube 5-8 mm long, corolla 1-2 mm long,
acute. Fruits becoming 1 1 mm long (not including the
calyx) and 8-10 mm diam., globose or ovoid, deep blue,
persisting calyx ca. 3 mm long; seeds 0.5-1.3 mm long,
smooth or rugose, flattened and angular or lenticular.

Plants of evergreen or partly evergreen forest
formations in the central highlands and in the Ca-
ribbean lowlands, from near sea level to 900 m
elevation. Probably flowering throughout the year.
The species ranges throughout the American trop-
ics.

Coccocypselum herbaceum is recognized by the
small sessile groups of blue flowers, creeping habit,
bright blue fruit, and thin ovate leaves. This spe-
cies is very similar to C. hirsutum, which has pe-
dunculate inflorescences, but C. herbaceum is not
as common in Central America. It seems possible
that the two may prove to be conspecific, with C.
herbaceum having priority.

Coccocypselum hirsutum Hurtling ex DC., Prodr.
4:396. 1830. Tontanea hirsuta (Bartling ex DC.)
Standl., N. Amer. Fl. 32: 147. 1921. C. glabrum
Bartl. ex DC., Prodr. 4: 397. 1830. Tontanea
glabra (DC.) Standl., J. Wash. Acad. Sci. 1 5:
104. 1925. C. hirsutum var. glabrum (Bartl. ex
DC.) L. O. Williams, Phytologia 25: 462. 1973.
Figure 2.

Herbs, prostrate or creeping, leafy stems 0.5-2 mm
thick (not including the pubescence), usually densely vil-
lous or hirsute with pale yellowish hairs 0.5-1.5 mm
long; stipules with narrow linear awns 3-5 mm long, ca.
0.3 mm broad at the base, sparsely to densely puberulent.
Leaves with petioles 4-16(-20) mm long, 0.4-0.8 mm
thick, usually densely pubescent; leaf blades 2-4 cm long,
1.4-2.6 cm broad (to 6 x 4 cm in northern Central
America), ovate to ovate-triangular or ovate-oblong, apex
obtuse and often with a slightly (0.4 mm) apiculate tip,
base obtuse to rounded and truncate, drying membra-
naceous to thin chartaceous and often dark green or dark

brown above, with thin straight or crooked hairs 1-1.5
mm long on the upper surface, the hairs somewhat short-
er beneath except along the midvein, 2 veins 6-8/side.
Inflorescences 1-2.5 cm long, capitula 1-1.5 cm broad,
usually 3-flowered (rarely with 1-5 flowers), peduncles
3-24 mm long, bracts ca. 5 mm long and 0.5 mm broad,
with slender hairs. Flowers with hypanthium ca. 1.5 mm
long, densely villous, calyx lobes 3-4 mm long, 0.5-1
mm broad, sparsely pubescent; corolla blue, white, or
white with purple markings, tube 5-7 mm long, 1.5-2
mm diam. near the mouth, corolla lobes 2-4 mm long
and 1.5 mm broad, triangular, anthers ca. 1.5 mm long.
Fruits 9-20 mm long, 7-12 mm diam., ovoid to broadly
ellipsoid, blue or purplish blue, with spongy exocatp;
seeds lenticular, 1-1.5 mm broad, rugose, brown.

Plants of evergreen forest formations, from 500
to 2500 m elevation. Probably flowering and fruit-
ing throughout the year. It ranges from Mexico to
South America (but see below).

Coccocypselum hirsutum is distinguished by its
low creeping habit, hirsute (less often glabrous)
rounded leaves, pedunculate heads of few flowers,
and blue fruits. The plants placed here may not
be specifically distinct from C. herbaceum, which
is the earlier name. Peduncle length seems to vary
greatly on the same plant, with subsessile and long-
pedunculate capitula nearby. We agree with Wil-
liams (in Standley & Williams, 1975) that the gla-
brous elements of this complex do not deserve
specific recognition (see synonymy above).

Coccocypselum lanceolatum (Ruiz & Pav.) Pers.,
Syn. PI. 1: 32. 1805. Condalia lanceolata Ruiz
& Pav., PI. Fl. Peruv. 1: 54. 1798. Coccocyp-
selum repens H.B.K., Nov. gen. sp. 3: 3 1 6. 1 8 1 9
(not C. repens Sw. 1788). Coccocypselum ca-
nescens Willd. ex Cham. & Schlechtend., Lin-
naea 4: 139. 1829. Tontanea canescens (Cham.
& Schlechtend.) Stand., N. Amer. Fl. 32: 146.
1921. Figure 2.

Herbs, prostrate to erect, 1 0-40 cm tall, leafy branches
1-3 mm thick, densely pubescent with whitish or pale
grayish hairs 0.2-0.8 mm long; stipules with central awns
3-6 mm long, 0.5-1 mm broad at the base, narrowly
linear for most of their length, pubescent. Leaves with
petioles 7-22 mm long, 0.5-1 mm thick, densely pu-
bescent; leaf blades 3-9 cm long, 1 .5-4 cm broad, ovate-
triangular to ovate-oblong or triangular-oblong, apex
gradually tapering and acute (rarely obtuse), usually
shortly (0.3 mm) apiculate at the tip, base obtuse to
subcordate, drying membranaceous or thin-chartaceous,
with thin appressed hairs ca. 0.5 mm long and parallel
with the secondary veins, 2 veins 7-1 I/side. Inflores-
cences 2-5 cm long, capitula ca. 1 cm diam., globose,
usually with more than 8 flowers and the flowers tightly
congested, peduncles (6-)10-55 mm long, 0.5-1 mm

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

105

thick, densely pubescent, bracts 1-3 mm long (often dif-
ficult to see). Flowers with the hypanthium ca. 3 mm
long, densely pubescent with hairs ca. 1 mm long, calyx
lobes unequal, 2-4 mm long and 0.7-2 mm broad; co-
rolla blue to bluish purple, lilac, or white marked with
blue, tube ca. 2 mm long, lobes ca. 3 mm long and 1-2
mm broad; anthers ca. 1 mm long. Fruits 8-1 5 mm long,
6-10 mm diam., ovoid to ellipsoid, bright blue, pilose,
persisting calyx ca. 3 mm long; seeds angular to tetra-
hedral, 0.8-1 .3 mm broad, with minutely rugose surface.

Plants of partly deciduous or evergreen (but sea-
sonally dry) forest formations, 1000-2000 m el-
evation. Probably flowering throughout the year
(mostly in January-July). The species ranges from
Guatemala through Central America to Bolivia
and Brazil.

Coccocypselum lanceolatum is recognized by its
narrower leaves with more numerous secondary
veins, somewhat taller erect stems, dense whitish
or grayish pubescence, globose heads with more
than eight densely packed flowers, bluish corollas,
and bright blue fruits. This is our most distinctive
species of Coccocypselum.

Coffea Linnaeus

Shrubs or small trees, branchlets subterete, often held
horizontally, usually glabrous; stipules interpetiolar, tri-
angular, often persisting. Leaves opposite or verticillate,
decussate or somewhat distichous, petiolate or subses-
sile; leaf blades entire, chartaceous to coriaceous, mostly
glabrous, domatia often present. Inflorescences of clus-
tered subsessile or short-pedicellate flowers in leaf axils,
bracts present and united to form a short cup at the base
of the pedicel. Flowers bisexual, monomorphic, usually
radially symmetrical, white to pink, hypanthium sub-
cylindrical to turbinate, calyx tube short, truncate to den-
tate or lobed, calyx lobes usually 5, small; corolla sal-
verform or funnelform, corolla tube short or long, glabrous
or villous at the throat, corolla lobes 4-8, oblong or
obtuse, convolute in bud; stamens usually 5 (4-8), in-
serted in the throat of the tube, filaments short or none,

anthers linear, included or exserted; ovary 2-locular, with
1 ovule in each locule attached to the middle of the
septum, style slender and glabrous, with 2 narrow stig-
mas (= style branches). Fruits drupaceous, oblong to
subglobose or ovoid, exocarp fleshy or dry, with 2 woody
nutlets (pyrenes) covered by a chartaceous or coriaceous
endocarp (the "parchment"); pyrenes 2 (the "nutlets" or
"beans"), convex abaxially and flattened and deeply
grooved on the inner (adaxial) face, oblong in outline;
seeds ellipsoid.

A tropical Old World genus of about 40 species,
mostly African. This genus is the source of coffee,
one of the most important agricultural commod-
ities in world trade. Coffea arabica is the primary
and the preferred source of coffee beans; see the
discussion in the Flora of Guatemala (Standley &
Williams, 1975, pp. 44-48). The quality of the
coffee beans is dependent on the environment in
which the plants grow, the ripeness of the fruit,
methods of gathering and drying, and the final
roasting of the beans (cf. J. W. Purseglove, Trop-
ical crops: Dicotyledons, vol. 2: 458-482. 1984).
Only two species are likely to be encountered in
Costa Rica, and they are keyed and described be-
low. In addition, Coffea canephora Pierre ex
Froehner, Notizbl. Konigl. Bot. Gart. Berlin 1:
237. 1897 (C. robusta Linden, Cat. pi. nouv. hort.
colon. 11.1 900), which grows well at lower ele-
vation and is rust-resistant, may be encountered.
It produces a quality of bean intermediate between
C. arabica and C. liberica. "Robusta" plants can
be distinguished from "arabica" by their larger
( 1 2-40 cm) leaves, rounded at the base, 8-17 pairs
of secondary veins, and more corrugated surface.
It is an important source of coffee in India and
Indonesia. This and many other important Coffea
cultivars have been grown for many years at the
Centre Agronomic Tropical de Investigation y
Ensenanza (CATIE) near Turrialba.

Key to the Common Species of Coffea

la. Flowers with 5 corolla lobes; leaf blades to 1 5(-l 8) cm long, pit domatia 0. 1-0.3 mm broad, usually
circular C. arabica

Ib. Flowers with 6-8 corolla lobes; leaf blades to 24(-30) cm long, pit domatia 0.2-0.7 mm broad,
circular to elongate C. liberica

Coffea arabica L., Sp. PI. 172. 1753.

Shrubs or small trees to 8 m tall, branches often held
horizontally or slightly drooping, leafy branchlets 1-4
mm thick, glabrous, nodes usually well spaced (4-7 cm)

and thickened (dried); stipules 3-7(-12) mm long, 2-5
mm broad, triangular to subulate, glabrous, deciduous.
Leaves with petioles 6-1 2(-l 5) mm long, 1-1 .6 mm thick,
glabrous; leaf blades 8-1 5(-l 8) cm long, 2.5-6(-7.5) cm
broad, elliptic-oblong to elliptic-obovate, apex acumi-
nate to caudate-acuminate, tip 1-2 cm long, base obtuse

106

FIELDIANA: BOTANY

to acute, drying chartaceous to subcoriaceous, glabrous
above and below, 2 veins 7-10/side, minute (0.2-0.3
mm) domatia often present at the vein axils beneath.
Inflorescences axillary, subcapitate or appearing verti-
cil laic, ca. 5 cm long (including the corollas), with 1-9
flowers per axil, flowers subsessile. Flowers 1 2-20 mm
long, white, calyx lobes 5, minute, corolla white, tube
10-14 mm long, lobes 14-20 mm long; anthers exserted,
8-12 mm long. Fruits 10-16 mm long, 8-13 mm diam.,
oblong and abruptly rounded at apex and base, green
becoming red (drying dark), glabrous, calyx scar 2-3 mm
broad; pyrenes ca. 10 x 7 x 3 mm, planoconvex.

Cultivated or rarely persisting in evergreen and
partly deciduous formations, 800-2000 m eleva-
tion. The flowering season is primarily in Febru-
ary, with fruiting in November-December. This
species, probably native to western Ethiopia, is
now cultivated throughout the tropics.

Coffea arabica is recognized by its glossy dark
green leaves (in life), lack of pubescence, axillary
clusters of aromatic white flowers, and distinctive
two-seeded fruit. This species is only occasionally
found outside of cultivation; birds and bats have
been described as dispersal agents. In addition to
the stimulant caffeine, coffee beans contain glu-
cose, dextrin, proteins, and the flavor-enhancing
volatile oil caffeol. Cultivation in cooler temper-
atures at higher elevations (ca. 1000-2000 m) is
an important factor in producing high-quality cof-
fee (cafe), and this may account for the excellent
reputation of Costa Rica's most valuable export
crop.

Coffea liberica Bull ex Hiern., Trans. Linn. Soc.,
Ser. 2, 1: 171, t. 24. 1876. C. excelsa A. Chev.,
Rev. cult, colon. 12: 258. 1903.

Shrubs or small trees to 5(-l 5) m tall, leafy branchlets
1.8-8 mm thick, glabrous; stipules 3-6 mm long, 4-8
mm broad. Leaves with petioles (4-)8-24 mm long, 1 .5-
4 mm thick, glabrous; leaf blades (8-) 1 2-24(-30) cm
long, (4-)5-12 cm broad, elliptic-oblong to elliptic-ob-
ovate, apex bluntly obtuse to bluntly short-acuminate,
base obtuse to acute, drying subcoriaceous, glabrous above
and below, 2 veins 7-10/side, pit domatia 0.2-0.8 mm
long at the vein axils beneath and often with a few short
hairs. Inflorescences 2-5 cm broad, with ca. 5-10 flow-
ers, bracts 3-8 mm long, narrowly elliptic to lanceolate.
Flowers 6-8-parted, 25-35 mm long, corolla tube 10-
14 mm long and ca. 1.5 mm diam., lobes 8- 12 mm long,
2-3 mm broad; anthers to 6 mm long. Fruits 1 .2-2.5 cm
long, oblong-rotund, yellowish red.

Cultivated or escaped trees of evergreen forest
formations, from near sea level to 1200 m ele-
vation. This species is native to coastal West Af-
rica and is now found cultivated around the world.

Coffea liberica is recognized by its larger leaves,
glabrous parts, white axillary flowers, and distinc-
tive fruit. Unlike C. arabica. which is essentially
a highland species, C. liberica grows well at lower
elevations. While C. liberica is more disease-re-
sistant and can do well at lower altitudes, the seeds
(beans) produce coffee of inferior flavor, and the
species is not an important crop in Central Amer-
ica.

( ondaminea DeCandolle

Shrubs or small trees, usually with few branches and
large leaves; stipules interpetiolar and intrapeliolar, unit-
ed above the leaf base to form a short sheath and with
a broad distal 2-parted blade (sometimes appearing as
4 free stipules at each node), persisting. Leaves opposite
and large, sessile or short-petiolate; leaf blades entire,
lacking domatia. Inflorescences terminal and solitary,
open paniculate with cymose or corymbose distal
branching, usually with 3 branches at apex of the pe-
duncle (lateral branches equaling the continuing rachis),
bracts small, bracteoles absent, pedicels short or none.
Flowers bisexual, large, hypanthium turbinate to cam-
panulate, calyx lobes 3-5 or none and the calyx tube
entire, corolla funnelform to salverform, thick-fleshy,
corolla tube barbate in the throat, corolla lobes 4-5,
valvate in bud, glabrous; stamens 5, filaments inserted
in the upper half of the tube, puberulent at the base,
anthers narrow, sagittate at the base, exserted; ovary 2-
locular, ovules many in each locule from axile placentas,
style narrow, stigmas oblong, exserted. Fruits woody
capsules, pyriform to turbinate, truncated at apex with
a circular calyx scar, with loculicidal basipetal dehiscence
forming 2 valves; seeds minute, attached horizontally,
testa reticulate.

A genus of four or five species ranging from
Costa Rica to Venezuela, Peru, and Bolivia. The
large, often subsessile leaves, deeply two-parted
stipules, lack of pubescence, very large terminal
open-branched inflorescences, greenish flowers,
and woody bivalved capsules with hundreds of
minute seeds make this a distinctive genus.

Condaminea corymbosa (Ruiz & Pav.) DC., Prodr.
4: 402. 1 830. Macrocnemum corymbosum Ruiz
& Pav., Fl. Peruv. 2: 48, pi. 189. 1799. Figure
29.

Shrubs or small trees, 2-5(-7) m tall, leafy branchlets
4-1 2 mm thick, essentially glabrous; stipules apparently
free and 4/node. united above the leaf base to form a
short (14 mm) sheath adnate to the stem (but difficult
to see), the free distal parts equal and 2-6(-9) cm long,
6-9(-l 2) mm broad, lanceolate, reddish brown, glabrous
or rarely minutely puberulent, with many parallel strong-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

107

ly ascending secondary veins, persisting. Leaves with
petioles 0-8 mm long, usually hidden by the auriculate
leaf base, glabrous; leaf blades 15-35(-50) cm long, 6-
14(-21) cm broad, obovate-oblong to oblanceolate-ob-
long or oblong, apex abruptly narrowed and short-acu-
minate, base gradually narrowed and slightly auriculate
or subcordate, drying subcoriaceous, glabrous, 2 veins
15-19/side, the minor (4) venation raised beneath and
forming a subreticulate ("wrinkled") surface. Inflores-
cences open panicles with opposite primary branching,
20-45 cm long and 15-28 cm broad, to 60 cm long in
fruit, primary peduncle 15-23 cm long, 4-12 mm thick,
primary branches 7-1 5(-25) cm long, secondary branch-
es 1-6 cm long, distal branches minutely puberulent,
bracts 1-3 mm long, pedicels 0-4 mm long. Flowers 2-
3 cm long, hypanthium 3-9 mm long, 3-5 mm diam.,
not differentiated from the pedicel, calyx tube 3-6 mm
long and 4-6 mm broad, lobes 4, 5, or none, usually
minute (0-0.5 mm); corolla salverform, white with pur-
plish tube, slightly fleshy, tube 10-15 mm long, 3-5 mm
diam., greenish white within, lobes 4-5, 6-10 mm long,
3-4 mm broad, oblong and rounded distally, becoming
reflexed, glabrous; stamens 5, anthers 4-7 mm long, ca.
1.3 mm broad; ovary 2-locular, style 15-25 mm long,
stigmas 2, 3-5 mm long and 0.7 mm thick, ellipsoid.
Fruits 10-18 mm long, 6-10 mm wide, obovoid to ob-
long-turbinate, drying brown with longitudinal veins
slightly raised, with a pale circular distal calyx scar 0.6
mm broad and 7-8 mm diam., glabrous or minutely
puberulent; seeds 0.5-1 mm long, 0.2-0.4 mm thick.

Small (?short-lived) treelets or large shrubs of
evergreen forest formations, between (10-)500 and
1 800 m elevation. This species has not been col-
lected below 300 m in Costa Rica. Flowering in
January-April and October; fruiting in January-
April. In Costa Rica this species has been collected
only near Turrialba, around the General Valley
Goto Brus region. The species ranges to Venezuela,
Peru, and Bolivia.

Condaminea corymbosa is recognized by the very
large subsessile leaves usually auriculate at the base,
the lack of pubescence on most parts, the large
terminal inflorescences (often with three equal
branches from apex of the peduncle), the woody
two-valved capsules with minute seeds, and char-
acters of the genus (see above). What appear to be
four large and distinct stipules at each distal node
also help to distinguish this species and genus.

C osmibuena Ruiz & Pav.,
nomen conservandum

REFERENCE C. M. Taylor, Revision of Cos-
mibuena (Rubiaceae: Cinchoneae). Ann. Missouri
Bot. Gard. 79: 886-900. 1992.

Small trees or shrubs, epiphytic or less often terrestrial,
stems glabrous and often succulent, becoming terete;
stipules interpetiolar and partly intrapetiolar, forming a
cap over the shoot apex in early stages, obovate to ob-
lanceolate and with many parallel veins, caducous. Leaves
decussate, petioles short and thick; leaf blades entire,
often coriaceous (semisucculent in life), domatia absent.
Inflorescences terminal, solitary, with few (3-1 1) cymose
flowers or the flowers solitary, bracts resembling the stip-
ules, flowers pedicellate. Flowers bisexual, usually large
and fragrant at night, monomorphic, glabrous externally,
hypanthium turbinate to cylindrical, calyx tube decid-
uous (circumscissile) or persistent (often varying within
a species), calyx lobes 5-6(-7) and subequal to strongly
unequal; corolla salverform and carnose, white or tinged
with pink (turning yellow or brown when old), corolla
tube long and slender, corolla lobes 5-6(-7), convolute
or imbricate in bud, rounded distally; stamens 5-6, fil-
aments short and attached near apex of tube, anthers
basifixed and sagittate, included; ovary 2-locular, ovules
many in each locule and borne on axile placentas, ver-
tical and imbricated, style long and slender, often pu-
berulent distally, stigmas 2 and papillate within. Fruits
cylindrical capsules, woody and often with conspicuous
white lenticels, dehiscing septicidally from apex into 2
valves, pericarp often separating from the papery en-
docarp; seeds many, elliptic and flattened, surrounded
by a papery or membranaceous marginal wing, erose to
fimbriate along the edge.

A genus of four species ranging from southern
Mexico to Peru. The large somewhat fleshy flowers
with long tubes, the glabrous coriaceous leaves,
the unusual stipules, the long narrow capsules with
many winged seeds, and the epiphytic habit dis-
tinguish this genus. These plants may be difficult
to distinguish from species of Hillia (seeds with
hairs), Ladenbergia (terrestrial, valvate corolla
lobes), and Posoqueria (fleshy globose fruits).

Key to the Species of Cosmibuena

la. Leaf blades mostly 1.5-3.5( 4) cm wide, narrowly obovate or oblanceolate, apex obtuse, drying
coriaceous and reddish brown to cinnamon brown; corolla tube and exterior of lobes white marked
with pink [capsules 40-100 mm long; 700-2300 m elevation] C. valerii

Ib. Leaf blades 2.5-10 cm broad, elliptic to oblong, apex otbuse to acute, drying coriaceous or sub-
coriaceous and grayish green; exterior of corolla tube and lobes pale green to white 2

108

FIELDIANA: BOTANY

2a. Capsules 40-65 mm long; leaf blades subcoriaceous to coriaceous, acute to obtuse at apex, secondary
veins straight to curved; 200-1000 m elevation c. grandiflora

2b. Capsules 62-1 15 mm long; leaf blades coriaceous, obtuse to broadly rounded at apex, secondary
veins straight; 0-500 m elevation c. macrocarpa

Cosmibuena grandiflora (Ruiz & Pav.) Rusby, Bull.
New York Hot. Gard. 4: 368. 1907. Cinchona
grandiflora Ruiz & Pav., Fl. peruv. prodr. 2: 54,
pi. 198. 1799. Cosmibuena skinneri (Oerst.)
Hemsley, Biol. centr. amer. Bot. 2: 12. 1881.
Buena skinneri 'Oerst., Vidensk. Meddel. Dansk.
Naturhist. Foren. Kjobenhavn 1852: 48. 1853.
Cosmibuena ovalis Standl., Contr. U.S. Natl.
Herb. 18: 137. 1916. Figure 28.

Trees or shrubs to 1 2 m tall, terrestrial or epiphytic,
leafy branchlets 3-6 mm thick, glabrous, quickly becom-
ing pale gray; stipules 8-30 mm long, to 1 2 mm broad,
obovate to oblanceolate, the intrapetiolar tube forming
V-t-Va of the length (reduced below inflorescences), round-
ed apically. glabrous. Leaves with petioles (5-) 1 040 mm
long, 1.2-2 mm thick, drying dark; leaf blades 7-19 cm
long, 4-12(-16) cm broad, broadly elliptic to broadly
oblong or obovate, apex bluntly acute to obtuse or short-
acuminate, base cuneate and slightly decurrent on pet-
iole, drying stiffly chartaceous to subcoriaceous, dark
brown or greenish brown above, glabrous above and
below, 2 veins 3-7/side (in Costa Rica). Inflorescences
with 3-5(-9) flowers, cymose to subumbellate, primary
peduncle 5-30(-40) mm long, 2-3 mm diam., glabrous,
pedicels (5-)10-20(-30) mm long and often merging im-
perceptibly into the calyx. Flowers 7-10 cm long, hy-
panthium 7-1 1 mm long, 4-6 mm diam., drying dark,
calyx tube 3-9(-l 1) mm long, entire or with lobes 0-8
mm long; corolla white, tubular-sal verform, tube 4-9(-10)
cm long, 2.5-5 mm diam., lobes 5-6, (10-)20-35(-40)
mm long, (6-)10-14(-20) mm broad, obovate; stamens
5-6, anthers 10-18 mm long; upper part of style densely
puberulent, stigmas 4-7 mm long, greenish. Fruits (3-)4-
6.5 cm long, 6-13 mm diam., oblong-cylindrical to ob-
long-ellipsoid; seeds 5-8 mm long, 0.5-1 mm broad,
linear fusiform, with thin narrow wing, body of the seed
1-2 mm long.

Trees of evergreen forest formations, 50-1000
m elevation. Flowering in April and July-Decem-
ber; fruits were collected in March. This species
ranges from southern Mexico to Peru.

Cosmibuena grandiflora is recognized by the
general lack of pubescence, large fleshy flowers with
long tubes, and unusual stipules. The thinner leaves
drying dark above and with the secondary veins
readily visible help to distinguish these plants from
material placed under C. macrocarpa. This species
is not often collected in southern Central America.
The flowers are aromatic in the morning and late
afternoon (Herrera 1072 CR).

Cosmibuena macrocarpa (Benth.) KJotzsch ex
Walpers, Repert. bot. syst. 6: 69. 1846. Buena
macrocarpa Benth., Bot. voy. Sulph. 104, t. 38.
1844. C. paludicola Standl., Contr. U.S. Natl.
Herb. 18: 137. 1916. Figure 28.

Shrubs or small trees to 12 m tall, trunks to 25 cm
dbh, epiphytic or terrestrial, leafy branchlets 3-7 mm
thick, smooth and glabrous, becoming pale gray; stipules
12-24 mm long, 6-12 mm broad, forming a tube ca. Vj
of the length but later splitting, obovate to rounded-
oblong, apex rounded to obtuse, glabrous and pale green
drying reddish brown. Leaves with petioles 8-20(-25)
mm long, 1.8-2.8 mm thick, glabrous; leaf blades 6-18
cm long, 2.5-8(-l 1) cm broad, obovate to obovate-ob-
long or subrotund-obovate, apex bluntly obtuse to
rounded, base cuneate and slightly decurrent on petiole,
drying coriaceous, often grayish, glabrous above and be-
low (or puberulous along the midvein beneath in young
leaves), 2 vein 4-5/side but difficult to see. Inflorescence
terminal, subumbellate with 3-8 flowers, peduncles 6-
15(-30) mm long, 2-4 mm diam. (peduncle and inflo-
rescence rachis to 3 cm long), pedicels 7-20 mm long.
Flowers 10-14 cm long, glabrous, hypanthium 6-1 5 mm
long but not clearly differentiated from the pedicel, 2.5-
4 mm diam., calyx tube 2-8 mm long, calyx teeth 0.5-
4 mm long, triangular to minute; corolla salverform and
fleshy, white or pale greenish, tube 5-9 cm long, 2.5-4.5
mm diam., lobes 5, 18-30 mm long, 9-12 mm diam.,
oblong; stamens 5, anthers 12-15 mm long, 1.5 mm
wide, included; style exserted, stigmas 6-9 mm long, 1 .8
mm broad. Fruits (4-)6-8(-l 2) cm long, 6-8 mm broad,
linear-cylindrical to linear-oblongoid, brown, borne on
pedicels 1-2 cm long, disc forming an elevated (2 x 2.5
mm) projection on the truncated apex of the capsule,
surface glabrous and drying dark, often with elongate (3
mm) lenticels; seeds 5-9(-13) mm long and ca. 1 mm
wide, thin, narrowly winged.

Trees of mangrove and evergreen lowland forest
formations on the Caribbean slope, from near sea
level to about 400 m elevation. In Costa Rica the
species probably flowers April-November; fruit-
ing in February-March and July-September. The
species ranges from Costa Rica to Peru.

Cosmibuena macrocarpa is recognized by the
large fleshy glabrous flowers with long tubes, co-
riaceous obovate leaves with rounded apices and
obscure venation, long woody capsules with small
winged seeds, and usually epiphytic habit. The
name Cosmibuena skinneri (Oersted) Hemsley has
been misapplied to this species in the past (cf.
Croat, 1978; Dwyer, 1980).

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

109

Cosmibuena valerii (Standl.) C. M. Taylor, Ann.
Missouri Bot. Gard. 79: 897. 1992. Hillia valerii
Standl., J. Wash. Acad. Sci. 16: 164. 1928. H.
ligulifolia Dwyer, Ann. Missouri Bot. Gard. 67:
218. 1980. H. chiriquiensis Dwyer, loc. cit. 216.
1980. Figure 27.

evations and with Hillia tetrandra with smaller
flowers, more greenish leaves (when dried), and
tufts of hairs at one end of the seed.

Coussarea Aublet

Shrubs or small trees, 2-15 m tall, usually epiphytic,
leafy stems 2.5-7 mm thick, semisucculent, glabrous,
drying reddish brown, older stems grayish; stipules 14-
42 mm long, 6-1 5 mm broad, enlarged beneath the flow-
ers, intrapctiolar and splitting along the sides, oblong to
obovate, bluntly obtuse to rounded distally, becoming
reddish, caducous. Leaves often closely clustered distal-
ly. petioles 3-12(-20) mm long, 1.5-3 mm thick, poorly
denned because of the decurrent leaf margins, glabrous;
leaf blades 3-8(-10) cm long, l-3(-4) cm broad, nar-
rowly obovate to narrowly obovate-oblong or oblanceo-
late, apex rounded to bluntly obtuse, base cuneate and
decurrent on petiole, drying coriaceous and reddish brown
to grayish green, glabrous above and below, 2 veins 3-
6/side, strongly ascending but obscure. Inflorescences
terminal, the flowers solitary, sessile or on peduncles to
3 mm long (and difficult to distinguish from the ovary
base), bracts short (2-1 1 mm) and triangular or long (20-
40 mm) and spatulate (enlarged stipules), caducous or
persisting. Flowers to 12 cm long, glabrous externally,
hypanthium 6-14 mm long, calyx lobes 4-18 mm long,
2-3 mm broad, triangular (when short) to narrowly Un-
gulate (when long), obtuse or rounded distally; corolla
salverform with a long tube, carnose, pale green to white,
pink or reddish where exposed in bud, tube 4.7-9 cm
long, 4-7 mm diam., lobes 5, 17-31 mm long, 10-18
mm broad, rounded distally; stamens 5, anthers 12-15
mm long; stigmas ca. 6 mm long. Fruits 4-10 cm long,
6-10(-14) mm thick, narrowly oblong or tubular, dark
brown with scattered lenticels; seeds 5-6 mm long and
ca. 0.5 mm broad, body of the seed 1.4-2 mm long,
margins erose.

Plants of evergreen montane forest formations,
from 700 to 2300 m elevation. Probably flowering
and fruiting throughout the year. This species
ranges from northwestern Costa Rica to central
Panama.

Cosmibuena valerii is distinguished by its epi-
phytic habit, smaller narrowly obovate stiffleaves,
long fleshy solitary flowers often marked with pink,
and montane habitats. The foliage often dries a
reddish brown. There are unusual variations in
the development of both the calyx lobes and the
floral bracts in this species, but neither seem im-
portant taxonomically. This species may be con-
fused with Cosmibuena macrocarpa of lower el-

Shrubs or small trees, rarely dioecious, glabrous or
less often puberulent, stems usually quadrangular in ear-
ly stages but becoming terete; stipules interpetiolar (in-
trapetiolar and sometimes forming a cap over the shoot
apex in a few species), obtuse to acute (never with aristate
or subulate appendages), deciduous or persistent. Leaves
opposite (rarely 3/node), petiolate or subsessile; leaf blades
entire, domatia present in a few species. Inflorescences
terminal, solitary, usually open paniculate with opposite
branching to elongate thyrsiform, racemiform, umbel-
liform or glomerulate, usually glabrous, flowers often in
distal cymes, bracts absent or minute (< 1 mm), flowers
sessile or pedicellate. Flowers bisexual (rarely unisexual),
hypanthium turbinate or obconic or ovoid, calyx tube
short and usually distally truncated (calyx lobes not clearly
developed or with 3-5 small lobes); corolla salverform
to tubular, white, corolla tube with glabrous throat, co-
rolla lobes 4(-5), valvate in bud, oblong to elongate or
triangular, often carnose; stamens 4(-5), borne near the
mouth or near the base of the tube, anthers subsessile,
linear, included or exserted; ovary 2-locular or incom-
pletely 1 -locular, ovules 2 and sometimes partly united,
erect from a short basal column. Fruits fleshy, coriaceous
or spongy drupes, usually longer than wide (ellipsoid to
globose), usually with only 1 ovule developing, becoming
blue-black or white; pyrene solitary (2), erect, without
or with longitudinal ribs dorsally.

A Neotropical genus of about 100 species, with
the largest number of species in South America.
The genus is characterized by its usually single-
seeded fruit, four-parted (less often five-parted)
white flowers, often bright white inflorescences with
minute bracts and bracteoles or lacking bracts en-
tirely, and whitish infructescences. The inflores-
cences are rarely more than 1 5 cm long. The stip-
ules are triangular to truncated and rarely bilobed.
Many of our species have spongy tissues in the
corolla and fruit; a number flower at night. Species
of this genus can be very difficult to distinguish
from some species of Psychotria, Rudgea, and Fa-
ramea. The pyrenes of Psychotria have hard walls
with ridges, in contrast to the thin smooth walls
of Coussarea. Closely similar species in these other
genera are referred to under individual species be-
low.

Key to the Species of Coussarea

la. Leaf blades conspicuously pubescent beneath, at least along the major veins 2

Ib. Leaf blades glabrous beneath, pubescent only in the vein axils beneath when domatia are present

. 4

110

FIELDIANA: BOTANY

2a. Calyx tube 7-12 mm long with lobes to 3 mm long; inflorescences with 3-15 distantly spaced

flowers, ovary and fruit pilose . c. enneantha

2b. Calyx tube 0.5-2 mm long with lobes to 1 mm long; inflorescences with 15-50 proximate

flowers, ovary and fruit glabrous .3

3a. Leaf blades 11-28 x 5-15 cm; corolla tube 10-13 mm long, distinctly pubescent; in the

Caribbean lowlands, 0-300 mm elevation c. hondensis

3b. Leaf blades 6-19 x 3-8 cm; corolla tube ca. 9 mm long, glabrous or minutely puberulent;

cloud forests, (600-) 1 200-2000 m elevation C. austin-smithii

4a. Leaves subsessile, domatia of tufted hairs or pits often present along the midvein 5

4b. Leaves with petioles usually more than 4 mm long, domatia absent along veins beneath (rarely

present in C. chiriquiensis 7

5a. Corolla usually minutely sericeous, calyx tube 2-3 mm long; domatia usually narrow de-
pressions along the midvein above the vein axils [leaf blades 7-18 x 3-9 C m; Caribbean

lowlands of northern Costa Rica] C impetiolaris

5b. Corolla glabrous or minutely papillate puberulent; calyx tube 1-2 mm long; domatia of shallow

puberulent depressions in the vein axils or absent 6

6a. Inflorescences 3-10 cm long, paniculate with distinct lateral branches; leaf blades 7-16 cm
long, domatia of puberulent pits in the vein axils; Caribbean lowlands of southernmost Costa

Rica C. sp. A aff. curvigemmia

6b. Inflorescences ca. 3 cm long, subcapitate (paniculate but with short, closely spaced, lateral
branches); leaf blades more than 1 5 cm long, domatia present or absent; lowland rain forest

C. sp. B aff. curvigemmia

7a. Flowering portion of the inflorescences elongate-racemiform, distinctly longer than broad ..... 8
7b. Flowering portion of the inflorescences open paniculate, corymbiform to umbelliform or pyramidal,

usually with length equaling breadth 10

8a. Stipules narrowly oblong to linear, to 2 cm long and 4 mm thick, acute at the apex and Ficus-
like, enclosing apices of stems or lateral branches in early stages; leaves usually less than 7

cm broad [corolla 1 5 mm long] C. caroliana

8b. Stipules not linear or narrowly oblong, not resembling the stipules of Ficus, flattened and

enclosing the shoot apex but quickly caducous; leaves usually more than 8 cm broad ... 9

9a. Calyx tube (limb) ca. 2 mm long; fruit 9-17 mm diam., ellipsoid to obovoid; leaf blades

chartaceous, short- to long-acuminate, petioles to 15 mm long; stipules usually bluntly obtuse

at apex C. talamancana

9b. Calyx tube 5-7 mm long; fruits 10-28 mm diam., ellipsoid-oblong; leaf blades subcoriaceous,
short-acuminate or rounded at apex, petioles to 40 mm long; stipules usually broadly rounded

distally C. latifolia

lOa. Plants of montane cloud forest formations (600-) 1 200-2000 m elevation; stipules often persisting

1 Ob. Plants of lowland evergreen formations, 0-600 m elevation; stipules usually caducous 13

1 la. Corolla tube ca. 6 mm long [leaf blades 3-5 m broad, 5-7 major 2 veins]

C. chiriquiensis

1 Ib. Corolla tube 9-18 mm long 12

12a. Leaf blades 2-4 cm broad and 4-6 major 2 veins; Chiriqui Highlands, Panama

C. nebulosa

12b. Leaves usually 3-8 cm broad and with 6-9 major 2 veins; Costa Rica . C. austin-smithii
13a. Leaves usually drying very dark or black, membranaceous to thin-chartaceous; flowers usually

4-parted; stipules often with 2 minute (0.3 mm) lobes on each side C. nigrescens

1 3b. Leaves usually drying greenish or brownish, usually stiffly chartaceous; flowers 5-parted; stipules

various 14

14a. First node of the inflorescence usually with 4 lateral branches; fruits with 2 seeds, dorsal surface

of seed with longitudinal ridges; corolla lobes 3-7 mm long; calyx lobes 0.3-1 mm long

Psychotria eurycarpa

14b. First node of the inflorescence usually with 2 lateral branches; fruits usually single seeded (rarely
2-seeded), seed smooth on the dorsal surface; corolla lobes 6-20 mm long; stipules without lobes;
calyx lobes present or absent C. psychotrioides

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 1 1 1

Coussareaaustin-smithiiStandl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 1286. 1938. Psychotria
tutensis Dwyer, Ann. Missouri Bot. Gard. 67:
434. 1980. Figure 47.

Coussarea caroliana Standl., Field Mus. Nat. Hist.,
Bot. Ser. 22: 178. 1940. C. veraguensis Dwyer,
Ann. Missouri Bot. Gard. 67: 134. 1980. Figure
47.

Shrubs or small trees 3-6(-10) m tall, leafy stems 1.5-
6 mm thick, glabrous or very minutely (0.05 mm) pa-
pillate in early stages, with thickened nodes and longi-
tudinally striate (dried); stipules 4-8 mm long, 4-7 mm
broad, triangular with an acuminate or narrowed apex,
rarely with a small (0.5 mm) U-shaped sinus at the tip,
glabrous, persisting or deciduous. Leaves opposite, pet-
ioles 8-30 mm long, 0.6-2 mm thick, glabrous or mi-
nutely puberulent; leaf blades (5-)7-19 cm long, (2-)3-
8.5 cm broad, elliptic to narrowly elliptic-oblong, ellip-
tic-obovate or narrowly ovate, apex acute to short-acu-
minate, base acute to obtuse and often slightly decurrent
on petiole, drying chartaceous, dark green or dark brown
above, glabrous above, glabrous or minutely (0.1-0.2
mm) puberulent on the veins beneath, 2 veins 6-9/side,
without domatia. Inflorescences 5-10 cm long, equally
broad, open paniculate or umbelliform (3-branched,
rarely with 4 branches from the first node), peduncles
12-20 mm long, 1-2 mm thick, glabrous or very mi-
nutely papillate-puberulent, primary branches 1-3 cm
long and opposite or alternate, flowers cymose or in distal
groups of (1) 2 or 3, bracts absent or minute (sometimes
borne 3-7 mm up along the lateral branches), pedicels
0-8 mm long, purplish. Flowers fragrant, hypanthium
1-1.5 mm long, ca. 1.5 mm diam., obconic-tubular, gla-
brous, calyx tube 0.5-1 mm long and ca. 3 mm broad,
broadly cupulate, calyx lobes to 0.5 mm high (and broad-
ly triangular) or not developed and the calyx entire; co-
rolla salverform, white, glabrous or minutely papillate-
puberulent externally, tube 9-10 mm long (to 1 5 mm in
life?), 1.2-2.2 mm diam., lobes 4(-5), ca. 7 mm long, 2
mm broad and fleshy, narrowly oblong; style branches
1.8 mm long. Fruits 10-12 mm diam., globose, greenish
white with pale longitudinal lines and becoming red,
purple, or black, persisting calyx less than 0.5 mm high.

Plants of evergreen montane cloud forest for-
mations of the Caribbean slopes and continental
divide, from (600-)1200 to 2000 m elevation.
Flowering in June-November (peaking in August);
fruiting in July and November-February. This
species is known only from the Cordilleras de
Guanacaste and Tilaran, the northern slopes of
the Meseta Central (San Ramon-Zarcero), and
above the Rio Reventazon, in northern and central
Costa Rica.

Coussarea austin-smithii is recognized by its
usually open-corymbiform or umbelliform inflo-
rescences without bracts, flowers with short broad-
ly cupulate calyx with poorly developed lobes, long
narrow corolla lobes, and its cloud forest habitat.
This is our only Coussarea species growing above
1 800 m elevation.

Shrubs or small treelets, 1 .5-4(-6) m tall, leafy branch-
lets 1 .2-5 mm thick, glabrous; stipules 1 2-20 mm long,
2-4 mm diam., united and forming a slender cap over
the shoot apex, glabrous, acute at apex, usually splitting
down one side and caducous. Leaves opposite, petioles
4-14(-18) mm long, 1-3 mm thick, glabrous, slightly
sulcate above; leaf blades 8-14(-17) cm long, 3-7(-9.5)
cm broad, elliptic-oblong, narrowly elliptic-oblong, el-
liptic to ovate-elliptic or elliptic-obovate, apex short-
acuminate, base acute, drying chartaceous and usually
grayish green, glabrous above and small linear cystoliths
often visible, glabrous beneath, 2 veins 7-10/side and
weakly loop-connected near the margin. Inflorescences
3-10 cm long, 3-5 cm broad, racemiform or spiciform,
peduncles 1-3 cm long, glabrous, the proximal 2 lateral
branches opposite or subopposite, to 6 mm long, distal
flowers or flower clusters sessile or subsessile, bracts ab-
sent or minute (0.3 mm), pedicels 0-3 mm long. Flowers
glabrous, hypanthium 1-2 mm long (not clearly differ-
entiated from the pedicel or calyx tube), calyx tube 1-
1.5 mm long, 1.8-2.5 mm broad at apex, entire; corolla
white, tube 6-10 mm long, 0.7-2 mm diam., lobes 4, 4-
8 mm long. Fruits 12-15 mm long, 7-10 mm diam.,
ellipsoid-oblong to oblong-obovoid, drying yellowish and
minutely white-lenticellate, the persisting calyx 0.5-1 mm
high.

Plants of evergreen rain forest formations on the
Caribbean slope and central Cordilleras, from 350
to 1500(-1800) m elevation. Flowering in Janu-
ary-April; probably fruiting throughout the year.
The species ranges from the Cordillera de Guana-
caste eastward to the western part of the Cordillera
de Talamanca and western and central Panama.

Coussarea caroliana is recognized by its lower
montane habitat, lack of puberulence, racemiform
inflorescence without bracts, entire calyx tube, and
distinctive fruit. The elongate Ficus-like stipules
forming a cap over the shoot apices are distinctive;
they are often seen at the base of an inflorescence
enclosing the apices of new lateral shoots. None
of our other species of Coussarea have such stip-
ules. The dried leaves are often a characteristic
grayish green beneath. Specimens with smaller
leaves and immature fruits may resemble Rudgea
cornifolia.

Several specimens with larger (18-26 x 6.5-12
cm) leaf blades and larger (30 x 13 mm) fruits on
long (14 cm) infructescences are tentatively placed
here. All are from the Caribbean lowlands: Gray-
urn et al. 8754 CR, Gomez- Laurito 8785 (sterile)
CR, and Opler 340 CR, F. They may represent an

112

FIELDIANA: BOTANY

unrecognized species or merely an extreme form
of C. caroliana.

Coussarea chiriquiensis (Dwyer) C. M. Taylor,
comb. nov. Rudgea chiriquiensis Dwyer, Ann.
Missouri Bot. Gard. 67: 476. 1980.

Shrubs ca. 4 m tall, leafy stems 0.7-4 mm thick, gla-
brous; stipules 2-4 mm long, broadly triangular or
rounded, entire distally or with 2-4 short stiff lobes,
usually with thickened teeth within, the base persisting.
Leaves with petioles 5-16 mm long, 0.5-1.3 mm thick,
glabrous; leaf blades (5-)6-12 cm long, (2-)3-5 cm broad,
ovate-elliptic to elliptic or lanceolate, apex acute to acu-
minate, base obtuse to acute, drying chartaceous, dark
above, glabrous, 2 veins 5-7/side. Inflorescences soli-
tary and terminal, (3-)5-7 cm long, (2-)4-9 cm broad,
peduncles 9-33 mm long, lateral branches of the first
node opposite, often longer than the peduncle and equal-
ing the rachis, glabrous, flowers sessile. Flowers glabrous,
hypanthium ca. 1 mm long and 0.9 mm diam., tubular,
calyx 0.5-1 mm long, broadly cupulate, calyx lobes mi-
nute or broadly triangular; corolla 6 mm long and 1.5
mm diam. in bud. Fruits not seen.

The original description of Coussarea chiri-
quiensis was based on a single collection (Croat
37071 MO) from about 1500 m elevation, above
San Felix, Chiriqui, Panama. The colleters or teeth
within the stipule may have been mistaken for the
distal stipular teeth that distinguish Rudgea, We
tentatively place Burger et al. 10702 (CR, F, MO)
here, which is similar but has pit-domatia and an
entire stipule sheath. It was collected on the Pacific
slope beneath Monteverde, Puntarenas, at ca. 1 400
m elevation. All this material is very similar in
overall appearance to Coussarea nebulosa and
Faramea ovalis.

Coussarea enneantha Standl., J. Wash. Acad. Sci.
18: 282. 1928. Figure 46.

Shrubs or small trees, 1.5-7 m tall, leafy stems 2-4
mm thick, glabrous or with thin pale brownish hairs 0.3-
0.7 mm long; stipules united and forming a short (2-3
mm) sheath with rounded to truncated distal margin.
Leaves opposite, petioles 14-36 mm long, 0.6-1.4 mm
thick, sparsely to densely pilose, leaf blades 9-17 cm
long, 3-8 cm wide, elliptic to elliptic-oblong, apex acute
to long-acuminate with tip 7-14 mm long, base obtuse
or acute, drying thin- to stiffly chartaceous and concol-
orous, glabrescent above, pubescent on the veins be-
neath, 2 veins 8-10/side and weakly loop-connected
near the margin. Inflorescences 2.5-8 cm long, to 8 cm
broad, peduncles to 6 cm long, ca. 0.7 mm thick and
with thin erect hairs 0.3-0.7 mm long, with 3-5 distant

flowers, peduncle often bearing 3 sessile or long-pedi-
cellate flowers (or with 2 lateral flowers and a slender
rachis bearing 3 remote flowers), bracts absent, pedicels
to 2 cm long, pilose. Flowers with hypanthium 1-3 mm
long and densely hirsute with erect or ascending yellow-
ish brown hairs 0.3-0.7 mm long, calyx tube 4-8 mm
long and 1.5-2.5 mm diam., sparsely pubescent, calyx
lobes 4, (3-)4-8 mm long, ca. 2 mm broad, narrowly
oblong; corolla salverform, white, tube 20-30 mm long
and 3-4 mm diam., with ascending hairs ca. 0.7 mm
long, corolla lobes 4, 12-15 mm long and 2 mm wide,
glabrous. Fruit to 3 cm long and 1 5 mm diam., ellipsoid,
densely hirsute, the persisting calyx 10-17 mm long.

Plants of evergreen rain forest formations, from
near sea level to 1000 m elevation. This species
is only known from Panama, but a collection from
the Fish Creek Mountains in Bocas del Toro Prov-
ince suggests that this species may be found in the
Talamanca Valley of Costa Rica. The pubescent
hypanthium, elongate calyx tube, and few-flow-
ered inflorescences with long slender pedicels are
very distinctive, but the flowers appear to vary
greatly in size. The open few-flowered inflores-
cences resemble those of Faramea occidentalis and
F. paucijlora.

Coussarea hondensis (Standl.) C. M. Taylor & W.
Burger, Selbyana 12: 138. 1 99 1 . Psychotria hon-
densisStandLJ. Wash. Acad. Sci. 18: 183. 1928.
P. ostaurea Dwyer & Hayden, Ann. Missouri
Bot. Gard. 54: 143. 1967. Figure 48.

Shrubs or small trees, 2-10(-15) m tall, trunks to 20
cm dbh, leafy stems 3-9 mm thick, with fine soft hairs
0. 1-0.4 mm long or minutely papillate-puberulent; stip-
ules 7-15 mm long and 4-8 mm broad at the base,
triangular to narrowly oblong, puberulent, acute or bi-
dentate with teeth to 2 mm long, often persisting. Leaves
with petioles 10-35(-70) mm long, 1-3 mm thick, pu-
bescent; leaf blades 11-29 cm long, 5-16 cm broad,
broadly elliptic to broadly elliptic-obovate or ovate-ob-
long, apex acute or short acuminuate apex, tip to 10(-14)
mm long, base acute to broadly obtuse, drying stiffly
chartaceous and brownish or greenish, glabrous above,
with short (ca. 0.2 mm) soft hairs on the veins and sur-
faces beneath, 2 veins 7-10/side. Inflorescences 6-15
cm long, to 10(-15) cm broad, paniculate or corymbi-
form with opposite, alternate or clustered lateral branch-
es, peduncles 4-10 cm long, 1 .5-2.5 mm thick, minutely
puberulent, bracts 0.5-1 mm long or caducous, flowers
in distal groups of 1-3, pedicels 0-3 mm long. Flowers
distylousand nocturnal, hypanthium 1-2 mm long, 1.3-
1.7 mm diam., turbinate, minutely puberulent, calyx
tube 0.7-2 mm long, cupulate or spreading and 3 mm
broad, lobes minute or absent; corolla salverform, white,
minutely puberulent externally, tube 10-18 mm long,
0.8-3 mm diam., glabrous within, lobes 4-5, (4-)7-12

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

113

mm long, 1.5-2 mm broad, narrowly oblong, acute; an-
thers ca. 5 mm long. Fruits 12-24(-20?) mm long, ca.
10(-15?) mm diam., ellipsoid-oblong, becoming red-
purple, persisting calyx less than 0.5 mm high.

Plants of evergreen rain forest formations in the
Caribbean lowlands, from near sea level to 500 m
elevation (to 900 m in Panama). Flowering in May-
September; fruiting in June-August and October-
January. The species ranges from Tortuguero in
northern Costa Rica and the Osa Peninsula, south-
ward to Code Province in Panama.

Coussarea hondensis is recognized by the mi-
nute soft puberulence on vegetative and flowering
parts, the large long-petiolate and broadly elliptic
leaves, the few- or many-branched and umbelli-
form inflorescences, broad and usually entire ca-
lyx, and the oblong fruit. An atypical collection
(Burger & Malta 4729 CR, F) with almost glabrous
leaves from above Golfito is tentatively placed
here.

Coussarea impetiolaris J. D. Smith, Bot. Gaz. 37:
418. 1904. Figure 46.

Coussarea impetiolaris is recognized by the sub-
sessile leaves with slightly auriculate leaf bases,
the few-branched glabrous inflorescences, puber-
ulent corolla tubes, and the unusual fruit drying
pale in color and with round or oblong wart-like
lenticels. The longitudinally elongate domatia with
hairs along the sides or hairs along the midvein
are a distinctive feature when present. This species
was misinterpreted in the past to include material
from Guatemala (now segregated as C. imitans L.
O. Williams with more puberulent hypanthium-
calyx and dark green fruit). Coussarea curvigem-
mia Dwyer of central Panama with smaller flowers
is also closely related; see the discussions under
Coussarea spp. A & B aff. C. curvigemmia. Ma-
terial from the Osa Peninsula shows considerable
variation and may indicate that the material placed
under Coussaria sp. B is only an extreme form of
C. impetiolaris; see the discussion under Coussa-
rea sp. B. This species may be mistaken for a
Rudgea.

Coussarea jiminezii J. D. Smith is a species of
Viburnum (Caprifoliaceae).

Small trees to 1 7 m tall, leafy branchlets 1-3 mm thick,
glabrous and drying greenish; stipules 2-4 mm long, tri-
angular, glabrous and coriaceous, deciduous. Leaves sub-
sessile, petioles 1-4 mm long, 0.7-1.8 mm thick, gla-
brous; leaf blades 7-1 8 cm long, 3-9 cm broad, narrowly
to broadly elliptic, elliptic-oblong or oblanceolate, apex
acuminate with tip 5-15 mm long, gradually narrowed
to an acute base and slightly auriculate on the petiole,
leaves drying chartaceous and greenish or grayish, gla-
brous above and below, but with hairs along the edges
of pit dormatia in or near the vein axils beneath, 2 veins
4-7 /side. Inflorescences 3-5 cm long, to 7 cm broad,
paniculate and often with 1 or 2(-3) pairs of opposite
branches and 9-1 5 or more flowers, peduncles 10-26(-40)
mm long, ca. 1.2 mm thick, glabrous, lateral branches
to 1 2(-l 5) mm long, flowers sessile, bracteoles 0.3-1 mm
long. Flowers with hypanthium ca. 2 mm long, calyx
tube little differentiated from the hypanthium, 2-3 mm
long, 2-3 mm diam. distally, glabrous or minutely pa-
pillate-puberulent, calyx lobes 0.2-0.3 mm high; corolla
white, minutely sericeous externally, tube (8-)l 1-16(-20)
mm long, lobes 4, 6-9 mm long, 2 mm broad at the
base; anthers ca. 8 mm long. Fruits 1 5-20 mm long and
14-15 mm diam., broadly ellipsoid and slightly flattened
laterally, surface smooth and pale yellowish white with
distinctive white (becoming brown) tuberculate lenticels
0.5-1.4 mm long, persisting calyx ca. 2 mm high.

Plants of the lowland Caribbean rain forest for-
mations, from 15 to 500 m elevation. Flowering
in March-July; fruiting in January, March, and
June-November. The species is found in the Ca-
ribbean lowlands, the Osa Peninsula, and Panama.

Coussarea latifolia Standl., J. Wash. Acad. Sci. 1 8:
281. 1928. Figure 48.

Small trees, 6-1 5 m tall, perhaps dioecious, leafy stems
4-12 mm thick, glabrous, quadrangular; stipules ca. 5
mm long, rounded distally, glabrous, deciduous. Leaves
opposite, petioles 9-30 mm long, 2-4 mm thick, terete,
glabrous; leaf blades 17-30 cm long, 9-19 cm broad,
broadly elliptic to broadly elliptic-obovate or ovate-el-
liptic, apex abruptly narrowed to the short-acuminate,
tip 4-10 mm long, base obtuse, drying stiffly chartaceous
or subcoriaceous, grayish green, glabrous above and be-
low, 2 veins 7-9/side, domatia absent. Inflorescences
2-6 cm long, 3-6 cm broad, racemose in form with cy-
mose flower clusters on short (6-1 5 mm) lateral branches
or with pedicellate flowers from the central rachis, pe-
duncles 2-30 mm long, 2-3.5 mm thick and glabrous,
bracts minute, pedicels 1-5 mm long. Flowers glabrous,
hypanthium 3-4 mm long, obovoid, poorly differenti-
ated from the calyx tube, calyx tube 5-7 mm long, 4-5
mm diam., calyx lobes not developed; corolla salver-
form, yellowish white, tube ca. 10 mm long (perhaps not
fully expanded), lobes 4, 13-15 mm long; anthers 4-5
mm long. Fruits 25-45 mm long (including persisting
calyx 2-6 mm long), 10-28 mm diam., ellipsoid-oblong,
green with white spots and becoming yellow.

Plants of evergreen rain forest formations of the
Caribbean lowlands, from 5 to 300 m elevation
(to 1 000 m in the central highlands of Panama).
Flowering in April-May; fruiting in July, Septem-
ber-October, and December. The species ranges

114

FIELDIANA: BOTANY

from central Costa Rica (Reventazon valley) to
Colombia.

Coussarea latifolia is recognized by its large
broadly elliptic leaves on prominent thick petioles
and with relatively few secondary veins, the lack
of pubescence, the racemose inflorescences, and
long calyx tube. We have seen only the following
collections from Costa Rica: Grayum et al. 8754
MO, Shank & Molina 4422 F, and Tonduz 9574
us holotype. This species is difficult to separate
from the much more often collected C. talaman-
canum in the absence of flowers or mature fruit.

Coussarea nebulosa Dwyer, Ann. Missouri Bot.
Card. 67: 131. 1980.

Shrubs, ca. 3 m tall, leafy stems 1-5 mm thick, gla-
brous, becoming grayish, terete and smooth; stipules 1-
3 mm long, 2-3 mm broad, with a broad U-shaped sinus
and 2 small (0.7 mm) lobes, deciduous. Leaves with
petioles 4-16 mm long, 0.5-1 mm thick, glabrous; leaf
blades 6-13 cm long, 2-4 cm broad, elliptic-lanceolate
to narrowly elliptic-oblong, apex tapering gradually to
the acuminate, base acute, drying stiffly chartaceous and
dark olive green above, glabrous above and below, 2
veins 4-6/side. Inflorescences solitary and terminal, 2.5-
7 cm long, 1.5-6 cm broad, paniculate with opposite
lateral branches, peduncle 6-46 mm long, 0.7-1.3 mm
thick, glabrous, bracts ca. 2 mm long, subulate, flowers
mostly sessile in distal triads. Flowers glabrous exter-
nally, hypanthium 0.8-1.3 mm long, ca. 1 mm diam.,
calyx tube 0.5-0.8 mm high, lobes 0.2-0.5 mm long;
corolla white, tube 10-1 8 mm long, lobes ca. 6 mm long.
Fruits unknown.

Coussarea nebulosa is a species of the Chiriqui
Highlands known only from near Boquete at about
1200-1600 m elevation. The smaller leaves, un-
usual stipules (for the genus), glabrous parts, and
small inflorescences are distinctive. There are two
other similar small-leaved species in the Chiriqui
Highlands: Coussarea chiriquensis and Faramea
ovalis.

Coussarea nigrescens C. M. Taylor & Hammel,
Selbyana 12: 134. 1991.

Shrubs or small treelets, 2-7 m tall, leafy stems 1.2-
4 mm thick, glabrous or minutely (0.05 mm) papillate-
puberulent, drying dark; stipules united to form a short
sheath 0.8-2 mm long, at first broadly triangular but
becoming truncated or with 2 minute lobes ca. 0.3 mm
long, glabrous, the base persisting as a short collar above
the node. Leaves with petioles 6-35 mm long, 0.7-1.7
mm thick, glabrous or minutely papillate-puberulent.

drying black; leaf blades 8-2 1 cm long, 3-9 cm broad,
elliptic to elliptic-oblong or slightly ovate-elliptic, apex
abruptly narrowed and short-acuminate with tip 2-8 mm
long, base cuneate to obtuse, drying membranaceous to
thin-chartaceous and blackish above, glabrous and lus-
trous above, glabrous or minutely puberulent along the
midvein beneath, 2 veins 6-10/side, weakly loop-con-
nected near the margin in the distal half of the leaf.
Inflorescences solitary and terminal, 4-1 1 cm long, 3-
1 1 cm broad, open umbelliform panicles with 2 or 4
branches at the first node, peduncle 15-50 mm long,
0.7-2.2 mm thick, glabrous or papillate-puberulent, dry-
ing black, bracts subtending the 1 branches 2-6 mm
long, flowers sessile in distal cymes or glomerules of 3-
7, distal bracts ca. 1 mm long. Flowers minutely (0.05
mm) papillate-puberulent externally, hypanthium 0.6-1
mm long, calyx cup only 0.5 mm long, becoming rotate,
lobes minute; corolla tubular, white, tube 14-24 mm
long and 1.2-2 mm diam., lobes 4 or 5, 6-10 mm long,
lanceolate; ovary with well-developed septum and 2 loc-
ules. Fruits 9-26 mm long, 6-18 mm diam., purple-
black, glabrous; pyrenes 1 or 2, globose to ellipsoid.

Plants of evergreen rain forest formations of the
Caribbean lowlands and southern Pacific low-
lands, from 20 to 800 m elevation. Flowering in
July and October-November; fruiting in Febru-
ary. This species has been collected in the de-
partment of Zelaya, Nicaragua, near La Selva and
Puerto Viejo de Sarapiqui, Cerro Nara east of Que-
pos, and Corcovado National Park in Costa Rica.
This species is known only from Costa Rica and
Nicaragua.

Coussarea nigrescens is recognized by the many
parts drying dark or blackish, the very thin leaves,
short collar-forming stipules, and slender corolla
tubes that are minutely papillate-puberulent. This
species may be related to C. nebulosa of the Chi-
riqui Highlands with smaller stiffer leaves that do
not dry so dark, larger calyx lobes, lack of puber-
ulence, and higher-elevation habitat.

Coussarea psychotrioides Taylor & Hammel, Sel-
byana 12: 135. 1991. Figure 47.

Shrubs or small trees, 2.5-7(-12) m tall, trunks to 20
cm dbh, leafy stems 1.5-4 mm thick, glabrous, nodes
often conspicuously thickened: stipules 0.3-2 mm long,
truncate or slightly bilobed, quickly caducous and leav-
ing a short cupulate ring around the stem just above the
distal nodes, glabrous. Leaves with petioles 6-20 mm
long, 0.6-1.7 mm thick, glabrous, sulcate above; leaf
blades 8-18 cm long, (2.5-)3-8.5 cm broad, narrowly
elliptic to elliptic-oblong or elliptic-ovate, apex short-
acuminate (rarely acute or obtuse), tip 5-12 mm long,
base obtuse to acute, drying chartaceous and often yel-
lowish green or greenish brown above, glabrous above
and below, 2 veins 6-9/side and loop-connected near

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

115

the margin in the distal half of the lamina. Inflorescences
3-10 cm long and 4-10 cm broad, broadly paniculate to
umbelliform, peduncles 1-3 cm long, 1-2 mm thick,
glabrous, first branching node usually with 2 branches
and these often coequal with the continuing rachis (=
umbelliform), bracts 0.4-3 mm long or absent, flowers
usually subsessile in distal 3-7-flowered cymes or glom-
erules, pedicels 0-3.5 mm long. Flowers glabrous, noc-
turnal and distylous, hypanthium 0.7-2 mm long, 0.5-
1.5 mm diam., obconic or turbinate, calyx tube 0.2-1
mm long, calyx lobes not clearly developed; corolla white
or tinged with pink, salverform or slightly funnelform,
tube 12-18 mm long, 1-2 mm diam. near the base and
2-3 mm near the mouth, often curved, lobes 5(-6), 6-
12(-20) mm long, 0.8-2 mm broad. Fruits 14-20 mm
long, 10-15 mm diam., ooblong or ovoid and abruptly
rounded at apex and base, blue-black in life, persistent
calyx not elevated or less than 0.5 mm high and 4 mm
diam.; pyrenes 1 or 2, smooth or sulcate adaxially.

Plants of poorly drained areas in lowland rain
forest formations, 50-600(-900) m elevation.
Flowering in February-July; fruiting in Septem-
ber-February. The species is known only from the
Caribbean lowlands of northern Costa Rica and
the Osa Peninsula.

Coussarea psychotrioides is recognized by its re-
stricted lowland habitat, glabrous parts, somewhat
umbelliform inflorescences, and slightly pink flow-
ers with long corolla lobes. The leaves are quite
variable in shape but tend to dry greenish. This
species is common at La Selva, where crushed
leaves are said to have a slight odor of wintergreen.
This species resembles Psychotria eurycarpa, which
has larger stipules and calyx lobes, shorter corolla
lobes, and an earlier flowering period (at La Selva).
This species was studied by Bawa and Beach (1983)
and referred to as Coussarea sp. (voucher JHB
1467); that Beach collection is also the type (ho-
lotype DUKE). Specimens from the Osa Peninsula
often have longer ( 1 6-20 mm) corolla lobes.

Coussarea talamancana Standl.. Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 1288. 1938. Figure 48.

Shrubs or small trees, 2-5 m tall, dioecious, leafy stems
2-8 mm thick, glabrous, often drying pale yellowish green;
stipules 6-16 mm long (to 22 mm below the inflores-
cences), to 1 2 mm broad, united to form a sheath around
the shoot apex, broadly obtuse to rounded distally, gla-
brous, coriaceous, caducous. Leaves opposite, petioles
4-13 mm long, 1-2.5 mm thick, glabrous; leaf blades
12-28(-34) cm long, 4-18(-22) cm broad, broadly ellip-
tic to elliptic-obovate or elliptic-suborbicular, apex
abruptly narrowed to the acuminate, tip 6-18(-25) mm
long, base broadly obtuse to acute, drying chartaceous,
grayish green, glabrous above and below, 2 veins 7-10/
side, domatia absent. Inflorescences 3-7(-10) cm long,
3-5 cm broad, racemose panicles with short opposite

lateral branches, peduncles 5-13 mm long, 1.2-3.5 mm
thick, glabrous, bracts minute or absent, pedicels 0-3
mm long. Flowers functionally unisexual, glabrous ex-
ternally, hypanthium ca. 1-2 mm long (not clearly dis-
tinguished from the calyx tube), calyx tube ca. 2 mm
long, becoming 3-5 mm broad, cupulate, calyx lobes
minute or not developed; corolla salverform, white, tube
4-6 mm long, 1.5-2 mm diam., lobes 4, 4-6 mm long
and 1.2 mm broad, narrowly oblong. Fruits 14-22 mm
long, 9-1 7(-24) mm diam., ellipsoid to obovoid, becom-
ing white with spongy exocarp in final stages; pyrene
solitary.

Plants of evergreen formations of the Caribbean
lowlands and the southern Pacific slope, from near
sea level to 700 m elevation. Flowering in Janu-
ary-July; fruiting in every month but May. The
species ranges along the Caribbean lowlands from
northern Costa Rica to Bocas del Toro Province
in Panama and in southern Puntarenas province.

Coussarea talamancana is recognized by the
large broad leaves on short petioles, glabrous parts,
united ovate-elliptic stipules, short racemiform in-
florescences, smaller unisexual flowers, and larger
spongy-white fruit. The type material (Cooper
10466 F) has long (to 25 mm) narrow drip tips,
whereas some other Costa Rican material has
shorter (to 1 mm) tips, but other characteristics
are very similar and suggest that the material placed
here is conspecific.

Coussarea sp. A aff. C. curvigemmia Dwyer, Phy-
tologia 38: 215. 1978. Figure 47.

Shrubs or small trees, 2-8 m tall, leafy stems 1.5-3.5
mm thick, glabrous, grayish or yellowish green when dry;
stipules 0.5-3 mm long, ovate and rounded distally or
reduced to an entire ridge, glabrous. Leaves subsessile
or with petioles 1-3 mm long, 0.5-1.5 mm thick, gla-
brous; leaf blades 7-16 cm long, 2.5-5 cm broad, nar-
rowly elliptic-oblong to elliptic-obovate, apex acumi-
nate, tip ca. 1 6 mm long, gradually narrowed to the acute
or cuneate base and usually slightly auriculate at the
petiole, drying thin-chartaceous and grayish green, gla-
brous above and below, with tufts of hairs in depressions
(domatia) in the vein axils, 2 veins 4-7/side. Inflores-
cences 4-7 cm long, 2-6 cm broad, paniculate with a
single main rachis and short (3-12 mm) opposite or
subopposite lateral branches, peduncles 1-2.2 cm long,
glabrous and drying pale yellowish, flowers in distal pairs
or triads, bracts subtending the flowers absent or less
than 0.5 mm long. Flowers glabrous externally except
for short (0.1 mm) erect whitish hairs on the ovary,
hypanthium ca. 1 mm long, calyx tube 1-2 mm long,
lobes 0.2-0.5 mm long, narrowly dentate; corolla nar-
rowly tubular-salverform, white, tube 6-10 mm long,
0.7-1.5 mm diam., lobes 4, ca. 6-7 mm long; anthers
4-5 mm long, linear. Fruits not known (probably similar
to those of C. curvigemmia: 10-13 x 5-8 mm, oblong
and slightly flattened, whitish).

116

FIELDIANA: BOTANY

Plants of evergreen rain forest of the Caribbean
lowlands, from near sea level to 300 m elevation.
Flowering in April and June (Barringer et al. 2642
&3615cR,F, Gdmez-Laurito 8388 CR). This spe-
cies is known only from near Suretka in the Tal-
amanca Valley in southern Limon Province.

Coussarea sp. A aff. C. curvigemmia is recog-
nized by the thin subsessile leaves often slightly
auriculate at the base and with weakly defined
domatia, delicate whitish inflorescences, and small
flowers with puberulent ovary and slender gla-
brous corolla tubes. Coussarea curvigemmia, of
central Panama, differs from the material placed
here in having clearly outlined ellipsoid pit-doma-
tia with few or no hairs, glabrous ovaries, and
minutely papillate-puberulent corollas that dry
dark. Both taxa have distinctive thin, slightly
curved corolla tubes, and it may be that they are
conspecific. This material also resembles Cous-
sarea impetiolaris with more robust inflorescences
and some species of Faramea.

Coussarea sp. B aff. C. curvigemmia. Figure 48.

Another species with very similar flowers may
be represented by Burger & Gentry 8960 F from
the Osa Peninsula and Bunting & Licht 793 F from
the lower Rio San Juan, Nicaragua. Both of these
collections have smaller (4 cm) compact capitate
inflorescences and larger (20 cm) subsessile ob-
ovate leaves with long (20 mm) narrow drip tips
and with slightly auriculate leaf bases. However,
the Nicaraguan collection has a pedunculate inflo-
rescence and the Osa collection has a subsessile
inflorescence, which may be immature. In con-
trast, Liesner 3225 and Hammel et al. 18604 (all
at CR) are intermediate with typical C. impetiolaris
and indicate that the unusual specimens may be
bridged by intermediates, in which case the de-
scription given for C. impetiolaris needs to be ex-
panded to include the collections placed here.

Coutarea Aublet

REFERENCE A. Aiello, A reexamination of
Portlandia (Rubiaceae) and associated taxa. J. Ar-
nold Arbor. 60: 38-124. 1979.

Shrubs or small trees, branchlets terete, glabrous or
puberulent, often with conspicuous elongate lenticels;
stipules interpetiolar, short and acute, persisting. Leaves
decussate or somewhat distichous, petiolate; leaf blades
chartaceous, entire, some species with domatia. Inflo-

rescences terminal or apparently axillary (terminal on
short axillary shoots with poorly developed leaves), flow-
ers usually in open cymose groups of 3 or solitary, pe-
duncles short, pedicels subtended by narrow bracts.
Flowers bisexual, monomorphic, large and showy, bi-
laterally symmetrical due to curvature of the corolla tube
and asymmetric stamens, hypanthium turbinate, calyx
lobes 5-6(-8), narrow, often unequal: deciduous; corolla
funnelform to campanulate and often inflated on the
lower side, white to rose or purple, corolla tube slightly
curved, with a glabrous throat, corolla lobes 5-6(-8),
imbricate in bud; stamens 5 or 6, inserted near the base
of the corolla tube, filaments long and often twisted in
bud, anthers basi fixed, linear, exserted or included: ovary
2-locular, placentas borne on the septum, ovules many
in each locule. Fruits capsules, ovoid to obovoid or ob-
long, flattened, coriaceous or woody, 2-locular, dehiscing
loculicidally from the apex (down the center of the broad
face of the capsule) to form 2 valves; seeds many, im-
bricate and ascending, flattened and broadly winged with
a thin margin around the circumference.

A genus of 6-10 species ranging from southern
Mexico to Argentina; only 1 species is found in
Mexico and Central America. The large curved
corolla tubes, long free filaments, and the capsules
splitting down the middle of their flattened sides
make the genus distinctive. Only a few species of
Rubiaceae in our flora have similarly large flowers
(cf. figs. 15 and 31).

Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl.
Bras. 6, pt. 6: 196. 1889. Portlandia hexandra
Jacq., Enum. PI. Carib. 16. 1760; Sel. Stirp. 63,
pi. 1 82, f. 20. 1 763. C. speciosa Aubl., PI. Guian.
1: 314, pi. 122. 1775. Figure 31.

Shrubs or small trees, 2-8(-18) m tall, leaf branchlets
1.7-4 mm thick, glabrous or sparsely puberulent, be-
coming dark brown with elongate whitish lenticels; stip-
ules 1.5-4(-5) mm long, ca. 2 mm broad at the base,
intrapetiolarand forming a short (0.5- 1.5 mm) tube above
the petioles, distally triangular and acute, glabrous in
Central America. Leaves with petioles 2-10(-15) mm
long, 0.7-1 .5 mm wide, glabrous or minutely puberulent;
leaf blades 5-1 5 cm long, 2-9 cm broad, ovate to broadly
elliptic or ovate-oblong, apex acute, short-acuminate or
caudate-acuminate, base obtuse to rounded and sub-
truncate (acute), drying thin -chartaceous or membra-
naceous, glabrous or minutely puberulent on the veins
above, glabrous or minutely (0.2-0.4 mm) puberulent
beneath, 2 veins (4-)6-10/side, with tufts of minute
hairs (domatia) in vein axils beneath. Inflorescences few-
branched and with (l-)3-9 flowers, peduncles 3-30 mm
long, bracts 3-5 mm long, pedicels 2-1 5 mm long, merg-
ing gradually with the base of the flower, glabrous or
sparsely and minutely puberulent. Flowers to 10 cm long
and 4 cm broad, mostly glabrous (in Central America),
hypanthium 4-7 mm long, 2-3 mm diam., calyx tube

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

117

1.5-2.5 mm long, 4-5 mm broad, calyx lobes 4-6, 4-
9(-12) mm long, lanceolate to linear, deciduous; corolla
funnelform-campanulate, white or tinged with pink, gla-
brous externally, tube 45-80 mm long, 1 0-20 mm diam.
distally, slightly curved, inflated before anthesis, lobes
4-6, 1-2 cm long, 7-10 mm broad, apex ovate-triangular
and obtuse to acute; stamens 6, filaments to 8 cm long,
anthers 14-19 mm long, 0.5-1 mm thick, yellow, ex-
serted; style to 85 mm long. Fruits 25-45 mm long, 15-
28 mm broad, ca. 8 mm thick, oblong-obovate, woody
when mature, surfaces dark brown and glabrous, with
or without small white punctate lenticels, the valves
slightly split in 2 at apex; seeds 7-14(-20) mm long, 6-
9 mm abroad, oblong to suborbicular, body of the seeds
3-4 mm diam., wing pale brown.

Trees and shrubs of both deciduous and ever-
green forest formations, from near sea level to 900
m elevation. Flowering in late June-October;
fruiting in November-April in southern Central
America. This species appears to be much more
common in seasonally deciduous forests than in
evergreen forests in Costa Rica. The species ranges
from southern Mexico and Belize to Argentina.

Coutarea hexandra is recognized by its thin
short-petiolate leaves, few-flowered inflorescences
with large flowers, the curved corolla tube, and the
large flattened capsules with winged seeds. The
fruits are unusual in that they split down the center
of the broadly flattened halves (fig. 31). The flow-
ers appear to be filled with gas before anthesis, and
they are often pendulous at anthesis. The floral
biology has been discussed by Haber and Frankie
(1989). In northern Central America, bitter prin-
ciples in the bark have been used medicinally,
especially for malaria (Mabberley, 1987).

Crusea Schlectendal & Chamisso

REFERENCE W. R. Anderson, A monograph of

the genus Crusea (Rubiaceae). Mem. New York
Bot. Gard. 22: 1-128. 1972.

Annual or perennial herbs, sometimes woody at the
base, decumbent or erect, stems terete or 4-angled with
longitudinal ribs, pubescent; stipules interpetiolar and
united to adjacent petioles to form a thin sheath, with a
distal cross- vein and 2-12 setae, persisting. Leaves op-
posite and decussate, sessile or petiolate, petioles adnate
to the stipular sheath; leaf blades mostly ovate to lan-
ceolate with strongly ascending pinnate venation, entire,
domatia absent. Inflorescences terminal or axillary, cap-
itate, verticillate or of congested dichasial cymes, sessile
to long-pedunculate, subtended by 2, 4, or 8 (more) leaf-
like bracts, pedicels short (to 2 mm) or absent. Flowers
bisexual and radially symmetrical, homostylous, usually
4-parted, calyx lobes 4 or reduced to 2-3, with minute
glands in the base of the sinuses between the lobes; co-
rolla funnelform to campanulate, white to pink, red, or
purple, corolla lobes 4, valvate in bud; stamens 4, fila-
ments adnate to the upper half of the tube and free be-
neath apex of the tube, anthers dorsifixed, exserted; ovary
2-locular, placenta elongate from the center of the sep-
tum, with 1 ovule in each locule, stigma 2-lobed or sub-
capitate. Fruits of 2 dry 1 -seeded mericarps (cocci) borne
on the sides of and separating from a persisting bifid or
fenestrated carpophore, mericarps indehiscent and ecos-
tate, calyx dehiscing circumscissily or persisting; seeds
with the persisting placenta on the adaxial face.

A genus of 13 species ranging from Arizona,
U.S.A., through Mexico and Central America to
western Panama. The herbaceous habit, congested
subsessile flowers, often lanceolate leaves with
strongly ascending secondary veins, narrow co-
rolla tube (in our species) and unusual fruit help
distinguish this genus. The mericarps are easily
mistaken for seeds because of their smooth round-
ed brownish surfaces and longitudinal adaxial sul-
ci. The genus reaches its southern limit in Costa
Rica and western Panama and is represented by
only a few collections from Costa Rica. This treat-
ment is based on Anderson's detailed monograph.

Key to the Species of Crusea

la. Flowers bright pink to magenta; leaf blades 1-5 cm broad, to 13 cm long 2

Ib. Flowers white or white-tipped with pink; leaf blades 0.5-3.5 cm broad, to 8 cm long 3

2a. Corolla tube 20-38 mm long, stigma lobes 1-4 mm long; secondary veins arising from the
proximal half of the midvein; 1 800-3000 m elevation C. coccinea

2b. Corolla tube 5-18 mm long, stigma lobes 0-0.5 mm long; secondary veins arising from the

proximal '/: or 2 /3 of the midvein; 0-200 m elevation C. hispida

3a. Corolla tube 5-1 1 mm long, stigma lobes 0.2-0.6 mm long; plants usually found around 2000 m

elevation in Costa Rica C. longiflora

3b. Corolla tube 1.84 mm long, stigma lobes 0.1-0.3 mm long; plants found in deciduous and partly

deciduous areas below 1 200 m elevation C. parviflora

118

FIELDIANA: BOTANY

Crusea coccinea DC., Prodr. 4: 567. 1830. C. coc-
cinea var. chiriquiensis W. R. Anderson, Mem.
New York Hot. Card. 22: 45. 1972. Figure 31.

Decumbent or low perennial herbs to 1 m tall, often
rooting at the nodes, leafy stems 1-3.5 mm thick, gla-
brous or sparsely puberulent, quadrangular or terete;
stipule sheath 4-14 mm long (including the setae), 1-6
mm wide, glabrous or puberulent, setae 3-12 and to 12
mm long and linear, evenly spaced or in a central group
with fused bases. Leaves with petioles 4-20(-25) mm
long, glabrous or puberulent; leaf blades 2.2-10(-l 3) cm
long, l-4(-5) cm broad, narrowly elliptic to elliptic or
ovate, apex acute to short- or long-acuminate, base acute
and decurrent on petiole, drying stiffly chartaceous and
dark, often slightly scabrous with the major veins im-
pressed above, glabrous or sparsely puberulent above,
sparsely hispidulous with short (0.3 mm) hairs along the
veins beneath, 2 veins 3-6/side, strongly ascending, sub-
parallel and arising from the proximal half of the mid-
vein, minutely punctate on both surfaces. Inflorescences
bracteate/involucrate heads or with verticillate flowers
in the node below the terminal head, 10-15 mm diam.,
with 15-100 flowers in the head but few-10 flowering at
the same time, involucral bracts 2 or 4, 2-4 cm long,
leaf-like and often sessile within the expanded petioles
and st i pular sheath of the subtending node, often with
additional smaller bracts. Flowers with hypanthium 1 .4-
3 mm long, glabrous or puberulent distally, calyx lobes
to 8 mm long, narrowly triangular, corolla deep red to
pink, purple, or lavender, funnelform, tube (8-)20-38
mm long, gradually expanded from a narrow (3 mm)
base, lobes (3.5-)5-l 1 mm long; stamens with free por-
tion of the filaments ca. 4-10 mm long, filiform, anthers
(2-)2.5-3.8 mm long; style as long as the corolla tube.
Fruits with a broad flat carpophore, 4-7 mm long with
lobes 0.7-2 mm long, cocci (1.7-)3-6 mm long, 2-2.7
mm broad, rounded-turbinate to turbinate, brown, calyx
often coming off as a complete whorl (circumscissile).

Crusea hispida (Miller) Robinson, Proc. Amer.
Acad. Sci. 45: 409. 1910. Crucianella hispida
Miller, Gard. Diet. ed. 8, no. 4. 1768.

Erect annual herbs to 0.6 m tall, many-branched, leafy
stems 0.5-4 mm thick, terete with stiff" unicellular trans-
parent or whitish hairs 1.5-3 mm long, spreading or
retrorse from a thickened base, smaller (0.2 mm) hairs
sometimes also present; stipule sheath 2-5 mm long, 4-
10 mm broad, with 3-7 setae, conspicuously hispid.
Leaves opposite, often with smaller axillary leaves from
the same node, petioles 5-25 mm long, hispid; leaf blades
4-9(-l 1) cm long, l-3.6(-4.8) cm broad, narrowly ovate-
elliptic to lanceolate or narrowly ovate, apex tapering
gradually and acuminate, base acute to obtuse, drying
thin-chartaceous, both surfaces with thin whitish hairs
ca. 1.3 mm long, 2 vein 5-6/side, strongly ascending.
Inflorescences solitary terminal bracteate capitulae with
40-100 closely crowded sessile flowers, subtended by 8
(4) leaf-like bracts and many linear hispid bracteoles.
Flowers with glabrous hypanthium, calyx tube 0.3-1 mm
long, lobes 2-6 mm long, subulate, margins with stiff
hispid hairs; corolla salverform, light pink to dark red
or purple, tube 5.5-12(-18) mm long, ca. 0.3 mm diam.
(dried), lobes 2-5 mm long; anthers 1-1.5 mm long,
exserted. Fruits with cocci 2-3.5 mm long, 1.1-2.3 mm
broad, yellowish to dark brown.

A species of open grassy sites, ranging from
Mexico to El Salvador and collected only recently
in Costa Rica at a single locality: Costa de Pajaros,
Bahia de Nicoya, Puntarenas (C. M. Taylor 249
and Wilbur 31715, both at DUKE). This population
was in flower in July; it is variety hispida (with
shorter corolla tubes). The broader petiolate leaves,
colorful flowers, and unusual pubescence distin-
guish this species.

Herbs of montane evergreen forest formations
of the Pacific slope and central highlands, from
1 800 to 3000 m elevation. Probably flowering and
fruiting mostly in the wet season and beginning of
the dry season: May-January. The species ranges
from western Mexico to western Panama.

Crusea coccinea is recognized by the herbaceous
habit, short stipular sheath with long narrow setae,
narrow leaves with subparallel secondary veins,
and large pink to lavender flowers. Plants of Costa
Rica and Chiriqui, Panama, belong to variety chi-
riquensis W. R. Anderson. This variety is similar
to variety coccinea in having larger corollas, an-
thers, and mericarps, but variety chiriquensis dif-
fers in having leaves with major veins deeply im-
pressed above, a greater number of stipular setae,
and pink to magenta flowers. While often collected
in the Chiriqui Highlands, these plants have rarely
been collected in Costa Rica.

Crusea longiflora (Willd. ex Roem. & Schult.) W.
R. Anderson, Mem. New York Bot. Gard. 22:
89. 1 972. Spermacoce longiflora Willd. ex Roem.
& Schult., Syst. Veg. 3: 531. 1818. C. brachy-
phylla Schlechtend. & Cham., Linnaca 5: 165.
1830.

Erect annual herbs to 50 cm tall, stems terete or less
often quadrangular, pubescent to pilose with hairs 1-2
mm long; stipule sheath 1 .4-6.5 mm long, 2-7 mm broad,
with 3-7 distal setae, and 0-2 inconspicuous sessile lat-
eral colleters, longest setae 0.5-3(-5) mm long, equaling
or shorter than the sheath, with thin hairs 0.5-2 mm
long. Leaves sessile or subsessile with petioles to 5 mm
long; leaf blades 8-50 mm long, 3-2 1 mm wide, narrowly
to broadly elliptic, or ovate, apex acute to obtuse, base
abruptly narrowed, drying chartaceous and scabrous,
sparsely hispidulous, 2 veins 2-3/side, strongly ascend-
ing. Inflorescences 1-2 cm diam., small terminal brac-
teate heads (or with lateral heads reduced to an axillary
cluster of 1 or a few flowers), with up to 75(-l 00) flowers

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

119

in a head, involucral bracts 4 or 8. Flowers sometimes
cleistogamous and resembling small unopened flower
buds, hypanthium 0.7-1 . 1 mm long, glabrous, calyx tube
0.5-1 mm long, lobes 1-3.5 mm long and 0.2-0.8 mm
wide, broadly to narrowly triangular, often ciliate along
the edge; corolla white (rarely pink or lavender), tube 5-
1 1 mm long, narrow at the base and only 0.8 mm broad
at apex, papillose externally, with long straight hairs within
distally, lobes 1 .6-3.6 mm long, 0.8-1 .5 mm broad, nar-
rowly elliptic, becoming strongly reflexed, with few long
hairs at the base within; stamens with filaments 1.5-3.8
mm long, anthers 0.6- 1.3 mm long; style 5-16 mm long,
glabrous, stigmatic lobes 0.2-0.6 mm long. Fruits with
a bifid carpophore to 1 .5 mm long, the mericarps (cocci)
1.1-2.3 mm long, ellipsoid-cylindrical, whitish to yel-
low-brown or dark brown, calyx often coming off as a
whorl.

Herbaceous plants of montane evergreen for-
mations, from ca. 1000 to 2700 m elevation. Flow-
ering in July-December; fruiting in September-
December (over the entire range). This species
ranges from northern Mexico through Guatemala,
with isolated occurrences in the highlands of Hon-
duras and Costa Rica.

Crusea longiflora is distinguished by its erect
annual habit, short stipular sheath with setae, small
and narrow (often subsessile) leaves, the narrow
corolla tube, and the small rounded mericarps.
The long (1-2 mm) slender unicellular hairs also
help distinguish this species. It has been collected
infrequently in the Central Highlands at about 2200
m elevation. This species is common in Mexico.

Crusea parviflora Hook. & Arnott, Bot. Beechey
Voy. 430, pi. 99. 1840. Figure 4.

Erect, trailing or decumbent herbs to about 50 cm high
and 1 m long, annual or perennial, leafy stems 0.5-3.5
mm thick, with 4 longitudinal ridges or wings in early
stages, with thin whitish hairs 0.3-1 .2 mm long but often
glabrescent; stipule sheath 1-3 m long, 1.5-4 mm wide,
often with minute spots, with 3-5 setae 1-7 mm long
and with hairs to 1 mm long. Leaves with petioles 3-
9(-15) mm long, with lateral wings continuous with the
lamina margin, with thin hairs on both surfaces; leaf
blades 2.4-7(-9) cm long, 1-3.6 cm broad, narrowly to
broadly elliptic or lanceolate, apex bluntly to sharply
acute, base gradually (abruptly) narrowed and decurrent
on petiole, drying thin-chartaceous and often grayish
green, pilose on both surfaces with thin whitish hairs
0.3-1 mm long, 2 veins 3-5/side. Inflorescences brac-
teate heads, with 10-many flowers per head and a ma-
jority of the flowers in bloom at one time, the heads
terminal or axillary to distal leaves, subglobose and 1-
2 cm broad, often on long (to 1 5 cm) peduncles, invo-
lucral bracts to 3 cm long and leaf-like, pedicels short.
Flowers with small (0.5-1 mm) glabrous hypanthium,
calyx tube 0.5-1.3 mm long, lobes 0.6-2.5 mm long,
0.2-0.5 mm broad and linear-triangular, with thin hairs

on the outer surface, calyx persisting on the fruit; corolla
white or the lobes tipped with pink, funnelform, tube
\.8-4 mm long, 0.7-1.4 mm broad at the throat, usually
glabrous externally and with longer hairs on the distal
portion inside, lobes 1.3-2.8 mm long, 0.5-1 mm broad,
narrowly triangular, erect to spreading (not reflexed),
usually with small hairs externally at the tips; stamens
with filaments 1.3-4.5 mm long, glabrous and becoming
retracted into the corolla after anthesis; style 2.7-8 mm
long, glabrous, stigmatic lobes 0.1-0.3 mm long. Fruits
with mericarps 0.8-1.2 mm long, 0.6-0.9 mm broad,
oblong-cylindrical or subglobose, surface slightly pitted.

Weedy plants of deciduous and evergreen for-
mations, from 50 to 1600(-2000) m elevation in
Central America. Flowering in October-April;
mature fruits in November-April. The species
ranges from western Mexico along the Pacific slope
to isolated localities in Honduras, Nicaragua, and
Costa Rica.

Crusea parviflora is recognized by its short weedy
habit, quadrangular or slightly winged young stems,
short stipular sheath with few setae, capitate in-
florescences with usually only 4 broad bracts, and
unusual carpophore and seed-like mericarps. The
short-pedicellate flowers contrast with the sessile
flowers of similar-looking species of Mitracarpus
and Spermacoce. In our area it has only been found
in Guanacaste Province below 500 m elevation.

Declieuxia Humboldt, Bonpland & Kunth

REFERENCE J. H. Kirkbride, Jr., A revision of
the genus Declieuxia (Rubiaceae). Mem. New York
Bot. Gard. 28: 1-87. 1976.

Herbs or subshrubs, perennial and often woody at the
base, branches terete or angular, glabrous or puberulent;
stipules interpetiolar, subulate or reduced to a line bear-
ing 1-3 linear lobes. Leaves opposite or whorled, sessile
or short-petiolate, leaf blades entire and usually small,
linear to elliptic, deltoid or orbicular, usually coriaceous,
domatia absent. Inflorescences of terminal or axillary
panicles (compound cymes), often spike-like or race-
miform, solitary to 3 at the end of the stem, pedunculate,
branching often dichotomous, flowers in distal cymose
groups, bracts and bracteoles present or absent. Flowers
bisexual and radially symmetrical, small, mostly gla-
brous, hypanlhium turbinate to subglobose or obovoid,
slightly compressed laterally, calyx lobes 4 (2), equal or
unequal, small and persisting; corolla funnelform to tu-
bular, white to blue or purple, tube pilose-villose in the
throat, corolla lobes 4, short, spreading or reflexed, val-
vate in bud; stamens 4, inserted in the corolla throat or
between the lobes, filaments slender, anthers dorsifixed
and versatile, partly or completely exserted; ovary
2-locular, ovule solitary in each locule, erect from a near-
basal placenta, style slender, stigmas 2. Fruits drupa-
ceous, dry or fleshy, laterally compressed and 2-parted
or with 2 prominent rounded lobes, black at maturity.

120

FIELDIANA: BOTANY

A genus of about 40 species in tropical America,
with the largest number of species in Brazil. Our
representative of this genus is recognized by the
very short woody stems, stiff subsessile leaves, cy-
mose branching of the short inflorescences, two-
lobed fleshy fruits, and restriction to open grassy
habitats in deciduous or partly deciduous forma-
tions.

rocky or savanna-like habitats, the stiff narrow
sessile leaves often three at a node, the short di-
chotomously branched panicles, the small flowers,
and the sessile fleshy deeply two-lobed or rounded
fruit (laterally compressed when dry). Chacon
(2258 CR, MO) stated that the flowers are white
with blue-lavender stamens. Our material belongs
to variety mexicana, which is distinguished by its
narrow leaves and lack of pubescence.

Declieuxia fruticosa (Willd. ex Roem. & Schult.)
Kuntze, Rev. gen. pi. 1: 279. 1891. Houstonia
fruticosa Willd. ex Roem. & Schult., Syst. Veg.
3: 527. 1818. D. mexicana DC, Prodr. 4: 479.
1830. D. fruticosa var. mexicana (DC.) Standl.,
Field Mus. Nat. Hist., Bot. Ser. 12: 378. 1936.
Figure 1.

Erect subshrubs or herbaceous 20-70 cm tall, often
from a hard woody rootstock, with vertical simple or
few-branched stems, leafy stems 0.7-3 mm thick, gla-
brous (in variety mexicana) to pubescent, with 2 or 4
longitudinal ridges or wings (0.5 mm high), becoming
terete; stipules 2-5 mm long, linear to linear-subulate,
glabrous (in variety mexicana) or pubescent, deciduous.
Leaves 2 or 3/node, sessile or subsessile (petiole to 1 mm
long); leaf blades 20-^0(-50) mm long, 4-15(-22) mm
broad, narrowly elliptic-oblong to linear-oblong or ob-
long, apex acute or obtuse, base cuneate, drying stiffly
chartaceous to subcoriaceous, glabrous and often lus-
trous above, glabrous (in variety mexicana) or puberu-
lent beneath, 2 veins (2-)3-6/side. Inflorescences ter-
minal or axillary, 1-4 cm long, 1.5-5 cm broad, solitary
or 3 at the ends of branchlets, open cymose paniculate
with dichotomous nodes (and a sessile flower at the di-
chotomy), primary peduncle 5-15(-40) mm long, bracts
1.5-3 mm long, linear, pedicels 0-1 mm long. Flowers
heterostylous, hypanthium 0.3-1 mm long, calyx lobes
0.3-0.8 mm long, linear-oblong, glabrous (in variety
mexicana) to villous, corolla white to blue or purple,
4.5-6 mm long, tube 3-4.5 mm long, cylindrical, lobes
4, ca. 2 mm long; anthers ca. 1 mm long, linear; style
3-4.5 mm long, stigmas 0.3 mm long. Fruits slightly
fleshy and prominently 2-lobed (when both ovules de-
velop), ca. 2 mm long and 3 mm broad, sessile, the lobes
suborbicular, the fruit subglobose when only 1 ovule
develops and ca. 2 mm diam., drying black, glabrous.

Small subshrubs of open grassy savanna-like sites
in deciduous and semideciduous forest forma-
tions, from ca. 20 to 1200(-1800?) m elevation.
In Costa Rica the species is restricted to the Pacific
slope and is common in Guanacaste and the Bue-
nos Aires area of the General Valley. Flowering
throughout the year (primarily in July-Septem-
ber). The species ranges from southern Mexico to
Brazil.

Declieuxia fruticosa is recognized by the short
stature from a woody base, restriction to open

Deppea Chamisso & Schlechtendal

REFERENCE D. H. Lorence and J. D. Dwyer, A
revision of Deppea (Rubiaceae). Allertonia 4: 389-
436. 1988.

Slender shrubs and small trees, stems sparsely to densely
puberulent; stipules interpetiolar, small, triangular, per-
sistent. Leaves opposite (rarely in whorls of 3), opposing
leaves of the same node often unequal, petiolate, leaf
blades entire and pinnately veined, drying thin-charta-
ceous, domatia sometimes present. Inflorescences axil-
lary or terminal, scorpioid, umbellate or corymbiform
to thyrsoid, the flowers in cymose groups (or rarely of
solitary flowers), pedunculate, flowers pedicellate, brac-
teolate. Flowers bisexual and radially symmetrical, hy-
panthium hemispheric to turbinate or cylindrical, calyx
lobes 4, minute or large, equal or unequal, usually with
a small gland in each sinus; corolla short-funnelform to
salverform, yellow (less often white, orange, or purple),
corolla tube glabrous within, usually shorter than the
lobes, corolla lobes 4, spreading or erect at anthesis,
comoluatc in bud; stamens 4, inserted near the base of
the tube, filaments short and linear, anthers dorsifixed,
oblong to narrowly ellipsoid, exserted or partly included,
a nectariferous disc present; ovary 2-locular, placentas
elongate and peltate on the septum, ovules many in each
locule and longitudinally imbricate, style slender, stigma
entire or bilobed. Fruits a small dry capsule, turbinate
to obovoid, coriaceous to chartaceous, usually with (6-)8
longitudinal costae, bisulcate and dehiscing loculicidally
from apex, valves cleft, calyx persisting; seeds many and
minute, angulate, testa foveolate and reticulate.

A genus of about 25 species, centered in Mexico
and ranging through Central America ( 1 species)
to southeastern Brazil ( 1 species). This account is
based on the recent revision by Lorence and Dwyer
(see reference above). Deppea can be confused with
Hamelia and Hoffmannia, which have fleshy fruits.

Deppea grandiflora Schlcchtend., Linnaea 19: 748.
1847. D. costaricensis Polak., Linnaea 41: 566.
1877. D. floribunda Hemsl., Diagn. PI. Nov.
Mexic. 3 1 . 1 879. D. longipes Standl., Contr. U.S.
Natl. Herb. 18: 138. 1916. Figure 38.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

121

Shrubs or small trees, 1-4 m tall, leafy branchlets 1-
4 mm thick, hirtellous with short (0.3 mm) brownish
hairs, at first with longitudinal ridges but becoming terete
and glabresent; stipules 0.5-1 mm long, 2-6 mm wide,
broadly deltoid, glabrate to densely hirtellous, the inner
surface or margin with 4-8 dark brown digitate colleters.
Leaves opposite and subequal or unequal at the same
node (with 1 up to 2 times as long as the other), petioles
6-35(-50) mm long, 0.5-2 mm thick, glabrate or densely
hirtellous on the adaxial side; leaf blades (3-)5-17(-21)
cm long, (1.5-)2-7(-8.5) cm broad, elliptic to narrowly
elliptic, narrowly ovate or lanceolate, apex tapering grad-
ually and acuminate, base attenuate to acute (obtuse),
drying chartaceous, glabrous to sparsely puberulent with
minute (0.2 mm) thin whitish hairs above, minutely
strigillose along the veins beneath, occasionally with tuft-
ed domatia in the vein axils beneath, 2 veins 6-1 I/side
and weakly loop-connected near the margin. Inflores-
cence terminal (rarely axillary), 3-12 cm long and 3-15
cm broad, dichasial and corymbiform, with 15-100
flowers, peduncles 1.2-7 cm long, densely and minutely
strigillose-hirtellous, primary branches 1-3 cm long and
with up to 3 additional orders of branching, distal cy-
mules of 2-6 flowers, pedicels 1-4 mm long, bracteoles
present or absent. Flowers with hypanthium 1-2 mm
long, 0.5-1 mm diam.. obconic or turbinate, with lon-
gitudinal costae, calyx cup 0. 1-0.2 mm deep, lobes 0.3-
1 mm long, deltoid; corollas funnelform or rotate, yellow,
glabrous, tube 1-2 mm long, corolla lobes 4, 47 mm
long, 3-4 mm wide, obtuse; stamens 4, filaments 2 mm
long, anthers 3-4 mm long, exserted, basally sagittate;
style 3.5-4.5 mm long, stigmas 1.5-2 mm long, entire.
Fruits 2-5 mm long, 2-4 mm diam., obconical to subglo-
bose, with 6-8 prominent longitudinal costae, opening
at the top; seeds 0.5-0.7 mm long, discoid and often
angulate, testa foveolate.

Shrubs of evergreen montane forest formations,
from 1 600 to 2700 m elevation. Flowering in Jan-
uary-July, with a peak in April. Fruiting in Jan-
uary-September. The species has been collected
around Monteverde, Volcan Barva, and in the
western Cordillera de Talamanca (to above San
Isidro del General) in Costa Rica. The species
ranges from central and eastern Mexico through
the highlands of Guatemala and Honduras to the
Chiriqui Highlands of Panama.

Deppea grandiflora is recognized by its restric-
tion to higher montane forest formations, small
shrubby habit, thin leaves unequal at a node and
gradually tapering at both ends, bright yellow gla-
brous corollas with very short tube and broad lobes,
and small costate capsules opening at the top. Plants
with pseudoaxillary inflorescences and unopened
flowers may resemble species of Hoffmannia. Ma-
terial of Chiococca is also similar.

Didymaea Hooker f.

Perennial herbs, scandent or procubent, stems brittle,
usually much-branched and with long slender inter-

nodes, glabrous or rarely puberulent; stipules interpetio-
lar and 2-lobed or apparently free (with 4/node), decid-
uous or persisting and becoming recurved. Leaves
petiolate; leaf blades entire and pinnately or subpal-
mately veined, membranaceous to thin chartaceous,
domatia absent. Inflorescences of solitary flowers in the
axils of leaves, the pedicels long but not articulate, be-
coming recurved in fruit. Flowers bisexual and radially
symmetrical, hypanthium turbinate-globose, calyx en-
tire; corolla campanulate to rotate, yellowish to greenish
brown or purple, glabrous, corolla lobes 4, triangular and
subacute, valvate in bud; stamens 4, inserted between
the corolla lobes, filaments short and subulate, anthers
dorsifixed; ovary 2-locular, ovules solitary in each locule
and attached on the lower half of the septum. Fruits
2-parted or 2-lobed, somewhat fleshy, becoming dark
blue or black and lustrous, the lobes rounded, 1 lobe
usually smaller and lacking a fully developed seed; py-
renes rounded.

A small genus of two to five species, ranging
from Mexico to Panama. The plants are recog-
nized by their herbaceous climbing Galium-like
habit, small thin leaves, minute flowers solitary in
the leaf axils, and fleshy, often 2-parted rounded
fruit. The circumscription of species in Central
America is not resolved (see below).

Didymaea alsinoides (Cham. & Schlechtend.)
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
18: 1291. 1938. Nertera alsinoides Cham. &
Schlechtend., Linnaea 6: 413. 1831. D. alsi-
noides var. australis Standl., loc. cit. 1 292. 1 938.
D. alsinoides var. mollis Standl., loc. cit. 1292.
1938. D. australis (Standl.) L. O. Williams,
Fieldiana Bot. 24, pt. 11:61. 1972. Figure 3.

Herbs or weak-stemmed climbers, leafy stems 0.3-2
mm thick when dry, quadrangular or with 2 or 4 lon-
gitudinal ridges or wings ca. 0.5 mm high, sparsely and
minutely (ca. 0. 3 mm) puberulent or glabrescent; stipules
0.5-2.5 mm long, narrowly triangular or linear, decid-
uous. Leaves with petioles ( l-)2-8 mm long, 0.2-0.5 mm
broad; leaf blades (5-) 7-30 mm long, (3-)4-14 mm broad,
ovate to ovate-elliptic or narrowly ovate (rarely lanceo-
late), apex gradually narrowed and acute with sharp tip,
base abruptly narrowed or rounded and obtuse to trun-
cate, decurrent on petiole, drying membranaceous
(translucent), glabrous or minutely puberulent above the
midvein on the upper surface, sparsely pubescent be-
neath with thin hairs ca. 0.3 mm long, 2 veins 1-3/side,
often with subpalmate venation in broadly ovate leaves.
Inflorescence of solitary flowers in leaf axils, usually only
1 flower per node, pedicels 1-5 mm long, glabrous. Flow-
ers ca. 3-4 mm long, hypanthium ca. 0.7 mm long, calyx
to 0.2 mm long, truncate; corolla ca. 3 mm long, greenish
purple, tube ca. 1.5 mm long. Fruits 4-6 mm long, sub-
globose when 1 -seeded, deeply 2-parted and 6-8 mm
broad when 2-seeded, becoming fleshy and blue at ma-
turity, lustrous, drying black, usually glabrous; seeds of-
ten curved and reticulate.

122

FIELDIANA: BOTANY

Climbing plants of the shaded understory in
montane evergreen wet forest formations, from
1500 to 2800(-3100) m elevation. Flowering and
fruiting in September-June. The species (in a wide
sense) ranges from Mexico to the Chiriqui High-
lands of Panama.

Didymaea alsinoides is recognized in Costa Rica
by its slender-stemmed clambering habit, thin,
usually ovate leaves, minute flowers usually soli-
tary at each node, and small fleshy blue-black fruit
that are subglobose or deeply two-lobed. The pre-
ceding description is based on Costa Rican and
Panamanian collections referred to as Didymaea
australis by Williams. They differ from the more
northerly collections in having more ovate leaves
often truncate at the base. Recognizing the various
morphological and geographic forms of D. alsi-
noides as separate species seems unwise, insofar
as nearly all live in the same kind of montane
habitats and there is considerable morphological
variation in any one area; compare Williams'
treatment (in Standley & Williams, 1975, pp. 60-
63). Compare Nertera granadensis with smaller
leaves and orange fruit.

Diodia Linnaeus

Annual or perennial herbs or small shrubs, erect or
scandent, usually woody at the base, stems often much-
branched near the base, terete or 4-angled, glabrous or
pubescent; stipules interpetiolarand united with the leaf
bases to form a broad sheath, usually bearing 2-12 slen-

der awns from the truncated or rounded distal edge of
the sheath, persisting. Leaves opposite or pseudoverti-
cillate (with smaller axillary leaves), sessile or short-pet-
iolate: leaf blades mostly narrow and often scabrous,
entire or serrulate with minute scabrous hairs along the
margin. Inflorescences axillary and sessile, capitulate or
fasciculate (rarely of solitary axillary flowers) and often
verticillate, long-spicate when the distal subtending leaves
are reduced, subtended by the leaves and stipular sheaths,
flowers usually subsessile. Flowers bisexual and radially
symmetrical, small or minute, hypanthium obovoid to
turbinate, calyx lobes 2^*; corolla funnelform to cam-
panulate, white to pink or purplish, corolla tube short
or long, throat glabrous to villous, corolla lobes 4(3, 5-
6), valvate in bud; stamens 4 (3, 5-6), inserted in the
corolla throat, filaments filiform, anthers dorsifixed. lin-
ear-oblong; ovary 2-locular (rarely 3- or 4-locular), ovule
solitary and ascending in each locule, affixed to the center
of the septum, style filiform and exserted, bilobed or
with 2 short branches. Fruits splitting into 2 mericarps
(cocci), crustaceous to slightly woody, septicidal from
apex (rarely splitting at the base), without a central axis,
each mericarp indehiscent or opening slightly at the base;
seeds ellipsoid, longitudinally sulcate on the inner face,
rounded abaxially.

A genus of about 35 species in the tropical and
subtropical Americas and with a few species in
Africa. The weedy growth habit, awned stipular
sheaths, narrow leaves, small sessile flowers, and
fruit of two one-seeded indehiscent mericarps help
to distinguish this genus. Species of this genus may
resemble species of Crusea and Spermacoce, but
the leaves do not become pseudoverticillate. This
treatment has benefited from the annotations and
advice of C. D. Adams (pers. comm., 1991).

Key to the Species of Diodia

la. Plants essentially glabrous, often prostrate and restricted to the Caribbean seashore; leaves often

closely clustered on short lateral branches (1-4.5 cm long); rarely collected D. serrulata

Ib. Plants minutely to conspicuously pubescent, rarely prostrate and not restricted to the Caribbean

shore; leaves not closely clustered on short lateral branches 2

2a. Largest leaves usually less than 3 cm long, stems erect, stipular sheaths with usually glabrous awns

2b. Largest leaves more than 3 cm long, stems erect or clambering, stipular sheaths with awns with thin

whitish hairs distally or glabrous 5

3a. Leaves usually petiolate, to 2 cm long, thin-chartaceous and drying dark, often verticillate;

evergreen formations, 600-1200 m elevation D. brasiliensis

3b. Leaves sessile, to 3(-4) cm long, subcoriaceous and drying grayish, opposite; savannas in sea-
sonally very dry deciduous forest areas, 0-300 m elevation 4

4a. Top of the fruit with erect stiff hairs 0.2-0.5 mm long, back of the fruit smooth; plants annual

(collections often with slender fibrous roots); corolla 3-4 mm long; a common species

D. teres

4b. Top of the fruit glabrous or with few minute white hairs, back of the fruit with 3 longitudinal
ribs; plants mostly perennial (collections with thick taproots); corolla 8-10 mm long; rare . . .

D. apiculata

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

123

5a. Fruits dehiscent, rounded and indehiscent at the apex or opening slightly at the apex but mericarps
then often separating from the base and opening slightly at the base; stipular awns mostly glabrous;
leaves usually chartaceous; commonly collected Spermacoce ocymifolia

5b. Fruits (mericarps) indehiscent or opening slightly near the top; stipular awns with thin hairs; leaves
usually drying stiffly chartaceous; rarely collected D. sarmentosa

Diodia apiculata (Willd. ex Roem. & Schult.) K.
Schum. in Engler, Bot. Jahrb. 10: 313. 1889.
Spermacoce apiculata Willd. ex Roem. & Schult.,
Syst. Veg. 3: 531. 1818. D. rigida (Willd. ex
Roem. & Schult.) Schlechtend. & Cham., Lin-
naea 3: 301. 1828. Spermacoce rigida Willd. ex
Roem. & Schult., Syst. Veg. 3: 531. 1818, not
S. rigida Salisb. 1 796. Figure 6.

Herbs or subshrubs 9-40 cm tall, stems erect or pro-
cumbent, perennial and usually woody and branched at
the base, leafy stems 0.5-3 mm thick, hirsutulous to
hispidulous with whitish hairs ca. 0.3 mm long or with
sparse longer hairs to 1 mm long (rarely glabrous); stipule
sheath 1-2 mm long, with 6-9 setae 3-10 mm long.
Leaves opposite, often closely spaced, sessile; leaf blades
10-30 mm long, 1.5-7 mm broad, linear-lanceolate to
narrowly linear-oblong, broadest near the base, apex
gradually narrowed and acute with slender tip, base ob-
tuse to subtruncate, margins usually revolute, drying
subcoriaceous, hispidulous or hirsute above and below,
2 veins 2-3/side or obscure. Inflorescences fasciculate,
ca. 5 mm broad, with 2-8 flowers at a node, flowers
sessile. Flowers ca. 10 mm long, hypanthium 2-2.5 mm
long, 1-1.5 mm diam. (3-4 mm at the mouth), calyx
lobes 1.8-2.2 mm long, unequal, subulate-lanceolate,
erect, green; corolla funnelform to campanulate, white
to rose, glabrous on the exterior, tube 4-8 mm long and
1-1.5 mm diam., lobes 2-5 mm long, broadly ovate to
triangular, 1.5-3 mm broad at the base; stamens with
filaments 0.5-0.8 mm long, anthers 0.7-1.7 mm long;
style 5-8 mm long. Fruits 2.5-3(-4) mm long, glabrous
or puberulent, mericarps with 3 longitudinal costae (ribs)
on the curved dorsal (abaxial) side, obovoid, flat on the
inner (adaxial) side, 1.5-2.2 mm broad, calyx lobes to 2
mm long.

Plants of seasonally very dry deciduous for-
mations, from near sea level to 300 m (to 1 600 m
elevation in Honduras and to 2000 m in Guate-
mala). Probably flowering throughout the year in
northern Central America. It is primarily found
in the region around Liberia, Guanacaste, and is
collected in the wet season in Costa Rica. The
species ranges from Mexico and the West Indies
through Central America to Brazil.

Diodia apiculata is distinguished by the setose
stipular sheaths, narrow sessile stiff scabrous leaves,
the small axillary flowers, and the mericarps with
three longitudinal ribs. The mericarps are often
sparsely puberulent, in contrast to the very similar
D. teres.

Diodia brasiliensis Spreng., Syst. Veg. 1 : 406. 1 824.
D. polymorpha Cham. & Schlechtend., Linnaea
3: 344. 1828. D. brasiliensis var. angulata
(Benth.) Standl., Carnegie Inst. Wash. Publ. 46 1 :
90. 1935. Triodon angulatum Benth., PI. Hartw.
70. 1840. Figure 1.

Small shrubs to 1 m tall, much-branched, leafy stems
0.3-2(-5) mm thick, glabrous (minutely puberulent), with
4 longitudinal ribs and 4-angled; stipule sheath small
(0.5 mm), setae to 2 mm long. Leaves opposite or pseu-
doverticillate with 4, 6, or 8 small leaves at a node on
reduced axillary shoots (sometimes appearing to be an-
isomorphous with pairs of leaves differing in size), pet-
ioles 0-4 mm long; leaf blades 4-15(-20) mm long, 1-
4(-5) mm broad, oblong to elliptic, apex bluntly acute,
base acute and decurrent on petiole, drying chartaceous
and dark, scabrous along the margin, 2 veins 2/side or
obscure. Inflorescences often spiciform with flowers ver-
t id Hate in the axils of greatly reduced distal leaves, cap-
itulae 3-5 mm broad, flowers sessile. Flowers 3-4 mm
long, hypanthium 0.3-0.5 mm long, calyx lobes 4, ca.
0.5 mm long; corolla white, tube 1-2 mm long, lobes
4(-5), ca. 1 mm long. Fruits broadly turbinate, 2 mm
long (including the calyx lobes), 2 mm diam., glabrous,
the persistent calyx lobes ca. 0.7 mm long, the mericarps
usually remaining attached to each other at the base,
smooth on their abaxial surfaces.

Small plants of open or forested sites in ever-
green formations, from 600 to 1000 m elevation
in most of Central America (near sea level in Be-
lize). Probably flowering and fruiting throughout
the year. The species ranges from Mexico to Brazil.

Diodia brasiliensis is recognized by its many-
branched erect stems and the small leaves often
pseudoverticillate and drying black. This species
is only known from collections by Brenes near San
Ramon and it may not be native to Costa Rica.
Some of this material had been annotated as C.
polymorpha, now considered to be a synonym of
D. brasiliensis.

Diodia sarmentosa Sw., Prodr. Veg. Ind. Occ. 30.

1788.

Herbs, stems procumbent to scandent, to 4 m long,
sometimes forming tangles, leafy stems 0.7-4 mm thick,
with 4 longitudinal ridges, hispidulous with hairs 0.3-
0.6 mm long; stipule sheaths 1-3 mm long, bearing more

124

FIELDIANA: BOTANY

than 6 brown setae 4-8 mm long on each side, setae with
minute thin hairs distally. Leaves opposite, petioles 0-
3 mm long; leaf blades 3-6 cm long, 0.8-2.5 cm broad,
lanceolate to oblong-lanceolate, narrowly oblong-elliptic
or narrowly ovate, apex acuminate, base obtuse, margins
minutely aculeate-serrulate, drying subcoriaceous, sca-
brous and hispid u 1 1 nis above and below, with short (0.1-
0.2) scabrous hairs and longer 0.2-0.5 mm hairs, 2 veins
3-5/side, deeply impressed above and prominent below,
strongly arcuate-ascending. Inflorescences glomerules of
1-5 flowers in each axil, 5-1 5 mm broad, often becoming
verticillate in fruit, bracts linear-lanceolate. Flowers with
hypanthium ca. 2 mm long and 1 mm diam., calyx lobes
usually 2 large and 2 small, to 2.5 mm long, persistent;
corolla white, tube 1-1 .5 mm long, lobes 1-1 .5 mm long.
Fruits splitting into 2 mericarps, 3.5-5 mm long, 2-2.8
mm broad, obovoid, persisting sepals 1-2 mm long,
sparsely puberulent with short (0.2 mm) straight hairs,
abaxial surface without ribs; seeds ca. 3 x 1.6 mm, dark
brown, smooth.

Scandent plants of evergreen or partly decidu-
ous forest formations, from near sea level to 900
m elevation (to 1 500 m in Guatemala). Flowering
and fruiting in December-May. The species is
rarely encountered in southern Central America,
though it occurs on Cocos Island. The species rang-
es from Mexico and the West Indies into northern
South America, and it occurs in Africa.

Diodia sarmentosa is recognized by the thin
stipular setae, stiff subsessile leaves with deeply
impressed and strongly ascending secondary veins,
very small axillary flowers, and small two-parted
fruit.

Diodia serrulate (P. Beauv.) G. Taylor in Exell,
Cat. S. Tome 220. 1940. Spermacoce serrulata
P. Beauv., Fl. Oware 1: 39, t. 23. 1805. D. mar-
itirna Thonning ex Schumacher, Beskr. Guin.
PI. 75. 1827.

Prostate or clambering herbs, sometimes forming mats,
stems to 1.5 m long, leafy stems 1.2-4 mm thick, gla-
brous and brownish, at first with prominent wings but
becoming quadrangular or terete; stipule sheath 1 .5-2.5
mm long, to 3 mm broad, with 3-5 linear awns 0.5-3
mm long, glabrous or with a few minute hairs. Leaves
opposite or sometimes appearing verticillate (4), often
crowded on short lateral branches, subsessile or with
short (2 mm) winged petioles; leaf blades 10-45 mm
long, 4-13 mm broad, narrowly elliptic to lanceolate,
oblong-lanceolate or narrowly elliptic-oblong, apex acute,
gradually narrowed to the cuneate base and decurrent
on petiole, drying stiffly chartaceous and grayish brown
above, glabrous above, slightly scabrous beneath, 2 veins
3-4/side. Inflorescences of solitary axillary flowers (2/
node), subsessile on pedicels ca. 1 mm long. Flowers
glabrous externally, calyx lobes 4; corolla white, 6-7 mm
long, lobes 4, 2 mm long; anthers 0.9 mm long. Fruits

5-6 mm long, ca. 3 mm wide, splitting into 2 indehiscent
mericarps, glabrous and persisting calyx lobes to 2 mm
long; seeds 2.1-2.3 mm long, 1.4-1.6 mm broad, dark
reddish brown and smooth.

Rarely collected plants restricted to areas close
to the Caribbean seashore. Probably flowering and
fruiting throughout the year. The species ranges
from British Honduras and the West Indies to
Colombia; it also occurs on the west coast of Af-
rica.

Diodia serrulata is distinguished by its ocean-
side habitat, often prostrate habit, glabrous parts,
awned stipular sheath, and solitary flowers. The
leaf edges are entire and quite scabrous but not
serrulate. We have seen only the following from
Costa Rica: Gomez- Laurito 12109 CR, Playa Co-
des near Pto. Viejo, and Shank & Molina 4336 F,
Playa del Parismina, Limon.

Diodia teres Walt., Fl. Carol. 87. 1788. D. pros-
trata Sw., Prodr. Veg. Ind. Occ. 30. 1788. Fig-
ure 6.

Annual herbs to 40 cm tall, usually stiffly erect, stems
simple or branched near the base, leafy stems 0.7-2.3
mm thick, with 4 longitudinal ribs, hispidulous and sca-
brous with short (0.1-0.3 mm) and longer (1-2 mm)
hairs, internodes usually 0.5-3 cm long; stipule sheaths
1-2 mm long, with 6-9 conspicuous glabrous awns 2-8
mm long. Leaves sessile, leaf blades(4-)10-30(-45) mm
long, l-6(-8) mm broad, linear to linear-elliptic or lin-
ear-oblong, broadest near the base, apex acute, tip 0.5-
1 .8 mm long, usually revolute along the thickened mar-
gin, scabrous-hispid along the margin, drying subcoria-
ceous, scabrous or hirsute above, scabrous and hispidu-
lous beneath with thin white hairs 0.7-1.7 mm long, 2
veins usually obscure. Inflorescences of sessile solitary
or clustered flowers in leaf axils, the glomerules 3-6 mm
broad and with 2-4 flowers at each node. Flowers with
hypanthium 1.5-2 mm long, calyx lobes often unequal,
0.5-3 mm long and 0.1-0.8 mm broad, green; corolla
pink to purplish (white), tube 3-4.5 mm long, 0.5-0.7
mm diam. near the base, glabrous to sparsely puberulent,
lobes 1 .5-2.5 mm long, 0.5-1 .5 mm broad; stamens with
filaments 0.5 mm long, anthers 0.5-0.7 mm long, style
3-5 mm long. Fruits 2-5 mm long, with erect thin hairs
at apex and persisting calyx ca. 1 mm long, mericarps
(cocci) 1.8-2.5 mm broad, abaxial surface without lon-
gitudinal ribs (in Central America), minutely pubescent.

Plants of seasonally very dry deciduous for-
mations (especially sandy grass savannas) on the
northern Pacific slope, 0-300 m elevation (to 1 400
m in Guatemala). Flowering and fruiting in June-
January. The species ranges from the eastern Unit-
ed States to South America.

Diodia teres is recognized by its short height,

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

125

stipular setae, stiff narrow sessile scabrous leaves
(drying grayish), very small axillary pink flowers,
and distinctive mericarps (cocci).

Duroia Linnaeus f.

Small trees or shrubs, dioecious, branchlets tetrago-
nous or terete, with expanded areas housing ants in a
few species, glabrous or puberulent; stipules interpetiolar
and intrapetiolar, forming a cap over the shoot apex,
circumscissile and deciduous. Leaves opposite or ver-
ticillate with 3-5 leaves at a node, sessile or petiolate,
entire and pinnately veined, drying thin-chartaceous to
coriaceous. Inflorescences terminal, 6 flowers subcapi-
tate, umbellate, to corymbose or cymose, sessile or pe-
dunculate, 2 flowers 1-3 at the tip of the stem, sessile or
pedunculate, the flowers usually short-pedicellate. Flow-
ers unisexual, radially symmetrical, usually large, hy-
panthium oblong to hemispheric, calyx cupular to tu-
bular, truncated and entire or with 5-9 lobes, corolla
salverform, white to yellowish white, often thick or fleshy,

sericeous on the outer surfaces, glabrous or pilose in the
tube within, corolla lobes 5-9(-12), convolute in bud;
stamens 5-9, inserted in the corolla tube, subsessile or
sessile, anthers dorsifixed, linear, acute, included; ovary
1-5-locular, with 5-6 parietal placentas sometimes joined
in the center, ovules many and biseriate, stigmas 2. Fruits
baccate, globose to oblong, large with a thick fleshy cor-
tex, with 1-4 locules; seeds large and horizontal, flattened
and suborbicular, immersed in pulp, testa thin.

A tropical American genus of about 25 species,
with 2 or 3 species in Central America. Durroia
hirsuta (Poepp. & Endl.) Schumann and D. petio-
laris Hook. f. have swollen elongated areas of the
stem with longitudinal slits, and D. saccifera Hook,
f. has saccate ant domatia at the base of its leaf
blades. Our species have no such structures and
are not known to have an association with ants.
Specimens may resemble species of Amaioua,
Hippotis, and Randia.

Key to the Species of Duroia

la. Leaves subsessile, obovate, with hairs ca. 2 mm long, minor venation lineolate; calyx ca. 4 mm
long, male corolla lobes 5-8 mm long D. costaricensis

Ib. Leaves petiolate, broadly oblong to elliptic-obovate, with hairs ca. 0.5 mm long, minor venation
not lineolate; calyx ca. 8 mm long, male corolla lobes ca. 10 mm long D. utleyorum

Duroia costaricensis Standl., Contr. U.S. Natl.
Herb. 20: 208. 1919. Figure 25.

diam., oblong to ovoid, covered with long hairs but with
the surface visible beneath the hairs.

Shrubs or small trees, 5-10 m tall, leafy branchlets 2-
6 mm thick, hirsutulous with straight thin ascending
hairs ca. 2 mm long, becoming reddish brown and gla-
brescent; stipules 1 545 mm long, caducous, densely hir-
sute-sericeous on the outer surfaces. Leaves closely
crowded at the tips of branchlets, subsessile with petioles
2-6(-10) mm long and ca. 2 mm thick, densely hirsute;
leaf blades (7-)9-19 cm long, (3-)4-7.5 cm broad, ob-
long-obovate to narrowly obovate, widest at or above
the middle, apex abruptly narrowed and slender acu-
minate or caudate-acuminate, the narrow (ca. 2 mm) tip
5-13 mm long, gradually narrowed to the cuneate base,
drying thin-chartaceous, often brown, with long ( 1 .5-2.5
mm) thin straight or slightly crooked hairs on upper and
lower surfaces, 2 veins 5-8/side, 3 veins subparallel
between the secondaries. Male flowers 8-15 and sub-
capitate or fasciculate-cymose (tightly clustered at the
tips of stems), 1 5-20 mm long, outer surfaces of calyx
and corolla densely sericeous with ascending hairs, calyx
4-5 mm long, calyx lobes 6-7 and equalling the tube;
corolla 11-15 mm long, white, corolla lobes 6-8, 5-8
mm long, equalling or longer than the tube. Female flow-
ers l(-3) subsessile at apex of branchlets, hypanthium
ca. 8 mm long, densely sericeous, with hairs 2-3 mm
long, calyx lobes ca. 4 mm long, linear. Fruits 1-3 at the
tips of branchlets, subsessile, ca. 22 mm long, 1 2 mm

Small trees of lowland rain forest formations of
the Golfo Dulce region, from 10 to 200 m ele-
vation. Flowering in March and May-June; fruit-
ing in July-August and October. This species is
known only from southernmost Costa Rica.

Duroia costaricensis is recognized by the sub-
sessile thin hirsute obovate leaves clustered at the
ends of stems, the terminal clusters of subsessile
sericeous flowers, and the hirsute subsessile fruits
at the tips of branches. The type collection (Pittier
6803 us) came from Sierpe, but most of the other
collections come from near Rincon de Osa. Duroia
hirsuta (Poepp. & Endl.) Schumann of South
America has inflated stems, longer petioles, gen-
erally longer leaves, pedunculate male flowers, and
larger fruit. Duroia genipifolia, now Randia geni-
pifolia (Standl. & Steyerm.) Lorence, of Guatemala
has less hirsute leaves with better denned petioles
and more elliptic blades. All three species are sim-
ilar in appearance. This species also resembles
Costa Rican material of Hippotis.

126

FIELDIANA: BOTANY

Duroia utleyorum Dwyer, sp. nov.

Frutices vel arbores 1.5-7 m altae. Foliae lamina ob-
longa vel oblongo-elliptica, 10-23 cm longa, 6-14 cm
lata; venis lateralibus 7-1 1, hirsutulis; petiolis 6-23 mm
longis. Flores lobis calycinis ca. 1 mm longis; corollae
tubo ca. 12 mm longo, lobis ca. 10 mm longis. Fructus
immaturi solitarii terminales.

TYPUS Liesner /774(holotypuscR, isotypus MO), from
Rincon de Osa, Costa Rica.

Shrubs or small trees, (1.5-)4-7 m tall, leafy stems 2-
5 mm thick, densely pubescent with thin straight yel-
lowish hairs 0.3-0.7 mm long; stipules ca. 9 mm long
and 4 mm broad, ovate-lanceolate with a sharp acu-
minate apex, united (intrapetiolar) to form a short (1
mm) basal tube, with thin brown margins and hirsute.
Leaves with petioles 6-23 mm long, 1-3 mm thick, hir-
sutulous with yellowish hairs; leaf blades 10-23 cm long,
6-14 cm broad, broadly oblong to broadly obovate or
elliptic-oblong, apex short-acuminate with tip 5-1 5 mm
long, base obtuse to rounded and subtruncate, drying
chartaceous, brown, pubescent on the major veins above,
densely hirsute on the veins beneath with yellowish hairs
ca. 0.7 mm long, sparsely hirsutulous between the veins,
2 veins 7- 11 /side. Inflorescences of solitary terminal
female flowers, on pedicels ca. 8 mm long; male inflo-
rescences of 3-7 terminal sessile flowers. Flowers whitish
pilosulous to sericeous; 3 flowers with calyx tube 5-6
mm long, calyx teeth ca. 1 mm long, remote; corolla
(preanthesis) white and probably salverform, tube 12
mm long, lobes 10 mm long, convolute and apparently
lanceolate. Fruits solitary and terminal, the immature
fruit ca. 1 5 mm diam. and globose with persisting calyx
10 mm high and 8 mm diam., densely hirsutulous with
hairs ca. 0.5 mm long.

A species of lowland (10-300 m) rain forest for-
mations on both the Caribbean and Pacific slopes
of Costa Rica. Young flowers were collected in
February on the Osa Peninsula (Jimenez et al. 650
CR, Liesner 1774 & 1853 CR, MO); young fruits
were collected along the Rio Sarapiqui in May
(Hartshorn 1486 CR).

Duroia utleyorum is recognized by its larger
broad leaves, sessile terminal flowers, and lowland

evergreen forest habitat. This poorly known spe-
cies is more likely to be confused with species of
Randia than with its local cogener. Many impor-
tant collections have been made in Costa Rica by
Kathleen and John Utley.

Elaeagia Weddel

Trees or large shrubs, branchlets terete and puberulent;
stipules interpetiolar (also intrapetiolar in some species),
often covering the shoot apex, caducous or deciduous.
Leaves opposite, petiolate or subsessile; leaf blades entire
and pinnately veined, usually puberulent, thin-charta-
ceous to coriaceous, domatia present or absent. Inflo-
rescences terminal, paniculate (rarely racemose), many-
flowered, pedunculate. Flowers bisexual, radially
symmetrical, usually small, hypanthium hemispheric to
turbinate, often sulcate on opposite sides, calyx tube cu-
pular and spreading or inflated, calyx lobes 5 or none,
short or inconspicuous; corolla campanulate to funnel-
form, corolla white to yellow-white, tube usually short
and broad, corolla lobes 5, oblong, rotate to reflexed,
convolute to slightly imbricate in bud; stamens 5, in-
serted between the corolla lobes, filament densely pu-
berulent at the base, anthers dorsifixed, oblong, exserted;
ovary 2-locular, placentas peltate and attached to the
septum, ovules many in each locule, crowded and ver-
tical, style short, stigmas 2 with obtuse tips. Fruits small
woody capsules, crowned or encircled by the persistent
calyx, loculicidally and basipetally dehiscent into 2 valves,
the valves splitting at the top; seeds many, minute, elon-
gate with membranous or winged testa.

A genus of about 1 2 species in Mexico, Central
America, Cuba, and tropical South America. Elae-
agia utilis Weddel of Colombia is the source of a
protective lacquer. The genus was misspelled as
Elaeagnia in the Flora of Panama (1980). Three
mesoamerican species of Elaeagia were compared
by Lorence (in Bol. Soc. Bot. Mexico 45: 65-69.
1 983). The small rounded capsules resemble those
of Chimarrhis.

Key to the Species of Elaeagia in Costa Rica

la. Leaf blades usually 9-25 cm wide, often subsessile and sometimes auriculate at the base, with 1 1-
19 pairs of secondary veins; inflorescences usually much-branched panicles E. auriculata

Ib. Leaf blades usually 3-9 cm wide, clearly petiolate and gradually narrowed to the base, with 6-12
pairs of secondary veins; inflorescences with few lateral branches (paniculate with many branches
in E. myriantha) 2

2a. Calyx cup entire distally; flowers separate and borne on long (4 mm) slender pedicels

E. nitidifolia

2b. Calyx cup with undulate or minutely lobed margin; flowers crowded and subsessile in small groups
of2-5 . . 3

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

127

3a. Corolla tubes 1-1.5 mm long, lobes ca. 1.7 mm long; leaf blades with 7-9 major 2 veins per side,
without tufted domatia along the 2 veins; Cordillera de Tilaran E. uxpanapensis

3b. Corolla tubes 2.2-3.4 mm long, lobes 0.4-1 mm long; leaf blades with 9-12 major 2 veins per side,
often with tufted domatia along the 2 veins; General Valley E. myriantha

Elaeagia auriculata Hem si.. Diagn. PI. Nov. Mex-
ic. 32. 1879. Figure 39.

Shrubs or small trees, 3-10 m tall, leafy branchlets 3-
8 mm thick, puberulent or glabrescent; stipules 20-50
mm long, to 20 mm broad, oblong and rounded at apex,
reddish brown and with subparallcl venation. Leaves
with petioles 0-10 mm long, 1.5-3 mm thick, often dif-
ficult to see on auriculate leaves; leaf blades ( 1 2-) 1 5-40
cm long, (6-)9-25 cm broad, elliptic-obovate to broadly
elliptic or obovate, apex abruptly narrowed or rounded
and obtuse to short-acuminate, gradually narrowed to
the cuneate or auriculate base, drying chartaceous to
subcoriaceous, dark brown above, upper surface gla-
brous to short (0.2-0.5 mm) pubescent, lower surfaces
sparsely to densely pubescent with short stiff hairs, 2
veins 1 l-16(-19)/side, 3 veins usually subparallel and
the 4 veins raised beneath. Inflorescences 12-20 cm
long, to ca. 20 cm broad, many flowered and much-
branched panicles, solitary or 3 from the terminal leaf-
bearing node, primary peduncles 3-5 cm long, with 3-
4 pairs of 1 opposite lateral branches, puberulent, flow-
ers in racemose or cymose distal branches, pedicels 1-5
mm long. Flowers 5-7 mm long, hypanthium ca. 1 mm
long, calyx cup ca. 1 mm long (including the lobes), calyx
lobes 0.5-1 mm long and 1.5 mm broad, glabrous; co-
rolla white, tube 1-2 mm long, lobes 1-2 mm long and
becoming reflexed; stamens with filaments 2 mm long,
glabrous distally, anthers ca. 1.5 mm long. Fruits 3-5
mm long and 3-4.5 mm broad, short-ovoid and broadly
rounded, calyx persistent and visible on the lower part
of the capsule, opening at the top (but not splitting to
the base), becoming dark brown or black; seeds 0.6-1
mm long, body of the seed 0.2-0.4 mm long, with a thin
membranous wing at opposite ends.

Trees of evergreen lower montane cloud forest
and rain forest formations, along the central high-
lands, from Volcan Tenorio in the Cordillera de
Guanacaste to San Vito de Goto Brus, and on the
Caribbean slope, ranging from 350 to 1700 m el-
evation. Flowering in November-March; fruiting
in February-June. The species ranges from Gua-
temala and Honduras to eastern Panama.

Elaeagia auriculata is recognized by the large
oblong stipules, short petioles, large broad leaves
often auriculate at the base, large terminal pani-
cles, small flowers with truncated calyx lobes and
very short corolla tubes, and short rounded cap-
sules opening only at the top. The very large leaves
with an auriculate base obscuring the petiole are
very distinctive, but not all specimens have this
kind of leaf base (see below). The small capsules
make this species look like species ofChimmarhis.

Elaeagia karstenii Standley was recorded for
Costa Rica's flora (Standley, 1938) on the basis of
a single sterile collection: Valeria 1665 F, from
Tapanti at 1300 m. The longer (10-14 mm) and
slender (1.5-2 mm) petioles and the smaller
sparsely puberulent leaves do resemble some ex-
amples ofE. karstenii from South America. How-
ever, longer petioles with cuneate (not auriculate)
lamina bases do occur in specimens that otherwise
appear to be typical of E. auriculata: Lent 3734
and Molina et al. 17346 (note: both of the latter
collections come from below 1000 m elevation).
Thus, it is possible that Valeria 1665 is an aberrant
juvenile shoot of an E. auriculata plant. Alter-
natively, expanding the concept of E. auriculata
to include material with longer petioles, cuneate
leaf bases, and smaller more glabrous leaves may
require the inclusion of South American speci-
mens and synonymizing E. karstenii.

Elaeagia myriantha (Standl.) C. M. Taylor &
Hammel, Novon (in press). Sickingia myrian-
tha Standl., Publ. Field Columb. Mus., Bot. Ser.
7: 27. 1930. Holtonia myriantha (Standl.)
Standl., Trop. Woods 30: 37. 1932. Simira myr-
iantha (Standl.) Steyerm., Mem. New York Bot.
Gard. 23: 306. 1972. Deppea panamensis Dwyer,
Ann. Missouri Bot. Gard. 67: 145. 1980.

Trees, 8-25 m tall, leafy stems 2-7 mm thick, gla-
brous, drying brownish or grayish; stipules 3-5 mm long,
short-tubular with rounded lobes, often splitting between
the leaf bases and remaining entire above the petioles
(ligulate), translucent, persisting. Leaves with petioles 5-
20(-30) mm long, 0.8-2 mm thick, glabrous, drying red-
dish brown to dark brown; leaf blades (5-)9-20 cm long,
(2-)3.5-7(-8) cm broad, elliptic-oblong to elliptic or el-
liptic-obovate, apex acuminate with tip 10-15 mm long,
base gradually narrowed and acute or cuneate and de-
current on petiole, drying chartaceous, greenish to dark
brown, glabrous above, glabrous or minutely (0.05 mm)
papillate-puberulent on the veins beneath, 2 veins 9-
12/side, domatia with short hairs present in the vein
axils. Inflorescences 10-22 cm long, 12-18 cm broad,
pyramidal with usually opposite 1 branches, peduncles
8-40 mm long, 1.5-2.5 mm thick, minutely puberulent,
larger (2 cm) narrow leaf-like bracts present or absent,
with smaller (0.5-3 mm) bracts and bracteoles subtend-
ing branches and flowers, flowers usually subsessile in
groups of 2-5. Flowers glabrous externally, hypanthium
1-1.5 mm long, obconic, calyx cup 0.1-0.3 mm long.

128

FIELDIANA: BOTANY

1.3 mm diam., calyx lobes 0.2-0.4 mm long, broadly
triangular; corolla tubular-campanulate, cream white,
tube 2.2-3.4 mm long, ca. 2 mm diam., lobes 5, 0.4-1
mm long, triangular, slightly imbricate in bud; filaments
exserted 0.5-2 mm, anthers ca. 1.5 mm long; stigmas 1-
2 mm long. Fruits ca. 3 mm long, 3 mm broad at the
top when opened, dark brown; seeds 0.3-0.5 mm long,
broad, reticulate.

Collected with flowers in January in the General
Valley at 975 m elevation (Skutch 2387 us, the
only Costa Rican collection). This species is also
known from the Andes of Colombia and Vene-
zuela.

Elaeagia myriantha is distinguished by the large
pyramidal terminal panicles of small white flowers
with broad cylindrical corolla tube and minute
lobes and the almost glabrous leaves with pubes-
cent domatia in the vein axils. These are unusual
in that they are found in the axils of some 3 veins
as well as along the midvein. The Colombian type
(H. H. Smith 1810F) has smaller leaves than most
other collections.

Elaeagia nitidifolia Dwyer, Ann. Missouri Hot.
Gard. 67: 157. 1980. Chiococca jefensis Dwyer,
loc. cit. 67: 88, f. 19. 1980. Figure 39.

Trees to 1 2 m tall, leafy stems 2-4 m thick, glabrous,
becoming terete; stipules 3-5 mm long, to 4 mm broad,
cylindrical at first but splitting, rounded above the pet-
iole and deeply split between the petioles (becoming lig-
ulate), glabrous. Leaves closely clustered distally, petioles
6-17(-25) mm long, 1-2 mm thick, glabrous; leaf blades
10-23 cm long, 3.5-8 cm broad, oblanceolate to nar-
rowly oblong-obovate or elliptic-oblong, apex short-acu-
minate, tip 5-8 mm long, gradually narrowed to the
cuneate base and decurrent on petiole, drying charta-
ceous, subglabrous above, minutely (0.05-0. 1 mm) pa-
pillate-puberulent on the veins beneath, 2 veins 8-12/
side. Inflorescences solitary or 3, terminal, 8-1 6 cm long,
open paniculate with distant cymose flower groups, pe-
duncles 3-5 cm long, glabrous, pedicels 3-10 mm long,
slender. Flowers with hy pan t h i urn and calyx continuous,
ca. 3 mm long and 3-4 mm broad distally, calyx cup
entire distally; corolla white, tube 1-2 mm long, lobes
5, 3-4 mm long, 1.3-2 mm broad, oblong; anthers 1.5-
2 mm long; stigma lobes 2-2.5 mm long. Fruits to 6 mm
long and 10 mm broad, turbinate, truncated distally.

Trees of wet evergreen forest formations of the
Caribbean slope at 400-800 m elevation. The spe-
cies is only known from the Rara Avis site in
Heredia and was collected flowering in September
(O. Vargas 128 CR, MO). The species is also known
from central and eastern Panama.

Elaeagia nitidifolia is distinguished by its un-
usual ligulate stipules, the open few-flowered in-

florescences, long pedicles, and cupulate calyx with
entire margin. Specimens may resemble Rustia oc-
cidentalis (with porate anthers) and Simira myri-
antha (with smaller corolla lobes and domatia).

Elaeagia uxpanapensis D. Lorence, Bol. Soc. Bot.
Mex. 45: 66. 1983 (1984). Figure 39.

Trees, 15-40 m tall, 55-80 cm dbh, leafy stems 3-9
mm thick, flattened in early stages, glabrous and becom-
ing terete; stipules 8-16 mm long. 3-7 mm broad, ovate
lanceolate with overlapping margins, acute at the apex,
glabrous and drying dark, deciduous. Leaves with peti-
oles 3-12 mm long, ca. 1.5 mm thick, glabrous, drying
dark; leaf blades 6-15(-20) cm long, 3-7(-9) cm broad,
broadly elliptic to broadly oblong or slightly elliptic-
obovate, apex abruptly narrowed and bluntly obtuse,
base cuneate and slightly decurrent on petiole, drying
stiffly chartaceous to subcoriaceous, dark grayish brown
above, glabrous above, with a few thin yellowish hairs
0.3-1 mm long along the side of the midvein beneath
and in leaf axils (= domatia?), 2 veins 7-9/side. Inflo-
rescences solitary and terminal or axillary to distal leaves,
6-14 cm long (perhaps enlarging in fruit), narrowly pa-
niculate, peduncles 2.5-5 cm long, glabrous, proximal
lateral branches 1-6 cm long, usually minutely ap-
pressed-puberulent, flowers sessile in opposite or ter-
minal glomerules of 2-5 flowers, bracts 0.5-3 mm long.
Flowers ca. 4 mm long, yellowish green, hypanthium 1-
1.8 mm long, minutely puberulent at the base, calyx
lobes 0.5-1 mm long; corolla white, salverform to short -
funnelform, glabrous externally, tube 1-1.5 mm long,
scarcely exceeding the calyx lobes, lobes 4-5, ca. 1 .7 mm
long and 1 mm wide, bluntly rounded distally, with long
hairs at the mouth of the tube and base of lobes within;
stamens 4-5, filaments 2-4 mm long, anthers 0.6-0.7
mm long, oblong; style and stigma 3-4 mm long, stig-
matic lobes 1 mm long. Fruits not seen (probably ca. 2
mm long and 2.5 mm broad).

Only known from the wet Caribbean slopes of
the Cordillera de Tilaran at ca. 900 m elevation.
Immature flowers were collected in July (Haber &
Bella 1928 CR, MO) and mature flowers in May
(Herrera 600 CR, MO). This species is known only
from the Rio Penas Blancas valley below Mon-
teverde and the Reserva Forestal de San Ramon
(Gomez- Laurito 12065 CR), in Alajuela Province.
The species is also known from southern Mexico.

Elaeagia uxpanapensis is recognized by the op-
posite subsessile flower clusters along the branches
of the inflorescence, the very small flowers, and
the two-locular ovule with placentas borne on the
septum. The large size of the trees, flattened young
leafy stems, and occasional domatia along the
midvein are additional distinctive characteristics.
When first discovered in Costa Rica, specimens
were thought to be a species related to Warszewic-
zia schwackei Schum. of South America.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

129

Exostema L. Richard

Shrubs or trees, branchlets usually terete, glabrous or
puberulent; stipules interpetiolar, small, entire or bifid,
deciduous or persisting. Leaves petiolate or subsessile;
leaf blades entire, drying membranaceous to subcoria-
ceous, domatia sometimes present. Inflorescences ter-
minal or axillary panicles with the distal flowers usually
in groups of 3, or of solitary axillary flowers, with or
without bracts, pedicels present. Flowers bisexual, ra-
dially symmetrical (or somewhat bilaterally symmetrical
by curvature of the corolla tube), small to large, hypan-
thium cylindrical or obovoid, calyx with (4-)5(-6) lobes,
lobes broad or narrow; corolla short or long, salverform,
tube often long and narrow, throat glabrous or barbate,
corolla lobes (4-)5, oblong or linear, imbricate in bud
with 2 exterior; stamens 5 inserted at the base of the
corolla tube, filaments essentially free, elongate and fi-

liform, anthers basifixed, linear, usually exserted; ovary
2-locular, placentas peltate on the septum with many
ascending ovules, style filiform, usually exserted, stigma
capitate or bilobed. Fruits capsular, woody, 2-locular,
ellipsoid to cylindrical, septicidal from apex, bivalved,
valves entire or bipartite from the apex; seeds many,
imbricate, flattened and oblong, testa forming a thin
marginal wing around the body of the seed.

A genus of ca. 35 species of the American trop-
ics, especially abundant in drier vegetation of the
West Indies. The flowers with narrow corolla lobes
almost equalling the length of the slender corolla
tube and the long linear exserted anthers make the
flowers quite distinctive. Our species are confined
to deciduous or partly deciduous vegetation.

Key to the Species of Exostema

la. Flowers solitary in the axils of leaves, 6-8 cm long, corolla lobes 4-5; fruit ca. 1 cm broad, erect;
leaf blades usually clearly decurrent on the petiole, domatia absent E. caribaeum

Ib. Flowers more than 1, borne in small panicles axillary to distal leaves or terminal, 2-3 cm long;
corolla lobes 6; fruit 34 mm broad, pendulous; leaf blades very slightly decurrent on the petiole,
domatia often present E. mexicanum

Exostema caribaeum (Jacq.) Roem. & Schult., Syst.
Veg. 5: 19. 1819. Cinchona caribaeum Jacq.,
Enum. PI. Carib. 16. 1760. E. longicitspeOeTsl.,
Vidensk. Meddel. Dansk. Naturhist. Foren.
Kjobenhaven 1852: 48. 1853. Figure 31.

Shrubs or small trees to 8 m tall, leafy branchlets 1 .2-
3.5 mm thick, glabrous; stipules l-3(-5) mm long, 2-3
mm broad at the base, triangular to subulate, glabrous
on the abaxial surface but slightly ciliate along the distal
edge, persistent. Leaves evenly spaced along the stems,
petioles 6-1 2(- 16) mm long, 0.4-0.8 mm thick; leaf blades
4-1 1 cm long, 1.5-5 cm broad, elliptic-ovate to ovate,
apex gradually narrowed and acuminate, tip 1-2 cm long,
base obtuse to acute and decurrent on petiole, drying
thin-chartaceous and dark, glabrous above, glabrous or
pubescent with thin curved whitish hairs ca. 0.4 mm
long beneath, often with short (0.5 mm) hairs (domatia)
in the vein axils beneath, 2 veins 4-5/side. Inflores-
cences absent, the flowers solitary in distal leaf axils,
bracts ca. 1 mm long, pedicels 4-10 mm long, ca. 0.5
mm thick (dry), glabrous. Flowers 6-8 cm long, hypan-
thium 3-5 mm long, 2-3 mm diam., glabrous, calyx
lobes 0.5-1 mm long, broadly triangular, corolla white,
glabrous, tube 23-45 mm long, 2-3 mm diam., often
curved, lobes 25-40 mm long, ca. 2 broad and linear,
becoming recurved; stamens 5. long-exserted, anthers
16-26 mm long, 0.2-0.3 mm broad (dry); stigma 2 mm
long. Fruits 7-15 mm long, 6-14 mm broad, oblong-
ellipsoid, smooth and dark brown externally; seeds 3-6
mm long and 3-4 mm broad, with a thin brownish wing
on all sides and slightly longer at the 2 ends, body of the
seed ca. 5 mm long and 3 mm broad.

Trees of deciduous and partly deciduous forests
of the Pacific slope, from near sea level to 300 m
elevation (to 1300 m in Guatemala). Rarely col-
lected in Costa Rica (Guanacaste and adjacent
Puntarenas provinces). In Central America flow-
ering in June-August; fruiting in October-January
and April. The species ranges from central Mexico
to northern Costa Rica and the West Indies.

Exostema caribaeum is recognized by its re-
striction to seasonally very dry vegetation, the long
flowers solitary in leaf axils, the long narrow co-
rolla tube, long corolla lobes, and long slender ex-
serted anthers. The woody bivalved fruit splitting
down the broader side and the seeds surrounded
by a short thin wing are also distinctive. We have
not seen Oersted's type of E. longicuspe from Pun-
tarenas (cf. Standley, 1938, p. 1295). This species
is called "caribee bark tree" and "princewood" in
the Caribbean, where the wood is used for making
handles. The bark is used for treating fevers and
malaria in Mexico.

Exostema mexicanum A. Gray, Proc. Amer. Acad.
Sci. 5: 180. 1 86 1 . Figure 40.

Small to medium-sized trees to 20 m tall with trunks
to 40 cm dbh, leafy branchlets 1.3-4 mm thick, glabrous

130

FIELDIANA: BOTANY

to sparsely puberulent, lenticels conspicuous, stems be-
coming grayish; stipules l-3(-4) mm long, triangular and
acuminate to short-tubular and cuspidate, deciduous.
Leaves well spaced along the stems, petioles 4-10(-15)
mm long; leaf blades (5-)6-15(-18) cm long, (2-)3-
7.5(-10) cm broad, ovate-oblong to elliptic-oblong, apex
long-acuminate, base obtuse to rounded and subtrun-
cate, usually glabrous above and below except for the
minute (0.2 mm) hairs (domatia) in the vein axils be-
neath, 2 veins 6-9/side. Inflorescences 3-6(-l 2) cm long,
4-7(-10) cm broad, terminal or axillary to distal leaves,
primary peduncles 1.5-4 cm long, glabrous or minutely
puberulent, trichotomous, lenticellate, bracts 1-2 mm
long, distal flowers in groups of 3, often crowded, ped-
icles 2-4 mm long, minutely (0. 1 mm) puberulent. Flow-
ers ca. 2 cm long, hypanthium 1 .5-2.4 mm long, glabrous
or sparsely and minutely puberulent, calyx lobes 5-6,
very small (0.5 mm) and deltoid/acute; corolla white to
yellowish white or yellow, subglabrous or with thin curved
whitish hairs, tube 7-10 mm long, 0.7-1.3 mm diam.,
lobes 6, ca. 9-12 mm long, 1 .5 mm broad, oblanceolate-
linear; stamens 6, filaments to 1 6 mm long, anther 3-4
mm long, 0.2-0.3 mm broad; style to 2 cm long, stigmas
clavate, 0.6 mm long. Fruits 10-14 mm long, 3-4 mm
broad, obovoid-clavate to narrowly oblong-obovoid,
surface glabrous and dark brown with whitish lenticels,
the 2 valves each splitting in 2 at the top; seeds 6-9 mm
long, 2-3 mm broad, body of the seed ca. 2.5 x 1 .5 mm,
oblong, wing often lobed on 1 end.

Trees of deciduous and partly deciduous forests
of the Pacific slope, from near sea level to 900 m
elevation (to 1 300 m in Guatemala). Flowering in
July-September; fruiting in July and September-
February. The species ranges from Tamaulipas,
Mexico, to central Panama.

Exostema mexicanum is distinguished by its de-
ciduous habitat and thin leaves, close clusters of
narrow-tubed flowers in short axillary or terminal
panicles, the long narrow corolla lobes, exserted
linear anthers, and woody bivalved capsules with
winged seeds. The bark has been boiled to make
a treatment for malaria and fevers. This species is
rarely collected in Costa Rica.

trapetiolar, often forming a tubular sheath above the leaf
base, short-triangular to long-aristate at apex (with 2
awns per node), persistent or deciduous. Leaves often
held in a single plane, petiolate (rarely subsessile); leaf
blades entire, usually oblong to lanceolate, usually gla-
brous, lacking domatia, pinnately veined. Inflorescences
terminal (rarely axillary), 1 -many-flowered, corymbose
to umbellate (rarely capitate), pedunculate, peduncle and
branches often with a pale blue or purple color like the
flowers, pedicels usually present. Flowers bisexual and
radially symmetrical, monomorphic or distylous, hy-
panthium small, ovoid to turbinate, terete or angular,
calyx cupular to short-tubular, truncate and entire to
4-lobed, persistent; corolla usually salverform, white to
blue or lavender, tube short or long, usually narrow,
glabrous, corolla lobes 4(-5), linear to lanceolate, valvate
in bud, spreading or reflexed; stamens 4 (5. 6), inserted
on the tube or at the throat, anthers linear, dorsifixed.
linear, included or exserted; ovary 1-locular (rarely
2-locular in early stages), ovules 2 (less often 1), erect
from a basal placenta, style filiform, with 2 short branch-
es. Fruit baccate or drupaceous, transversely oblate (reni-
form) to subglobose, often broader than long, smooth or
costate, deep blue to blue-black, 1-locular and 1- (rarely
2-) seeded; seed (pyrene) horizontal, transversely in-
dented (excavated) and somewhat reniform, with thin
testa.

A large genus of about 1 30 species, ranging from
Mexico, Central America, and the West Indies
through tropical South America. The unusual fruit
(usually broader than long) and single seed help
to distinguish this genus. A striking sky blue to
lavender color of both the flowers and inflores-
cences characterizes a number of our species. The
usually four-parted flowers, short-tubular stipules
with only a single apex on each side, and a well-
developed submarginal vein are additional char-
acters found in many species. The fruits tend to
have a leathery exocarp in Faramea, in contrast
to the succulent exocarps of Coussarea and Psy-
chotria. Despite these unusual traits, specimens of
some species may look very similar to some spe-
cies of Psychotria, Coussarea, and Rudgea.

Faramea Aublet

Shrubs or small trees, branchlets terete, 4-angled or
flattened, usually glabrous; stipules interpetiolar and in-

Key to the Species of Faramea

la. Flowers solitary or the inflorescence with 2-3 flowers (l^t flowers per node); rarely collected .

Ib. Flowers few to many in pedunculate open inflorescences; commonly collected species 4

2a. Leaf blades 12-20 cm long; peduncles 2-5 cm long, calyx and hypanthium ca. 12 mm long

[Caribbean slope at 20-1 100 m elevation in Costa Rica] F. pauciflora

2b. Leaf blades 1-4 cm long; peduncles less than 2 cm long, calyx and hypanthium less than 5
mm long 3

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

131

3a. Inflorescences terminal; shrubs 3-4 m tall; Pacific slope, 1200-1700 m elevation

F. capulifolia

3b. Inflorescences axillary; herbaceous subshrubs ca. 1 m tall; Caribbean slope, ca. 300 m ele-
vation F. myrtidfolia

4a. Leaves sessile or subsessile and the leaf blade usually rounded at the petiole, petioles 0-5 mm long
5

4b. Leaves definitely petiolate or if subsessile the leaf blades not rounded at the petiole, petioles 4-20

mm long 7

5a. Leaf blades narrowly oblanceolate, 10-16 cm long and 1.5-3.5 cm broad; Chiriqui Highlands

at ca. 2000 m elevation F. scalaris

5b. Leaf blades usually oblong, 15-27 cm long and 6-17 cm broad; 0-500 m elevation .... 6
6a. Leaves strongly tripliveined, secondary veins united by a linear (melastome-like) lateral

submarginal vein; southern Caribbean lowlands F. trinervia

6b. Leaves not strongly tripliveined, secondary veins loop-connected near margin; Golfo Dulce
area F. sessifolia

7a. Leaves strongly tripliveined (like that of Melastomaceae); flowers and fruit bright blue; 0-800(-1000)
m elevation F. suerrensis

7b. Leaves not strongly tripliveined and lacking strong lateral veins near the margin (sometimes present
in F. eurycarpa), the 2 veins often loop-connected near the margin with the submarginal vein
arcuate; flowers blue or white 8

8a. Inflorescences umbellate or fasciculate in the leaf axils, without lateral branches; flowers usually
white 9

8b. Inflorescences paniculate, corymbose or cymose, with conspicuous lateral branches from the pri-
mary peduncle or central rachis; flowers blue or white 10

9a. Inflorescences fasciculate, some flowers subtended by ovate bracts 5-1 1 mm long; flowers

not drying black; leaf blades to 1 7 cm long, usually oblong; 0-300 m elevation

F. parvibracteata

9b. Inflorescences umbellate, lacking conspicuous bracts; flowers drying black; leaf blades to 1 1
cm long, elliptic; 1000-2300 m elevation F. ovalis

lOa. Flowers with the corolla tube 12-22 mm long and corolla lobes 8-18 mm long, white and usually
drying black; fruit slightly broader than long, often with persisting calyx tube; stipular awns 4-18
mm long [0-800 m elevation] F. occidentalis

lOb. Flowers with corolla tubes 4-13 mm long, corolla lobes 3-7 (-10) mm long, blue or white and
rarely drying black; fruit distinctly broader than long, a persisting calyx tube rarely present; stipular
awns 1-6 mm long 11

1 1 a. Leaf blades usually drying yellowish green beneath, with a prominent submarginal vein; inflores-
cences robust with branches ca. 1 mm thick when dry; stipules early deciduous; corolla tube 6-
1 mm long; 500-1 700 m elevation F. eurycarpa

1 1 b. Leaf blades usually drying grayish or pale green beneath; inflorescences delicate with primary
branches ca. 0.5 mm thick when dry; 0-1600 m elevation 13

1 2a. Stipules clearly tubular and persisting on the leafy stems; corolla tubes 6-12 mm long, calyx lobes
0.2-0.4 mm long F. multiflora

\ 2b. Stipules quickly deciduous, tubes short and inconspicuous; corolla tubes 2-6 mm long, calyx lobes
0-0.7 mm long 13

13a. Corolla tube slender, 0.7-1.5 mm diam., blue; 0-400 m elevation F. stenura

13b. Corolla tube broad, 1.5-2 mm diam., white; 1300-1400 m elevation F. hondurae

Faramea capulifolia Dwyer, sp. nov. fructum solitarium gerenti. Floras non visi. Fructus glo-

bosus, 6-9 mm longus, glaber, niger.

TYPUS Folsom 4262 (holotypus MO, isotypus ?PMA),

Frutices ad 4 m alti. Foliae lamina lanccolata. 2-6 cm from Cerro Pirre, ridgetop at 1200 m elevation, Darien,
longa, 0.5-1.6 cm lata, acumine ad 10 mm longo, venis Panama,
lateralibus 10-16; petiolis 2-4 mm longis; stipulis 2-8

mm longis, vagina 1-5 mm longa. Inflorescentiae ter- Shrubs, 1-4 m tall, leafy stems 0.5^4 mm thick, gla-

minales, pedunculo nullo vel 1 5-20 mm longo, glabro, brous with 2 or 4 barely distinct longitudinal ridges;

132 FIELDIANA: BOTANY

stipules 2-8 mm long, sheathing tube 1-5 mm long,
truncated distally with 1 small narrow lobe (2/node),
glabrous, deciduous. Leaves with petioles 24 mm long,
ca. 0.5 mm thick, glabrous, sulcate above; leaf blades 2-
6 cm long, 0.5-1.6 cm broad, lanceolate to narrowly
ovate-elliptic, apex acuminate, base acute and slightly
decurrent on petioles, drying chartaceous, olive green
above, glabrous above and below, 2 veins 5-9/side and
loop-connected ca. 1 mm from the leaf margin (major
and intermediate secondaries often difficult to distin-
guish). Inflorescences terminal and solitary, sessile or
with peduncles to 2 cm long in fruit (after the loss of the
distal leaf pair), peduncles ca. 0.4 mm thick and glabrous,
pedicels 0-2 mm long, fruit solitary. Flowers solitary,
ca. 10 mm long, glabrous, corolla ca. 8 mm long, rose-
colored, tube ca. 5 mm long, ca. 0.9 mm diam. Fruits
6-9 mm diam., glabrous, persisting calyx 0.5-1 mm high,
surfaces smooth, brilliant blue or becoming black.

Understory plants in wet cloud forests from 1 000
to 1700 m elevation. Flowering in May; fruiting
in June-July and December. This species is known
only from the Cordillera de Tilaran area in Costa
Rica and eastern Panama.

Faramea capulifolia is recognized by its small
leaves, glabrous parts, and solitary terminal flow-
ers and fruits. The Costa Rican material has some-
what smaller leaves (to 4.5 cm long) and blue fruits,
whereas the type has leaves to 6 cm long and fruits
becoming black. The fruits are subtended by a
node where the leaves have fallen, with the distal
internode appearing to be a peduncle. Flowering
data was added in proof, based on Gomez- La urito
12431 F. Also placed here are Barringer et al. 4194
CR, F, Dryer 1711 F, and Zamora et al. 647 CR, F.

Faramea eurycarpa J. D. Smith, Bot. Gaz. 44: 1 1 3.
1907. F. bocaturensis Dwyer, Ann. Missouri Bot.
Gard. 67: 163. 1980. Figure 45.

Shrubs or small trees to 4(-8) m tall, leafy branchlets
1.5-5 mm thick, glabrous and drying yellowish green;
stipules to 1 6 mm long, united to form a tubular sheath
3-8 mm long, 2-4 mm broad, with awns 1-3 mm long,
deciduous. Leaves with petioles 3-7(-10) mm long, 1-
1.5 mm thick, glabrous; leaf blades (5-)9-16(-19) cm
long, (2-)3-5.5 cm broad, narrowly oblong to elliptic-
oblong, apex obtuse or rounded and acuminate to cau-
date-acuminate, tip 10-14(-20) mm long and ca. 2 mm
broad, base cuneate (or slightly rounded at the petiole),
drying chartaceous, usually yellowish green below with
the midvein yellow, glabrous above and below, 2 veins
9-14/side and loop-connected near the margin to form
an arcuate submarginal vein 2-4 mm from the edge, with
less distinct secondary veins paralleling the major sec-
ondaries. Inflorescences terminal and solitary, to 14 cm
long and 1 2 cm broad, pyramidal panicles with opposite
bracteolate branches, glabrous, peduncles 2-5 cm long,
stout and 1.3-3 mm thick (dried), the slender pedicels

4-10 mm long. Flowers with hypanthium 0.7-1.5 mm
long, calyx cup 0.5-1.5 mm long, calyx teeth 4 or 5, 0.3-
1 mm long, narrow distally; corolla salverform, blue,
purple, or white with purple apex, tube 6-10 mm long,
1-1.5 mm diam., lobes 2-4 mm long, narrowly ovate;
stamens 4, attached near the middle of the tube and
subsessile, anthers 3-4 mm long; style 4-8 mm long,
stigmas ca. 0.7 mm long. Fruits 6-8 mm long and 9-13
mm broad, transverse-reniform to subglobose, blue, pur-
ple, or black, drying dark, smooth or slightly rugose when
dried, persisting calyx less than 1 mm high.

Plants of the very wet evergreen cloud forests
of the Caribbean slope, from (300-)500 to
1200(-1700) m elevation. Flowering in March-
June, September, and December; probably fruit-
ing throughout the year. The species ranges from
Costa Rica and Panama to Colombia and Ecua-
dor.

Faramea eurycarpa is recognized by the oblong
leaves with slender "drip tips," greenish yellow
color of the veins beneath when dried, deciduous
stipules, arcuate submarginal vein, and small blue
flowers with small corolla lobes. The inflorescence
branches and deciduous tubular stipules are much
thicker in texture than those of/", multiflora, which
shares many of the same habitats as F. eurycarpa.
Collections from the Chiriqui Highlands have a
greater range of variation than those found in Cos-
ta Rica, but they do not appear to be specifically
distinct. The name F. bocaturensis Dwyer applies
to the distinctive Chiriqui Highland material.

Faramea hondurae Standl., J. Wash. Acad. Sci. 18:
169. 1928. Figure 45.

Shrubs, 3-4.5 m tall, leafy branchlets 1 .3-3 mm thick,
glabrous; stipules forming a short (3-4 mm) tube, ca-
ducous and leaving a transverse whitish scar. Leaves with
petioles 6-16 mm long, 0.8-1.4 mm broad, glabrous;
leaf blades (6-) 1 0- 1 9 cm long, ( 1 . 5-)3-5 cm broad, nar-
rowly oblong to narrowly elliptic-oblong, apex long-acu-
minate to caudate-acuminate, tip 12-20 mm long, base
acute, drying chartaceous and dark green above, glabrous
above and below, 2 veins ca. 14/side (but with less
prominent 2 veins between the major), an arcuate sub-
marginal vein present near (ca. 2 mm) the margin and
connecting the secondaries. Inflorescences terminal or
axillary to distal leaves, solitary or 3, 5-8 cm long, to 6
cm broad, paniculate with 2 or 3 pairs of opposite
branches, primary peduncles 20-30 mm long, bracts 2-
6 mm long and mucronate, distal bracteoles 0.5-3 mm
long, pedicels 2-5 mm long, slender, glabrous. Flowers
glabrous, hypanthium ca. 1 mm long, turbinate, calyx
tube 0.5-1 .5 mm long, campanulatc. with 4 minute lobes;
corolla salverform, white, tube 2-4 mm long, 1 .5-2 mm
diam.. lobes 1-2 mm long, obtuse; anthers ca. 1.5 mm
long, style and stigma ca. 2.3 mm long. Fruits unknown.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

133

A problematic species known only from two
collections (Standley 36534 us, 37890 us the type)
collected between 1 300 and 1 700 m elevation near
Bajo La Hondura, San Jose, in March 1924. The
short, relatively broad white corolla tubes make
this species unique among our species of Faramea
and make the generic placement doubtful. How-
ever, a dissection of an ovary (Standley 36534)
showed two ovules in a single locule, consistent
with Standley's placement of this species in Fara-
mea.

pearance to F. stenura but the tubular stipules of
F. multiflora are persisting, the calyx lobes are less
well developed, and distal inflorescence branches
lack the small bracts characteristic of F. stenura.
Central American collections differ in minor ways
(generally smaller thinner leaves and inflores-
cences) from South American collections, but all
appear to be part of the same wide-ranging species.

Faramea myrticifolia Dwyer, sp. nov.

Faramea multiflora A. Rich., ex DC., Prodr. 4:
497. 1830. F. talamancarum Standl., Publ. Field
Columb. Mus., Hot. Ser. 4: 332. 1929. Figure
45.

Shrubs or small trees, (l-)2-5(-6) m tall, leafy branch-
lets 1-3.5 mm thick, terete and glabrous; stipules 4-
8(-l 3) mm long, 2-3 mm broad but broader beneath the
inflorescence, the tubular basal sheath 2-5 mm long,
awns l-5(-8) mm long, persisting. Leaves with petioles
3-7(-l 1) mm long, 0.7-1.8 mm thick; leaf blades (6-)8-
13(-17) cm long, 1.5-4(-7) cm wide, oblong to oblong-
obovate or narrowly elliptic-oblong, apex gradually or
abruptly narrowed and caudate-acuminate or acuminate,
tip 7-15 mm long and straight or falcate, base acute or
obtuse, leaves drying thin-chartaceous to chartaceous
and dark greenish above, 2 veins 6-12/side (and lesser
parallel 2 veins between the major), united near the
margin by a slender arcuate submarginal vein 2-4 mm
from the leaf edge. Inflorescences terminal, solitary (or
3), 5-1 4 cm long, to 9 cm broad, paniculate with opposite
branches 1-2 cm long, bright blue, glabrous, primary
peduncles 2-3(-6) cm long, basal branches subtended by
small leaves or broad bracts 8-22 mm long, distal bracts
absent, pedicels 3-7(-10) mm long, slender. Flowers gla-
brous externally, hypanthium 0.5-1 mm long, calyx tube
0.2-0.4 mm long, calyx lobes 4, 0. 1-0.4 mm long; corolla
bright sky blue, salverform, tube 6-12 mm long, 1.2-2
mm diam.. lobes 4, 4-9 mm long, 1.3-2.5 mm broad,
ovate-oblong; anthers 2-3 mm long. Fruits 6-8 mm long,
(8-) 10- 13 mm broad, oblate or transversely reniform,
laterally compressed (oblong in cross-section), surface
smooth and without costae, blue-black at maturity; py-
renes solitary.

Understory shrubs of wet evergreen forest in-
teriors and forest edges, from 20 to 1600 m ele-
vation. Flowering in every month but October
(flowering is mainly in May-June at La Selva);
fruiting throughout the year. The species ranges
from northern Costa Rica to Brazil and Bolivia.

Faramea multiflora is recognized by the thin
smaller leaves (often drying grayish green be-
neath), narrow acuminate apices, persisting tu-
bular stipule sheaths, and bright blue flowers and
inflorescences. This species is very similar in ap-

Suffrutices 0.3-0.4 m alti; ramulis multis glabris. Foli-
ae lamina elliptica 2-4 cm longa, 8-15 mm lata, glabra,
venis lateralibus 6-8; petiolis 3-6 mm longis; stipulis 4-
5 mm longis. Inflorescentiae axillares, flores 2-5 geren-
tes, pedunculis 24 mm longis; pedicellis 1-2 mm longis.
Flores glabri; calycis cupula ca. 0.5 mm longa, lobis ca.
0.5 mm longis, lilacinis; corolla viridi-alba, ca. 4 mm
longa. Fructus ca. 8 mm lati, globosus, azureus, glaber.

TYPUS L. D. Gomez et al. 23401 (holotypus CR, iso-
typus MO), from Las Brisas de Pacuarito, Limon, Costa
Rica.

Herbaceous subshrubs, ca. 40 cm tall, main stem un-
branched for 20 cm and with many slender distal op-
posite horizontal branches, leafy stems 0.5-1 .3 mm thick,
glabrous, prominently 2- or 4-ridged; stipules 4-5 mm
long, with a short (0.5-1 mm) truncated tube and single
filiform central awn 3-5 mm long, persisting. Leaves
with petioles 3-6 mm long, 0.3-0.9 mm thick, glabrous;
leaf blades 2-4 cm long, 8-15 mm broad, elliptic to
elliptic-oblong, apex acute or short-acuminate, base acute
and decurrent on petiole, drying chartaceous, dark green
above, glabrous above and below, 2 veins 6-8/side and
loop-connected 1-1.5 mm from the margin. Inflores-
cences axillary to leaves, 1 or 2/node, cymose umbellate
with 2-5 flowers on a short (2-4 mm) glabrous peduncle
drying dark, glabrous, bracts ca. 2 mm long and linear,
pedicels 1-2 mm long. Flowers glabrous and drying
blackish, hypanthium 0.5-1 mm long, calyx tube ca. 0.5
mm long and truncated with linear lobes 0.5 mm long;
corolla greenish white, ca. 4 mm long and 0.7 mm diam.
in bud. Fruit globose, ca. 8 mm diam., intense blue but
drying black, usually I/node.

Distinctive little plants with many (ca. 1 2) distal
lateral horizontal branches, small leaves, unusual
stipules, and minute flowers. Presently known only
from the type, collected near Siquerres, Limon, 1 8
April 1985, at about 300 m elevation. It appears
to be related to F. cobana J. D. Smith of Honduras,
but that species has larger (8- 1 cm) leaves, longer
peduncles (to 1 5 mm), and a higher-elevation hab-
itat.

Faramea occidentalis (L.) A. Rich., Mem. Fam.
Rubiac. 96. 1830. Ixora occidentalis L., Syst.
Nat. ed. 2: 893. 1759. F. zeteki Standl., Contr.

134

FIELDIANA: BOTANY

Arnold Arbor. 5: 147. 1933. F. belizensis Standl.,
Carnegie Inst. Wash. Publ. 461: 90. 1935. F.
standleyana L. O. Williams, Phytologia 26: 490.
1973. Figure 44.

Shrubs or small trees, 2-6(-10) m tall, to 20 cm dbh,
leafy branchlets 1.5-5 mm thick, glabrous, terete, often
dichotomously or trichotomously branched; stipule
sheath 2-8 mm long, 5-8 mm wide at the base, with
awns 4-12(-18) mm long, deciduous. Leaves with pet-
ioles 6-15 mm long, 1-2 mm thick, glabrate; leaf blades
8-18(-21)cm long, (2.5-)3.5-9.5(-l l)cm broad, oblong
or elliptic to narrowly elliptic-obovate (less often ovate-
oblong), apex rounded to obtuse and acuminate to cau-
date-acuminate, the narrow (2 mm) tip 4-17 mm long,
base acute to obtuse or rounded and subtruncate, drying
chartaceous to stiffly chartaceous, glabrous above, gla-
brous or minutely puberulent beneath, 2 veins 6- 107
side and only loosely loop-connected near the margin (a
definite submarginal vein absent), with 1-3 more weakly
defined secondaries between the major secondary veins.
Inflorescences terminal or less often axillary, solitary or
several, 5-12 cm long, equally broad, umbelliform or
trichotomous. few-branched and with 3-9 flowers, often
drying black, peduncles 1.5-6 cm long, lateral branches
opposite and few, distal flowers usually in groups of 3,
bracts 4 mm long, linear, pedicels 3-12(-20) mm long.
Flowers white but drying black, sweet scented in life,
probably nocturnal, hypanthium ca. 2 mm long, oblong,
calyx cup 1.5-3 mm long, teeth absent or minute; corolla
salverform, tube 10-19(-22) mm long, 2-3 m diam.,
narrowly cylindrical, lobes 4, 8-16(-25) mm long, 1.3-
3 mm wide near the base, lanceolate to narrowly ovate;
stamens 4, filaments 0.5 mm long, anthers ca. 8 mm
long, slightly exserted; style as long as the corolla tube
or 'Ath as long, stigma 2.5-5.5 mm long. Fruits 6-9 mm
long, 9-14 mm diam., globose to subglobose-oblate. per-
sisting calyx 1-2 mm long and 2-3 mm diam., drying
black.

Trees and shrubs of evergreen forest formations,
from near sea level to 1 000 m elevation. Flowering
in February-July and October in southern Central
America (primarily in June in Costa Rica); prob-
ably fruiting throughout the year. The species
ranges from southern Mexico, Central America,
and the West Indies through tropical South Amer-
ica.

Faramea occidentalis is recognized by the large
slender flowers drying black, relatively few-
branched and few-flowered inflorescences also
drying black, and rounded fruit slightly shorter
than broad. This species varies greatly in leaf form
and it does not have a well-developed submarginal
vein. Though wide-ranging, it has not been col-
lected very often in Costa Rica. This species is
closely related to F. luteovirens with larger flowers.
Fruiting and sterile material may be difficult to
separate from F. eurycarpa.

Faramea ovalis Standl., Contr. U.S. Natl. Herb.
18: 138. 1916. F. quercetorum Standl., J. Wash.
Acad. Sci. 18: 168. 1928. Figure 45.

Small trees or shrubs, 2-6(-13) m tall, to 20 cm dbh,
leafy branchlets 1-4 mm thick, glabrous, becoming gray-
ish; stipules 3-13 mm long, the basal cupulate tube 1-
2 mm high, with a single slender awn 3-10 mm long,
often persisting. leaves not closely congested, petioles
3-10 mm long. 0.7-1.8 mm thick, glabrous; leaf blades
3-1 1 cm long, 1.5-4.8 cm broad, elliptic-oblong to ob-
long or narrowly elliptic, apex abruptly narrowed and
acuminate or caudate-acuminate, the tip 6-1 3 mm long,
base acute to obtuse, drying stiffly chartaceous and gray-
ish green to dark green above (slightly paler beneath),
glabrous above and below, 2 veins (3-)5-8/side and
loop-connected distally, with a vein-like edge along the
leaf margin. Inflorescences terminal (axillary), 3-7 cm
long, to 4 cm broad, umbellate with 3-5(-9) flowers,
peduncles 8-24 mm long, 0.5 mm diam., glabrous and
drying dark, pedicles 4-18 mm long, ca. 0.5 mm thick.
Flowers glabrous externally, drying dark, hypanthium 1-

2 mm long, calyx tube 0.5-2 mm high, teeth to 0.5 mm
long; corolla salverform to somewhat funnelform, white
or tinted with pink-purple, tube (7-)9-14 mm long, 1 .2-

3 mm diam., lobes 4-6 mm long; stamens 4. Fruit 6-10
mm diam., globose, blue drying black and smooth, per-
sisting calyx tube 1-2.5 mm long and ca. 2 mm diam.

Plants of moist evergreen cloud forests, 1 000-
2300 m elevation (400-600 m on Volcan Orosi).
Flowering in March-June and December; prob-
ably fruiting throughout the year. The species is
found in the northwestern cordilleras, eastern parts
of the Cordillera de Talamanca, and the Chiriqui
Highlands.

Faramea ovalis is recognized by its cloud forest
habitat, lack of pubescence, smaller often caudate-
acuminate leaves, umbellate inflorescences with
relatively few large flowers, and globose black fruits.
The species resembles smaller-leaved specimens
of F. occidentalis and some species of Coussarea.

Faramea parvibractea Steyerm., Mem. New York
Bot. Gard. 17: 376. 1967. Figure 45.

Shrubs or small trees 2-8 m tall, leafy branchlets 1-4
mm thick, glabrous and drying greenish, strongly qua-
drangular; stipules 6-15 mm long, 0.5-1 mm broad,
encircling the shoot apex, quickly caducous. Leaves with
petioles 5-14(-20) mm long, ca. 1 mm thick, glabrous;
leaf blades 7-17 cm long. 1.5-6(-7.5)cm broad, elliptic-
oblong, ovate-oblong, to oblong-lanceolate, apex acu-
minate to caudate-acuminate, tip 6-12 mm long, base
acute to obtuse, drying chartaceous, glabrous above and
below, 2 veins 6-10/side. weakly loop-connected 3-5
mm from the margin. Inflorescences fasciculate or pseu-
doumbellate from distal nodes, to 6(-10) cm long and
equally broad, with 6-ll(-19) primary peduncles 11-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

135

20(-30) mm long, flowers usually borne in distal groups
of (l-)2, 3, or 5 at apex of the primary peduncles, a few
peduncles with broadly ovate leaf-like bracts 5.5-16 mm
long and 3-12 mm broad at their apex, bracts subcordate
and with petiole-like base ca. 1 mm long, white in life,
pedicles 1-3 mm long. Flowers with hypanthium 0.5-
1.5 mm long, calyx tube 0.5-1 mm long, calyx lobes
small and usually unequal (ca. 0.1 mm and 0.5 mm),
corolla funnelform, white, 7-12(-16) mm long, tube 5-
8 mm long, 1.5-3 mm diam., lobes 4, 5-10 mm long,
2-3.5 mm broad, lanceolate to narrowly oblong, obtuse;
stamens exserted, filaments 3-3.5 mm long, inserted at
or below the middle of the tube, anthers 3.5-4 mm long,
linear, bluntly rounded; style 4-6 mm long. Fruits 5-7
mm long, 8-12 mm broad, oblate and rounded in cross-
section, smooth, the ribs obscure or prominent when
dried; pyrene solitary.

Plants of wet lowland rain forest formations of
the Caribbean slope, from 4 to 600 m elevation.
Flowering in January-June and October; fruiting
in February and May-August. The Costa Rican
collections are mostly from between Tortugero and
Limon, with a few from the Osa Peninsula. This
species is also known from Panama and Vene-
zuela.

Faramea parvibractea is recognized by the clus-
ters of few-flowered peduncles at distal nodes, the
ovate-subcordate bracts subtending some flower
groups, the elliptic-oblong leaves with "drip tips,"
the narrow stipules enclosing the shoot apices, the
glabrous white flowers, and the fruit often in um-
bellate groups at the ends of leafy stems. The un-
usual bracts do not appear to be developed be-
neath all the flower groups, but they are large and
contradict the specific name.

Faramea pauciflora Dwyer, Ann. Missouri Bot.
Gard. 67: 172. 1980.

Understory shrubs or small trees, 2-5(-10) m tall,
branches at right angles to main stem, leafy stems 1.5-
4 mm thick, glabrous; stipules 8-15 mm long and 4 mm
broad, triangular to tubular with 2 slender awns 3-5 mm
long, covering the terminal bud, glabrous, drying black,
caducous. Leaves held in a single plane, petioles 4-7(-l 2)
mm long, 0.9-1.5 mm thick, glabrous, drying dark; leaf
blades 11-22 cm long, 4-11 cm broad, elliptic-oblong
to narrowly oblong or elliptic-obovate, apex acuminate
or caudate-acuminate, tip 7-17 mm long, base acute,
drying stiffly chartaceous, dark brown or blackish above,
glabrous above, glabrous or sparsely and minutely pa-
pillate-puberulent beneath, 2 veins 8-12/side. Inflores-
cences ca. 10 cm long with 1-3 flowers, terminal or ax-
illary, usually I/node, peduncles 2-5 cm long, 0.7-1.5
mm thick, often with 1-2 pairs of stipule-like bracts near
the base and terminated by several minute bracts, ped-
icels 1 5-55 mm long, 0.5 mm thick, glabrous and drying

black, merging with the hypanthium. Flowers glabrous
externally and drying black, hypanthium and calyx 10-
1 5 mm long, 2.5-4 mm diam., tube ca. 4 mm long, lobes
few or unequal, 1-8 mm long; corolla salverform, white,
tube 10-22 mm long, lobes 4, 13-30 mm long, rotate
and with a narrow tip 4-9 mm long often held at 90.
Fruits 12-14 mm long, 10-14 mm diam., with a per-
sisting calyx tube 7-9 mm long and 3-4 mm diam., dark
blue drying black, pendulous.

Plants of the wet Caribbean slope at elevations
from 20 to 1100 m. Flowering in March-April;
fruiting in March-April and August-September in
Panama. The species ranges from near Nuevo Ar-
enal southward to Colombia.

Faramea pauciflora is unusual because of its
glabrous parts, usually axillary inflorescences with
one to three long white flowers and long pedicels.
Two Costa Rican collections have been seen: Her-
rera 2556 and A. Smith 1644. This species is close-
ly related to F. luteovirens Dwyer of Panama with
thicker leaves, fewer 2 veins, and shorter calyx.
Compare F. occidentalis and the pubescent but
similarly few-flowered Coussarea enneantha (fig.
46).

Faramea scalaris Standl., Contr. U.S. Natl. Herb.
13: 139. 1916.

Shrubs or small trees, 1.5-6 m tall, leafy branchlets
1.5-4 mm thick, terete or quadrangular, drying brown-
ish, leaves distant (3-6 cm) along the stems; stipules 4-
8 mm long, with a short-tubular sheath, truncated or
deltoid at apex, with or without a short (1-2 mm) awn,
persistent. Leaves subsessile with petioles l-3(-5) mm
long; leaf blades (5-) 7- 15 cm long, (1-) 1.3-4 cm broad,
narrowly elliptic-oblong to narrowly oblanceolate, apex
acuminate, tip 7-15 mm long and straight or falcate,
base gradually narrowed but often slightly rounded
and subauriculate at the petiole, drying chartaceous and
greenish, glabrous above and below, 2 veins 9-13/side,
these arising at almost 90 angles and united by a linear
or slightly arcuate lateral vein 2-5 mm from the leaf
edge, a slender submarginal vein also present 0.3-1 mm
from the margin in fully developed leaves. Inflorescences
terminal and solitary, 3-6 cm long, paniculate with short
opposite branches, primary peduncles 1 5-20 mm long,
glabrous, bracts caducous, pedicels 2-4 mm long. Flow-
ers glabrous, hypanthium ca. 1.5 mm long, calyx tube
ca. 0.5 mm long, lobes 0.1-0.3 mm; corolla salverform,
blue becoming white, tube 7-10 mm long, 1-1.5 mm
diam., lobes 3-4 mm long, ca. 3 mm broad, ovate, disc
1 mm long; style ca. 7 mm long. Fruits unknown.

Plants of montane wet forest formations, from
1800 to 2300 m elevation. Flowering in January
and March. The species is known only from the
Chiriqui Highlands in western Panama.

136

FIELDIANA: BOTANY

Faramea scalaris is distinguished by its narrow
subsessile leaves with prominent lateral (mela-
stome-like) venation, the small inflorescences, and
the restricted high-elevation habitat. The unusual
leaf venation suggests that this species is closely
related to F. trinervia and F. suerrensis of lowland
formations.

Faramea sessifolia P. Allen, Rain Forests of Golfo
Dulce 409. 1956. Figure 44.

Shrubs or small trees, 3-7 m tall, leafy branchlets 2-
7 mm thick, the nodes to 10 mm thick, rectangular in
cross-section (flattened and 4-angular), glabrous, drying
yellowish; stipules 10-25 mm long, 3-5 mm broad at
the base, triangular-subulate with a narrow distal tip,
caducous. Leaves somewhat dimorphic with smaller and
narrower leaves often subtending the inflorescences, ses-
sile or subsessile, petioles 0-^4(-6) mm long, ca. 2 mm
thick; larger leaf blades 17-27 cm long, 6-17 cm broad,
narrowly oblong to ovate-oblong, smaller leaf blades 8-
1 7(-l 9) cm long and 2-4(-6) cm broad, narrowly oblong-
lanceolate to narrowly oblong, apex abruptly narrowed
(in larger leaves) or gradually narrowed (in smaller leaves)
and acuminate, tips 8-17 mm long, base rounded and
subtruncate, drying chartaceous and yellowish green, gla-
brous above and below, 2 veins 12-16/side and loop-
connected 2-5 mm from the margin. Inflorescences ter-
minal, solitary or 3, 10- 15 cm long and 10-1 2 cm broad,
paniculate with opposite many-flowered branches, pri-
mary peduncles (2-)6-7 cm long, 0.7-2 mm thick, gla-
brous, whitish in life and yellowish when dried, bracts
0.5-1 mm long, pedicels 1-2 mm long above small (0.5
mm) bracteoles. Flowers glabrous, hypanthium 0.5-1
mm long, calyx tube 0.2-0.5 mm long, lobes 0.2-0.4
mm long; corolla salverform, bright blue or bluish pur-
ple, tube ca. 6-7 mm long and 1 mm diam., lobes 3-4
mm long, to 2 mm broad at base. Fruits 4-5 mm high,
7-9 mm broad, oblate, circular or oblong in cross-sec-
tion, with 8 longitudinal costae, pale grayish brown be-
coming black.

Plants of the lowland rain forests of the Osa
Peninsula region, 20-400 m elevation. Flowering
in April-May (Allen 5539 F, us type); fruiting in
March, August-September, and November. The
species is known only from southwestern Costa
Rica.

Faramea sessifolia is recognized by its large sub-
sessile leaves with many secondary veins and ar-
cuate submarginal vein, leafy stems rectangular in
cross-section, lack of pubescence, blue flowers, and
restricted distribution. The veins on the lower leaf
surface and inflorescences dry yellowish. This name
should not be confused with F. sessilifolia (H.B.K.)
A. DC. of South America.

Faramea stenura Standl., Publ. Field Columb.
Mus., Hot. Ser. 4: 331. 1929. Figure 45.

Small trees to 7 m tall, leafy branchlets 1.7-4 mm
thick, glabrous, drying greenish; stipules 6-10 mm long,
3-4 mm broad at the base, triangular to ovate, acute or
mucronate at apex, caducous. leaves with petioles 5-10
mm long, 0.6-1.5 mm thick; leaf blades 9-19 cm long,
2.5-5.5 cm wide, narrowly oblong to narrowly oblong-
obovate, apex acuminate, tip 5-23 mm long, base acute
to cuneate, drying chartaceous, dark grayish green above,
glabrous above and below, 2 veins 9-12/side, arising at
almost 90 and weakly loop-connected near the margin,
with thinner secondaries present and parallel with the
larger. Inflorescences usually terminal and solitary, pa-
niculate pyramidal, 3-5.5 cm long, 4-7 cm broad, blue,
primary peduncles 1.5-5(-8) cm long and 1-1.5 mm
thick, bracts 3-5 mm long, linear or triangular, purplish,
pedicels 2-4 mm long. Flowers glabrous externally, hy-
panthium ca. 0.6 mm long, calyx tube 0.2-0.3 mm long,
lobes 4, 0.3-0.7 mm long; corolla salverform, pale to
deep blue, glabrous, tube 5-8 mm long, lobes 4, 3-5 mm
long, 1.2-1.5 mm broad, elliptic to ovate, acute at apex.
Fruits 4-8 mm long, 12-15 mm broad, transversely reni-
form (oblong in cross-section), rounded basally and flat-
tened or depressed above, smooth and usually drying
green.

Plants of lowland rain forest formations, 30-
600 m elevation (but see below). Flowering in Jan-
uary-September; probably fruiting throughout the
year. It ranges along the Caribbean coast, from
Veracruz, Mexico, to western Panama, and in the
Golfo Dulce area.

Faramea stenura is distinguished by its small
blue corollas, caducous stipules, oblate fruit, and
narrowly oblong leaves with long "drip tips." This
species is very similar to F. multiflora, but their
stipules help differentiate most collections. At La
Selva F. stenura grows in low swales, whereas F.
multiflora grows on the ridgetops. There are very
similar plants growing at higher elevations (ca.
1200 m) in Chiriqui with larger (10 x 14 mm)
ribbed fruit that dry whitish. It is not clear whether
they are a high-elevation subspecies or an unde-
scribed closely related species.

Faramea suerrensis (J. D. Smith) J. D. Smith, Bot.
Gaz. 44: 1 1 2. 1 907. Faramea trinervia K. Schum.
& J. D. Smith var. suerrensis J. D. Smith, Bot.
Gaz. 31: 115. 1901. Figure 44.

Shrubs or small treelets, 2-6 m tall, leafy branchlets
2-6 mm thick (to 12 mm broad at the node), usually
quadrangular in cross-section, glabrous; stipules 5-10
mm long, 3-8(-12) mm wide at the base, often united
to form a short tube, entire or rounded with a small ( 1

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

137

mm) mucronate tip. Leaves with petioles 4-18 mm long,
1-2 mm thick; leaf blades 9-20(-28) cm long, 3-8(-12)
cm broad, narrowly elliptic-oblong to oblong or narrowly
oblong-oblanceolate, apex short-acuminate to long-cus-
pidate, tip to 1 8 mm long in some specimens, base acute
to rounded and subtruncate, drying chartaceous to sub-
coriaceous, pale grayish green to yellowish green, gla-
brous above and below, the major veins often becoming
impressed above, 2 veins 8-16(-25)/side and with less
prominent parallel 2 veins, 2 veins arising at nearly 90
from the midvein and joined near the margin by a prom-
inent linear lateral vein (2-)5-7(-14) mm from the leaf
edge, with a smaller submarginal vein 0.5-2 mm from
the leaf edge. Inflorescences terminal and solitary (or 3),
corymbose panicles with opposite branching, 9-12(-20)
cm long, to 16 cm broad, bright blue, many-flowered,
peduncles 2-5(-9) cm long, 2-3 mm thick, glabrous, ped-
icels 2.5-9 mm long. Flowers distylous, glabrous, hy-
panthium ca. 1.2 mm long, calyx tube a small (0.3 mm)
rim, lobes minute or 0.5 mm long and triangular; corolla
salverform, brilliant pale blue to deep blue, tube 6-10
mm long, 0.8-1.5 mm diam., lobes 4, 3-5 mm long, ca.
1.7 mm broad; stamens 4, filaments attached near the
middle of the tube, anthers 2.2 mm long, included. Fruits
6-1 1 mm long, 12-16 mm broad, transversely reniform
and rounded-oblong in cross-section (somewhat flat-
tened laterally and flat or slightly depressed distally),
longitudinal ribs slightly developed or obscure, exocarp
spongy and deep blue in life; pyrenes solitary.

Plants of lowland rain forest formations on both
the Pacific and Caribbean slopes, from near sea
level to 800(-1000) m elevation. Flowering in Jan-
uary-August and November; fruiting in every
month except December. The species ranges from
southern Nicaragua through Costa Rica and Pan-
ama to northwestern Colombia.

Faramea suerrensis is distinguished by its larger
leaves with prominent melastome-like venation,
brilliant bluish inflorescences and flowers, lack of
pubescence, and lowland rain forest habitat. The
unusual quality of the blue coloring of the inflo-
rescence and the leaf venation make this one of
our most distinctive species of Rubiaceae. This
species is very closely related to F. trinervia (q.v.);
it is possible that the two may be conspecific as
Schumann and Smith originally thought. The
breeding system was studied by Bawa and Beach
(1983).

Faramea trinervia K. Schum. & J. D. Smith, Bot.
Gaz. 31: 115. 1901. F. bullata Standl., Publ.
Field Mus. Nat. Hist., Bot. Ser. 4: 294. 1929.
Figure 44.

Shrubs or small trees, leafy stems 2-6 mm thick, gla-
brous; stipules 6-10 mm long, the basal tube 2-4 mm
long with rounded or acute apex and narrow awn 3-5
mm long. Leaves sessile or subsessile with petioles 2-4

mm long; leaf blades 14-31 cm long, 4-12 cm broad,
narrowly oblong to narrowly oblong-obovate, apex acu-
minate or caudate-acuminate, somewhat narrowed be-
low the middle, base abruptly rounded and subcordate,
drying stiffly chartaceous to subcoriaceous, grayish green,
glabrous above and below, 2 veins 10-1 7(-22)/side (dif-
ficult to separate from the less prominent secondaries),
united near the margin by a linear lateral vein 5-10 mm
from the leaf edge, a smaller submarginal vein also pres-
ent 1-3 mm from the edge. Inflorescence solitary and
terminal (or 3 with 2 axillary), 4-7 cm long, to 12 cm
broad, paniculate with opposite or trichotomous branch-
ing at the apex of the peduncle, basal bracts (small leaves)
to 2 cm long, peduncles 3-6 cm long, 2.2-3.4 mm thick.
Flowers glabrous externally, hypanthium ca. 1 mm long,
obconic, calyx limb ca. 0.3 mm long with minute (0.2
mm) lobes; corolla color not known, salverform, tube
6-7 mm long, 0.5-0.9 mm diam., lobes 4, ca. 4 mm long
and 1 mm broad. Fruits 7-8 mm long and 12-16 mm
broad, oblate-reniform, flat or depressed centrally above,
drying smooth and black.

A species of lowland Caribbean rain forest for-
mations, 10-600 m elevation. Flowering in March
(Tonduz 8690 us) and April (Tonduz 9583 us the
type); fruiting in April (Barringer et al. 2632 CR,
F) and July (/. Chacon 209 CR). This species is
known only from the Talamanca valley region,
southeastern Costa Rica, and Bocas del Toro
Province in Panama.

Faramea trinervia is recognized by the stiff nar-
rowly oblong leaves with melastome-like vena-
tion, coupled with the short petioles and rounded
auriculate leaf bases. This species is very similar
to and may prove to be conspecific with material
placed under F. suerrensis (a later name). The
thicker subsessile leaves rounded at the base are
a unique character combination within the more
common and more widely ranging F. suerrensis,
and we treat the two as distinct species, although
they appear identical in most other respects. The
type of F. bullata (Cooper 507 F from Panama)
has short petioles but with a rounded lamina base,
and we interpret it as an aberrant form of F. tri-
nervia.

Ferdinandusa Pohl

Trees or shrubs, stems glabrous or puberulent; stipules
interpetiolar, triangular, deciduous or caducous. Leaves
opposite or verticillate, decussate or distichous, petio-
late, glabrous or puberulent, entire, pinnately veined,
without domatia. Inflorescences terminal or axillary to
the distal leaf pair, paniculate with opposite branching
and cymose to corymbose in form (rarely fasciculate or
umbellate), bracteate, flowers pedicillate. Flowers bisex-
ual and usually radially symmetrical (corolla tube some-
times curved), monomorphic, calyx cupular, calyx teeth

138

FIELDIANA: BOTANY

short, deciduous or persistent; corolla salverform to fun-
nelform, white to reddish or greenish, corolla lobes 4(-5),
convolute or broadly imbricate in bud, rounded distally;
stamens 4(-5), filaments attached in the upper part of
the tube, anthers versatile, exserted or included; ovary
2-locular with few to many ovules vertical on the axile
placentas, stigmas subcapitate. Fruits a cylindrical to
oblong (subglobose) woody capsule with septicidal de-
hiscence from apex; seeds few to many, elliptic and flat-
tened, with entire to lacerate marginal wings.

A genus of 20-25 species with 1 species in Costa
Rica and Panama and the others in South Amer-
ica. The elongate capsular fruit with small winged
seeds and broadly overlapping corolla lobes dis-
tinguish this genus. Compare this genus with ma-
terial placed in Ladenbergia (with valvate corolla
lobes) and Macrocnemum.

Ferdinandusa panamensis Standl. & L. O. Wil-
liams, Ceiba 3: 34. 1952. Figure 40.

Trees to 20 m tall, major branches held at 90 angles
or drooping, leafy stems 2-5 mm thick, terete or slightly
quadrangular, glabrous or with erect hairs 0.5-1 mm
long, stems slightly resinous where the hairs are broken
off; stipules 5-20 mm long, narrowly triangular, acu-
minate, glabrous, caducous. Leaves opposite or 3/node,
smaller in size beneath the inflorescence, petioles 4-10
mm long, 1.5-3 mm thick, glabrous or pubescent; leaf
blades (6-)10-18(-22) cm long, (4-)6-10 cm broad, ob-
long to elliptic-oblong or ovate-oblong, apex abruptly
short-acuminate, tip 3-10 mm long, base obtuse to
rounded or truncate (subcordate), leaves drying stiffly
chartaceous, glabrous above and below or with erect
brownish hairs 0.5-1 mm long on the veins beneath and
more sparsely on the surfaces beneath, 2 veins 6-9/side,
arcuate-ascending near the margin but loop-connected
only near apex, domatia absent. Inflorescences terminal
(apparently axillary when leaves develop at the first
branching node of the panicle), solitary or 3, 6-15 cm
long, 6-22 cm broad, primary peduncles 2-5 cm long,
bracteolate, bracts ca. 1 mm long, pedicels 5-12 mm
long and merging with the flower base. Flowers glabrous,
hypanthium ca. 2 mm long, calyx cup 0.5-1 mm long,
2-4 mm diam., calyx teeth 4 or 5, ca. 0.5 mm long;
corolla funnelform, yellowish green or white, fleshy, tube
(4-)6-18(-25) mm long, 2-3 mm diam., slightly ex-
panded at the base, and much expanded distally, lobes
5, 4-6 mm long, 3-6 mm broad at the base; stamens 5,
filaments linear, attached near the middle of the tube,
anthers sometimes dimorphic (long and short); style to
11 mm long, stigmas ca. 1.5 mm long. Fruits (2-)3-6
cm long, 6-1 2 mm thick, narrowly oblong or cylindrical,
abruptly rounded at the base and apex, with a short (0.5-
1 mm) persistent calyx, surface smooth and with obscure
longitudinal ribs, brown; seeds 1-2 cm long, elliptic.

Trees of lowland Caribbean rain forest forma-
tions, from near sea level to 200 m elevation.
Rowers have been collected in January-May and

November-December; fruits were collected in
February-July. The species is known only from
Costa Rica and near Chiriqui Lagoon in Panama
(von Wedel 2232 F the type).

Ferdinandusa panamensis is recognized by the
elongate woody capsules with small winged seeds,
fleshy flowers with broadly overlapping corolla
lobes, broad leaves often rounded at the base, and
the occasional presence of stiffbrownish hairs. The
names cafe macho and cafecillo have been used
for this species in Costa Rica. The wood is very
hard and young trees are used for boat poles.

Galium Linnaeus

REFERENCES L. Dempster, The genus Galium
(Rubiaceae) in Mexico and Central America. Univ.
Calif. Publ. Bot. 73: 1-33. 1978. The genus Galium
(Rubiaceae) in South America, IV. Allertonia 5:
283-345. 1990. F. Ehrendorfer, Revision of the
genus Relbunium (Endl.) Benth. & Hook. (Rubi-
aceae-Galieae). Bot. Jahrb. Syst. 76: 516-553.
1955.

Herbs or slender vines, annual or perennial, often
woody at the base (rarely shrubs), erect, decumbent or
climbing, monoecious or dioecious, stems usually slen-
der and with 4 prominent longitudinal ridges (square in
cross section), glabrous or puberulent with thin hairs;
stipules apparently absent but represented by leaf-like
parts, indistinguishable from the true leaves (except that
they lack axillary buds). Leaves in whorls of (3-)4-8(-10
to many) at each node, comprising the true leaves and
the transformed stipules, sessile or subsessile, often nar-
rowly oblong, entire or with sharp retrorse trichomes
along the edge (serrulate), with 1 or 3 major veins, with-
out domatia. Inflorescences terminal or axillary, often
paniculate with dichasial or trichotomous branching, or
with 3 terminal flowers from the distal node, inflores-
cences often with bract-like reduced leaves and thin-
divaricate branching (with a 4-parted involucre in spe-
cies formerly placed in Relbunium), pedicels articulate
beneath the flower. Flowers bisexual or unisexual, ra-
dially symmetrical, usually very small, calyx tube minute
or lacking; corolla rotate to campanulate or urceolate,
white to yellow, green, pink, or red, corolla lobes (3-)4(-5),
valvate in bud; stamens (2-)3-4(-5), filaments attached
to the short tube of the corolla, anthers versatile and
exserted; ovary 2-locular with 1 ovule borne on the sep-
tum in each locule, styles 2 (sometimes united at the
base), stigmas capitate. Fruits usually 2-lobed or 2-parted
(singular when 1 ovule fails to develop), dry or fleshy,
smooth and glabrous to tuberculate or densely hispidu-
lous, small, finally separating into 2 1 -seeded mericarps,
the mericarps rounded and indehiscent; seeds convex
dorsally, attached to the pericarp, testa membranous.

A large genus of some 300-400 species, es-
pecially well represented in the North Temperate

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

139

zone and at higher elevations in the tropics.
Dempster listed 4 1 species in Mexico and Central
America (3 of which are probably early introduc-
tions); the genus is especially well represented in
Baja California and the central highlands of Mex-
ico. Galium, as a genus, is easy to recognize with
its small whorled leaves on slender herbaceous

clambering 4-angled stems, minute flowers on
slender pedicles, and 2-lobed little fruit. Material
of Galium may be mistaken for species ofDidym-
aea and Nertera. This treatment is based in large
part on the annotations and publications of Laura-
may Dempster.

Key to the Species of Galium

la. Leaves in whorls of 6 or 8(-10); fruit covered with ascending curved hairs G. mexicanum

1 b. Leaves in whorl of 4 at each node; fruit glabrous or with uncinate (hooked at the tip) hairs ... 2
2a. Flowers solitary in the leaf axils and subtended by a whorl of 4 leaf-like or calyx-like bracts

G. hypocarpium

2b. Flowers rarely solitary in leaf axils, rarely subtended by 4 bracts 3

3a. Fruits glabrous, leaves with 1 primary vein and no lateral veins [usually glabrous; stems with minute

(0. 1-0.2 mm) hairs] G. aschenbornii

3b. Fruits covered with ascending uncinate hairs; leaves with 1 primary vein and 2 lateral veins usually

visible 4

4a. Leaf blades usually ovate to elliptic; stems and leaves with few to many longer (0.4 mm) hairs;

inflorescences usually with few (ca. 5) flowers; corollas often hispidulous G. uncinulatum

4b. Leaf blades usually oblong-lanceolate to elliptic, stems and leaves sparsely short-puberulent (ca. 0.2

mm); inflorescences usually with more than 5 flowers; corollas glabrous G. orizabense

Galium aschenbornii Schauer, Linnaea 20: 701.
1 847. Relbunium aschenbornii (Schauer) Hemsl.,
Biol. centr. amer. Bot. 2: 62. 1881. Figure 3.

Creeping or climbing herbs to 1 .2 m long, with slender
woody or herbaceous stems arising from a small root-
stock, sometimes rooting from distal nodes, with both
long (5 cm) and short (4 mm) internodes, leafy stems
0.3-1 mm thick, glabrous or with short retrorse hairs
0. 1-0.2 mm long; stipules leaf-like. Leaves usually 4/node,
petioles 0. 1-0.5 mm long; leaf blades 4-8(-14) mm long,
1 .2-3(-4) mm broad, narrowly oblong or elliptic-oblong
(to lanceolate or ovate-oblong), apex obtuse to acute with
a short (0.3 mm) tip, base acute to obtuse (rarely round-
ed), margin entire or with a few retrorse aculeolate tri-
chomes, midvein prominent, secondary and lateral veins
obscure or weakly developed. Inflorescences often of 3
terminal flowers subtended by a whorl of 4 reduced leaves,
or of several flowers in compound dichasia, pedicels 1-
5 mm long, filiform. Flowers ca. 1.5 mm long, greenish
yellow, said to be <J, 2, and bisexual on the same plant
or on different plants, hypanthium 0.5-0.7 mm long,
calyx ca. 0.3 mm long; corolla rotate, lobes 0.5-1 mm
long, white to yellowish or red. Fruits 3-4 mm long, 4-
7 mm broad when dry, 2-lobed distally, orange becoming
black, glabrous and wrinkled when dried, borne on slen-
der pedicels 2-5 mm long.

Plants of evergreen montane forest formations,
from 1 200 to 2900(-3400) m in Central America.
Probably flowering and fruiting throughout the

year. The species ranges from the states of Jalisco
and San Luis Potosi in Mexico to western Panama.
Galium aschenbornii is recognized by the usu-
ally narrow leaves in whorls of four, short petioles,
and glabrous fruit. It is infrequently collected in
Costa Rica.

Galium hypocarpium (L.) Clos in Gay, Fl. Chil. 3:
186. 1847. Vaillantia hypocarpia L., Syst. Nat.
ed. 10: 1307. 17 59. Relbunium hypocarpium (L.)
Hemsl., Biol. cent. amer. Bot. 2: 63. 1881. G.
hypocarpium (L.) Fosberg, Sida 2: 386. 1966.
Figure 3.

Herbs to 60 cm high, prostrate to procumbent or
climbing over low objects, leafy stems 0.4-1 .3 mm thick,
with 4 prominent longitudinal ridges, nodes usually well
spaced, puberulent with thin whitish hairs 0.3-1 mm
long; stipules leaf-like. Leaves 4/node, sessile; leaf blades
3-1 3(-22) mm long, 2-4(-8) mm broad, oblong to ovate-
oblong or broadly oblong-obovate, apex obtuse to
rounded, with a minutely apiculate tip, base obtuse, dry-
ing chartaceous to subcoriaceous, both surfaces with stiff
ascending hairs to 0.7 mm long or the surfaces glabrous
and the margins hirsutulous. 2 veins 2-3/side (or the
venation obscure). Inflorescences of solitary flowers in
the leaf axils, usually with 2 flowers per node, peduncles
5-15 mm long, with an involucre of 4 bracteoles sub-
tending each flower, bracteoles subequal, 2-4 mm long,

140

FIELDIANA: BOTANY

ovate to oblong or lanceolate, hirsutulous. Flowers small,
hypanthium rounded, calyx tube and teeth undeveloped;
corolla campanulate, white, 1.5-2.5 mm long, tube cy-
lindrical, lobes 4, shorter than the bracteoles and alter-
nating with them, surfaces glabrous and with marginal
hairs; stamens 4, filaments short, anthers small. Fruits
2-3 mm long, to 3.5 mm broad, broadly 2- or 3-lobed,
or with only 1 seed and globose, orange or reddish or-
ange, glabrous to puberulent.

Plants of moist evergreen montane forest for-
mations and high elevation paramos, from 1800
to 3400 m elevation. Probably flowering and fruit-
ing throughout the year. The species ranges from
Veracruz, Mexico, southward through the higher
elevations of Central America into the Andes
mountains of South America as far south as north-
ern Chile and Argentina.

Galium hypocarpium is recognized by its slender
stems with four leaves at each node, creeping or
clambering habit, the solitary axillary flowers sub-
tended by an involucre of four leaf-like bracteoles.
These plants closely resemble our other species of
Galium, but the other species rarely have single
flowers subtended by a four-parted involucre.

Galium mexicanum H.B.K., Nov. gen. sp. 3: 337
(quarto). 1818. Figure 3.

Trailing or climbing herbs to l(-2.4?) m long, leafy
internodes 0.3-2.8 mm thick, with minute (0. 1-0.3 mm)
retrorse aculeolate hairs, often with dense longer (0.5
mm) whitish hairs at the node; stipules leaf-like. Leaves
usually 6 or 8(-12)/node, essentially sessile; leaf blades
(4-)8-20(-25) mm long, 1-3 mm broad, narrowly ob-
lanceolate to narrowly oblong, apex obtuse (rounded),
usually with an apiculate tip ca. 0.5 mm long, base grad-
ually narrowed, with curved aculeolate retrorse hairs (ca.
0.2 mm long) along the margin and midvein beneath,
mostly glabrous above and on the flat surfaces beneath,
primary vein prominent, 2 weakly denned lateral veins
often present. Inflorescences usually terminal, paniculate
arrangements of small cymose groups of 3-7 flowers sub-
tended by reduced leaf-like bracts, with divaricate
branching and slender pedicels 1-3 mm long. Flowers
1.5-2 mm long, bisexual, hypanthium 0.5-0.7 mm long,
turbinate, densely hirsutulous with minute (0. 1 mm) as-
cending hairs, calyx reduced; corolla 1-1 .5 m long, white
to pink or red, campanulate or rotate, glabrous exter-
nally, puberulent within. Fruits dry, ca. 3 mm broad,
covered with ascending slightly curved hairs or with a
few uncinate hairs at the tip (in Costa Rica).

Plants of evergreen montane forest formations,
from ( 1 200-) 1 600 to 3 1 00 m elevation. Flowering
in April-July and December-January in southern
Central America. Rarely collected in Costa Rica
but apparently common in the Chiriqui High-

lands. The species ranges from the southwestern
United States to Panama.

Galium mexicanum is recognized by the slender
clambering stems with whorls of six or eight nar-
rowly oblong or oblanceolate leaves and the small
fruit with dense pubescence of curved hairs. The
aculeolate trichomes on stems and leaves help the
plants to climb and makes them adhesive and very
difficult to disentangle from shrubbery or clothing.

Galium orizabense Hemsley, Diagn. PI. Nov.
Mexic. 3: 54. 1878. Figure 3.

Erect or spreading herbs, 20-75 cm long, with several
to many stems from a small root stock, leafy stems 0.3-
1 mm thick, with thin white curves hairs 0.1-0.3 mm
long; stipules leaf-like. Leaves 4/node, subsessile or short-
petiolate, petioles to 2 mm long; leaf blades 6-18(-25)
mm long, 2-4(-5) mm broad, narrowly oblong to nar-
rowly elliptic-oblong or lanceolate, apex obtuse or short-
apiculate, drying thin-chartaceous or membranaceous,
with thin hairs ca. 0.3 mm long, on the upper surface,
margin and major veins beneath, with a prominent 1
vein and 2 straight lateral veins (usually readily appar-
ent). Inflorescences paniculate, with divaricate branches
subtended by slightly reduced leaves, often with 5-15
flowers on open lateral branches, pedicels 1-10 mm long.
Flowers 1-1.5 mm long, hypanthium ca. 0.5 mm long,
covered with minute hairs; corolla rotate, lobes ca. 0.4
mm long and equally broad, white or greenish yellow.
Fruits dry, 1-2 mm broad, covered with minute uncinate
hairs, borne on thin (to 0.5 mm) pedicels.

Plants of evergreen montane forest formations,
from 1 500 to 2500 m in southern Central Amer-
ica. Flowering in March, July-August, and De-
cember-January in southern Central America. The
species ranges from eastern and central Mexico to
Panama.

Galium orizabense is recognized by the uncinate
hairs on the fruit and the narrow verticillate leaves.
This species is very similar to G. uncinulatum and
the two may be conspecific. However, most spec-
imens can be differentiated by the key, and we
follow Dempster's treatment.

Galium uncinulatum DC., Prodr. 4: 600. 1830.
Figure 3.

Prostrate or procumbent herbs 1 5-90 cm long, leafy
stems 0.3-1 .5 mm thick, with thin whitish hairs 0.3-0.6
mm long, slightly scabrous; stipules leaf-like. Leaves 4/
node, sessile or subsessile with petioles to 1 (-2) mm long;
leaf blades (4-)6-12(-22) mm long, (2-)2.5-6(-13) mm
broad, ovate to narrowly ovate or narrowly oblong, apex
obtuse (rounded) with a short (0.5 mm) tip, base obtuse,

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

141

drying membranaceous or chartaceous, margin and sur-
faces with thin ascending or spreading hairs 0.2-0.4 mm
long, with 1 vein and 2 well-defined lateral veins, the
lateral veins 0.5-1 mm from the margin in larger leaves.
Inflorescences terminal or axillary, with opposite or cy-
mose branching, to 3 cm long, flowers usually few (3-5)
on the slender peduncles, often subtended by whorls of
reduced leaves, pedicels 1-10 mm long, 0.1-0.2 mm
thick when dried. Flowers 1-1.5 mm long, hypanthium/
ovary ca. 0.5 mm long, covered by minute hairs that will
expand in fruit, calyx reduced; corolla campanulate to
rotate, white to greenish or yellow, usually puberulent
externally. Fruits ca. 1.5 mm long and (l-)2-3 mm broad,
usually 2-lobed and rounded, dry and covered by pale
yellowish or whitish hooked (uncinate) hairs ca. 0.3-0.4
mm long.

Plants of evergreen montane forest formations
from 1000 to 2800(-3300) m elevation. Flowering
collections have been made in all months of the
year except May and October-November in Cen-
tral America. The species has been little collected
in Costa Rica. The species ranges from southern
Arizona and Texas (U.S.A.) through Mexico and
highland Central America to Panama.

Galium uncinulatum is recognized by having
four, often broad, little leaves at each node, small
few-flowered inflorescences with thin peduncles
and pedicels, and fruits densely covered with thin
uncinate hairs. This species is very similar to G.
orizabense(q.v.), which appears to live in the same
habitats but tends to have narrower leaves and
shorter puberulence. Specimens referred to as Ga-
lium obovatum H.B.K. by Standley, both in the
herbarium and in his flora (1938), are G. uncin-
ulatum.

Gardenia Linnaeus

Trees or shrubs, branches terete, glabrous or puber-
ulent; stipules interpetiolar and intrapetiolar, triangular,
apex acute to acuminate, often forming a short sheath
at the base. Leaves opposite or in whorls of 3, subsessile
to short-petiolate, coriaceous to chartaceous, entire, pin-
nately veined, domatia often present. Inflorescences ter-
minal or axillary, of 1 or 3 flowers (rarely more and
corymbose), sessile or short-pedicellate. Flowers radially
symmetrical, bisexual, usually large, hypanthium ovoid
to ellipsoid or obconic, calyx tube short (sometimes
spathe-like), calyx lobes 5-8 when present; corolla sal-
verform to campanulate or funnelform, white or yellow,
corolla tube glabrous or puberulent, corolla lobes 5-11,
convolute in bud, spreading or recurved; stamens 5-9,
inserted on the upper half of the tube, filaments short or
absent, anthers dorsifixed, linear to linear-oblong, in-
cluded or partly exserted, disc annular to crenate; ovary
1 -locular (rarely 2- or 6-locular at apex), ovules many
and horizontal on parietal placentas, style linear and

terete, stigma linear to clavate, 1 - or 2-lobed. Fruits ob-
long to ovoid, pyriform or globose, terete or costate, the
outer wall fleshy to leathery or woody, rupturing irregu-
larly or the endocarp breaking into 2-5 valves; seeds
very many, imbedded in a fleshy pulp, horizontal, an-
gulate, embryo small.

A genus of about 200 species in the tropics and
subtropics of the Old World. A few species are
important as ornamental trees in warm climates;
they are also grown under glass in cold climates
for their large aromatic flowers, which are often
used for corsages. The genus is similar to Genipa.
The genus has not become naturalized in Central
America, where one species is commonly seen in
gardens.

Gardenia augusta (L.) Merr., Interpr. Herb. Am-
boin. 485. 1917. Varneria augusta L., Amoen,
Acad. 4: 136. 1759. G. jasminoides Ellis, Phil.
Trans. 51, pt. 2: 935. 1761. G.florida L., Sp. PI.
ed. 2: 305. 1762.

Shrubs or many-branched small trees to 5 m tall, leafy
branchlets 1.5-5 mm thick, sparsely and minutely (0.2
mm) puberulent, glabrescent; stipules 5-10 mm long, at
first enclosing the apex and splitting down one side to
become spathe-like, with a short tube 2-4 mm long at
the base, persisting. Leaves 2(-3)/node, petioles 14 mm
long and little differentiated from the leaf base; leaf blades
3-12 cm long, 1.5-5 cm broad, elliptic-obovate to ellip-
tic-oblong or broadly elliptic (in smaller leaves), apex
acuminate, base acute and decurrent on petiole, glabrous
above and below (but sometimes with pit domatia and
a few hairs in the leaf axils), 2 veins 6-9 /side. Inflores-
cences often of 3 terminal flowers, or solitary flowers in
the axils of near-terminal leaves, pedicels ca. 1 mm
long. Flowers large (6-10 cm long), apparently differing
in size in different cultivars, sweetly aromatic, hypan-
thium ca. 10 mm long, calyx lobes 8-30 mm long, nar-
rowly oblong, spur-like; corolla white, tube 2-5 cm long,
corolla lobes usually 6 (some cultivars with 2 series), ca.
25 mm long and 18 mm broad, obovate; anthers ca. 18
mm long.

Plants of parks and gardens cultivated for their
large white sweetly aromatic flowers. These plants,
native to Asia, are called jazmin, jazmin delcabo,
and "gardenia."

Genipa Linnaeus

Trees, branchlets usually thick, terete, glabrous or pu-
berulent; stipules interpetiolar and intrapetiolar, connate
to form a short tube, caducous or deciduous with the
leaves. Leaves opposite and decussate, subsessile or pet-

142

FIELDIANA: BOTANY

iolate, often large, pinnately veined, chartaceous to co-
riaceous, without domatia. Inflorescences terminal or
axillary, flowers solitary or in few-flowered cymes or
subcapitate, pedicels present and continuous with the
hypanthium base. Flowers radially symmetrical, bisex-
ual (rarely unisexual and dioecious), 5- or 6-parted, hy-
panthium turbinate to campanulate, calyx tube truncat-
ed and entire or with 5-6 short lobes; corolla salverform
to funnelform, carnose, white to yellowish white, tube
short to long, glabrous or puberulent externally, barbate
in the throat within and at the base of the lobes, corolla
lobes 5-6, convolute in bud, spreading; stamens 5-6,
inserted in the upper part of the tube, anthers subsessile,
dorsifixed, linear, partly exserted; ovary 1-locular or be-
coming 2-locular, style thick, stigmas fusiform, placen-
tation parietal, ovules many and horizontal in vertical
files. Fruits baccate, large, ovoid to subglobose or ob-
ovoid, calyx tube persistent at apex of the fruit, pericarp
thick, fleshy to coriaceous; seeds many, large, com-

pressed (with two parallel flattened sides), the testa slight-
ly fibrous.

A genus of 5-10 species, ranging from southern
Florida (U.S.A.) and Mexico through Central
America into tropical South America. The genus
is distinguished by its few-flowered terminal and
subterminal inflorescences, large flowers with thick
corolla lobes, parietal placentation, and many large
horizontal seeds in vertical files within the large,
often solitary fruit. Cenipa vulcanicola Stand!, of
Mexico and Guatemala has been transferred to
Glossostipula concinna (Standl.) Lorence; it has
axile placentation.

Key to the Species of Genipa

la. Corolla densely sericeous distally, peduncles to 25 mm long; stipules acute at apex, persisting with
the leaves; leaves glabrous or pubescent, with 7-18 major secondary veins on each side; widespread
G. americana

Ib. Corolla glabrous on the outer surfaces, peduncles to 10 mm long; stipules broadly ellipsoid and
rounded at apex, usually caducous; leaves glabrous above and with appressed hairs on the veins
beneath, with 69 major secondary veins on each side; not recorded north of southern Costa Rica

. G. williamsii

Genipa americana L., Syst. Nat. ed. 10. 2: 931.
1759. G. oblongifolia Ruiz & Pav., Fl. Peruv.
Chil. 2: 67, pi. 220. 1798. G. caruto H.B.K.,
Nov. gen. sp. 3: 407 (quarto). 1820. G, ameri-
cana var. caruto (H.B.K.) Schum. in Mart., Fl.
Bras. 6(6): 352. 1889. G. codonocalyx Standl.,
Contr. U.S. Natl. Herb. 1 7: 446. 1 9 1 4. G. venosa
Standl., J. Wash. Acad. Sci. 18: 168. 1928. Fig-
ure 26.

Small to large trees 4-27 m tall, often with a spreading
hemispheric crown, trunk to 50 cm thick, bark smooth
and lenticellate, leafy branchlets 4-9 mm thick, densely
pubescent in early stages or glabrous; stipules 10-25 mm
long, triangular, the basal sheathing tube 1-3 mm long,
acute, deciduous with the leaves. Leaves with petioles
2-13 mm long, 2-3 mm thick, glabrous or pubescent;
leaf blades 1 2-42 cm long, (4-)6-19 cm broad, obovate
to elliptic-obovate or broadly oblanceolate, apex acu-
minate to obtuse or rounded, gradually narrowed to a
cuneate or slightly decurrent base, drying chartaceous
and often very dark above, glabrous and lustrous above,
glabrous to densely pubescent beneath with thin soft
hairs ca. 0.5 mm long, 2 veins 9-1 8/side. Inflorescences
terminal or subterminal, 4-10 cm long and with 1-9
flowers, cymose, peduncles to 25 mm long, glabrous,
pedicels 4-12 mm long. Flowers 2.5-4 cm long, appar-

ently bisexual but perhaps functionally unisexual, hy-
panthium difficult to distinguish from the calyx tube and
together 7-17 mm long, calyx tube to 10 mm diam.
distally, truncate or with broad short lobes, glabrous on
the exterior and puberulent within; corolla ca. 4 cm broad,
salverform, carnose, densely descending-sericeous ex-
ternally (except at the base of the tube), white or yellow-
ish white, darkening with age, tube 5-15 mm long, 4-7
mm diam. (to 10 mm at the lobes), lobes 5-6, 11-28
mm long, 5-12 mm broad, obovate and rounded at apex;
anthers 6-14 mm long, becoming recurved between the
lobes; stigmas ca. 5 mm long. Fruits 4-1 1 cm long, 3-
1 1 cm diam., obovoid to subglobose, smooth and grayish
brown or yellowish brown, the persisting calyx 3-6 mm
long, crateriform on apex of the fruit and 8-10 mm
diam., pedicels up to 5 cm long in fruit; seeds 6-12 mm
long, 4-7 mm broad, ca. 2.3 mm thick.

Trees of both wet evergreen rain forests and
seasonally very dry deciduous forest formations
in the Caribbean and Pacific lowlands, from near
sea level to 900 m elevation. Flowering occurs
primarily in March-August; fruiting throughout
the year. This species ranges from southern Flor-
ida and the West Indies, through Mexico and Cen-
tral America through tropical South America to
Paraguay.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

143

Genipa atnericana is recognized by the larger
leaves with short petioles clustered at the ends of
stems, usually solitary large fruit with many hor-
izontal seeds in vertical files, and large sericeous
flowers with short corolla tubes and large lobes.
This is a common and distinctive tree, especially
conspicuous in deciduous forest formations in the
dry season because of its large terminal fruit (but
compare Alibertia edulis). This species may be
confused with species of Borojoa, but those tend
to have sessile terminal flowers and stipules with
parallel venation. Guaitil, caruto, jagua, andjagua
negro are common names for this species. The
juice of the young pulpy fruit turns black or dark
blue and is used by Native Americans as a dye or
body paint. The species is sometimes cultivated,
and the fruit is eaten. The wood is easy to work
but strong and resistant; it is used for making fur-
niture and carts and in building construction.

Genipa americana is here interpreted to be a
very variable species, following Dwyer (1980) and
Steyermark (1974). The types of Standley's G. co-
donocalyx(Pittier 12085 us) and G. venosa (Stand-
ley & Valeria 45269 us) appear to represent no
more than unusual forms of G. americana and
were described when the full pattern of variation
in G. ameriana was not apparent. The type of G.
venosa has prominent petioles (3.54 cm long) and
unusually long fruit, but it seems better to treat it
as a variant of G. americana rather than as a dis-
tinct species. Collections with the leaves densely
pilose beneath have been referred to variety caruto
(H.B.K.) K. Schum.

Genipa williamsii Standl., J. Wash. Acad. Sci. 8:
642. 1918. Figure 26.

Small to medium-sized trees, 4-20 m tall, with boles
ca. 25 cm dbh, leafy stems 3-7 mm thick, glabrescent,
becoming pale brown; stipules 10-32 mm long, 6-18
mm broad, ovate from a narrowed base, flattened, ap-
pressed-sericeous. Leaves with petioles 7-22(-60) mm
long, 1 .5-2.7(-4) mm thick, glabrous, often drying black-
ish and lustrous; leaf blades 8-17(-33)cm long, 5-10(-16)
cm broad, obovate-oblong, to broadly elliptic or elliptic-
oblong, apex rounded and lacking a narrowed tip, base
obtuse to acute and slightly decurrent on petiole, drying
stiffly chartaceous to subcoriaceous and dark reddish
brown above, glabrous above, appressed-sericeous on
the major veins beneath with hairs ca. 0.3 mm long, 2
veins 7-9/side and weakly loop-connected distally, 3
venation obscure. Inflorescences of ca. 3 (4-7) terminal
flowers subtended by 2 ovate-lanceolate bracts (stipules)
ca. 1 2 mm long with glabrous surfaces but ciliolate along
the edge, peduncles 5-10 mm long, pedicels 8-10 mm
long and continuous with the hypanthium, drying black,

bracteoles 3 mm long or reduced to ridges. Flowers gla-
brous externally, drying black, hypanthium 3-4 mm long,
4-5 mm broad at apex, obconic, calyx tube 0.5-1 mm
high, entire or slightly undulate; corolla salverform, white
and carnose, corolla tube 1.8-3 cm long, 4-5 mm diam.,
lobes 5, 18 mm long, 8-12 mm broad distally, obovate-
oblong and rounded distally; anthers sessile, stigmas 4,
to 4 mm long, unequal. Fruits subglobose, ca. 7 cm
diam., drying black; seeds ca. 10 mm long, 5-6 mm
broad, imbedded in white pulp.

This species has been collected only in south-
ernmost Limon Province at 450-650 m elevation
(Hammel et al. 1 7597 CR, MO, Herrera 3208 CR.MO)
in Costa Rica. Flowering in July; fruiting in April-
June and October in Panama. The species ranges
to Colombia.

Genipa williamsii is recognized by the leaves
rounded distally, clavate flower buds, and both the
fleshy flowers and the large fruits that dry black.
It is similar to species of Ladenbergia, but those
have domatia and valvate corolla lobes.

Geophila D. Don

REFERENCE L. O. Williams, Geophila (Rubi-
aceae) in North America. Phytologia 26: 263-264.
1973.

Creeping perennial herbs, stems slender and puberu-
lent or glabrous, rooting at the nodes; stipules interpetio-
lar, small, rounded-ovate to triangular, entire to shal-
lowly bilobed, persisting. Leaves usually with long
petioles; leaf blades rounded and often cordate at the
base, membranaceous to thin-chartaceous, venation pin-
nate, domatia absent. Inflorescences terminal or pseu-
doaxillary, few-flowered heads or cymes, peduncles short
or long, flowers subtended by an involucre of small bracts,
pedicels short or absent. Flowers radially symmetrical
and bisexual, usually 5-parted (less often 4-7-parted),
calyx tube with 4-7 narrow lobes, persistent; corolla fun-
nelform to salverform, white, corolla tube narrow, pilose
in the throat, corolla lobes 4-7, valvate in bud, spreading
or recurved; stamens 4-7, filaments filiform and inserted
in the floral tube, anthers dorsifixed, linear, half exserted;
ovary 2-locular, ovules solitary in each locule and basal,
style slender with 2 stigmas. Fruits a juicy berry, usually
containing 2 1 -seeded pyrenes (nutlets); pyrenes plano-
convex and smooth or costate on the dorsal surface, with
a ventral sulcus.

A genus of 20-30 species native to the American
tropics, Africa, and Asia. The slender creeping
stems, long-petiolate leaves with rounded blades,
few-flowered inflorescences, and fleshy, two-seed-
ed fruits characterize this genus. These plants may
be mistaken for species ofCoccocypselum, but that
genus has many-seeded fruit.

144

FIELDIANA: BOTANY

Key to the Species of Geophila

la. Fruit black at maturity, pyrenes weakly costate; peduncles 2-10 cm long; basal lobes of the leaf

blades separated by a sinus (cordate with non-overlapping lobes) G. macropoda

Ib. Fruit red at maturity (if fruit are blue go to the genus Coccocypselum), pyrenes strongly costate;

peduncles 0.2-10 cm long; basal lobes of the leaf blades separate to overlapping 2

2a. Ovary, fruit and leaves conspicuously pilose with thin hairs 0.5-2 mm long [leaf blades subcordate

with a small basal sinus; peduncles to 7 cm long in fruit] G. cordifolia

2b. Ovary and fruit glabrous, leaves glabrous or puberulent with short (0.1-0.3 mm) hairs; peduncles

to ca. 2 cm long 3a

3a. Leaf blades ovate and slightly longer than broad, often narrowed at apex, cordate but usually without

a visible basal sinus (because the lobes overlap slightly) G. repens

3b. Leaf blades ovate-triangular and distinctly longer than broad, usually acute at apex, cordate to

subcordate at the base and with a small sinus G. gracilis

Geophila cordifolia Miq., Stirp. Surin. Sel. 176.
1 850. Mapouria trichogyne Muell.-Arg. in Mart.,
Fl. Bras. 6(5): 426. 1881. Geophila trichogyne
(Muell.-Arg.) Standl., Publ. Field Columb. Mus.,
Bot. Ser. 7: 423. 1931. Figure 2.

Creeping herbs, leafy stems 0.7-1 .5 mm thick, densely
pubescent with slender pale straight or crooked hairs 0. 5-
2 mm long; stipules 2-4(-6) mm long, 1.5-4 mm broad
(broadest beneath the inflorescences), usually glabrous,
persisting. Leaves with petioles 3-7(-l 3) cm long (short-
er on leaves subtending the inflorescences), conspicu-
ously pubescent with slender crooked or straight mul-
licellular hairs 0.7-1.5 mm long; leaf blades 3-7(-ll)
cm long, 2-6(-8.5) cm broad, broadly ovate to oblong-
ovate or narrowly ovate (triangular-ovate), apex obtuse
to short-acuminate, base cordate with rounded lobes 8-
35 mm broad, basal sinus 3-15 mm deep, drying thin-
chartaceous and brownish, both surfaces covered with
thin usually crooked hairs 0.7-2 mm long, 2 veins 3-
5/side, not usually loop-connected near the margin. In-
florescences terminal, capitate, 1-3 cm long (to 6 cm in
fruit), with 5-1 7 flowers, peduncles 5-1 5 mm long, elon-
gating in fruit, densely pubescent, bracts 3-10 mm long,
lanceolate and pubescent, pedicels 0.5-3 mm long. Flow-
ers with an urceolate hypanthium, calyx tube 0.5 mm
long and 1 .5 mm diam.. lobes ca. 3 mm long, with slen-
der hairs ca. 1 mm long; corolla 5-6 mm long, white
often tinged with pink distally, tube 2-4.5 mm long,
glabrous externally, with a short collar of hairs at the
point of filament attachment within, lobes 5, 1 .5-2.5 mm
long; stamens with anthers 0.8-1 mm long, included.
Fruits ca. 8 mm long, ovoid-globose, orange to red, with
scattered slender hairs, pyrenes ca. 4 mm long and 3 mm
broad, with 3-5 prominent longitudinal dorsal ribs.

Plants of evergreen forest formations, from near
sea level to 500(-1100) m elevation. Probably
flowering and fruiting primarily in the wet season
(May-December). The species ranges from Belize
along the Atlantic slope of Central America to
Colombia, Venezuela, and the Amazon basin in
Brazil and Peru.

Geophila cordifolia is recognized by the long,
often crooked, thin multicellular hairs that cover
almost all parts of the plants. The long-petiolate
leaves with cordate bases, short inflorescences
elongating in fruit, and glabrous stipules are ad-
dition distinctions. In Costa Rica the species, is
known only from the La Selva area and from Vol-
can Rincon de la Vieja.

Geophila gracilis (Ruiz & Pav.) DC, Prodr. 4: 537.
1 830. Psychotria gracilis Ruiz & Pav., Fl. Peruv.
2: 63, pi. 211, f. C. 1799. G. croatii Steyerm.,
Phytologia35:401. 1977.

Creeping herbs to 20 cm tall or slender-stemmed vines
to 1.5 m long, leafy stems 0.3-1.3 mm thick, glabrous,
horizontal internodes 3-7 cm long, with adventious roots
near the nodes; stipules 2-4 mm long. 1-2 mm broad,
usually rounded at apex, curving outward in age and
persisting. Leaves with petioles 2-8 cm long (but shorter
below the inflorescences), 0.3-1 mm thick, glabrous
abaxially and with 2 rows of short (0.2-0.5 mm) stiff
retrorse or erect hairs along either side of the adaxial
sulcus; leaf blades 1.8-5 cm long, 1-3.5 cm broad, tri-
angular-ovate to ovate, apex gradually narrowed and
acute (or obtuse), base cordate to subcordate, sinus 3-8
mm deep, the basal lobes usually separate but occasion-
ally overlapping and the sinus obscured, drying thin-
chartaceous, glabrous above or with few thin hairs 0.3-
0.8 mm long, usually glabrous beneath, 2 veins 3-47
side, usually loop-connected near the margin. Inflores-
cences terminal, 12-15 mm long, capitate with ca. 3-7
flowers, peduncles 2-8 mm long (apparently longer when
the leaves of the subtending node are reduced and bract-
like, bracts 4-7 mm long, 1-1.5 mm broad, united at
the base and persisting, pedicels 0-1 mm long. Flowers
with hypanthium ca. 2 mm long, calyx lobes ca. 3 mm
long and 0.5 mm broad, narrowly oblong, persistent and
enlarging in fruit; corolla white. Fruits red at maturity,
3-5 mm long, subglobose. pyrenes ca. 3.5 mm long, with
3 prominent longitudinal dorsal ridges.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

145

Plants of evergreen forest formations, from near
sea level to ca. 500 m elevation. Flowering in May-
December (primarily in June and July in central
Panama; Croat, 1978). The species is known from
southeastern Nicaragua, the Canal area of Pana-
ma, and the upper Amazon basin of Brazil, Peru,
and Bolivia.

Geophila gracilis is recognized by its slender
stems rooting at most nodes, triangular-ovate leaf
blades with little or no puberulence, and short
terminal capitate inflorescences subtended by
bracts fused at the base. Variation in specimens
from South America clearly encompass the dis-
tinctions used to separate G. croatii. This species
is apparently common on Barro Colorado Island,
Panama, and has been collected in Nicaragua, but
it has yet to be collected in Costa Rica.

Geophila macropoda (Ruiz & Pav.) DC., Prodr. 4:
537. 1830. Psychotria macropoda Ruiz & Pav.,
Fl. Peruv. 2: 63, pi. 21 1, f.6. 1799. Figure 2.

Creeping herbs, leafy stems 0.8-2 mm thick, glabrous
or very minutely (0.1 mm) puberulent, often with 2
prominent longitudinal ridges; stipules 2-6 mm long,
ovate-oblong, glabrous, deciduous or obscured by the
adventitous roots. Leaves with petioles (2-) 3-1 4 cm long
(shorter below the inflorescences), 0.7-1.2 mm thick,
glabrous abaxially but with 2 adaxial ridges with short
(0.1-0.4 mm) dense hairs; leaf blades 3-9 cm long, 2.5-
8 cm broad, broadly ovate to ovate-orbicular, apex
rounded to broadly obtuse, base cordate with lobes 1-4
cm broad, basal sinus 2-15 mm deep, drying membra-
naceous or thin-chartaceous and often grayish green, gla-
brous above, glabrous beneath except for the minute
puberulence on the major veins near the base, 2 veins
3-5/side and weakly loop-connected near the margin.
Inflorescences usually axillary, 2-4 cm long and elon-
gating in fruit, capitate with 3-7 flowers, peduncle 1.5-
5(-7) cm long, minutely puberulent with whitish hairs
ca. 0. 1 mm long, bracts 3-6 mm long, united at the base,
pedicels to 2 mm long. Flowers with hypanthium ca. 1.5
mm long, essentially glabrous, calyx lobes 2-3 mm long;
corolla 3-7 mm long, tube 3-4 mm long, corolla lobes
5, 2-3 mm long. Fruits black or blue, sessile, 5-10 mm
long, 3-7 mm diam.. ellipsoid to ovoid; pyrenes 4-7 mm
long, 2-3 mm broad, without raised longitudinal ribs
(costae) on the convex surface.

Plants of the lowland Caribbean rain forest for-
mations, from near sea level to 600 m. Flowering
in April-November. The species ranges from
southern Mexico through Central America to Bo-
livia and Paraguay.

Geophila macropoda is recognized by the axil-
lary and long-pedunculate inflorescences, closely
clustered flowers, black fruit, and pyrenes without
prominent longitudinal costae.

Geophila repens (L.) I. M. Johnston, Sargentia 8:
281. 1949. Rondeletia repens L., Syst. ed. 10:
928. 1759. Psychotria herbaceajacq., Enum. PI.
Carib. 16. 1760. Geophila herbacea (Jacq.)
Schumann in Engl. & Prantl., Nat. Pflanzenfam.
4,4: 119. 1891. Figure 2.

Creeping herbs to ca. 10 cm high, leafy stems 0.5-1
mm thick, glabrous or very sparsely and minutely pu-
berulent; stipules 0.5-2 mm long, 1-2 mm broad, broad-
ly ovate, glabrous, persisting or deciduous. Leaves with
petioles l-6(-8.5) cm long (sometimes shorter in leaves
subtending the inflorescences), 0.4-1.3 mm thick, gla-
brous on the abaxial surface but with short (0.2-0.5 mm)
retrorse or crooked hairs along the adaxial (upper) side;
leaf blades 1.2-5.5 cm long, 1-5 cm broad, broadly ovate
to ovate-suborbicular, apex bluntly obtuse or rounded-
obtuse, base cordate with lobes 5-25 mm broad, basal
sinus 1-7 mm deep and usually obscured by the over-
lapping basal lobes, drying membranaceous, upper sur-
face glabrous or with a few short hairs near the margins,
glabrous beneath or with a few short (0. 1-0.3 mm) hairs
on the veins near the petiole, 2 veins 3-5/side, cystoliths
visible (as short whitish lines) or obscure on the lower
surface. Inflorescences solitary and terminal on short
leafy shoots (apparently 2-3 and axillary when directly
subtended by small leaves), with (l-)2-5 flowers, pe-
duncles 5-35 mm long, puberulent with short retrorse
hairs, subtending bracts 3-6 mm long, united at the base
and lanceolate distally, pedicels 0-2 mm long. Flowers
white or becoming pink in age, hypanthium ca. 1 mm
long, calyx tube 1-1.5 mm long, glabrous, calyx lobes
1-3 mm long, 0.3-0.7 mm broad, glabrous; corolla 8-
14 mm long, glabrous or puberulent, funnelform, tube
6-9 mm long and 1-1.5 mm diam., lobes 5, 3-5 mm
long, 1.5-2.5 mm broad, ovate and obtuse to acute; sta-
mens with filaments ca. 0.5 mm long, attached near the
middle of the tube, anthers ca. 2 mm long; style 5-7 mm
long. Fruits 8-10 mm long, ovoid or globose, bright red
at maturity, subsessile or short (1-2.5 mm) pedicellate;
pyrenes 3.5-5 mm long, 2.5-3 mm broad, with 3 slightly
raised longitudinal ribs (costae) on the convex face.

Plants of the shaded forest floor in evergreen or
partly deciduous forest formations of both the Ca-
ribbean and Pacific lowlands of Costa Rica, from
near sea level to 800 m elevation. Flowering in
June-October; fruiting in July-November. The
species ranges from Mexico and the West Indies
to Peru and Bolivia in the New World; it is also
found in West Africa, the Philippines, and the
western Pacific.

Geophila repens is recognized by its creeping
habit and short stature, small cordate leaves with
the sinus obscured, petioles with puberulence along
one side, bright red fruit, and pyrenes with weakly
developed costae. This is our most commonly en-
countered species of Geophila; it has been col-
lected at La Selva and from near Canas in Guana-
caste Province to the Golfo Dulce area along the

146

FIELDIANA: BOTANY

Pacific. Lechuga is a name used for this species in
the Golfo Dulce region. Breeding biology was stud-
ied by Bawa and Beach (1983).

Gonzalagunia Ruiz Lopez & Pavon

Shrubs or small trees, distal branches often curved,
pendant or scandent, leafy stems usually slender and
pubescent, terete, nodes usually thickened; stipules in-
terpetiolar, usually broad at the base and triangular with
a narrow distal awn (rarely intrapetiolar and tubular).
Leaves distichous, petiolate or subsessile; leaf blades of-
ten thin-chartaceous, usually narrow and with ascending
secondary veins, domatia absent or obscure. Inflores-
cences solitary, usually terminal and narrowly long-ra-
cemiform, spiciform or thyrsiform, the flowers solitary,
cymose or fasciculate on short lateral branches of the
central axis, bracteoles present, flowers sessile or pedi-
cellate. Flowers radially symmetrical, bisexual, mono-
morphic or distylous, small, 4- (less often 5-) parted,
hypanthium urceolate to rounded, calyx tube very short,
calyx lobes 4 or 5, small, equal or unequal, persisting in
fruit; corolla salverform to funnelform, white or pink,
corolla tube narrow, villose in the throat, corolla lobes

4(-5), imbricate or valvate; stamens 4(-5), filaments short
or absent, anthers dorsifixed, 2-lobed at the base, ex-
serted or partly exserted; ovary 2-or 4-locular, ovules
numerous, placentation peltate on the septum, style with
2 or 4 stigmatic lobes. Fruits baccate, fleshy or spongy,
subglobose, 2- or 4-locular and usually with 2 or 4 lobes
or sulci, with 2 or 4 hard cocci; seeds 4-many within
the cocci, minute.

A genus of 25-35 species, ranging from Mexico,
Central America, and the West Indies through
tropical South America. The long narrow spike-
like distal inflorescences, slender drooping distal
stems, small narrow-tubed flowers and baccate
fruits with 2-4 pyrenes make this a very distinctive
genus. Some species of Rondeletia with long-nar-
row inflorescences may be confused with species
of Gonzalagunia in the absence of fruit; Rondeletia
has capsular fruit. Several of our species are weedy
shrubs of open secondary growth and closely re-
lated; they can be difficult to distinguish in the
absence of mature flowers or fruit. A few species
resemble species of Buddleia (Loganiaceae).

Key to the Species of Gonzalagunia

la. Stipules united or overlapping above the petioles to form a short tubular sheath or broad tube-like

base 4-12 mm long 2

Ib. Stipules not forming a tube above the node, or the broad margins not overlapping at the base, stipule

only 1-3 mm long before being narrowed into the awn-like apex 3

2a. Stipular sheath to 1 cm long; flower clusters and lateral branches of the inflorescences subtended

by caducous bracts 4-12 mm long and 1-2 mm broad, corolla 34 mm long; leaf blades with

5-8 pairs of major secondary veins; small treelets of wet forest understory .... G. bracteosa
2b. Stipular sheath 0-5 mm long or the stipule margins slightly overlapping; bracts less than 2 mm

long, corolla 4-6 mm long; with 10-14 pairs of secondary veins; trees to 18 m tall

Rondeletia brenesii

3a. Petioles usually less than 4 mm long; corolla tube usually less than 7 mm long 4

3b. Petioles usually more than 4 mm long: corolla tube usually more than 7 mm long 7

4a. Leaf blades with 4-7 pairs of major secondary veins, laminae thin-textured; flowers thin-textured

and often solitary, corolla glabrous externally G. rudis

4b. Leaf blades with 7-1 3 pairs of major secondary veins; thinly to stiff-chartaceous; flowers thick-
textured, corolla densely sericeous externally 5

5a. Leaf blades subsessile, narrowly lanceolate, 9-26 cm long, with 9-15 pairs of secondary veins

arising at angles of about 3040 [only known from the Golfo Dulce area] G. brenesii

5b. Leaf blades subsessile or short petiolate, ovate to elliptic and rarely lanceolate, to 1 6(-30?) cm

long, with 4-12 pairs of secondary veins arising at angles of 40-60 5

6a. Corolla lobes ca. 1.5 mm long; leaf blades with 4-1 1 pairs of major secondary veins, ovate to

ovate-elliptic, thinly chartaceous; Caribbean and Pacific slope, 0-1200 m elevation

G. ovatifolia

6b. Corolla lobes 3-4 mm long; leaf blades with 7-12 pairs of major secondary veins, ovate-elliptic

to lanceolate, stiffly chartaceous; wet Caribbean slope 300-1200 m elevation

G. stenostachya

7a. Mature fruit becoming blue-black; corollas white or white tinged with pink, corolla tubes 8-13 mm
long; inflorescences with the flower clusters sessile; 0-1200(-1500) m elevation . . G. panamensis

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

147

7b. Mature fruit white; corollas reddish to pink, corolla tubes 6-10 mm long; inflorescences with the
flower clusters on short (1-3 mm) peduncles (but note that some inflorescences may only have
solitary flowers and no apparent secondary peduncles); 900-2200 m elevation G. rosea

Gonzalagunia bracteosa (J. D. Smith) B. L. Rob-
inson, Proc. Amer. Acad. Arts 45: 405. 1910.
Gonzalea bracteosa J. D. Smith, Hot. Gaz. 33:
252. 1902. Figure 21.

rescences appear to be restricted to northeastern
Costa Rica. This species differs greatly from our
other members of the genus; it resembles Psy-
chotria pilosa.

Shrubs or small treelets, 1.5-4 m tall, leafy branchlets
2-4(-6) mm thick, terete, with many ascending strigose
hairs 1-1.5 mm long; stipules 8-18(-24) mm long, 3-6
mm broad, with a tubular sheath to 12 mm long, dark
brown and pubescent along the midvein, acute to acu-
minate and with a slender tip 1-7 mm long. Leaves with
petioles 2-10 mm long, 1-2 mm thick, broad, pubescent;
leaf blades 6-19(-22) cm long, 2.5-7.5(-8.5) cm broad,
narrowly elliptic-obovate to obovate-oblong, oblong or
elliptic, usually broadest above the middle, apex acu-
minate (acute), tip to 1 5 mm long, gradually narrowed
to the acute or obtuse base, leaves usually drying char-
taceous and dark brown above (rarely subcoriaceous),
sparsely pubescent above with thin appressed hairs 0.5-
1 mm long (densely strigulose on the midvein), more
densely appressed-pubescent beneath with brownish hairs
0.5-1.3 mm long, 2 veins 5-7 /side. Inflorescences sol-
itary, axillary or terminal, 6-25 cm long, spike-like or
thyrsiform panicles with small (5-15 mm) alternate or
opposite flower clusters 2-14 mm distant along the rachis
(rarely with lateral branches to 4 cm long), primary pe-
duncles 2-8 cm long, with ascending hairs to 2 mm long,
several bracts 5-12 mm long and 1-2 mm broad sub-
tending the flower clusters, flowers sessile. Flowers with
hypanthium ca. 1 mm long, densely hirtellous, calyx tube
minute, calyx lobes ca. 0.5 mm long, difficult to see
among the hairs, glabrous on the inner surface; corolla
funnelform, greenish white to white, with few straight
hairs 0.5 mm long externally, tube 2-3 mm long, corolla
lobes 5, 1-2 mm long; stamens 5, anthers ca. 1 mm long;
style ca. 2.5 mm long, stigma 0.4 mm long. Fruits 3-5
mm long, 3-5 mm broad, depressed globose, becoming
blue or blue-black, pubescent, sessile; pyrenes 2-4.

Plants of lowland rain forest formations on the
Caribbean and Pacific slopes in Costa Rica, from
10 to 850 m elevation. Flowering in January-Sep-
tember; fruiting throughout the year. The species
ranges from northeastern Costa Rica to Colombia.

Gonzalagunia bracteosa is characterized by the
hirsutulous pubescence on many parts, obovate-
oblanceolate leaves drying dark above, narrow in-
florescences with short flower clusters subtended
by conspicuous bracts, small sessile flowers, and
tubular stipules. Two collections are noteworthy
because the inflorescences have lateral branches
2-4 cm long and with many bracts along their
length: Folsom 9778 (DUKE, F) and Zamora & San-
chez 469 (CR, F). Specimens exhibiting such un-
usual variation in bract development and inflo-

Gonzalagunia brenesii Standl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 1302. 1938. Figure 20.

Shrubs, 1.5-3 m tall, leaf branchlets 1.5-6(-8) mm
thick, rounded-quadrangular in cross-section, sparsely
pubescent with appressed hairs ca. 0.5 mm long or gla-
brous, drying reddish brown; stipules 4-8 mm long, 2.5-
7 mm broad at the base, with a very short (1-2 mm)
broadly triangular base and a slender awn 4-6 mm long.
Leaves subsessile, petioles 0-3(-5) mm long; leaf blades
9-26 cm long, 2.7-6 cm broad, narrowly lanceolate to
narrowly elliptic-oblong or elliptic-oblanceolate, apex ta-
pering gradually and long-acuminate, base obtuse, drying
chartaceous and dark brown above, lustrous above in
life, glabrous or sparsely pubescent above, with thin
whitish ascending hairs beneath, the hairs longer (ca. 0.4
mm) on the major veins beneath, 2 veins 9-1 3/side and
weakly loop-connected distally, 3 veins subparallel. In-
florescences solitary and terminal, 20-50 cm long, spi-
ciform with short (1-5 mm) lateral branches bearing 2-
6 flowers, peduncles 0-6 cm long, 1.5-3 mm thick, with
ascending appressed hairs, bracts 1-3 mm long, linear,
pedicels 1-2 mm long. Flowers 6-12 mm long, hypan-
thium 0.5-1 mm long, sericeous at the base, calyx tube
ca. 0.5 mm long, calyx lobes 0.2-0.3 mm long, glabrous;
corolla white or pink, tube 7-10 mm long and 0.7-1 mm
diam., pubescent, lobes 3 mm long and 1.8 mm broad,
obtuse; ovary 4-locular. Fruits 3-5 mm long, 3-6 mm
broad, usually 4-lobed, becoming white, minutely and
sparsely puberulent.

Plants of rain forest formations in Costa Rica's
southern Pacific lowlands, from near sea level to
500 m elevation. Flowering in April-December;
fruiting in January and August-September. This
species is endemic to Costa Rica, ranging from the
forests of the Pacific slope above Quepos eastward
to the Osa Peninsula.

Gonzalagunia brenesii is distinguished by its re-
stricted range, long narrow subsessile leaves, long
spicate inflorescences, and short narrowly tubular
white or pink flowers. This species appears to be
related to the G. panamensis-G. rosea complex.

Gonzalagunia ovatifolia (J. D. Smith) B. L. Rob-
inson, Proc. Amer. Acad. Arts 45: 405. 1910.

148

FIELDIANA: BOTANY

Gonzalea ovatifolium J. D. Smith, Bot. Gaz. 27:
336. 1899. Figure 20.

Shrubs to 3(-4) m tall, leafy branchlets 1.3-4 mm
thick, with thin ascending brownish hairs ca. 0.5 mm
long, terete, glabrescent; stipules 6-10 mm long, subulate
with a short (1-2 mm) base and long (4-7 mm) awn-like
apex. Leaves subsessile with petioles l-4(-5) mm long,
ca. 1.3 mm thick, pubescent; leaf blades 5-12(-16) cm
long, 2-6(-7) cm broad, ovate to ovate-elliptic, apex
short-or long-acuminate, tip 3-20 mm long, abruptly
narrowed to rounded at the obtuse base, unequal at the
base, the leaves drying thin-chartaceous and dark above,
pale grayish to pale greenish beneath, essentially glabrous
above, appressed-pubescent on the veins beneath with
thin short (ca. 0.5 mm) hairs, 2 veins 6-1 I/side. Inflo-
rescence solitary and terminal, 12-45 cm long, narrowly
spike-like with flowers in distant (3-10 mm) clusters,
rachis slender 0.5-1 mm thick, with thin whitish as-
cending hairs ca. 0.5 mm long, flowers in sessile or sub-
sessile groups of 1-3, subtended by linear bracts 2-4 mm
long, flowers usually solitary in the distal half of the
inflorescence, pedicels to 1 mm long. Flowers with hy-
panthium ca. 1 mm long, densely sericeous, calyx lobes
1-2 mm long, glabrous; corolla white, tube 4-5 mm long,
lobes 1-2 mm long; anthers ca. 1 mm long. Fruits 2-3
mm long and 3-4 mm broad (dried), white, usually
4-lobed, pubescent with thin hairs ca. 0.3 mm long.

Shrubs of wet evergreen forest formations of
both the Caribbean and Pacific slopes in Costa
Rica, from near sea level to 1200 m elevation.
Flowering in January-April and July-August;
fruiting in the same months and in October and
December. The species ranges from Nicaragua to
Colombia.

Gonzalagunia ovatifolia is recognized by its sub-
sessile ovate acuminate leaves, the long inflores-
cences with few-flowered cymules or solitary flow-
ers along its length, and the small corollas. This
species may be difficult to separate from some
specimens of G. rosea (q.v.).

Gonzalagunia panamensis (Cav.) K. Schum. In
Mart., Fl. Bras. 6(6): 292. 1889. Buena pana-
mensis Cav., Anales Hist. Nat. 2: 279. 1800.
Gonzalea panamensis (Cav .) Spreng., Syst. Veg.
1:417. 1825. Figure 20.

Shrubs to 3(-5) m tall, erect or scandent, leafy branch-
lets 0.8-4 mm thick, terete or slightly quadrangular,
sparsely to densely sericeous with thin whitish ascending
hairs ca. 0.5 mm long; stipules 4-7 mm long, the broad
base 0.5-2 mm long, subulate with a narrow awn-like
tip, pubescent on the midrib and edges. Leaves with
petioles 6-27 mm long (shorter on young axillary shoots),
ca. 1 mm thick, pubescent, with lateral margins contin-
uous with the decurrent leaf margin; leaf blades 5-15
cm long, 1-6 cm broad, narrowly lanceolate to lanceo-

late-elliptic or ovate-lanceolate, apex tapering gradually
and acute or acuminate, base acute and often decurrent
on petiole, drying thin-chartaceous, densely pubescent
on the veins and more sparsely between the veins with
appressed hairs ca. 0.3 mm long above and below, 2
veins 5-7/side, ascending. Inflorescences solitary and
terminal (or axillary by later lateral growth of side shoots),
6-22(-40) cm long, peduncles 1-5 cm long, 1-2 mm
thick, pubescent, the flower clusters ca. 5 mm diam.,
essentially sessile (but solitary flowers pedicellate), bracts
to 4 mm long and linear, pedicels 0-2 mm long. Flowers
4-parted, hypanthium 0.7-1 mm long, sparsely pubes-
cent, calyx ca. 1 mm long, calyx lobes 0.5-1 mm long
and 0.5 mm broad at the base; corolla white or white
tinged with pink, tube (8-)10-13 mm long, 0.7-1 mm
diam., sparsely pubescent, lobes 2-3 mm long, ca. 2 mm
broad at the base, puberulent within; stamens 4, anthers
2-2.2 mm long; style 7-10 mm long, stigma ca. 0.7 mm
long. Fruits 2.5-4 mm long, 3-8 mm broad, depressed
globose to 4-lobed, glabrous or sparsely puberulent, red
becoming purple black or black.

Shrubs of evergreen or partly deciduous forest
formations of the Pacific slope in Costa Rica, from
near sea level to 1 200 m elevation. Flowering in
all months except March-April and November;
fruiting in January, February, September-Octo-
ber, and December. The species ranges from Mex-
ico to Colombia and in the West Indies.

Gonzalagunia panamensis is recognized by the
usually lanceolate leaves on slender well-devel-
oped petioles, the flower clusters sessile on the
inflorescence rachis (or the flowers solitary and
pedicellate), the long narrow white corolla tube,
and the fruit turning red or black. Collections from
higher elevations have broader leaves and may
represent introgression from another species; com-
pare G. ovatifolia and G. rosea.

Gonzalagunia rosea Standl., Ann. Missouri Bot.
Gard. 25: 836. 1938. G. longithyrsa Fosberg,
Sida 2: 387. 1966. Figure 20.

Shrubs or small trees, 2-4(-6) m tall, branches erect
or scandent, leafy branchlets 1-4 mm thick, terete, with
stiff appressed-ascending pale yellowish or grayish hairs
0.2-0.5 mm long; stipules 3-7 mm long, ca. 3-4 mm
wide at base, broadly triangular with a short (1-4 mm)
narrow tip, pubescent on the edge and midrib. Leaves
with petioles 4-20 mm long, ca. 1 mm thick, densely
pubescent; leaf blades 7-18(-22) cm long, 2-6(-8) cm
broad, narrowly ovate-elliptic to lanceolate, lanceolate-
oblong or narrowly elliptic-oblong, apex gradually or
abruptly acuminate (acute) with tip 5-1 5 mm long, base
acute to obtuse, drying thin-chartaceous to chartaceous,
dark grayish brown above, puberulent on the upper sur-
face (denser on the major veins) with short (0.2-0.3 mm)
straight hairs, more densely puberulent beneath with hairs
to 0.5 mm beneath, 2 veins 5-9(-l l)/side, strongly as-
cending (and not loop-connected near the margin). In-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

149

florescences solitary, terminal (axillary by further growth
of lateral branches), 12-35 cm long, flowering part ca.
2.5 cm broad, peduncles 5-35 mm long, 1-2.5 mm thick,
densely short hirsute, flowers in groups of l-3(-5) and
borne on secondary peduncles 1-5 mm long, alternate
or opposite on the rachis and 2-10 mm distant, bracts
1-3 mm long and linear, pedicels l-3(-4) mm long.
Flowers with hypanthium 0.5-1.5 mm long, ca. 1 mm
diam., densely pubescent, calyx tube ca. 0.5 mm long,
calyx lobes 4 or 5, 0.5 mm high, triangular; corolla red
in early bud, becoming rose red or pink, tube (6-)8-10
mm long, 0.7-1.6 mm diam., slender and widening be-
low the lobes, sparsely to densely puberulent, lobes 2-3
mm long, ca. 2 mm wide at the base, glabrous distally
on the inner surface and villous near the mouth; stamens
4, anthers 2-2.5 mm long, anthers partly exserted; style
9-1 1 mm long, stigma 0.3-0.7 mm long. Fruits 3-5 mm
long, 3-6 mm broad, white, usually 4-lobed, glabrous or
sparsely puberulent.

Common shrubby plants of lower montane ev-
ergreen forest formations, from (900-)1 100-2200
m elevation. Probably flowering and fruiting
throughout the year, but flowering primarily in
February-September and fruiting primarily in
June-September. The species ranges from the Cor-
dillera de Tilaran in Costa Rica to eastern Panama.

Gonzalagunia rosea is recognized by the thin-
petiolate narrowly ovate to lanceolate leaves, the
long inflorescences with pedunculate flower clus-
ters, the pink corollas, and the spongy white fruit.
This is a common weedy shrub of open habitats.
Some specimens of this species may be difficult to
distinguish from G. panamensis and G. ovatifolia,
and it is possible that hybridization occurs.

Gonzalagunia rudis (Standl.) Stand 1.. J. Wash.
Acad. Sci. 17: 170. 1927. Duggenia rudis Standl.,
Contr. U.S. Natl. Herb. 18: 125. 1916.

Shrubs, 1.5-4 m tall, leafy stems 0.7-3 mm thick, with
minute (0.2 mm) thin appressed-ascending whitish hairs,
terete and glabrescent; stipules 4-7 mm long, with a short
(1-2 mm) broad base and long narrow awn. Leaves with
petioles 1-3 mm long, ca. 1 mm thick, hertellous; leaf
blades 2.5-10 cm long, 1-4 cm broad, narrowly ovate
to ovate-oblong or lanceolate, apex tapering gradually
and acute to long-acuminate, base acute to obtuse, drying
thin-chartaceous, with scattered thin appressed hairs ca.
0.5 mm long on both surfaces, with denser pubescence
on the veins beneath, 2 veins 4-7/side. Inflorescences
terminal or axillary, solitary, 4-14 cm long, spicate, pe-
duncles 1-3 cm long, 0.3-0.7 mm thick, densely pubes-
cent with ascending hairs, flowers usually solitary or in
groups of 2-3 ca. 5 mm diam., subtended by linear bracts
2-3(-5) mm long, pedicels 1-2 mm long. Flowers
5-parted, hypanthium ca. 1 mm long, urceolate, densely
puberulent, calyx lobes 1-2 mm long, linear, corolla white,
tube ca. 6 mm long and 0.7 mm diam., lobes ca. 5 mm
long and 1 mm broad, glabrous externally, puberulent

within; stamens included. Fruits 2-4 mm long, 2.5-5
mm broad, white, depressed globose, 4-lobed, with thin
erect hairs 0.5 mm long.

Plants of evergreen forest formation on the Pa-
cific slope of southern Costa Rica, from near sea
level to 500 m elevation (to 1 000 m in Panama).
Flowering primarily in the wet season (June-Sep-
tember). This species ranges from about 84W in
Costa Rica to eastern Panama.

Gonzalagunia rudis is recognized by its smaller
thin lanceolate leaves, the slender spike-like inflo-
rescences with mostly solitary subsessile flowers,
and the unusual calyx and corolla with long narrow
lobes. Costa Rican collections differ somewhat in
their narrower more lanceolate leaves, but collec-
tions with such leaves are also found in Panama.

Gonzalagunia stenostachya (Standl.) W. Burger,
comb. nov. Rondeletia stenostachya Standl.,
Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1372.
1938. Arachnothryx stenostachya (Standl.) Bor-
hidi, Acta Bot. Hung. 33: 303. 1987.

Shrubs to 3 m tall, leafy stems 2.3-6 mm thick, with
short dense yellowish or reddish brown hairs ca. 0.5 mm
long; stipules 4-15 mm long, triangular, acute, with yel-
lowish hairs along the midrib. Leaves with petioles 2-4
mm long, ca. 1.5 mm thick; leaf blades 7-16 cm long,
2-6 cm broad, elliptic, elliptic-oblong to lanceolate or
oblanceolate, apex acute or short-acuminate, base grad-
ually narrowed and cuneate base (rounded in Dryer 1274),
drying stiffly chartaceous and dark reddish brown above
(grayish green beneath), with thin hairs 0.4 mm long or
glabrescent above, densely puberulent on the midvein
beneath, 2 veins 7-12/side, ascending. Inflorescences
(7-) 15-25 cm long, 1.3-2.5 cm broad, spiciform thyrsoid
panicles, peduncles 1.3-6 cm long, 1.2-2 mm thick, stri-
gose, lateral cymules sessile and separate along the rachis,
of (l-)2-5 flowers, bracts ca. 3 mm long, linear, pedicels
0-4 mm long. Flowers 4-parted, hypanthium ca. 1.5 mm
long and 1.5 mm diam., densely sericeous, calyx lobes
1-2 mm long, narrowly triangular; corolla white, densely
sericeous with hairs 0.5-0.9 mm long, tube 4-9 mm long,
slender, lobes 3-4.5 mm long, oblong. Fruits ca. 3 mm
long and 4 mm broad (?immature) with sericeous hairs
ca. 0.5 mm long.

Plants of wet evergreen forest formations of the
Caribbean slope, 300-1200 m elevation. Flower-
ing in February-March, July, and October; fruit-
ing in October. This endemic species is known
from near Monteverde and the La Selva-Braulio
Carillo area.

Gonzalagunia stenostachya is recognized by its
subsessile leaves with many ascending secondary
veins, the long slender spikes with sessile and well-

150

FIELDIANA: BOTANY

separated cymules, and the sericeous flowers. The
Panamanian G. kallunkii Dwyer (?= G. veraguen-
sis Dwyer) has rather similar inflorescences but
the petioles of that species are well developed and
the leaves have fewer secondary veins.

Guettarda Linnaeus

Trees or shrubs, branchlets terete, puberulent or gla-
brous, occasionally with spines; stipules interpetiolar,
simple or slightly connate above the petioles (intrapetio-
lar), often slightly overlapping above the node, triangular
and acuminate to rounded distally, persisting or decid-
uous. Leaves opposite (rarely 3 or 4/node), petioles short
to long; leaf blades entire, with pinnate venation, the
distal secondaries often strongly ascending and the 3
veins often subparallel, domatia sometimes present. In-
florescences solitary and axillary (1 or 2/node), usually
pedunculate and with cymose branching, branches of the
inflorescences often dichotomous (bifurcate) and with
sessile flowers along 1 side, bracts and bracteoles present
or reduced. Flowers bisexual (rarely unisexual), radially
symmetrical, 4-9-parted (usually 5- or 6-parted), hy-
panthium ovoid to globose or tubular, calyx tube cu-
pulate to campanulate or short-tubular, calyx entire or
with 2-9 poorly developed lobes/teeth; corolla funnel-
form or salverform, white, yellowish, purple or bluish,

corolla lobes 4-9, imbricate or subvalvate, the margins
often undulate; stamens 4-9, anthers narrow, sessile or
subsessile, dorsifixed, included; ovary 2-9-locular, with
1 pendulous ovule from apex of each elongate-tubular
locule, stigma capitate or lobed. Fruits drupaceous, glo-
bose to elongate, rounded or angulate in cross-section,
the exocarp fleshy but thin, endocarp woody to stony,
2-9-locular.

A genus of 60-80 species in the New World
tropics, with a few species in the southwest Pacific
and a species widespread on tropical coasts (G.
speciosa L.). Some species of Guettarda have in-
florescences with two equal cincinnoid branches;
these are scorpioid cymes (cincinni) in which the
sessile flowers are all in a close line along a single
side of the rachis. The leaves of Guettarda are
generally thin, often clustered at the ends of
branchlets, and with the distal secondaries strong-
ly ascending. Domatia are often present, and the
3 veins are usually subparallel. The inflorescences
are always axillary, and the flowers are all salver-
form in Costa Rican species. This treatment ben-
efited from the annotations made by Alfredo Gri-
jalva in 1982.

Key to the Species of Guettarda

la. Inflorescences subsessile; plants of Cocos Island and the evergreen Pacific lowlands . . .G. conferta

Ib. Inflorescences short-to long-pedunculate; plants of mainland Central America 2

2a. Bracts subtending the flowers 3-6 mm long, thin-brownish, narrowly ovate-oblong; Pacific slope

and lowlands 3

2b. Bracts subtending the flower absent or less than 3 mm long and caducous 4

3a. Inflorescences with peduncles 2.5-6 cm long; leaf blades usually rounded at the base; Gulf
of Nicoya G. brenesii

3b. Inflorescences with peduncles 1-3 cm long; leaf blades acute at the base; western Costa Rica

G. foliacea

4a. Inflorescences with short (to 1 cm) branches, the branches not cincinnoid (scorpioid-cymose) in

appearance 5

4b. Inflorescences with conspicuous lateral cincinnoid (scorpioid) branches more than 1 cm long (with

the flowers all along 1 side 6

5a. Corolla tubes 8-12 mm long, peduncles less than 3 cm long; petioles 4-30(-40) mm long;
fruit ca. 12 mm long, subglobose; Pacific and Caribbean lowlands G. macrosperma

5b. Corolla tubes 13-18 mm long, peduncles more than 3 cm long; petioles 20-70 mm long;

fruit ca. 20 mm long, oblong, truncated distally; Golfo Dulce and Panama

G. sanblasenesis

6a. Trees to 30 m tall; leaves and twigs glabrous; flowers becoming 2-4 mm distant on the rachis

[corolla tubes 20-30 mm long] G. turrialbana

6b. Trees to 10(-25) m tall; leaves and stems densely to sparsely puberulent; flowers 0-2 mm distant

on the rachis 7

7a. Stipules glabrous, often broadly overlapping; midvein sparsely appressed strigose along the sides

beneath [corolla tube 16-20 mm long; 1500-2500 m elevation] G. poasana

7b. Stipules pubescent (at least along the midrib); midvein densely pubescent over the entire surface

on the lower side of the leaf blade . 8

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

151

8a. Leaf blades narrowly oblong and coriaceous, densely tomentulose beneath; corolla tube 4-7 mm

long [1200-1900 m elevation] G. tornefortiopsis

8b. Leaf blades elliptic to broadly ovate, drying chartaceous, lacking a densely matted tomentum

beneath; corolla tubes 6-18 mm long 9

9a. Leaf blades rounded and often truncate at the base, to 30 cm long; peduncles to 10 cm long [corolla

tubes 15-18 mm long]; northern Caribbean lowlands G. combsii

9b. Leaf blades usually acute to obtuse at the base, to 22 cm long; peduncles to 4 cm long; evergreen

formations 300-1700 m elevation 10

lOa. Leaf blades with 6-10 pairs of major secondary veins, 3 veins clearly differentiated from the

smaller 4 veins; corolla tubes 13-18 mm long; Central Highlands and evergreen Pacific lowlands

G. crispiflora

lOb. Leaf blades with 3-5 pairs of major secondary veins, 3 and 4 veins little differentiated and parallel

with each other; corolla tubes 6-9 mm long; Caribbean slope Chomelia venulosa

Guettarda brenesii Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 18: 1303. 1938. Figure 33.

Trees to 10 m tall, leafy branchlets 2-4 mm thick, at
first with straight ascending hairs 0.5-1 mm long, terete,
soon glabrescent and becoming very dark with numerous
narrow lenticels ca. 0.5 mm long; stipules 7-12 mm long,
narrowly ovate-triangular, acute, sericeous along the
midrib and base, deciduous. Leaves clustered at the end
of branchlets, petioles 5-12 mm long, ca. 1 mm thick,
with thin ascending hairs 0.5-1.3 mm long; leaf blades
(4-)6.5-2 1 cm long, (3-)4.5-l 2 cm broad, broadly ovate
to broadly elliptic, apex obtuse or subrotund, base obtuse
(and rounded at the petiole) to broadly rounded and
subcordate, drying chartaceous and dark above, sparsely
hispidulous above with hairs ca. 0.5 mm long, more
densely pubescent beneath with thin whitish hairs to 1
mm long, 2 veins 6-8/side, with denser tufts of hairs
(domatia) in the vein axils, 3 veins subparallel basally
and joining with the opposing tertiary veins at an angle.
Inflorescences 3-10 cm long, equally wide, axillary, pe-
duncle 2-6.5 cm long, ca. 0.8 mm thick, pubescent, usu-
ally bifurcate with 2 main branches and further cymose
branching, bracts and bracteoles 4-8 mm long, lanceo-
late, pubescent along the midrib, distal axes to 4 mm
long often bearing single flowers and resembling pedicels
(but with bracteoles at their apex beneath the flowers).
Flowers with hypanthium ca. 1 mm long and 1 mm
broad, short-tubular and sericeous, calyx tube ca. 1 mm
long, entire and more sparsely pubescent distally; corolla
white, tube (9-) 14-1 8 mm long, 0.5-0.8 mm diam., with
minute appressed-ascending whitish hairs externally,
lobes 5, ca. 3 mm long and 1.7 mm broad, margin broad-
ly rounded and subentire. Fruits unknown.

Trees of lowland deciduous forest formations
near the Bay of Nicoya. Immature inflorescences
were collected in June (Brenes 15694 the type),
and inflorescences with falling corollas were col-
lected in July. Fruiting in July-August and Oc-
tober-November. The species is known only from
along the Pacific coast of central Costa Rica.

Guettarda brenesii is recognized by the bracteate
inflorescences, the long narrow corollas, the broad-
ly ovate leaves, and the rocky, seasonally very dry

seaside habitat. This species is probably related to
G. foliacea, which shares characters of the inflo-
rescence. The type appears to have immature leaves
and inflorescences; hence, Standley's description
represents minimal measurements. A highly re-
stricted range and short flowering season may ex-
plain the paucity of collections.

Guettarda combsii Urban, Symb. Ant. 6: 48. 1 909.

Small to large trees (shrubs), 5-30 m tall, trunks to 60
cm dbh, leafy branchlets 2-6 mm thick, with soft whitish
erect or ascending hairs 0.5-1 mm long but soon gla-
brescent, terete; stipules 6-14 mm long, ovate to lan-
ceolate, acuminate, pubescent along the midrib. Leaves
often clustered at the ends of branchlets, petioles 2-9 cm
long, 1.3-2 mm thick, minutely pubescent; leaf blades
7-20(-28) cm long, 5-13 (-20) cm broad, very variable
in shape, from broadly ovate to ovate-oblong or subor-
bicular, apex abruptly narrowed and obtuse or short acu-
minate, base broadly obtuse to rounded and truncate to
subcordate, drying thin-chartaceous, subglabrous or
sparsely pubescent above with thin hairs ca. 1 mm long,
more densely pubescent beneath with thin whitish hairs
0.5-1 mm long, 2 veins 8-1 I/side, 3 veins parallel and
prominent. Inflorescences 12-18 cm long, with long (5-
1 5 cm) peduncles and 2 or 4 distal cincinnoid branches
2-5 cm long, minutely (0.1-0.4 mm) pubescent, bracts
2-4 mm long and ca. 1 mm broad, flowers sessile and
closely spaced. Flowers with hypanthium ca. 1 mm long
and 1.3 mm diam., with a dense greenish white or pale
grayish white tomentum, calyx tube 1.5-2.5 mm long,
entire or 2-lobed; corolla greenish white to cream white,
tube 15-18 mm long, 1-1.5 mm diam., with dense re-
trorse sericeous hairs, corolla lobes 5-7, ca. 3-4 mm long;
ovary 4- or 5-locular. Fruits 6-8 mm long, subglobose,
covered with a dense minute (0.05 mm) tomentum, gray-
ish green, the surface becoming wrinkled.

Trees of evergreen Caribbean rain forest for-
mations, from near sea level to 900 m elevation.
Flowering in May; fruiting in September in Belize.
The species ranges from Belize to southeastern
Nicaragua.

152

FIELDIANA: BOTANY

Guettarda combsii is recognized by the broad
pubescent leaf blades usually rounded and trun-
cate to subcordate at the base, the long-pendun-
culate inflorescences with four short cincinnoid
branches, and the subglobose fruit. The fact that
these plants become very tall trees may account
for the paucity of collections in southern Central
America. A sterile collection from a tall tree at
Minefields. Nicaragua (Proctor et al. 27130 F), sug-
gests that this species is also likely to occur in
northern Costa Rica.

Guettarda conferta Benth., Hot. voy. Sulph. 106.
1845.

Trees to 10 m tall, with dense, ferruginous hairs on
the branchlets, petioles, peduncles, and nerves of the
leaves; stipules broadly obovate, 12 mm long, hirsute
externally at base, otherwise glabrous, about equaling
the petioles, deciduous. Leaves with blades 10-20 cm
long, 6-9 cm broad, ovate, apex acuminate, base acute,
hirsute on both sides. Inflorescences 2.5-3 cm long, cy-
mose, subsessile, the branches recurved and 2.5 cm long
or less. Flowers ca. 8 mm long, calyx tube 1-2 mm long,
shallowly 3- or 4-dentate; corolla white, sericeous-hir-
tellous, tube ca. 10 mm long, corolla lobes 4, ca. 4 mm
long, obtuse crispate. Fruits ovoid-tetragonous, 4 mm
long, 3 mm diam.. hirsute, 4-locular.

Plants of Cocos Island and the Pacific slope of
southern Costa Rica and the Osa Peninsula, 0-
300 m elevation. Flowering in August and Decem-
ber; fruiting in January.

Guettarda conferta is distinguished by its short
sessile inflorescences. In general aspect this species
resembles G. crispiflora.

Guettarda crispiflora Vahl, Ecolog. Amer. 36: pi.
6. 1797. G. chiriquiensisSlandl., Ann. Missouri
Hot. Gard. 25: 838. 1938. Figure 32.

Small to medium-sized trees 4-20 m tall, leafy branch-
lets 1.5-6 mm thick, at first quadrangular but soon be-
coming terete, with short (0.3 mm) thin appressed-as-
cending hairs but glabrescent; stipules 8-18(-22) mm
long, to 1 cm broad, ovate-elliptic to slightly obovate,
apex acute to acuminate, with thin ascending sericeous
hairs along the midrib and at the base. Leaves clustered
distally, petioles 2-7 cm long, 1.3-2 mm thick, minutely
puberulent and with longer hairs along the adaxial side;
leaf blades (6-)8-22 cm long, (3-)5-l 1 cm broad, ovate-
elliptic or ovate-oblong to ovate-lanceolate or broadly
ovate, apex tapering gradually and acuminate or acute,
base obtuse to slightly rounded or attenuate and acute,
drying stiffly chartaceous, glabrous or minutely (0.2 mm)
puberulent above, pubescent beneath with larger (0.4-1
mm) hairs on the major veins and smaller (0.3 mm) thin

whitish hairs on the 3 veins, tufts of hairs (domatia)
sometimes present in the vein axils beneath, 2 veins 6-
10/side, distal 2 veins strongly ascending, 3 veins often
parallel. Inflorescences 2-6(-8) cm long, equally wide,
with a short (4-25 mm) peduncle and 2 diverging cin-
cinnoid branches 1 5-30(-60) mm long and enlarging in
fruit, the rachis minutely grayish white tomentulose, the
flowers sessile and closely (0-4 mm) spaced. Flowers
sweet-scented, hypanthium 1.5-2.5 mm long, 1.5-2 mm
diam., densely grayish white tomentulous, calyx tube ca.
0.5 mm long, calyx lobes 4, ca. 0.5 mm high; corolla
white or pinkish, tube 13-18 mm long, 1-1 .5 mm diam..
narrowly tubular, densely short-sericeous with retrorse
or spreading hairs, lobes 5-6 mm long, the lobes with
smaller undulate marginal lobes. Fruits ca. 8 mm long
and 6 mm diam., oblong and with 4 prominent longi-
tudinal ribs, becoming purple and with a white pulp.

Trees of the Caribbean slope cloud forest for-
mations and southern Pacific wet forest forma-
tions, from 300 to 1700 m elevation. Flowering
in January, April, and June-September, with Jan-
uary and August collections being most frequent;
fruiting in March and JuneJanuary. In our area
the species is known from the Caribbean slope of
the Central Highlands (from near Monteverde
eastward to Tapanti, Cartago, and San Joaquin de
Dota, San Jose), on the Osa Peninsula, near San
Vito, and in the Chiriqui Highlands. The species
also occurs in the Lesser Antilles and Trinidad.

Guettarda crispiflora is recognized by the short-
pedunculate inflorescences with bifurcate cincin-
noid branching, the white corollas with crisped
and undulate corolla lobes, the four-angled fruit,
longer petioles, and the subparallel (almost lineo-
late) minor venation. Guettarda poasana is closely
related to G. crispiflora, and material of the two
species should be compared when making iden-
tifications. Guettarda chiriquiensis was distin-
guished by its more densely pubescent vegetative
parts, but there are a few intermediate collections
in Costa Rica. Nevertheless, the distinctive pop-
ulations of the Chiriqui Highlands and adjacent
Costa Rica may be worthy of subspecific rank.

Guettarda foliacea Standley, Contr. U.S. Natl.
Herb. 18: 139. 1916. Figure 33.

Shrubs or small trees, 3-6(-10) m tall, often branching
from the base and with clambering branches, leafy stems
0.9-4 mm thick, at first strigose with thin ascending hairs
0.3-1 mm long, glabrescent, becoming brown and terete,
spines often present; stipules 3-5(-12) mm long, trian-
gular to lanceolate, strigulose. caducous. Leaves oppo-
site, petioles 3-25 mm long, 0.4-1.2 mm thick, ap-
pressed strigose to sericeous; leaf blades 3-16 cm long,
2-7 cm broad, elliptic to ovate or obovate, apex acute
to short-acuminate, base cuneate to slightly rounded or

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

153

subtruncate, drying thin-chartaceous and greenish, with
thin whitish hairs 0.2-0.9 mm long on both surfaces
(dense only on the major veins), 2 veins 4-8/side, 4
veins often parallel, domatia of dense hairs present in
the vein axils beneath. Inflorescences 2-7 cm long, to 5
cm broad, peduncles 5-30 mm long, 0.5-1.1 mm thick,
with thin ascending hairs, usually with a single pair of
dichotomous distal branches (each with 3-7 flowers),
subtended by lanceolate ciliate bracts 3-8 mm long, flow-
ers sessile and crowded in cymes. Flowers densely mi-
nutely sericeous externally, hypanthium ca. 1 mm long,
calyx tube 1-1.5 mm long, ca. 1.3 mm diam., entire;
corolla white, tube 12-20 mm long, lobes 4(-5), 2-5 mm
long, rounded distally. Fruits 1-3 cm diam., globose,
with a dense minutely velutinous surface, becoming red.

Plants of evergreen or partly deciduous forests,
0-300(-1000) m elevation. In central Panama
flowering occurs primarily in late June-early July
(Croat, 1978) and fruiting in September-Novem-
ber. This species ranges from the Cordillera de
Tilaran to Colombia.

Guettarda foliacea is recognized by its small in-
florescences, lack of calyx lobes, thin variable leaves
on slender petioles, and minor venation, often with
a small group of parallel veins (sublineolate). The
spines are rarely seen on herbarium collections.
The westernmost collection (8453'W, Haber et al.
4775 CR, MO) came from the edge of cloud forest
at 1 000 m elevation with immature flowers in May.

tomentulose, calyx tube 0.5-1 mm long, entire; corolla
white or yellowish, tube 8-13 mm long, 0.5-1.3 mm
diam., narrowly tubular, lobes 3-4 mm long, rounded
distally. Fruits 10-18 mm long, globose to oblong, yel-
lowish brown or grayish with a dense covering of minute
(0.1-0.2 mm) velvet-like or matted hairs.

Trees and shrubs of both evergreen and decid-
uous forest formations, from near sea level to 1 000
m elevation on the Pacific slope and from near sea
level to ca. 500 m on the Caribbean slope. Flow-
ering in March-November, with the majority col-
lected in May; fruiting in July-January. The spe-
cies ranges from southern Mexico to Panama.

Guettarda macrosperma is recognized by its
small cymose inflorescences, rounded fleshy fruit
usually over 1 cm diam., and often smaller leaves
with the subparallel 3 veins usually meeting at
angles between the 2 veins (> -shaped). Trees in
the Caribbean lowlands may reach 30 m in height;
the same trunk and bark characteristics are found
in trees of both seasonally dry forest and evergreen
rain forests (N. Zamora, pers. comm.). Some spec-
imens may resemble Chomelia panamensis. It ap-
pears that Guettarda divaricata (Roem. & Schult.)
Standl. of Mexico is closely related, and the two
may be part of a more broadly defined taxon. Ma-
lacahuite is a common name.

Guettarda macrosperma J. D. Smith, Bot. Gaz. 1 8:
204. 1893. Figure 33.

Shrubs or more often trees, 4-12(-30) m tall, trunks
often fluted or with deep depressions, with dark bark
exfoliating in patches, leafy branchlets 1.2-4 mm thick,
sparsely puberulent with thin ascending hairs 0.5-1.5
mm long, soon glabrescent and becoming dark brown
or blackish with short (0.3-1.2 mm) grayish lenticels;
stipules 3-8(-12) mm long, triangular-lanceolate, dense-
ly sericeous on the back with longer hairs, caducous.
Leaves often crowded at the ends of branchlets, petioles
(4-)10-45 mm long, 0.8-1.7 mm thick, with straight
ascending hairs; leaf blades (4-)6-18 cm long, (2-)3-l 1
cm broad, broadly elliptic, broadly ovate-elliptic, obo-
vate-elliptic, or elliptic-oblong, apex abruptly narrowed
and short-acuminate to acute or obtuse, base obtuse to
rounded and subtruncate, drying chartaceous to stiffly
chartaceous, sparsely pubescent with thin short (0.3-0.7
mm) hairs and glabrescent, with thin short (ca. 0.3 mm)
ascending hairs along the 3 veins beneath and with lon-
ger and denser hairs along the major veins, 2 veins 4-
8/side, 3 veins rarely subparallel, often with tufts of hairs
(domatia) in the vein axils. Inflorescences 2-6 cm long,
primary peduncles 4-38 mm long, usually with 2 short
distal branches, densely ascending sericeous, bracts 2-3
mm long, flowers sessile and closely crowded (not clearly
cincinnoid in arrangement). Flowers with hypanthium
1-1.5 mm long and ca. 1 mm diam., densely whitish

Guettarda poasana Standl., J. Wash. Acad. Sci. 1 8:
182. 1928. Figure 32.

Small trees, 3-10(-15) m tall, trunk to 40 cm thick,
bark shredding off in oblong patches, leafy branchlets 2-
5 mm diam., somewhat flattened in early stages and
glabrous, drying dark but becoming grayish in age, len-
ticels difficult to see; stipules 12-20 mm long, 8-10 mm
broad, ovate and long-acuminate at apex, the stipules
overlapping on the sides, glabrous. Leaves clustered near
the ends of branches, petioles (2-)3-7(-10) cm long, 0.8-
1.6 mm thick, glabrous and drying dark; leaf blades 7-
14(-16) cm long, 2.5-6(-9) cm broad, broadly elliptic-
ovate to broadly elliptic or elliptic, apex gradually ta-
pering and short-acuminate, base obtuse to acute and
slightly decurrent on petiole, drying stiffly chartaceous,
glabrous and lustrous above, minor venation with thin
whitish ascending hairs ca. 0.3 mm long beneath, with
longer (0.5-1 mm) straight hairs on the mid vein and
secondaries beneath, 2 veins 5-8/side, the distal strongly
ascending, 3 veins often subparallel but not prominent
beneath, small tufts of hairs (domatia) often present in
the vein axils beneath. Inflorescences axillary, 3-6 cm
long and equally wide, peduncles 1-3 cm long, glabrous
or sparsely puberulent, with 2 cincinnoid branches 2-
3.5 cm long, the flowers 5-11 on each branch and sessile,
ca. 1-4 mm distant. Flowers sweet-scented, hypanthium
1.5-2 mm long, 1.3 mm diam., glabrous or sparsely
puberulent, calyx tube 0.5-1 mm long, lobes ca. 0.3 mm
long; corolla white, reddish, or lavender, tube 1 6-20 mm

154

FIELDIANA: BOTANY

long 1-1.4 mm diam., densely tomentulose externally,
lobes 4-5, 4-6 mm long, usually white with fringed-
undulate smaller lobes; tips of the anthers exserted 1-2
mm from the mouth of the tube. Fruits becoming 8 mm
long and 6 mm diam., oblong and with 4 prominent
longitudinal ridges, reddish purple.

Trees of evergreen cloud forest formations,
1300-2200(-2700?) m elevation (down to 1 100 m
on the northern vocanoes). Flowering in March
and May-November (mostly in June); fruiting
probably throughout the year. This species is en-
demic to Costa Rica and ranges from the Cordille-
ra de Guanacaste in the west to the eastern slopes
of Volcan Barva.

Guettarda poasana is recognized by its glabrous
stipules and stems, long-petiolate leaves, narrowly
tubular flowers with whitish fringed lobes, four-
angled fruit, and restricted cloud forest range. The
petioles dry dark and are sometimes pink in life.
This species is closely related to G. crispiflora and
might be considered a subspecific element of that
species. However, though their ranges overlap
slightly, G. crispiflora and G. poasana do not grow
in the same locality; G. poasana is generally found
at higher altitudes.

Guettarda sanblasensis Dwyer, Ann. Missouri Bot.
Gard. 67: 204. 1980. Figure 33.

Trees, 12-20 m tall, leafy branchlets 1.5-4 mm thick,
with short (0.3 mm) appressed-ascending hairs, quickly
glabrescent, terete; stipules 3-6 mm long, densely seri-
ceous with pale yellowish ascending hairs 0.5-1 mm long,
early caducous. Leaves 1-5 cm distant at the ends of
branchlets, petioles 2-7 cm long, 0.7-1 .3 mm thick, with
slender appressed-ascending hairs; leaf blades 1 1-22 cm
long, (4-)6-l 2 cm broad, broadly elliptic to broadly ovate-
elliptic, apex abruptly narrowed and short-acuminate (or
acute), base obtuse to slightly rounded, drying charta-
ceous and dark brown above, lustrous above and with
scattered thin appressed hairs 0.2-0.3 mm long, the hairs
more numerous and longer (0.3-0.5 mm) beneath, 2
veins 5-9/side, the distal arcuate-ascending, 3 veins
prominent above and below and paler in color beneath,
subparallel or > -shaped between the secondaries, with
minute tufts of hair (domatia) in the vein axils beneath.
Inflorescences 6-10 cm long, primary peduncles to 6 cm
long, ca. 1.2 mm thick and minutely appressed-puber-
ulent, bifid but the 2 branches with additional dichot-
omous branches (not scorpioid/cincinnoid), pedicels 0-
2 mm long, bracts minute (0.5 mm) and caducous. Flow-
ers with hypanthium ca. 1.5 mm long and 1.2 mm diam.,
with longitudinal ribs, calyx tube 2-3 mm long and ca.
2 mm diam., minutely velutinous, subentire distally;
corolla white or pink, tube 13-18 mm long, 1-1.3 mm
diam., minutely (0.2 mm) ascending-sericeous, lobes 5-
6, 4-5 mm long and 1-1.5 mm broad, oblong and entire,
stigma 0.5 mm long, subglobose. Fruits 2-2.2 cm long

and 8-12 mm thick, oblong-obovoid, abruptly rounded
(truncated) at apex, persisting calyx 1-3 mm long, 1.5-
2 mm diam., surface minutely velutinous and yellowish
or grayish brown.

Trees of evergreen forest formations of the Pa-
cific lowlands, from near sea level to 600 m ele-
vation and usually on limestone. Flowering in July-
September; fruiting in September-November. This
species is known only from a few collections in
the Golfo Dulce region of Costa Rica; it ranges to
eastern Panama.

Guettarda sanblasensis is recognized by the
broadly elliptic leaves on long slender petioles, the
long-pedunculate inflorescences with dichoto-
mous or cymose distal branching, long-tubular co-
rollas with five or six entire oblong perianth lobes,
and oblong fruit with truncated apex and usually
persisting calyx tube.

Guettarda tournefortiopsis Standl., Publ. Field
Columb. Mus., Bot. Ser. 7; 293. 1931. Tour-
nefortiopsis reticulata Rusby, Bull. New York
Bot. Gard. 4: 369. 1907, non G. reticulata Gri-
seb., 1863. Figure 32.

Small trees, 4-10 m tall, leafy branchlets 1.5-6 mm
thick, at first densely tomentose with soft wooly hairs to
2 mm long, 4-angular, soon .glabrescent and dark with
lenticels 0.3-0.7 long, becoming terete and pale grayish;
stipules 1 2-20 mm long, narrowly ovate-triangular and
acuminate, tomentose, usually early deciduous. Leaves
with petioles 13-35 mm long, 1.5-2.2 mm thick, gla-
brescent and drying dark; leaf blades 9-16 cm long, 2-
5 cm broad, lanceolate to narrowly oblong-lanceolate or
narrowly elliptic-oblong, apex acute to short acuminate,
base obtuse to acute and usually with the margin revo-
lute, drying subcoriaceous, dark brown above, glabrous
and usually lustrous above, with the major and minor
veins slightly impressed above and the surface slightly
rugose, densely tomentulous between the veins beneath
and whitish to pale brown in color, 2 veins 9-1 I/side,
domatia present. Inflorescences 2-5 cm long and equally
wide, becoming 8 cm long in fruit, peduncles 1 2-20 mm
long (to 30 mm in fruit), 1 .5 mm thick and tomentulous,
with 2 cincinnoid branches 2-6 cm long and each branch
bearing up to 20 flowers in 2 rows along 1 side, flowers
sessile and closely crowded, bracts absent. Flowers with
hypanthium 1 .5-2 mm long, covered by a dense tomen-
tum, calyx lobes ca. 1 mm broad, broadly obtuse and
difficult to see; corolla reddish to coral pink, white with-
in, tube 4-7 mm long, ca. 1.5 mm diam., densely pu-
berulent with retrorse hairs externally, lobes 5, 1-2 mm
long. Fruits 5-8 mm long, 4-6 mm diam., globose to
rounded-oblong, 5-angled during development, sessile,
becoming purple or black.

Trees of wet montane cloud forest formations,
from 800 to 1900 m elevation. Collections with

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

155

flowers (or flower buds) and fruits have been made
in January-March and August-October. Known
only from the Cordillera de Tilaran and the west-
ern parts of the Cordillera de Talamanca, prov-
inces of Cartage and San Jose, in Costa Rica. The
species is also known from the Chiriqui Highlands
and was originally described from Bolivia.

Guettarda tournefortiopsis is easily recognized
because of its stiff narrow leaves densely tomen-
tulose beneath (when young), the two-branched
scorpioid inflorescences (resembling those found
in the Boraginaceae), small tomentulose red flow-
ers, and small sessile fruits.

and long corolla tubes. The tall size of this dis-
tinctive species may explain why it had not been
collected before 1986. The collections from the
Osa Peninsula differ in a number of ways from the
type; more material is needed to assess the pattern
of variation.

Hamelia Jacquin

REFERENCE T. S. Elias, A monograph of the
genus Hamelia (Rubiaceae). Mem. New York Bot.
Gard. 26: 81-144. 1976.

Guettarda turrialbana Zamora & Poveda, Ann.
Missouri Bot. Card. 75: 1 157. 1988. Figure 30.

Trees to 30 m tall and with trunks 40 cm dbh, bark
exfoliating, leafy branchlets 1.5-6 mm thick, glabrous
and dark in early stages, terete, becoming pale grayish
and with few broadly ellipsoid lenticels 0.8-1 .5 mm long;
stipules ca. 1 2-25 mm long, triangular and acuminate,
overlapping, glabrous, deciduous. Leaves clustered at the
ends of branchlets, petioles 2-3.5 cm long, 1-2 mm thick,
glabrous and drying dark; leaf blades 9-26 cm long, 4-
1 2 cm broad, oblong to broadly elliptic-oblong or ovate-
oblong, apex abruptly narrowed and short-acuminate,
base abruptly narrowed or rounded and obtuse, drying
stiffly chartaceous, dark brown above, glabrous and lus-
trous above, subglabrous beneath with tufts of hairs
(domatia) in the vein axils, 2 veins 8-10/side, 3 veins
slightly raised above but not clearly subparallel. Inflo-
rescences 5-11 cm long, primary peduncles 3-5 cm long,
1-1.3 mm thick, glabrous and drying dark, cymose with
2 primary branches, the branches 2-3 cm long and with
3-5 flowers 2-5 mm distant on the rachis (or the branch-
es very short and all the flowers close together in Hammel
et al. 16848 CR, MO), pedicels short (1 mm) or absent,
flower subtended by short (0.5 mm) broad bracts. Flow-
ers glabrous externally, with the sweet odor of Coffea
flowers, hypanthium ca. 2 mm long and 1.2 mm diam..
calyx cup ca. 2 mm long, ca. 3 mm diam. distally and
entire; corolla white, tube 2040 mm long, 2-2.8 mm
diam., puberulent within, lobes 5, ca. 5 mm long and 2
mm broad, oblong and entire, distally rounded. Fruits
24-28 mm long, 6-12 mm diam., oblong, glabrous.

Trees of lowland rain forest formations; col-
lected at 600 m elevation, below the CATIE site
near Turrialba on the Caribbean slope, and on the
Osa Peninsula at 200-300 m elevation. Flowering
in May (Hammel et al. 16848 CR, F, MO) and June
(H err era 4246 MO, Zamora et al. 1263 CR holo-
type, F); fruiting in August and September. En-
demic.

Guettarda turrialbana is recognized by its tall
stature, mostly glabrous parts, oblong leaves, few-
flowered bifurcate inflorescences, entire calyx cup,

Shrubs or small trees, branchlets usually slender, terete
or 4-angled in cross-section, glabrous to densely puber-
ulent; stipules interpetiolar, usually small, entire or with
an awn (or trilobate), caducous or less often persistent.
Leaves opposite and decussate or in whorls of 3-5/node,
usually regularly spaced by well-developed internodes,
often long-petiolate, usually thin in texture, glabrous or
puberulent, pinnately veined, domatia present in some
species, conspicuous raphides visible on the dried leaf
surfaces in a few species. Inflorescences terminal or less
often axillary, 3-many-flowered, often thyrse-like with
cymose branches or with helicoid branches, pedunculate,
the flowers often all along 1 side of the distal branches,
sessile or short-pedicellate. Flowers bisexual and radially
symmetrical, monomorphic, hypanthium urceolate to
tubular, calyx tube minute or absent, calyx lobes 5, small,
rounded to elongate, usually persisting; corolla narrowly
tubular to funnelform or campanulate-urceolate (with a
short narrow base), bright yellow to orange, orange-red,
or deep red, with 5 longitudinal ribs alternating with the
lobes, corolla lobes 5, erect to recurved, slightly imbri-
cate in bud; stamens 5, filaments inserted at the base of
the corolla tube, flattened, anthers long-linear, sagittate
at the base and usually with the connective prolonged
distally, included or partly exserted; ovary (4-)5-locular,
with axile placentation and many ovules in each locule,
style narrowly cylindrical, stigmas 1-5. Fruit a fleshy
berry, oblong to ovoid or subglobose, with 5 longitudinal
ribs, terminated by the circular calyx scar and a conical
disc; seeds numerous, plano-convex or angled, foveolate.

Hamelia ranges from southern Florida, U.S.A.,
through Mexico, Central America, and the West
Indies into tropical South America. Elias recog-
nized 16 species in his fine monograph. The nar-
rowly tubular or funnelform yellow to orange or
red flowers are usually all aligned on the upper-
most sides of the inflorescence branches. In some,
the inflorescences have longer cincinnus-like
branches. Many of our species have three to four
leaves at distal nodes and a number have long-
petiolate leaves; most have small tufts of hairs
(domatia) in the vein axils beneath. Individual
plants may vary greatly within many species, and
this makes identification difficult. In addition, there

156

FIELDIANA: BOTANY

may be intermediates or hybrids between some of
our species.

The genus is divided into two subgenera. Sub-
genus Hamelia has narrowly tubular flowers that
do not enlarge distally at anthesis and are red,
orange, or yellow in color. Subgenus Amphituba
has the yellow corolla tube slightly to conspicu-
ously expanded distally at anthesis. However, the

flowers of subgenus Amphituba remain narrowly
tubular until just before anthesis and may be dif-
ficult to distinguish from those of subgenus Ha-
melia on this account. Species of Hoffmannia may
be similar, but they have consistently axillary in-
florescences; compare Deppea, with papery cap-
sules.

Key to the Species of Hamelia

la. Young stems and undersides of leaves conspicuously pubescent with hairs 0.2-0.5 mm long .. 2
Ib. Young stems and undersides of leaves glabrous or with minute (0.1-0.2 mm) inconspicuous hairs

2a. Stipules 7-17 mm long; leaf blades with 8-18 major 2 veins on each side, with hairs to 1 mm
long; corolla expanded distally at anthesis, corolla lobes 2-6 mm long [calyx lobes 0.5-2 mm

long] H. xerocarpa

2b. Stipules 2-5(-8) mm long; leaf blades with 4-9 major 2 veins on each side, with hairs to 0.5

mm long; corolla narrowly tubular, corolla lobes 1-2 mm long 3

3a. Calyx lobes 0-0.7 mm long; seeds 0.6-0.9 mm long; petioles 10-80 mm long; common wide-
spread plants, 0-1 500 m elevation H. patens

3b. Calyx lobes 24 mm long; seeds 1-1.2 mm long; petioles 3-20 mm long; uncommon plants of

the Caribbean lowlands, 0-300 m elevation H. rovirosae

4a. Corolla tube 8-1 3 mm long at anthesis; fruits 5-10 mm long 5

4b. Corolla tube 1 3-35 mm long at anthesis; fruits 7-16 mm long 6

5a. Inflorescences 3-9 cm long, corolla tube becoming expanded distally at anthesis; leaf blades
with 5-9 major 2 veins on each side, 5-1 7(-23) cm long, usually drying greenish to pale grayish;

fruits 4-7 mm long; Caribbean and evergreen Pacific lowlands H. axillaris

5b. Inflorescences 8-18 cm long, corolla tube narrowly tubular at anthesis; leaf blades with 7-12
major 2 veins on each side, 10-27(-32) cm long, usually drying reddish brown to pinkish gray;

fruits 6-10 mm long; evergreen Pacific slope H. magnifolia

6a. Leaf blades with 8-13 major 2 veins on each side; corolla tube distally widened (4-7 mm) at

anthesis; corolla lobes 2-4 mm long 7

6b. Leaf blades with 3-9 major 2 veins on each side; corolla tube narrowly (2-4 mm) tubular at anthesis

(in H. patens, often broader in H. calycosa)

7a. Leaf blades 3-6 cm broad; corolla tube 1 3-22 mm long; 0-800 m elevation ...//. xerocarpa

7b. Leaf blades 4-12 cm broad; corolla tube 25-35 mm long; 700-1600 m elevation

H. macrantha

8a. Leaf blades with 6-9 pairs of 2 veins, petioles 10-80 mm long; sepal lobes 0-0.8 mm long, corolla

lobes 1-2 mm long; common, 0-1 500 m elevation H. patens

8b. Leaf blades with 3-7 pairs of 2 veins, petioles 8-20 mm long; sepal lobes 1-2.5 mm long, corolla
lobes 3-6 mm long; not reported from Costa Rica H. calycosa

Hamelia axillaris Sw., Prodr. 46. 1788. H. lutea
Rohr ex Smith in Rees, Cyclop. 17.1811. Figure

42.

Shrubs, 1-5 m tall or small treelets to 5 m tall, leafy
branchlets 1 .2-4 mm thick, glabrous, with 4 longitudinal
ribs and 4-angular in cross-section but becoming terete;
stipules 2-6(-8) mm long, ca. 1 mm broad, triangular to
narrow with folded margins. Leaves opposite (rarely
4/node), petioles l-4(-7) cm long, ca. 1 mm wide, gla-

brous to sparsely and minutely papillate-puberulent; leaf
blades 5-17(-23) cm long, 2-8 cm broad, narrowly el-
liptic, elliptic-oblong or obovate to narrowly obovate-
oblong, apex acuminate with tip ca. 1 cm long, base
attenuate and decurrent on petiole, drying chartaceous
or membranaceous, glabrous above, glabrous or mi-
nutely (0. 1 mm) papillate-puberulent beneath, with tufts
of hairs (domatia) in the vein axils, 2 veins 5-9/side
and loop-connected near the margin, short (0. 1-0.3 mm)
linear cystoliths visible on the lower surface when dry.
Inflorescences axillary or terminal, 3-8 cm long, 3-8 cm

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

157

broad, compound dichasia with lateral branches 1-4.5
cm long, often scorpioid with 3-15 secund flowers on
the uppermost side, peduncles 5-15 mm long, glabrous
or minutely and sparsely puberulent, bracts 0.5-1 mm
long, narrow, flowers sessile or with pedicels. Flowers
with hypanthium 1.5-3 mm long, ca. 1.2 mm diam.,
glabrous, calyx lobes 0.5-1.5 mm long, 1 mm wide at
the base, glabrous to puberulent; corollas yellow, nar-
rowly urceolate to funnelform, tube 8-13 mm long, ca.
1 mm diam. near the base and 3-5 mm wide distally,
glabrous, lobes 1-2 mm long, broadly triangular; sta-
mens with filaments 4-5 mm long, anthers 5-8 mm long
with apical connective 0.5 mm long, style 8-10 mm long,
stigmas 3-4 mm long. Fruits 4.5-7 mm long, 3-4 mm
diam., ovoid-oblong to subglobose, disc 0.5 mm high
and 0.7 mm broad; seeds ca. 1 mm long.

Plants of the evergreen Caribbean slope and the
Osa Peninsula, from near sea level to 600(-1000)
m elevation. Flowering primarily in late June-Oc-
tober; fruiting in February and June-December.
The species is known from southern Mexico, Be-
lize, Guatemala, the larger West Indian islands,
and from Nicaragua southward to Venezuela and
Bolivia.

Hamelia axillaris is recognized by its short yel-
low corollas funnelform at anthesis, short rounded
fruit, general lack of pubescence, and usually com-
pact inflorescences with flowers along one side of
the distal branches. There may be intermediates
between this species and H. magnifolia on the Osa
Peninsula; compare the extreme upper-right figure
in Figure 42 (based on Utley & Utley 1208 F).

Hamelia calycosa J. D. Smith, Bot. Gaz. 12: 132.
1887.

Shrubs or small trees to 12 m tall, leafy branchlets 1-
3 mm thick, glabrous or glabrescent; stipules l-2.5(-4)
mm long, ca. 1 mm broad at the base, subulate or with
a narrow awn, minutely puberulent. Leaves opposite or
3-4/node on distal branches, petioles 6-20 mm long,
0.4-0.8 mm broad, glabrate; leaf blades 3-1 1(-15) cm
long, l-3.5(-5) cm broad, lanceolate to narrowly elliptic-
oblong or elliptic-oblanceolate, apex tapering gradually
and acuminate, tip ca. 7 mm long, base acute to cuneate
and decurrent on petiole, drying membranaceous to
chartaceous, glabrous above, glabrous or minutely (0. 1-
0.2 mm) pubescent beneath or rarely with longer (0.7
mm) thin hairs, with tufts of hairs (domatia) in the vein
axils beneath, 2 veins 3-7/side. Inflorescences terminal
or rarely axillary, 3-10 cm long, with 4-24 flowers, pe-
duncles to 2 cm long and often with 2 dichotomous
branches distally, usually minutely puberulent, bracts 1-
2 mm long, subulate or triangular, flowers secund and
with pedicels l-5(-8) mm long. Flowers with hypanthi-
um 2-4 mm long, oblong, glabrous or sparsely pubescent,
calyx lobes 1-3 mm long, ca. 0.5 mm wide, narrowly
oblong, caducous; corolla funnelform, yellow or pale or-
ange (striped with maroon), tube 1 5-24(-32) mm long,

constricted (1.5 mm) near the base and 8-10 mm diam.
distally, pubescent externally, lobes 3-6 mm long, ovate;
stamens 5, filaments 46 mm long, anthers 15-18 mm
long, 1 mm wide, distal rounded connective ca. 1 mm
long, style to 14 mm long, stigmas 5 and connate, ca.
1.2 mm long. Fruits 7-16 mm long, 4-8 mm thick, cy-
lindrical, disc conical and 1-3 mm long; seeds 1-1.2 mm
long.

Trees and shrubs of Caribbean rain forest for-
mations, from near sea level to 1500 m elevation.
Flowering is in April-September in northern Cen-
tral America. The species ranges from southern
Mexico, Belize, Guatemala, Honduras, Panama,
and Colombia to Peru.

Hamelia calycosa is recognized by the larger
sepal lobes, longer corollas expanded distally, ped-
icellate fruit, and smaller leaves with short peti-
oles. Although not yet collected in Costa Rica, this
species is likely to be present.

Hamelia macrantha Little, Carib. Forester 9: 274.
1948.

Shrubs or small trees to 8(-l 2) m tall and 1 3 cm dbh,
leafy branchlets 2-6 mm thick, 4-angular in cross-sec-
tion, glabrous or minutely puberulent; stipules 3-7 mm
long, 1-2.5 mm broad at the base, triangular to awl-
shaped, glabrous or rarely puberulent. Leaves opposite
but 3-4 at distal nodes, petioles (1 .5-)6-l 1 cm long, 1 .3-
2.7mm broad, glabrous or puberulent; leaf blades (7-) 1 2-
23(-27) cm long, (3-)4-12(-15) cm broad, broadly ob-
long or elliptic-oblong to ovate or obovate, apex obtuse
to abruptly short-acuminate, base rounded and subtrun-
cate to obtuse (cuneate), drying membranaceous to char-
taceous, glabrous above, glabrous to minutely (0. 1 mm)
papillate puberulent beneath in Costa Rica, domatia
sometimes present, 2 veins 8-12/side and loop-con-
nected near the margin. Inflorescences terminal or ax-
illary, 6-17 cm long and wide, to 15 cm broad, panic-
ulate, 20-many-flowered, peduncles 2-8 cm long, 1-3
mm thick, lateral branches to 6 cm long, bracts 0.4-0.7
mm long, ovate and acute, glabrate or puberulent, flow-
ers sessile to short (2 mm) pedicellate. Flowers with hy-
panthium 3-5 mm long, 1.5-2 mm diam., glabrous, calyx
lobes 0.7-1 .5(-2) mm long, 1-1.5 mm broad at the base,
triangular-subulate; corolla yellow, tubular-funnelform,
tube 23-35 mm long, constricted near the base and 5-

7 mm diam. distally, glabrous, lobes 3-5 mm long, ovate,
glabrous; stamens with filaments 6-8 mm long, anthers
16-20 mm long, included, connective prolonged ca. 1
mm beyond the thecae; style to 20 mm long, stigmas 5-

8 mm long. Fruits 11-15 mm long, 3.5-5 mm diam.,
ellipsoid, red becoming black and lustrous, ovarian disc
conical; seeds 0.8-1 mm long.

Trees and shrubs of evergreen cloud forest and
lower montane forest formations on both the Ca-
ribbean and Pacific slopes, from 700 to 1500 m
elevation in Costa Rica. Flowering in June-Sep-

158

FIELDIANA: BOTANY

tember in Costa Rica and Panama; fruiting in Au-
gust-September. The species ranges from Costa
Rica and Panama to Colombia and Ecuador.

Hamelia macrantha is recognized by the large
yellow corolla tube slightly widened distally at an-
thesis, larger leaves with many secondary veins
and often with long narrow petioles, and the mon-
tane habitat (in Costa Rica). Note that the widened
corolla tube is only seen at anthesis. The foliage
of this species resembles that of some specimens
of H. xerocarpa variety costaricensis, and the two
species may be difficult to separate.

Hamelia magnifolia Wernham, J. Bot. 49: 210.
1911. Figure 42.

Shrubs or small trees to 5(-6) m tall, trunks to 1 2 cm
dbh, leafy branchlets 2-7 mm thick, with 4 longitudinal
ridges and quadrangular in cross-section, glabrous or
minutely (0. 1 mm) papillate-puberulent; stipules 4-9 mm
long, to 2 mm wide at the base, narrowly triangular
(rarely bifid), glabrate. Leaves opposite (rarely 4/node),
petioles l-5(-7) cm long, 1.7-2.3 mm broad; leaf blades
10-27(-32) cm long, 4-ll(-15) cm broad, broadly ob-
long to ovate-oblong, elliptic-oblong or ovate-elliptic,
apex short-to long-acuminate with tip to 2 cm long, base
rounded and truncate to obtuse, drying stiffly charta-
ceous, glabrous above, glabrous beneath, 2 veins 7-
1 2(-l 5)/side, some 3 veins subparallel, domatia absent.
Inflorescences terminal, 8-12(-18) cm long, to 15 cm
wide, paniculate with opposite branching, with more than
60 flowers, peduncles 1-3 cm long, distal branches to 7
cm long and with secund flowers, minutely papillate-
puberulent or glabrous, bracts 0.5-1 mm long, narrow
pedicels 0-1 mm long. Flowers with hypanthium 2-3
mm long, 0.8-1 .5 mm diam., minutely puberulent, calyx
lobes 0.3-0.6 mm long, 1 mm broad at the base, deltoid
or broadly rounded, puberulent along the margin; corolla
narrowly tubular at anthesis (also somewhat narrowed
above the base and below the middle), yellow, tube 10-
13 mm long, 2-3 mm diam., lobes 1.2-3 mm long; sta-
mens with filaments 2-3 mm long, anthers 6-8 mm long,
apical connective ovate and ca. 1 mm long. Fruits 6-10
mm long, 3-4 mm diam., oblong to broadly ellipsoid,
glabrous, ovarian disc ca. 1 mm long, red; seeds 0.4-0.5
mm diam.

Plants of evergreen forest formations on the Pa-
cific slope, from 1 00 to 900 m elevation in Costa
Rica. Flowering in January-August; fruiting in
June-September and December-January. The
species is restricted to the evergreen formations of
the Pacific slopes of southern Costa Rica and ad-
jacent areas in Chiriqui Province, Panama.

Hamelia magnifolia is recognized by the small
narrowly tubular flowers in large inflorescences,
the large leaves often rounded at the base and with
many secondary veins, the smaller fruits, and the

restricted geographic range. Standley (1938) listed
zorillo Colorado as a common name.

Hamelia patens Jacq., Enum. PI. Carib. 16. 1760.
H. patens var. glabra Oersted, Vidensk. Meddel.
Naturhist. Foren. Kjobenhavn 1852: 42. 1853.
H. viridifolia Wernham, J. Bot. 49: 213. 1911.
Figure 42.

Shrubs or small trees, 2-7 m tall, leafy branchlets 1-
7 mm thick, glabrous or pubescent with crooked trans-
lucent or yellowish hairs to 0.6 mm long (in variety
patens), 4-angled in early stages but becoming terete;
stipules 2-6 mm long, narrowly oblong to linear, pu-
bescent, as many as the leaves at each node. Leaves
usually 3(-4) at distal nodes (less often opposite or rarely
5/node), often unequal at the same node, separated by
well-developed internodes, petioles (8-)20-55(-80) mm
long, 0.6-1.6 mm wide, glabrous to densely pubescent;
leaf blades 5-17(-23) cm long, l-7(-10) cm broad, el-
liptic-oblong, ovate-elliptic, ovate-oblong, or elliptic ob-
ovate-elliptic, apex usually short-acuminate, base acute
to attenuate (obtuse in larger leaves) and decurrent on
petiole, drying membranaceous to thin-chartaceous, gla-
brous (or sparsely pubescent in variety patens) above,
glabrous to densely villosulous beneath with thin straight
or curved whitish hairs 0.2-0.5 mm long (rarely with
scurfy yellowish hairs), tufts of hairs (domatia) often
present in vein axils beneath, 2 veins 6-9/side, small
(0.2 mm) linear raphides (cystoliths) resembling ap-
pressed hairs often visible on ,the dark upper surface of
dried leaves. Inflorescences terminal, solitary or 2-4, 4-
9(-15) cm long, to 12(-20) cm broad, an open panicle
with 2-3 nodes on the rachis and 2-4 branches at each
node, peduncles 1-3 cm long (to 5 cm in fruit), minutely
papillate-puberulent (less often with conspicuous hairs),
often orange to coral red, primary branches often ending
in a flower and 2 long distal secondary branches bearing
a row of sessile or subsessile flowers along the upper side,
bracts 0.3-1 mm long, triangular, distal flowers with
pedicels 1-5 mm long. Flowers with hypanthium 1.5-3
mm long, 1-2 mm diam., longitudinally ribbed, deep
red, glabrate to densely short-villous. calyx lobes 0.5-1
mm high, rounded or broadly triangular, persisting; co-
rolla narrowly tubular at anthesis, orange to reddish or-
ange, tube (12-)14-18(-23) mm long, 1.5-3 mm diam.,
lobes 1-2.5 mm long, 1-2 mm broad at the base, tri-
angular; stamens with filaments 5-7 mm long, anthers
8-12 mm long, included or slightly (3 mm) exserted. the
apiculatc connective 0.5 mm long; stigmas 3-5 mm long.
Fruits 6-1 3 mm long, 4-10 mm diam., oblong to ovoid-
oblong (subglobose), red becoming black or bluish, gla-
brous or with hairs ca. 0.2 mm long, raphides often
visible on the surface, with a ring around the top formed
by the calyx scar ca. 3 mm diam., disc to 1 mm high
and inconspicuous; seeds 0.5-1 mm long.

Common shrubs and treelets of open early sec-
ondary growth in evergreen and partly deciduous
forest formations, from near sea level to 1 600 m
elevation. Flowering and fruiting in all months of

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

159

the year in Costa Rica. The species ranges from
southern Florida, Mexico, the West Indies, and
Central America southward to northern Argentina
and Paraguay (the species is not found in the
Guianas, northeastern Brazil, the central Amazon
basin, or central Brazil).

Hamelia patens is recognized by its bright or-
ange or red-orange flowers with narrowly tubular
corollas with small lobes and included anthers and
the usually three- or four-leaved distal nodes. The
open inflorescences with dichotomous or dichasial
branching and with secund flowers often in a row
along the upper side of distal inflorescence branch-
es are also distinctive. These shrubs and little tree-
lets of open evergreen lowland secondary sites are
among the most common and conspicuous of Cos-
ta Rica's woody flora. They appear to germinate
only in open sunny sites. The breeding biology of
this species was studied by Bawa and Beach ( 1 983).
Standley ( 1 938) listed the common names anileto,
azulillo, coralillo, palo camaron, pissi, zorillo, and
zorillo real, and he cited Pittier for the Indian
names pili-tso (Guatuso) and tsus-krd (Brunka).

Hamelia patens variety patens has the leaves
sparsely to densely villous (especially on the lower
surface), and the flowers are sparsely to densely
villous externally. Variety glabra Oersted has the
leaves glabrous above and sparsely villous or pu-
berulent on the veins beneath, and the flowers are
usually glabrous externally. There seem to be no
ecological or geographic distinctions between the
varieties, and they may be no more than glabrous
and puberulent forms found within the same pop-
ulations.

Hamelia rovirosae Wernham, J. Bot. 49: 211. 1911.
Figure 34.

Shrubs or slender treelets to 5(-10) m tall, leafy
branchlets 0.8-3.5 mm diam., with curved or crooked
multiccllular hairs 0.3-1 mm long and often in longi-
tudinal rows, with 4 longitudinal ribs and quadrangular
in early stages but becoming terete and glabrescent; stip-
ules 2-6(-8) mm long, with a very short (1 mm) broad
base and long narrow linear awn, puberulent, caducous
or persisting with the leaves. Leaves usually 3/node (rare-
ly opposite), petioles 3-14(-20) mm long, 0.5-1 mm
thick, pubescent with short crooked hairs; leaf blades
(2.5-)5-15 cm long, (1.5-)2-6 cm wide, elliptic-oblong
to elliptic-ovate, elliptic or elliptic-obovate, apex acute
to short-acuminate, base acute to attenuate and slightly
decurrent on petiole, drying thin-chartaceous, glabrous
to sparsely pubescent above, sparsely to densely villous
beneath with thin straight or crooked hairs 0.2-0.5 mm
long, with denser tufts of hairs (domatia) in the vein
axils, 2 veins 3-7 /side and weakly loop-connected near

margin. Inflorescences 4-12 cm long, equally broad, pe-
duncles to 3 cm long, villous, floral rachis with dichot-
omous distal branches bearing 2-8 flowers, bracts to 0.5
mm long, caducous, flowers sessile or short (1-2 mm)
pedicellate. Flowers with hypanthium 2-4.5 mm long,
villous with curly hairs ca. 0.5 mm long, calyx lobes 2-
4 mm long and enlarging in fruit, 1 mm wide and oblong,
villous; corolla narrowly tubular at anthesis, reddish or-
ange to bright red or dark red, tube 16-22 mm long, 2-
3 mm diam., minutely villous with hairs ca. 0.5 mm
long (often in longitudinal rows), lobes 1-2 mm long, 1-
1.5 mm broad at the base, ovate; stamens with filaments
7-9 mm long, anthers 10-12 mm long, slightly exserted,
connective appendage ca. 0.5 m long; stigmas connate,
2-3 mm long. Fruit 8-14 mm long, 4-8 mm diam.,
cylindrical to oblong-ellipsoid, red becoming black, cov-
ered with small (ca 0.5 mm) crooked hairs, persisting
sepals to 6 mm long and 2 mm broad; seeds ca. 1 mm
long.

Shrubs of Caribbean lowland evergreen forest
formations, from near sea level to 200 m elevation.
Flowering in February, April, and July-October
in Central America. The species ranges along the
Caribbean lowlands, from Tabasco, Mexico, to
Bocas del Toro, Panama.

Hamelia rovirosae is recognized by the unusual
crooked multicellular hairs, corollas narrowly tu-
bular at anthesis, puberulent fruit with prominent
calyx lobes, and restriction to the Caribbean low-
lands. This species is frequently confused with H.
patens.

Hamelia xerocarpa Kuntze, Rev. gen. pi. 1: 284.
1 89 1 . H. costaricensis Standl., Contr. U.S. Natl.
Mus. 20: 207. 1919. H. panamensis Standl., loc.
cit. 208. 1919. H. rowlei Standl., J. Wash. Acad.
Sci. 15: 7. 1925. H. storkii Standl., loc. cit. 7.
1925. Figure 42.

Shrubs or small trees to 5 m tall, leafy branchlets 1.5-
5 mm thick, with 4 longitudinal ridges and quadrangular
in cross-section, glabrous to sparsely pilose with stiff
erect hairs to 0.9 mm long; stipules 6-1 3(-l 7) mm long,
cuspidate and often with 2 small lateral teeth, glabrate
or pubescent along the edge, drying black. Leaves usually
3 or 4/node, petioles (10-15)-85 mm long, about 1 mm
broad, glabrous to densely pubescent; leaf blades 8.5-
1 7(-37?) cm long, 3.5-9(-l 4?) cm broad, ovate to broad-
ly elliptic-oblong or ovate-rotund, apex acute to acu-
minate, base obtuse to cuneate and decurrent on petiole,
leaves drying thin-chartaceous, glabrate above, usually
minutely papillate-puberulent or with straight or curved
hairs 0.2-0.9 mm long beneath, 2 veins (7-)9-13(-18)/
side and loop-connected near the margin, 3 veins often
subparallel, domatia present or absent. Inflorescences
terminal on short lateral branches (and apparently ax-
illary), 5-15 cm long and equally wide, peduncles 1-
2.5(-5) cm long, the dichasia to 6(-l 5) cm long and with
3-9(-26) flowers along 1 side, glabrous or with yellowish

160

FIELDIANA: BOTANY

hairs to 1 mm long, bracts 0.6-0.8 mm long, the secund
flowers sessile or subsessile. Flowers with hypanthium
2-3 mm long, glabrous to pilose, calyx lobes 0.5-2 mm
long, ovate, glabrous to pilose; corolla funnelform (but
narrowly tubular until anthesis), yellow, tube 1 8-36 mm
long, 4-7 mm diam., glabrous to densely puberulent with
hairs 0.2-0.4 mm long, lobes 2-5(-8) mm long, 2-5 mm
broad at the base, ovate and acute; stamens with fila-
ments 8-16 mm long, anthers 10-16 mm long, connec-
tive little ( 1 mm) extended distally; style ca. 20 mm long,
stigmas 5, to 5 mm long. Fruits 11-14 mm long and 3-
5 mm diam., oblong to ovoid-oblong; seeds 0.5-1 mm
long.

Plants of evergreen lowland formations and gal-
lery forests in deciduous areas, from near sea level
to 800 m elevation. Flowering in May-December
(throughout the year in Panama). The species
ranges from Nicaragua to northern Colombia.

Hamelia xerocarpa is recognized by the broader
corolla tube (at anthesis) with longer corolla lobes,
the unusual yellowish puberulence (when present),
the lowland evergreen habitat, and the leaves usu-
ally with many secondary veins and often with
long petioles. Unfortunately, few herbarium col-
lections exhibit the broader funnelform corollas
in anthesis, and it is easy to misidentify this spe-
cies. This species may be difficult to separate from
H. macrantha. The breeding biology was studied
by Bawa and Beach (1983).

Hamelia xerocarpa variety xerocarpa is distin-
guished by the conspicuous hairs on many parts
of the plant and the leaves with 11-18 pairs of
secondary veins. Variety costaricensis (Standl.)
Elias is recognized by the lack of pubescence and
leaves with 9-12 pairs of secondary veins.

1 1 illia Jacquin

REFERENCE C. M. Taylor, Revision of Hillia
subg. Ravnia (Rubiaceae: Cinchonoideae). Sel-
byana 11: 26-34. 1989.

Shrubs, small trees, or lianas, epiphytic or less often
terrestrial, branchlets thick and terete, glabrous; stipules
interpetiolar and intrapetiolar but splitting apart along
the edges. Ungulate and blunt at apex, caducous. Leaves
opposite, equal or unequal at each node, subsessile to
short-petiolate; leaf blades elliptic to obovate, entire,
decurrent on the petiole, semisucculent and drying co-
riaceous, without domatia. Inflorescences of solitary ter-
minal flowers (or 3-flowered dichasia in H. thflora), bracts
reduced or absent (the flowers at first enclosed within
the large untied stipules), pedicels short or absent. Flow-
ers bisexual, monomorphic, often large, glabrous exter-
nally, hypanthium continuous with the pedicel, calyx
tube often absent, calyx lobes 2-5 and distant (or none),
sometimes with a secondary smaller set of calyx lobes
alternate with the larger lobes; corolla salverform to fun-
nelform or tubular, white to yellowish, pinkish, orange,
or red, semisucculent, corolla lobes (3-)5-7(-9), con-
volute in bud, becoming reflexed; stamens (4-)5-7, fil-
aments very short and inserted below the throat, anthers
basifixed, elongate-linear, obtuse at each end, included
(except in H. longifilamentosa); ovary 2-locular, ovules
many and ascending in each locule on scptal placentas,
style as long as the corolla tubes, stigmas subcapitate or
2. Fruits woody capsules, narrowly cylindrical to very
narrowly oblong, truncated distally, dark brown and
smooth, dehiscing septicidally and basipetally into 2 flat-
tened valves; seeds many and imbricated, rhombic and
flattened, with a minute circumferential wing, appen-
daged at the base and with a tuft of hairs at the distal
apex.

Hillia is a genus of about 20 species, ranging
from southern Mexico to Brazil and Peru. The
genus is recognized by the larger flowers with long
tubes, many ascending imbricated ovules, long tu-
bular ("cigar-shaped") capsules, and flattened
winged seeds with a tuft of hairs at one end. The
more colorful flowers of subgenus Ravnia (see fol-
lowing key) appear to be an adaptation to bird
pollination. Specimens lacking flowers or fruit can
be very difficult to identify to species. In addition,
one group of our species may be part of a poly-
morphic complex; see the discussion under H.
maxonii. Some of our species of Hillia are very
similar to Cosmibuena, but that genus has seeds
lacking the tufted hairs at one end.

Key to the Species of Hillia

la. Corolla pinkish to red or orange, funnelform or tubular with the central part of the tube inflated

and narrowed at both ends, flowers not scented; subgenus Ravnia .

Ib. Corolla greenish or white, long-tubular with rotate lobes or salverform, corolla tube expanded only
at the corolla lobes, flowers often sweet-scented; subgenus Hillia . 4

2a. Corolla tube rose red and inflated in the middle, 3-5 cm long, flowers often in groups of 3; a

commonly collected species " triflora

2b. Corolla tube yellow-orange to peach or rose, funnelform and widest at the mouth, 4-6.5 cm

long, flowers solitary; rarely collected species ....
3a. Corolla lobes 6-12 mm long; free portion of the filaments ca. 1 mm long H. allenii

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

161

3b. Corolla lobes 13-17 mm long; free portion of the filaments 9-10 mm long

H. longifilamentosa

4a. Plants of lower elevation forest, not known from above 300 m in Costa Rica; larger leaf blades

more than 10 cm long and 5 cm broad 5

4b. Plants of higher elevations, rarely collected below 600 m; larger leaf blades rarely more than 1 cm

long or 5 cm broad 6

5a. Leaf blades with thin texture, the 5-6 pairs of 2 veins arising at ca. 60 angles from the midvein;
corolla tubes 8-10 cm long H. macrophylla

5b. Leaf blades thick-textured, the 4-5 pairs of 2 veins arising at ca. 30 angles from the midvein;

corolla tubes 4-5 cm long H. grayumii

6a. Leaf blades 4-10 cm long, usually tapering gradually to a bluntly acute apex, broadest at the middle

or below; corolla tubes 4-5 cm long; seed hairs ca. 1 7 mm long H. loranthoides

6b. Leaf blades 0.7-3.7 cm long, usually bluntly obtuse to rounded at the apex; corolla tubes rarely

exceeding 4 cm in length; seed hairs ca. 10 mm long 7

7a. Leaf blades 12-37 mm long, stipules 6-16 mm long; corolla lobes suborbicular, corolla tubes 24-

42 mm long; seeds ca. 3 mm long; 1400-2400 m elevation H. maxonii

7b. Leaf blades 7-14 mm long, stipules 3-7 mm long; corolla lobes broadly to narrowly ovate, corolla

tubes 15-35 mm long; seeds ca. 2 mm long; 600-1600 m elevation H. panamensis

Hillia allenii C. M. Taylor, Selbyana 1 1: 32. 1989.
Ravnia panamensis Steyerm., Ceiba 3: 22. 1 952,
not Hillia panamensis Standl.

Epiphytic shrubs to 1.5 m tall, leafy stems 2.5-7 mm
thick, glabrous, terete; stipules 10-12 mm long, Ungulate,
quickly caducous. Leaves with short (14 mm), thick (2-
3 mm) petioles, glabrous and drying dark; leaf blades 4-

1 1 cm long, 1.5-5 cm broad, elliptic to narrowly elliptic-
oblong, apex slender acuminate, base cuneate or slightly
rounded, drying subcoriaceous, dark brown above, gla-
brous above and below, 2 veins 5-9/side, usually ob-
scure. Inflorescences of solitary terminal flowers borne
on short (34 mm) thick (2 mm) glabrous pedicels drying
black. Flowers glabrous externally, hypanthium 4-7 mm
long and 3.5 mm thick, calyx tube not developed, calyx
lobes 6, 6-14 mm long, 2-3 mm broad, narrowly spat-
ulate-oblong; corolla funnelform, pale red and pale yel-
low to salmon-pink, tube 25-40 mm long, lobes 6, 6-

1 2 mm long and 1 mm broad at the base, bluntly acute;
anthers 5-7 mm long on filaments ca. 1 mm long. Fruits
not seen.

Plants of montane cloud forest formations at ca.
1000 m elevation. Flowering in June and Septem-
ber. This species known only from the Cordillera
de Tilaran and in western Panama.

Hillia allenii is recognized by the brightly col-
ored funnelform corolla and the short filaments.
Fruiting material may be very difficult to distin-
guish from that of H. longifilamentosa.

Hillia grayumii C. M. Taylor, Selbyana 12: 137.
1991. Figure 28.

Epiphytic shrubs, ca. 1 m tall, leafy stems 4-7 mm
thick, slightly quadrangular, glabrous, brownish, mi-

nutely grooved; stipules ca. 40 mm long, 6-8 mm broad,
lanceolate, caducous. Leaves isophyllous, usually decus-
sate, petioles 6-20 mm long, 2-3 mm thick, articulated
at the stem; leaf blades 9-19 cm long, 3-6.5 cm broad,
elliptic to elliptic-oblong, apex acuminate with tip 1-2
cm long, base obtuse or acute, drying stiffly chartaceous
or subcoriaceous and dark brown above, glabrous above
and below, 2 veins 4-6/side and strongly ascending (ca.
30), obscure beneath. Inflorescences of solitary terminal
flowers, subtending stipules caducous, peduncles 2-3 mm
long, bracts 1-3 mm long, triangular, acute. Flowers gla-
brous, hypanthium 7-8 mm long, cylindrical, calyx limb
to 0.5 mm long, truncate or slightly lobed; corolla tu-
bular- funnelform, bright pale green to yellow, tube 43-
50 mm long, lobes 6, 8-9 mm long, triangular, obtuse
to rounded; filaments ca. 10 mm long, anthers 6, ca. 9
mm long. Fruits 10-12 cm long and 8-14 mm broad,
drying dark brown, stipe ca. 3 mm long; seeds 3 mm
long, 0.5 mm broad.

Plants of lowland rain forest and swamp forest
formations, collected from near sea level to 600
m elevation. Flowering in May-June; fruiting in
March and May-June. The species is known only
from the Caribbean lowlands of northern and cen-
tral Costa Rica.

Hillia grayumii is distinguished from its con-
geners by the lowland habitat, yellowish funnel-
form flowers, and large fruit. Among Costa Rican
species, it is similar to H. macrophylla, but that
species grows in cloud forests and has thin-tex-
tured leaves and tubular flowers.

Hillia longifilamentosa (Steyerm.) C. M. Taylor,
Selbyana 1 1: 32. 1989. Ravnia longifilamentosa
Steyerm., Ceiba 3: 21. 1952.

162

FIELDIANA: BOTANY

Epiphytic or clambering shrubs to 6 m tall, leafy stems
2-7 mm thick, glabrous, smooth, brown to gray; stipules
22-37 mm long, elliptic to oblanceolate, caducous. Leaves
with petioles 2-8 mm long, thick, glabrous; leaf blades
6-16 cm long, 2-7 cm broad, elliptic or elliptic-oblong,
apex acute or slightly acuminate, base obtuse to cuneate,
drying subcoriaceous, grayish green, glabrous above and
below, 2 veins 4-6/side, strongly ascending. Inflores-
cences of solitary terminal flowers, pedicles ca. 1 mm
long, bracts 2-3 mm long, triangular, acute. Flowers with
hypanthium 4-10 mm long, obconic to ellipsoid, calyx
lobes 6, 6-14 mm long, narrowly triangular to Ungulate
or oblanceolate; corolla tubular-rotate, orange-red to
salmon-pink, or white marked with pink, tube 32-43
mm long, ca. 3 mm diam., lobes 6, 13-17 mm long,
triangular to Ungulate; stamens 4-6, anthers ca. 5 mm
long, well exserted, dark green. Fruits 10 cm long.

Plants of montane cloud forest formations, 1 1 00-
1700 m elevation. Flowering in April, July-Au-
gust, and November. This species is known only
from a few collections, ranging from Zarcero, Ala-
juela, to Chiriqui, Panama.

Hillia longifilamentosa is distinguished by its
solitary terminal flowers with anthers extended 8-
10 mm beyond the throat of the tube. Compare
H. allenii.

Hillia loranthoides Standl., J. Wash. Acad. Sci.
18: 165. 1928. Figure 27.

Epiphytic shrubs, 0.4-1.5 m tall but with vining
branches, leafy stems 3-8 mm thick, glabrous, grayish,
terete or quadrangular; stipules 18-22 mm long (to 35
mm beneath the flowers), 6-10 mm broad, oblong-ob-
ovate, obtuse or rounded distally. Leaves closely clus-
tered or distant, decussate, petioles 4-8(-15) mm long,
1.5-2.8 mm thick; leaf blades (3-)4-10 cm long, (1 .3-)2-
4.5 cm broad, elliptic to elliptic-oblong, ovate or ovate-
lanceolate, apex bluntly acute or obtuse, base obtuse to
cuneate, drying coriaceous, dark grayish, 2 veins 4-57
side, strongly ascending or obscure. Inflorescences of sol-
itary terminal sessile flowers, subtended by a pair of
enlarged (8-25 mm) oblong-obovate bract-like reddish
stipules, 1-2 smaller (1-4 mm) pairs of triangular or
Ungulate bracteoles often present at the base of the short
pedicel. Flowers 6-7 cm long, hypanthium ca. 4 mm
long, calyx tube minute, calyx lobes 4, 8-10 mm long,
1-2 mm broad, linear-Ungulate; corolla tubular with ro-
tate lobes, white or cream, carnose, tube 40-60 mm long,
2-5 mm diam., lobes 4, 10-25 mm long, 8-16 mm broad,
rounded distally. Fruits 3-8 cm long, 7-10 mm diam.;
seeds ca. 2.5 mm long and 0.7 mm thick, distal hairs 6-
17 mm long.

Plants of evergreen montane forests, from
(300-)700 to 1400 m elevation. Flowering in Feb-
ruary-May; fruiting in January and May. In Costa
Rica, this species is known only from the Cordil-
lera de Tilaran, near San Ramon, and El Retiro

(Cartago) and from a single lower elevation col-
lection on the Osa Peninsula. The species is also
known from southern Mexico.

Hillia loranthoides is distinguished by its thick
leaves, usually tapering gradually to the apex, larg-
er flowers with slender tubes, and seeds with longer
bristles. Its distribution is unusual and may be an
artifact of the difficulty of collecting epiphytes.

Hillia macrophylla Standl., Publ. Field Columb.
Mus., Hot. Ser. 7: 201. 1931. Figure 28.

Epiphytic shrubs or vines, 3-7 m tall, leafy branchlets
3-7 mm thick, glabrous, pale brown and lenticelate; stip-
ules 10-35 mm long, 12-25 mm broad, narrowly lan-
ceolate (in Costa Rica) to oval-oblong, the basal sheath
1-2 mm long, caducous. Leaves decussate, petioles 6-
20 mm long, 2-3.5 mm thick, terete, clearly differenti-
ated from the lamina base; leaf blades 9-2 1 cm long, 6-
10 cm broad, ovate-elliptic to elliptic-oblong or elliptic,
apex acuminate (acute), base obtuse, drying membra-
naceous to chartaceous, dark brown above, glabrous
above and below, major veins with a rugose texture. 2
veins 6-9/side arising at ca. 60 angles and loop-con-
nected near the margin. Inflorescences of large solitary
sessile terminal flowers, subtended by elongated (20-50
x 5-24 mm) caducous stipules, bracteoles usually ab-
sent. Flowers with hypanthium 6-7 mm long, 3-3.5 mm
diam., cylindrical, calyx tube 0-0.5 mm long, subentire;
corolla tubular-salverform, white, tube 5.5-1 1 cm long,
2-4.5 mm diam., lobes 5 or 6, 2.5-5 cm long. ca. 8 mm
broad at the base and narrow (4 mm) distally, linear-
lanceolate to narrowly triangular. Fruits 7-12 cm long,
the opened valves becoming 14-20 mm broad, rounded
at the base, acute at apex, sessile; seeds 1.5-4 mm long
with distal hairs 14-18 mm long.

Plants of moist cloud forests from 800 to 1 800
m elevation. Flowering in March-June; a single
fruiting collection was made in December. This
species is known only from near Monteverde, Ca-
taratas de San Ramon, and above the Rio Gato,
Cartago, in Costa Rica. It is also found in Colom-
bia, Ecuador, and Peru.

Hillia macrophylla is unique among our species
of Hillia because of the larger thin-textured leaves,
more numerous ascending secondary veins, and
long-tubular flowers with corolla lobes that have
long (3-4 cm), narrow tips. It is a rarely collected
species. South American collections appear to have
larger (3-4 mm) seeds and larger stipules sub-
tending the flowers.

Hillia maxonii Standl., J. Wash. Acad. Sci. 18:
163. 1 928. H.palmana Standl. Joe. cit. 18: 164.
1928. H. hathewayi Fosberg, Sida 2: 387. 1966.
Figure 27.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

163

Epiphytic shrubs, 0.7-2. 5(-5) m tall, often pendant to
5 mm long, leafy stems 1-5 mm thick, dark or pale
grayish, glabrous, older nodes articulate with transverse
ridges; stipules 6-16(-32) mm long, 2-6 mm broad, ob-
long-obovate to obovate, rounded, larger and bract-like
beneath the flowers. Leaves decussate and often crowded
on short (3-15 mm) internodes, petioles 2-6(-15) mm
long; leaf blades (12-)18-37(-60) mm long, 6-20(-30)
mm broad, obovate to oblong-obovate or ovate-elliptic,
apex bluntly obtuse to rounded, base obtuse to cuneate,
decurrent on the petiole, drying subcoriaceous and dark
grayish or brownish, glabrous above and below, 2 veins
3-4/side, strongly ascending or obscure. Inflorescences
of solitary, terminal, sessile flowers subtended by a pair
of enlarged bract-like stipules rounded at apex. Flowers
glabrous externally, fragrant, hypanthium ca. 2.5 mm
long and 1 .8 mm thick, calyx tube very short, calyx lobes
absent or 4-6 mm long, 0.5-1.8 mm broad; corolla tu-
bular with rotate lobes, slightly fleshy, tube 24-42 mm
long, 1.4-4 mm diam., greenish or white, lobes 4, 10-
14(-22) mm long and usually equally broad or broader
than long, suborbicular and rounded distally, bright white
or yellowish white. Fruits (22-)30-60 mm long, 5-9 mm
thick, the valves to 8 mm broad when opened and ex-
panded; seeds 3-4 mm long, 0.5-1 mm diam., distal
hairs 6-13 mm long.

Plants of evergreen montane forest formations
from 500 to 2400 m elevation. Flowering in April-
September and December; fruiting in September
and November-March. This species has been col-
lected near Managua, Nicaragua, in the Cordillera
de Tilaran, Cordillera Central, in the western part
of the Cordillera de Talamanca, and in Chiriqui
and Veraguas, Panama.

Hillia maxonii is recognized by its usually epi-
phytic habit, smaller stiff leaves, bract-like stipules
subtending the flowers, and tubular corollas with
broadly rounded lobes. Smaller-leaved specimens
of this species may represent intermediates with
H. panamensis. Larger-leaved specimens resem-
ble H. tetrandra Sw. of the West Indies and north-
ern Central America. However, H. tetrandra has
larger (5-1 1 x 2.5-6 cm) leaves that are more
often obovate, quite unlike those of southern Cen-
tral America. All three taxa exhibit a wide range
of variation, and it is possible that they are ele-
ments of a single polymorphic species. (A similar
problem is found in the epiphytic species of Psy-
chotria; see the discussion under P. guadalupen-
sis.)

Hillia panamensis Standl., N. Amer. Fl. 32: 117.
1921. H. chiapensis Standl., J. Wash. Acad. Sci.
16: 16. 1926. Figure 27.

Epiphytic shrubs or small treelets to 4 m tall, leafy
stems 1-2.7 mm thick, glabrous and grayish, often de-

veloping opposite longitudinal sulci and expanded nodes;
stipules 3-7 mm long, 1-2 mm broad, oblong to narrowly
obovate and rounded distally, translucent. Leaves de-
cussate and usually closely crowded on short (0.5-5 mm)
internodes, petioles 1-4 m long; leaf blades 7-12(-14)
mm long, 4-8(-10) mm broad, elliptic to elliptic-oblong
or obovate, apex acute to bluntly obtuse or rounded,
base cuneate to obtuse, drying stiffly chartaceous to sub-
coriaceous, grayish to very dark, margins often becoming
slightly revolute, glabrous above and below, 2 veins 2-
3/side, strongly ascending or obscure. Inflorescences of
solitary terminal flowers, the distal stipule pair slightly
expanded (6 x 2.5 mm) and bract-like smaller (1-2 mm)
rounded bracts sometimes present at the base of the
sessile flowers. Flowers glabrous externally, hypanthium
ca. 1.5 mm long, 1 mm thick, tubular, calyx lobes to 1 1
mm long, 0.3-0.6 mm broad, linear, caducous; corolla
tubular with rotate lobes, white, slightly succulent, tube
15-30(-38) mm long, 0.9-1.5 mm diam., lobes 4, 5-7
mm long, 3.5-6 mm broad, ovate to lanceolate, bluntly
obtuse. Fruits 20-42 mm long, opened expanded valves
3-4 mm wide; seeds ca. 2 mm long and 0.6 mm thick,
distal hairs 10 mm long.

Plants of evergreen montane forest formations
from 600 to 1 600 m elevation. Flowering in May-
August; fruiting in December. This species is found
in southern Mexico-Guatemala, in the Cordilleras
de Guanacaste and Tilaran in Costa Rica, and in
western and central Panama.

Hillia panamensis is distinguished by its epi-
phytic habit, very small stiffclosely crowded leaves,
fragrant flowers with long slender corolla tubes,
and rotate lobes usually narrower than long. It has
been called jasmin del volcdn. This species may
not be specifically distinct from the very similar,
and partly sympatric, H. maxonii. But the differ-
ences used in the keys do seem to separate a great
majority of specimens. See the discussion under
H. maxonii.

Hillia triflora (Oersted) C. M. Taylor, Selbyana
1 1: 30. 1989. Ravnia triflora Oersted, Vidensk.
Meddel. Dansk Naturhist. Foren. Kjobenhavn
1852: 49. 1853. Ravnia pittien 'Standl., N. Amer.
Fl. 32: 1 14. \92l.LagenanthusparviflorusE\van,
Mutisia 4: 5. 1952. H. triflora var. pittieri
(Standl.) C M. Taylor, Selbyana 11:31. 1989.
Figure 27.

Epiphytic (rarely terrestrial) shrubs, 0.5-1 .5(-3) m tall
(branches to 2 m long and pendulous), leafy stems 1.8-
5 m thick, glabrous; stipules 15^43 mm long, 4-10 mm
broad, largest beneath the inflorescences, elliptic and
acute, glabrous and caducous. Leaves usually decussate,
isophyllous or anisophyllous at a node, petioles 3-7(-l 2)
mm long, 1.5-2 mm thick, poorly differentiated from
the base; leaf blades 5-1 3(-l 9) cm long, 1 .5-4.5 cm broad,
narrowly elliptic-oblong to oblanceolate, elliptic-oblong

164

FIELDIANA: BOTANY

or oblong (linear-lanceolate in variety pittieri), apex ta-
pering gradually and narrowly acuminate, tapering grad-
ually to the acute or cuneate base, drying coriaceous,
dark green or grayish green above, 2 veins 3-5/side.
Inflorescences of 1 or 3 terminal flowers borne on a very
short (0.5-2 mm) thick peduncle, bracts ca. 3 mm long
or absent, pedicels 1-6 mm long but difficult to distin-
guish from the hypanthium. Flowers glabrous externally,
hypanthium ca. 5 mm long, 1.2-1.8 mm diam. calyx
lobes 6 (5, 7), 4-14 mm long, 0.5-1 mm broad, linear-
oblong; corolla red, tubular with relatively small lobes,
tube 40-65 mm long, 1.5-10 mm broad, lobes 6 (5, 7),
3-4 mm long, to 5 mm broad at the base; stamens 5-6,
anthers 3-5 mm long, partially exserted. Fruits 5-10 cm
long, ca. 8 mm diam., valves becoming up to 12 mm
broad; seeds 0.8-2.5 mm long, distal hairs 15-30 mm
long.

Plants of montane and premontane evergreen
forest formations from (100-)800 to 2400 m ele-

vation. The lowest elevation collections are from
La Selva; this species has not been collected from
below 1000 m on the Pacific slope. Flowering in
all months, but most commonly February-Au-
gust; fruiting in October-February. The species
ranges from southern Mexico to northwestern Co-
lombia.

Hillia triflora is distinguish by its epiphytic hab-
it, the tubular reddish flowers usually in terminal
triads, long narrow capsules, and seeds with a dis-
tal tuft of hairs. There are few fruiting collections.
There may be some local differentiation between
the two varieties; at Monteverde, variety pittieri
is found at slightly lower elevations than is variety
triflora. The following key distinguishes the two
varieties.

la. Leaves at each node approximately equal in size at the same node on flowering shoots; mature
corolla tube not or only slightly inflated, up to 1.6 times broader in the broadest portion as at the
apex H. triflora var. triflora

Ib. Leaves at each node strongly differing in size on flowering shoots, larger leaves ca. 1.5 times longer
than the other leaf of the same node; mature corolla tube strongly inflated, ca. 1 .8 times broader
in center than at the apex H. triflora var. pittieri

Hippotis Ruiz Lopez & Pavon

Trees or shrubs, puberulent (in ours) or glabrous; stip-
ules interpetiolar, triangular to obovate, large and ca-
ducous. Leaves opposite, decussate, petiolate; leaf blades
usually thin, pinnately veined, without domatia, with
the minor venation parallel (lineolate) within the areolae
formed by the 3 and 4 veins. Inflorescences axillary to
leaves or undeveloped leaves and axillary in fruit, short
cymose and 2-3-flowered, capitate or of solitary flowers.
Flowers bisexual, monomorphic, radially symmetrical
or bilaterally symmetrical by curvature of the corolla,
usually large, sericeous externally, calyx bilobate or spa-
thaceous; corolla tubular or funnelform, white, rose, red,
or pale orange, corolla tube straight or curved, corolla
lobes 5, short and truncated or emarginate, plicate-val-
vate in bud; stamens 5, filaments usually unequal, borne
at the middle or lower half of the tube, anthers included;
ovary 2-locular, placentas peltate from the center of the
septum, with many horizontal ovules in each locule,
stigmas 2, subcapitate. Fruits baccate, ovoid to ellipsoid
or globose; seeds many, small, angular.

A genus of around 12 species in northern South
America, Ecuador, and Peru; 1 species reaches
northernmost Costa Rica. The larger thin hairy
leaves, unusual lineolate minor venation, few larg-
er pseudoterminal flowers, spathaceous calyx, and
hairy fleshy fruit with many small seeds help to
distinguish our species.

Hippotis albiflora Karst., Fl. Colomb. 1: 33, pi.
1 7. 1 858. Duroia panamensis Dwyer, Ann. Mis-
souri Bot. Gard. 55: 38. 1968. Figure 25.

Small trees, 3-17 m tall, leafy stems 2.5-8 mm thick,
densely pilose with yellowish hairs 1-3 mm long, terete
or slightly quadrangular, older stems glabrescent and pale
grayish; stipules 15-30 mm long, 5-12 mm broad, nar-
rowly ovate or lanceolate, with long yellowish hairs ex-
ternally, leaves with petioles 10-28(-35) mm long, 1.5-
2.5 mm thick, densely pilose with erect or ascending
hairs 1-2 mm long; leaf blades 14-30(-35) cm long, 6-
14(-18) cm broad, obovate to oblong-obovate, apex ta-
pering gradually or abruptly and acuminate or caudate-
acuminate, tip 1-2 cm long, base obtuse to rounded and
subtruncate, drying thin-chartaceous to chartaceous,
densely pilose on both surfaces with thin straight or curved
hairs 0.7-2 mm long, 2 veins 7-10/side, 3 veins often
subparallel, the smallest veins distinctly parallel (lineo-
late) in small groups. Inflorescences of 1-3 flowers in the
axils of the distal leaves but often appearing terminal,
peduncles absent or very short, bracts 3-5 mm long,
ovate, pedicels 3-8 mm long, 1-2 mm thick and densely
yellowish velutinous. Flowers 5-6 cm long, hypanthium
and calyx 3.3-3.8 cm long, ca. 8 mm broad, densely
pilose, spathe-like and split ca. 10 mm down 1 side;
corolla ca. 5 cm long and 2 cm broad distally, funnel-
form, white to cream, pilose externally, tube 35-45 mm
long, greenish white, straight or slightly curved, lobes 5-
8 mm long and 10 mm broad, obtuse. Fruits 3-4 cm
long (not including the persisting calyx), 1-2 cm diam.,

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

165

ellipsoid and hirsute, becoming purple; seeds 2 mm long,
foveolate.

Plants of rain forest formations in the Caribbean
lowlands, from 10 to 400 m elevation (to 1600 m
in Venezuela). Flowering in February, April, and
July-October; fruiting in May-September. This
species has been collected near Rio Colorado in
northern Costa Rica, from La Selva and the Hitoy
Cerere reserve. It is also known from the province
of Bocas del Toro, in Panama, and from Colombia
and Venezuela.

Hippotis albiflora is recognized by the conspic-
uous pubescence, larger thin leaves with lineolate
minor venation, few large pseudoterminal flowers,
spathaceous calyx, and larger, hirsute, many-seed-
ed fruit. Our material may differ in some details
from that described from Venezuela; it could be
that Costa Rican populations are worthy of sub-
specific recognition. Compare this species with
Duroia costaricensis.

Hoffmannia Swart/

Herbs, herbaceous subshrubs, or slender shrubs, main
vertical stems usually with few or no lateral branches,
glabrous or pubescent, terete or quadrangular, internodes
often hollow when dried; stipules interpetiolar, usually
small and triangular, often succulent and divergent, ca-
ducous. Leaves opposite and decussate or rarely verti-
cillate, equal at the node or sometimes slightly unequal,
petiolate or occasionally subsessile, petioles with inflated
chambers (ant vesicles) in a few species; leaf blades usu-
ally drying membranaceous to chartaceous (rarely sub-
coriaceous), entire, often decurrent on the petiole, pin-
nately veined, often with conspicuous raphides on the
dried leaf surfaces, domatia rarely present. Inflores-
cences axillary, sessile or pedunculate, with few to many
flowers, usually cymose or dichasial and often with heli-
coid branches, fasciculate to capitate in a few species,
flowers usually pedicellate, bracts and bracteoles rarely

present. Flowers bisexual and monomorphic, glabrous
or pubescent externally, hypanthium often with longi-
tudinal ribs, calyx tube usually very short, calyx lobes
4(-5), usually short and often triangular, persisting; co-
rolla salverform to funnelform or rotate, white, yellow,
orange, or rose to deep red or purple, corolla tube short
and usually narrow, glabrous on the interior, corolla lobes
4(-5), imbricate along the edges or apparently valvate,
apex usually acute; stamens 4 (3-5), borne within the
tube of the corolla, filaments very short, anthers linear
to narrowly oblong, usually white; ovary 2- (3, 4) locular,
each locule with many ovules borne longitudinally on
bilamellate axile placentas, style slender, stigmas bilo-
bate or clavate. Fruits baccate, usually small, 2- (3, 4)
locular, often becoming enlarged and spongy at maturity,
with many minute multiseriate horizontal seeds; seeds
angulate, their surfaces often reticulate to foveolate.

A genus of about 1 00 species, ranging from Mex-
ico and the West Indies to South America. A ma-
jority of the species are found in Mexico and north-
ern Central America. The usually single-stemmed
habit, presence of raphides in many parts, small
triangular stipules (often caducous), axillary inflo-
rescences, four-parted flowers, and many-seeded
fleshy fruits help to distinguish the species of Hoff-
mannia from other Rubiaceae in Costa Rica. Most
of the Costa Rican species are restricted to areas
with very high rainfall (none are found in decid-
uous vegetation), and relatively few species grow
below 800 m elevation.

Hoffmannia is probably Costa Rica's taxonom-
ically most difficult genus of Rubiaceae. These
semisucculent subshrubs do not dry well, and the
flowers, often in small dense clusters, are also not
well preserved. In addition, there appears to be
great variation from plant to plant within many
species. The genus is related to Hamelia and Dep-
pea; it is currently being studied by Dr. John Dwyer
(MO), and the review presented here should be con-
sidered no more than tentative.

Key to the Species of Hoffmannia

la. Petioles with lateral longitudinal inflated vesicles; plants to 1.2 m tall, surfaces with longer (1.5
mm) multicellular hairs H. vesiculifera

Ib. Petioles without lateral inflated structures; plants 0.5-3 m tall, glabrous to villous with multicellular
hairs 2

2a. Inflorescences sessile and capitate in the leaf axils in early stages (rarely becoming pedunculate or
branched in later stages), often verticillate at the nodes 3

2b. Inflorescences sessile and fasciculate to pedunculate and paniculate or cymose, neither densely

capitate nor verticillate 4

3a. Leaf blades neither bullate nor areolate, to 30( 40) cm long, glabrous or pubescent; corolla
ca. 7 mm long; commonly collected plants H. congesta

166

FIELDIANA: BOTANY

3b. Leaf blades bullate or areolate, to 20(-25) cm long, pubescent; corolla ca. 4.5 mm long; rare

H. areolata

4a. Leaves usually 3/node, leaf blades cuneate at the base with long-decurrent lateral margins on the
petiole and these slightly expanded near the stem, lateral margins of the petiole 2-4 mm broad
[leaf blades with 1 1-18 pairs of 2 veins]; Vara Blanca to the upper Rio Grande de Orosi, 1400-

1600 m elevation H. amplexifolia

4b. Leaves opposite (rarely with 3 leaves per node), leaf blades obtuse to cuneate at the base, if long-
decurrent on the petiole never forming a lateral margin 2-4 mm wide along the petiole and not

expanded near the base; species widely distributed, 10-2300 m elevation 5

5a. Plants of Cocos Island 6

5b. Plants of mainland Central America 7

6a. Petioles 2-5 cm long; corolla glabrous on the exterior H. piratarum

6b. Petioles 6-8 cm long; corolla puberulent on the exterior H. nesiota

7a. Young stems glabrous or subglabrous, or with very small (0.2 mm) hairs in longitudinal lines along

the young stern 8

7b. Young stems densely pubescent with crooked multicellular hairs, the hairs varying 0.3-2 mm long,

older stems pubescent or glabrescent 19

8a. Inflorescences short, the 1 peduncle and rachis usually less than 3 cm long (if plants grow
below 600 m elevation and have petioles 2-6 cm long and leaves 4-15 cm broad, go to

dichotomy 12) 9

8b. Inflorescences small to large, peduncle and rachis of some inflorescences usually exceeding

4 cm in length 12

9a. Corolla tubes 3-4 mm diam. and 3-4 mm long; leaf blades 15-35 cm long with 9-15

pairs of 2 veins, cuneate basally and long-decurrent on the petiole H. dolae

9b. Corolla tubes 1-3 mm diam. (dried) and 1-6 mm long; leaf blades 5-15 cm long with
5-9 pairs of 2 veins, abruptly narrowed at the base (but leaves to 23 cm long, with 7-

12 pairs of 2 veins, and cuneate long-decurrent bases in H. hamelioides) 10

lOa. Corolla 9-14 mm long, corolla lobes 3-6 mm long; calyx lobes ca. 1.5 mm long, ovary
2-3 mm long; flowers fasciculate (inflorescences rarely pedunculate), usually more than

1 5/node; fruit elongate when dried H. psychotriifolia

lOb. Corolla less than 9 mm long, corolla lobes 1.5-4 mm long; calyx lobes 0.5-1 mm long,
ovary 1-2 mm long; flowers fasciculate or borne on slender pedunculate; fruit rounded

when dried 11

1 la. Inflorescences compact, fasciculate or with short stiff peduncles, often less than 2 cm
long; leaf blades to 1 5 cm long, rarely tapering gradually at the base and decurrent on

the petiole; a commonly collected species H. longipetiolata

lib. Inflorescences open with thin peduncles and pedicels, usually more than 2 cm long;
leaf blades to 23 cm long, usually tapering gradually at the base and decurrent on the

petiole; uncommon plants H. hamelioides

12a. Sepal lobes 2-4 mm long; corolla tube 1-3 mm long, corolla lobes 3-7 mm long; plants not
collected from about 700 m elevation [leaf blades usually with an arcuate submarginal vein

and abruptly narrowed at the base] H. liesneriana

12b. Sepal lobes rarely more than 1 .8 mm long; corolla tube usually more than 2 mm long (except
in H. laxa), corolla lobes 2-6 mm long; plants rarely found below 800 m elevation (except

H. pallidiflord) 13

1 3a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein; in-
florescences with a prominent erect peduncle 4-10 cm long and usually terminated by 3

branches; 1200-1500 m elevation H. davidsoniae

13b. Secondary veins loop-connected only in the distal part of the lamina, not forming an arcuate
submarginal vein; inflorescences often pendant, with shorter slender peduncles (400-)800-

2300 m elevation -I 4

14a. Corolla tubes 4-8 mm long; inflorescences short to long and with many lateral branches; leaf
blades drying stiffly chartaceous and often large, 12-37 cm long with 9-15 pairs of 2 veins;
1000-2200 m elevation ! 5

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 167

14b. Corolla tubes 1 .5-4 mm long; inflorescences short (to 6 cm) with few lateral branches [corolla
tube 1.54 mm long]; leaf blades drying membranaceous to thin-chartaceous, 7-20(-25) cm

long with 7-10 pairs of 2 veins; (400-)800-1 500 m elevation 17

1 5a. Leaves usually drying grayish and stiffly chartaceous, petioles 2-10 cm long; leaf blades
cuneate and slightly decurrent on the petiole; (corolla tubes 4-8 mm long); fruit rose-
red to white; central Costa Rica H. leucocarpa

1 5b. Leaves usually drying dark brown and chartaceous, petioles 2-5 cm long; leaf blades
usually cuneate and long-decurrent at the base; fruit red; central Costa Rica to the

Chiriqui Highlands (note: the following 2 species may intergrade) 16

16a. Corolla tubes 6-10 mm long; calyx lobes 1.2-3 mm long; leaf blades often narrowly
obovate and long attenuate at the base; central Costa Rica to Panama, 1000-2500 m

elevation H. arborescens

16b. Corolla tubes 3-7 mm long; calyx lobes 0.5-1.5 mm long; leaf blades not usually
narrowly obovate and long attenuate at the base; westernmost Costa Rica and the

Chiriqui Highlands, 1 100-1 700 m elevation H. pittieri

1 7a. Inflorescences usually unbranched (sometimes bifurcate) [drying yellowish and not very slen-
der, probably erect; pedicels 0-3 mm long and ca. 0.5 mm thick (dried); corolla tube 2.54
mm long; 400-1300 m elevation on the Caribbean slope] H. pallidiflora

1 7b. Inflorescences usually branched 18

18a. Inflorescences with very thin (0.2-0.3 mm) branches that usually dry black, pendulous,
flowering portion 24 cm long, pedicels 4-9 mm long and 0.2-0.3 mm thick (dried); 1000-

1 500 m elevation H. laxa

18b. Inflorescences with thicker branches that do not dry black, flowering portion 1-2 cm long,

pedicels 2-6 mm long, not thin and black; 0-200 m elevation, Osa Peninsula

H. hammelii

1 9a. (from 7b) Leaf blades rounded to bluntly obtuse at the apex, often bullate, many 4 veins parallel
(sublineolate but difficult to see); inflorescences with few (3-7) subsessile flowers on long (3-7 cm)

peduncles; plants to 40 cm tall [200-1200 m elevation] H. bullata

1 9b. Leaf blades acute to acuminate (rarely obtuse) at the apex, never bullate, 4 veins partly parallel
only in H. aeruginosa; inflorescences with short (0-2 cm) peduncles or with more than 7 flowers

if peduncles are long, pedicels 0-2 cm long; plants usually more than 40 cm tall 20

20a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein [inflores-
cences subsessile and usually less than 2 cm long] 21

20b. Secondary veins loop-connected only in the distal part of the blade, a submarginal vein absent

(except in the distal third) 22

2 la. Leaf blades cuneate and slightly decurrent at the base, with 9-1 5 pairs of 2 veins; 400-1000

m elevation H. inamoena

21b. Leaf blades acute to obtuse at the base, not usually decurrent on the petiole, with 7-12 pairs

of 2 veins; 1000-1500 m elevation in north-central Costa Rica H. aeruginosa

22a. Plants not found above 900 m elevation; leaf blades usually less than 15 cm long, rarely cuneate

and long-decurrent at the base, with 6-10 pairs of 2 veins H. valerii

22b. Plants not found below 1 000 m elevation; leaf blades often exceeding 1 5 cm in length, usually

gradually cuneate and long-decurrent on the petioles, with 7-12 pairs of 2 veins 23

23a. Inflorescences with thick densely villous peduncles and pedicels; calyx lobes 1.5-3 mm long; corolla

tube 6-8 mm long (lobes 3-4.5 mm long) H. asclepiadea

23b. Inflorescences with slender glabrescent or sparsely puberulent peduncles and pedicels; calyx lobes
0.3-1.5 mm long; corolla tubes 1.5-4 mm long (note: the following 2 species may be synonymous)

24a. Inflorescences short but open and few-flowered; corolla lobes mostly 24 mm long

H. hamelioides

24b. Inflorescences short but crowded and the flowers clustered around the node; corolla lobes 3-7 mm
long H. decurrens

168 FIELDIANA: BOTANY

Hoffmannia aeruginosa Stand!. , Publ. Field Mus.
Nat. Hist., Hot. Ser. 18: 1313. 1938. Figure 9.

Herbaceous subshrubs or slender shrubs, 0.5-2.5(-4)
m tall, with few branches, leafy stems 2-6 mm thick,
quadrangular or rounded, densely pilose with reddish
brown crooked hairs 0.2-0.7 mm long; stipules 1-2 mm
long, 2-3 mm broad at the base, broadly triangular, vil-
lous, persisting. Leaves with petioles (10-)20-70 mm
long, 0.8-2 mm thick, densely villous, often with petioles
of the same node differing in length; leaf blades (6-)9-
20 cm long, (3-)4-10 cm broad, elliptic-oblong to ovate-
elliptic or broadly elliptic, apex acuminate or short-acu-
minate to acute, base obtuse to acute (not or only slightly
decurrent on the petiole); drying thin-chartaceous, gray-
ish or grayish brown above, glabrous above, puberulent
beneath with short (0.1-0.2 mm) hairs on 2 veins and
longer (0.5-2 mm) hairs on the midvein, 2 veins 7-12/
side and evenly loop-connected 2-4 mm from the mar-
gin, many 4 veins parallel (forming rectangular areolae).
Inflorescences solitary and axillary (2/node), 1-2 cm long
with 2-10 flowers, glomerulate, sessile or on short (3
mm) peduncles, unbranched, flowers sessile or subses-
sile. Flowers densely covered with curly-crooked hairs
0.3-0.5 mm long, dark reddish brown when dried, hy-
panthium 2.5-3 mm long, 2 mm diam. distally, obconic,
calyx lobes 1 .3-2.5 mm long, ca. 0.6 mm wide, narrowly
oblong; corolla rotate, reddish, tube 1-2 mm long, lobes
4-6 mm long, 2-3 mm broad; anthers ca. 2.5 mm long;
style and stigma white. Fruits not seen.

Plants of wet cloud forest formations on the
Caribbean slope from (?100-)800 to 1600 m ele-
vation. Flowering in April-August. The species is
known from north-central Costa Rica, near Ciu-
dad Quesada and Zarcero, and from western Pan-
ama.

Hoffmannia aeruginosa is recognized by the
dense reddish (in life) multicellular hairs on all
parts, long-petiolate leaves (with blades scarcely
decurrent), small dense inflorescences, and re-
stricted range. The petioles often unequal at a node,
the subparallel minor venation and arcuate sub-
marginal vein are also distinctive features. Com-
pare this species with material placed under H.
inamoena with less dense pubescence, especially
on the petioles.

Hoffmannia attinis Hemsley, Diagn. PI. Nov. 31.
1879.

TYPE Endres 150, without locality.

Branches terete, puberulent when young. Leaves
with petioles ca. 6 mm long; leaf blades 10-12.5
cm long, ovate-oblong, obtusely acuminate, atten-
uate at the base, minutely puberulent beneath. In-

florescences umbellate-cymose, fasciculate, with
ca. 6 flowers, peduncles 8-16 mm long, slender,
pedicels 2-4 mm long. Flowers 6-8 mm long, pu-
berulent. Based on Standley (1938), who had only
seen the description.

Hoffmannia amplexifolia Standl., J. Wash. Acad.
Sci. 15: 8. 1925. Figure 8.

Herbs or subshrubs, l-2(-3) m tall, leafy stems 3-10
mm thick, glabrous or minutely puberulent on the ridges
in early stages, with 3 longitudinal ridges; stipules 1-2
mm long and 2 mm broad at the base, triangular, pu-
berulent along the edge. Leaves 3 (4)/node, petioles ab-
sent or short (3 mm) when the decurrent leaf base is
reduced to a lateral ridge, minutely puberulent beneath;
leaf blades (11-) 16-37 cm long, 3-13 cm broad, nar-
rowly elliptic to oblanceolate or very narrowly elliptic,
gradually narrowed to the long-acuminate apex, gradu-
ally narrowed to the cuneate and long-decurrent base
with a margin 2-4 mm wide along the petiole, often
expanded (to 6 mm) and auriculate at the base, leaves
drying chartaceous and dark brown to grayish above,
glabrescent above, minutely (0.1-0.2 mm) puberulent
below, 2 veins 11-1 8(-22)/side, usually loop-connected
1-4 mm from the leaf edge. Inflorescences axillary to
leaves, 2-12/node, to 10 cm long, with short (3-15 mm)
or long (to 60 mm), thick (1.5-2 mm) peduncles and
long (to 7 cm) slender lateral branches from the apex of
the peduncle, minutely puberulent, pedicels 0-5 mm long
with hairs 0. 1-0.4 mm long. Flowers sparsely to densely
puberulent with minute crooked hairs that dry reddish,
hypanthium 2-3 mm long, 1.5-2 m diam. distally, tur-
binate to obconic, calyx reddish, calyx cup ca. 0.7 mm
deep, calyx lobes 1-2.5 mm long and 0.5-1 mm broad,
triangular to oblong; corolla rotate, greenish white to pale
yellow, tube 1-4 mm long, 1-2 mm diam., lobes 4-9
mm long, 1.5-3 mm broad; anthers 3-5 mm long and
0.7-1.5 mm broad, connivent, yellow. Fruits becoming
pink or red, to 8 mm diam. in life (4-5 mm long dried),
ellipsoid to subglobose.

Plants of the very wet forests of the Caribbean
slope, 800-2000 m elevation. Flowering in March-
July; fruiting in October-December and March.
The species is known only from central Costa Rica
(Vara Blanca de Sarapiqui to Tapanti).

Hoffmannia amplexifolia is recognized by the
nodes with three leaves, long-decurrent leaf base
forming a leafy lateral margin along the petiole
and often slightly expanded (auriculate) at the base
(appearing to be amplexicaul), distinctive puber-
ulence (but not on the stems), much-branched in-
florescences, and broad connivent anthers. Al-
meda el al. 6665 (CAS, F) from near Las Alturas,
Puntarenas, is tentatively placed here; it has very
pubescent inflorescences with corolla tubes 5-6

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

169

mm long. Pressed specimens can be similar to H.
subauriculata with two leaves per node.

Hoffmannia arborescens J. D. Smith, Bot. Gaz.
37: 417.1 904. H.josefina Standl., J. Wash. Acad.
Sci. 15: 8. 1925. (?//. steinworthii Fosberg, Sida
2: 388. 1966.) Figure 10.

Herbaceous subshrubs, 1.5-2 m tall, leafy stems 2-6
mm thick when dried, glabrescent; stipules ca. 2 mm
long. Leaves with petioles 3-8(-25) mm long; leaf blades
7-18 cm long, 2.5-7 cm broad, oblanceolate to narrowly
obovate, base acute to attenuate and long-decurrent on
petiole, glabrous above and below, drying stiffly char-
taceous, often dark reddish brown above, 2 veins 6- 127
side. Inflorescences 1-3 in each leaf axil (2-6/node), 3-
5 cm long, to 4 cm broad, open cymose with many
branches, peduncles 3-20 mm long, puberulent with stiff
hairs, flowers crowded distally, pedicels 2-6 mm long.
Flowers with hypanthium 2-3 mm long, ca. 2 mm diam.,
calyx lobes 1-3 mm long, 1 mm broad at the base, blunt
at apex; corolla funnelform, white to yellow or flushed
with red, glabrous or puberulent, tube 6-8(-l 0) mm long,
1-2 mm diam., lobes 3-6 mm long. Fruits ca. 8 x 4.5
mm, red.

In evergreen montane forests from 1 600 to 2500
m elevation. Flowering in February, April-Au-
gust, and November-December; fruits were col-
lected in February (immature fruits in August).
This species ranges from Costa Rica to western
Panama.

Hoffmannia arborescens is recognized by the
usually narrowly obovate leaves gradually long-
attenuate at the base, flowers on short inflores-
cences in leaf axils (or below), larger flowers with
prominent calyx lobes, long corollas, and the high-
er altitude range. Occasional individual plants may
be puberulent (Q. Jimenez 231 CR). This species
may resemble H. hamelioides, while large-leaved
specimens may be confused with the more com-
mon and closely related H. leucocarpa. The dif-
ferences between H. arborescens and H. pittieri are
minor, and the two species may intergrade in the
highlands of western Panama, where H. pittier is
common and distinctive.

This species was described from material orig-
inating from Sta. Rosa del Copey at ca. 1 800 m
elevation (Tonduz 12230 = J. D. Smith n. 8121
us holotype). An isotype labeled as H. arborescens
at CR lacks any indication of having the leaf-like
floral bracts 1 cm long mentioned in the original
description. This may have been an error on the
part of J. D. Smith, as no such bracts have been
seen in any Costa Rican material of Hoffmannia.
The type bears the same Herb. Nat. C.R. number

(12230) as does the type of H. decurrens Standl.
(q.v.), and they probably represent a mixed col-
lection of two different species.

Hoffmannia areolata Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 22: 52. 1940. Figure 9.

Herbaceous subshrubs or slender stemmed shrublets,
0.6-1 .5(-2) m tall, leafy stems 2-9 mm thick, glabrescent
or sparsely pilose with crooked hairs 0.3-0.7 mm long;
stipules 3-6 mm long and 2-4 mm broad at the base,
glabrous, ligulate, caducous. Leaves with petioles 3-
40(-90) mm long, difficult to distinguish from the de-
current lamina base; leaf blades (6-)8-20(-30) cm long,
3-8(-12) cm broad, elliptic to ovate-elliptic or oblan-
ceolate, apex tapering gradually and acute or short-acu-
minate, more abruptly narrowed to an obtuse or cuneate
base and long-decurrent on petiole, drying membrana-
ceous to stiffly chartaceous, dark greenish above, the
surface distinctly bullate and with short (0.2-0.5 mm)
scabrous hairs above, with thinner hairs ca. 0.3 mm long
beneath, 2 veins 1 1-16/side. Inflorescences verticillate
in leaf axils, glomerulate, to 2 cm long and 8-15 mm
broad, sessile, often obscuring the node, with pink hairs,
the flowers sessile or subsessile. Flowers closely con-
gested within the inflorescence, reddish tomentulous, ca-
lyx lobes 3-5 mm long and 0.5 mm broad, greenish to
magenta; corolla funnelform, white, rose or translucent,
ca. 4.5 mm long; tube ca. 2.5 mm long with lobes ca. 2
mm long; anthers ca. 1.5 mm long. Fruits 4-8 mm long,
3-6 mm diam. (to 10 mm in life), ellipsoid to subglobose,
rose to deep red, spongy.

A little-collected species of montane cloud forest
formation, 900-2100 m elevation. Flowering in
March-December; fruiting in April, July-Septem-
ber, and January. The species ranges from the Cor-
dillera de Tilaran eastward to the Chiriqui High-
lands of Panama.

Hoffmannia areolata is recognized by its rugose-
bullate leaves (areolate beneath), densely flowered
axillary sessile glomerulate (usually verticillate) in-
florescences, very small flowers, and hairs with
thickened bases. The leaves are very dark green
above in life. In general aspect, these plants re-
semble H. congesta and some Psychotria spp. with
axillary inflorescences.

Hoffmannia asclepiadea Standl., J. Wash. Acad.
Sci. 15: 7. 1925. Figure 10.

Slender herbaceous shrubs, 1.5-2(-3) m tall, usually
unbranched, leafy stems 2.5-7 mm thick, sparsely to
densely pilose with crooked hairs 0.3-0.5 mm long (?rarely
glabrous as in Williams el al. 28096 CR, F); stipules 2-4
mm long and 3 mm broad at the base, triangular, pu-
berulent, caducous. Leaves with petioles 1.5-5 cm long,

170

FIELDIANA: BOTANY

1-3 mm thick, densely to sparsely puberulent; leaf blades
6-21 cm long, 4-9(-ll) cm broad, elliptic to elliptic-
oblong or ovate-elliptic, apex acuminate with tip 1-2 cm
long, base rounded to obtuse to cuneate-decurrent, dry-
ing chartaceous, dark brown above (much paler be-
neath), glabrous or sparsely pubescent above with hairs
to 0.7 mm long, more densely pubescent beneath with
crooked reddish hairs, 2 veins 9-12/side, often loop-
connected near the margin. Inflorescences 2/node, open
cymose, 2-4 cm long, with many closely spaced flowers
on pubescent peduncles 3-10 mm long and ca. 0.7 mm
thick, usually with 3 major branches and small (0.6 mm)
bracteoles, pedicels 1-3 mm long, short villous. Flowers
pubescent, with 5 distal longitudinal lines of hairs on the
unopened buds, hypanthium 1-1.5 mm long, calyx lobes
0.5-1 mm long, narrowly triangular; corolla funnelform,
pale yellow, tube 6-8 mm long, ca. 1.5 mm diam., lobes
2-3(-4-5?) mm long, to 2 mm broad. Fruits said to be
subglobose and red.

Plants of wet cloud forest formations from 1 900
to 2300 m elevation. Flowering in April-July. The
species is known only from along the Cordillera
Central and western portion of the Cordillera de
Talamanaca in central Costa Rica.

Hoffmannia asclepiadea is recognized by the
dense puberulence on leaves and inflorescences,
compact many-flowered inflorescences on very
short peduncles, longer corolla tubes, and restric-
tion to higher-elevation cloud forests. This species
may be no more than a very puberulent form of
another species, such as H. arborescens.

6 mm long and 2 mm broad; anthers ca. 3.2 mm long.
Fruits becoming red or orange-red, 6-10 mm long and
4-9 mm diam., subglobose but with a truncated apex
and persisting calyx.

Plants of wet evergreen forest formations from
50 to 1200 m elevation in Costa Rica. Sometimes
forming a ground cover under the deep shade of
the forest. Flowering in April-August; fruiting Jan-
uary-July in Costa Rica. This species ranges from
Veracruz, Mexico, to central Panama.

Hoffmannia bullata is recognized by its smail
stature with roots at lower nodes, prominently bul-
late leaves that are obtuse or rounded apically,
few-flowered inflorescences with long peduncles,
and puberulence on many parts. The sublineolate
4 venation is distinctive but difficult to see in
some collections. The midvein is often white on
the dark green upper surface in living material.
This species displays much variation within its
range and within Costa Rica. Collections from the
Pacific slope and near the Panama border tend to
have smaller stature and narrowly elliptic-oblong
leaves with more deeply impressed venation and
acute apices (fig. 9, uppermost left). These differ-
ences are worthy of more detailed study. In ad-
dition, H. discolor (Lemaire) Hemsl. of Veracruz,
Mexico, is closely related, but the leaves dry darker,
the flowers have longer (2-4 mm) linear calyx lobes,
and the fruits are larger.

Hoffmannia bullata L. O. Williams, Fieldiana Bot.
36: 52. 1973. Figure 9.

Herbs or herbaceous subshrubs, 10-60 cm tall, erect
or decumbent, stems unbranched, lower nodes with roots,
leafy stems l-4(-7) mm thick, densely pilosulous with
curved hairs 0.1-0.3 mm long drying reddish brown,
terete; stipules 2-3 mm long, triangular, deciduous. Leaves
opposite, petioles 4-45(-55) mm long, 1-2.5 mm thick,
pilosulous with curved or crooked multicellular hairs;
leaf blades 4-22 cm long, 2-9(-12) cm broad, obovate
to oblong-obovate, apex rounded to bluntly obtuse, base
obtuse to cuneate (not or only slightly decurrent on pet-
iole), drying chartaceous, grayish or grayish brown above,
glabrous above, minutely puberulent on the veins be-
neath with curved reddish hairs ca. 0.2 mm long, 2 veins
5-12/side, many 4 veins subparallel (sublineolate) and
demarking narrow rectangular areas on the upper dried
surface (but often difficult to see). Inflorescences usually
solitary in leaf axils ( 1 -2/node), 3-8 cm long, glomerulate
or subumbellate, peduncles 2-7 cm long, 0.5-1 mm thick,
glabrous or sparsely pilose, with 3-9 flowers along the
same side of bifurcate branches, pedicels 1-5 mm long.
Flowers sparsely puberulent or subglabrous, hypanthium
2-3 mm long, 1.5-2 mm broad distally, urceolate or
turbinate, red, calyx lobes 1-2.5 mm long, 0.4 mm broad
and acute; corolla rotate, pale red or rose, 7-10 mm long
in bud, tube 3-4 mm long and 0.8 mm diam., lobes 3-

Hoffmannia congesta (Oerst.) Dwyer, comb. nov.
Xerococcus congestus Oerst., Vidensk. Meddel.
Dansk Naturhist. Foren. Kjobenhavn 1852: 52.
1853. Figure 9.

Herbs or subshrubs, 0.5-2(-3) m tall, stem erect and
usually unbranched, terete, leafy stems 3-15 mm thick,
glabrous or with a few large (1 mm) crooked hairs; stip-
ules 6-14(-20) mm long, 4-6 mm broad at the base,
triangular-ovate, with or without an awn, glabrous, co-
riaceous, deciduous. Leaves with petioles 2-8(-15) cm
long, 1.2-3 mm broad, glabrous or with crooked mul-
ticellular hairs to 2 mm long; leaf blades 15-30MO) cm
long, 8-15(-25) cm broad, broadly elliptic to broadly
ovate-elliptic or elliptic-obovate, apex short-acuminate,
somewhat rounded to abruptly narrowed at the obtuse
to acute base and slightly decurrent on petiole (rarely
long-decurrent or sometimes rounded to a truncated base
in large leaves), drying stiffly chartaceous, dark brown
or dark grayish above, glabrous or with scattered crooked
hairs, 2 veins 7-14(-17)/side. Inflorescences solitary in
leaf axils (2/node), sessile and usually glomerulate, many-
flowered, 1-2 cm long and to 6 cm broad (across the
node), subglobose to verticillate, reddish, pedicels 0-2.5
mm long. Flowers tightly congested, sparsely and mi-
nutely puberulent, hypanthium 3 mm long and 1-2 mm

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

171

diam.. calyx red, calyx lobes 3-6 mm long, 1-1.5 mm
broad, obtuse at the tip, with parallel venation; corolla
salverform, white, tube 2-3.5 mm long, lobes 2-3.5 mm
long and 1 mm broad. Fruit becoming white and spongy,
ca. 5 x 3 mm when dried, oblong.

Understory plants of the wet Caribbean slopes
from 700 to 1800(-2400?) m elevation. Flowering
in January-October; fruiting in October-March.
The species ranges from the Cordillera de Tilaran
eastward to Veraguas, Panama.

Hoffmannia congesta is characterized by its ses-
sile (often verticillate) glomerulate reddish inflo-
rescences, long calyx lobes, white fruit, un-
branched stems, large leaves, and prominent
stipules. This is one of our most commonly col-
lected and most distinctive species of Hoffmannia.
In the past it was assigned to its own genus (Xero-
coccus); specimens are often found in the uniden-
tified section of Rubiaceae collections.

Hoffmannia davidsoniae Standl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 22: 53. 1946. (?= H. lan-
cistigma Dwyer, Ann. Missouri Bot. Gard. 56:
279. 1969). Figure 7.

Herbs or slender subshrubs, 0.5-1 .7 m tall, stems usu-
ally unbranched, leafy stems 1.5-5 mm thick, glabrescent
(rarely densely puberulent, cf. Davidse et al. 29175 CR);
stipules 1.5-3 mm long, ca. 1 mm broad at the base,
narrowly triangular, caducous. Leaves with petioles 1-7
cm long, 0.8-2 mm thick, glabrous; leaf blades 8-17 cm
long, 3-9(-12) cm broad, broadly oblong to elliptic-ob-
long or ovate-oblong (rarely lanceolate), apex narrowed
abruptly and short-acuminate, base obtuse to acute and
slightly decurrent on petiole, drying grayish to dark brown
or dark grayish green above, glabrous above and below
or minutely puberulent, 2 veins 10-14/side, sometimes
arising from the midvein at nearly 90, loop-connected
distally and forming an arcuate submarginal vein 1-4
mm from the edge. Inflorescences borne on lower leafless
nodes, 1-2/node, 5-15 cm long, paniculate and with a
prominent peduncle 4-6(-12) cm long and 1-2.3 mm
thick, glabrous, with 3(-4) distal branches and each branch
with 7-17 flowers, pedicels 2-4(-6) mm long. Flowers
glabrous, hypanthium 2-3 mm long, ca. 1 mm diam.
distally, turbinate, calyx lobes 0.3-1 mm long, triangular;
corolla rotate to fimnelform, yellow or flushed with pur-
ple, tube 1-4 mm long, lobes 2-4(-6) mm long; anthers
to 3.8 mm long. Fruit ca. 8 mm long and 4 mm diam.,
ellipsoid to subglobose, becoming red.

Plants of wet cloud forest formations at 700-
2300 m elevation. Flowering in January, April,
July, and September; fruiting in September and
December-January. The species ranges from cen-
tral Costa Rica eastward to the Chiriqui High-
lands.

Hoffmannia davidsoniae is a little-collected spe-
cies characterized by the long stiff peduncles usu-
ally terminated by three branches and the long-
petiolate leaves with an arcuate submarginal vein.
This species is poorly known in Costa Rica, and
there appears to be great variation from collection
to collection. The secondary veins often arise at
nearly 90 from the midvein, but there are collec-
tions in which this is not the case. This species is
closely related to H. liesneriana.

Hoffmannia decurrens Standl., Contr. U.S. Natl.
Herb. 20:205. 1919.

Slender shrubs or subshrubs, 1.5-3 m tall, leafy stems
1.5-4 mm thick, densely puberulent with crooked yel-
lowish brown or reddish brown hairs 0.1-0.3 mm long;
stipules 1.5-3 mm long, ca. 2 mm broad at the base,
triangular, hirstulous, deciduous. Leaves with petioles
5-40 mm long, ca. 1 mm thick, difficult to distinguish
from the lamina base, densely hirsutulous with hairs 0.3-
0.5 mm long; leaf blades 7-21 cm long, 2-7 cm broad,
elliptic-obovate to oblanceolate or elliptic, apex acu-
minate, gradually narrowed to a narrowly cuneate base
and long-decurrent on petiole, drying membranaceous
or thin-chartaceous and dark greenish brown or grayish
green above, glabrous on the upper surface, pubescent
on the veins beneath with hairs 0.2-0.7 mm long, 2
veins 7-10/side. Inflorescences 1-3/axil, l-2(-3)cm long,
cymose or fasciculate with few (1-5) flowers or rarely
2-branched and scorpioid with 5-10 flowers, peduncles
5-10(-24) mm long, 0.2-0.3 mm thick, puberulent, ped-
icels 2-6 mm long, slender. Flowers sparsely to con-
spicuously pubescent, hypanthium 0.7-1 .3(-2) mm long,
0.6-1(-1.7) mm diam. distally, turbinate, calyx lobes
0.7-1.5 mm long, triangular; corolla yellow or whitish,
tube 2-4 mm long, lobes 3-7 mm long, ca. 2 mm wide
at the base. Fruits becoming 8-9 mm long, 5-6 mm
diam. (dried), ovoid to globose, red, borne on slender
peduncles and pedicels.

Plants of montane evergreen forest formations,
1 100-2000 m elevation. Flowering in April-June
and December; fruiting in December-February.
This poorly characterized species is endemic to
Costa Rica and presently known to range from
Monteverde to Sta. Maria de Dota.

Hoffmannia decurrens is recognized by the pu-
berulent young stems, narrowly obovate leaves with
cuneate long-decurrent base, and very short inflo-
rescences with very slender peduncle and pedicels.
This species is poorly characterized at present, with
a diverse array of specimens placed here. This
species may include material presently assigned to
H. hamelioides. The type, Tonduz 12230 us, bears
the same collection number as the type of H. ar-
borescens; these types (not seen) probably repre-
sent a mixed collection.

172

FIELDIANA: BOTANY

Hoffmannia dotae Standl., J. Wash. Acad. Sci. 18:
181. 1928. H. ramonensis Standl., loc. cit. 180.
1928. Figure 10.

Single-stemmed shrubs or herbaceous subshrubs,
(0.5-)l-3(-6?) m tall, leafy stems 2-10 mm thick, gla-
brous to pilosulous, older stems terete and hollow, woody;
stipules 3-4 mm long and 4 mm broad at the base,
glabrous, deciduous. Leaves 2 (rarely 3) at a node, pet-
ioles 4-20 mm long (to 5 cm if including the decurrent
lamina base), ca. 1.5 mm broad, glabrous; leaf blades
1 5-30(-35) cm long, (4-) 7-1 2 cm broad, elliptic-obovate
to obovate-oblong or oblanceolate, apex short-acumi-
nate, tapering gradually or abruptly to the cuneate and
long-decurrent base, the narrowed basal part of the lam-
ina to 6 cm long, drying membranaceous to thin-char-
taceous, usually dark brown or gray above, glabrous
above, pubescent in very early stages beneath and be-
coming glabrous or with reddish brown hairs 0.1-0.4
mm long along the major veins, 2 veins 9-15/side and
often loop-connected near the edge. Inflorescences ax-
illary or at older leafless nodes, l-3/axil(2-6/node), 1.5-
3(-5) cm long, ca. 1 5 mm broad, usually paniculate with

3 primary branches and 4-9 flowers, peduncles 0-5 mm
long (to 35 mm when the primary peduncle is reduced
and the 1 branches function as peduncles), minutely
puberulent, pedicels 3-6 mm long. Flowers conspicu-
ously (0. 1-0.3 mm) puberulent or glabrous, hypanthium
3-4 mm long, 2-3 mm diam. distally, obconic and red-
dish, calyx tube ca. 0.5 mm long and rotate, lobes (l-)2-

4 mm long, 1-2 mm broad at the base, bluntly triangular
to oblong, often held horizontally; corolla rotate, usually
red or orange at the base and orange or yellowish distally,
fleshy and glabrous, buds ca. 10 mm long, tube 2-5 mm
long, 2.5-4 mm diam., lobes 4-9 mm long, 2-3 mm
broad, yellowish within. Fruits 6-8 mm long and 4-6
mm diam. (dried), ca. 8 mm diam. in life, orange or
reddish.

Plants of wet montane cloud forest formations,
from 850 to 2100 m elevation. Flowering in Feb-
ruary-September; fruiting in September-January.
It is uncommon in Costa Rica, except for the San
Vito area. This species ranges from the Cordillera
de Tilaran to the Chiriqui Highlands of Panama.

Hoffmannia dotae is recognized by its larger leaf
blades with long-decurrent base forming narrowly
winged margins along the sides of the petiole (but
not auriculate at the base), the often cauliflorus
few-flowered inflorescences, and the short thick
corolla tube with larger stiff yellow petals. The
flower buds with their short conical corolla tubes
and broad (3-4 mm) base distinguish this species
from all other Costa Rican species, none of which
have such thick corolla tubes. Compare H. leu-
cocarpa with more numerous slender flowers and
leaves acute to cuneate at the base.

HofTmannia gesnerioides (Oerst.) Kuntze, Rev. gen.
pi. 285. 1891. Ophryococcus gesnerioides Oerst.,

Vidensk. Meddel. Dansk. Naturhist. Foren.
Kjobenhavn 1852: 53. 1852.

This species ranges from Guatemala to central
Nicaragua at elevations of 1000-1 500 m. The small
stature (to 60 cm), long reddish hairs, and few-
flowered cymes on peduncles that elongate to 3
cm in fruit help to distinguish this species, which
is not known from Costa Rica.

HofTmannia hamelioides Standl., J. Wash. Acad.
Sci. 15: 8. 1925. Figure 11.

Herbs or shrubs, (0.5-)1.5-3 m tall, stems often
branched, leafy stems 1.2-3 mm thick, usually densely
covered with crooked reddish brown multicellular hairs
0.3-1 mm long (sometimes glabrescent); stipules 2-3
mm long, triangular, drying reddish brown. Leaves with
petioles 6-25 mm long, 0.7-1 .5 mm thick, usually dense-
ly puberulent; leaf blades 8-23 cm long, 3-8 cm broad,
oblanceolate-elliptic to narrowly obovate or elliptic-ob-
long, apex acuminate, base gradually narrowed and acute
or long-cuneate, decurrent on petiole, drying membra-
naceous to thin-chartaceous, dark brown to blackish
brown above, glabrous or pubescent beneath with crook-
ed hairs 0.2-0.7(-1) mm long, 2 veins 7-12/side. Inflo-
rescences axillary or at older nodes, solitary (1-2/node),
1-3 cm long, usually on short (1-10 mm) slender pe-
duncles, with 3-7 flowers, pedicels 0.7-1.5 mm long,
0.2-0.4 mm thick, minutely (0.1-0.2 mm) puberulent.
Flowers with short (0.2-0.7 mm) crooked hairs, buds ca.
8 mm long, hypanthium ca. 2 mm long and 1.5-2 mm
diam., calyx lobes 0.7-1.5 mm long, 0.3-0.6 mm broad,
narrowly oblong to linear; corolla yellowish white or
yellow, tube 1.5-2 mm long, 1.5-2 mm diam., lobes 2-
4 mm long, 1.5-2 mm broad, apex acute, becoming re-
flexed. Fruits 4-8 mm long, 3-6 mm diam. (dried) red-
dish at maturity, with persisting calyx lobes.

Plants of evergreen forests in the central high-
lands, from 450 to 2200 m elevation. Flowering
in February and May-July; fruiting in July and
December. This species is known only from the
Central Volcanic highlands and the Cordillera de
Talamanca in Costa Rica.

Hoffmannia hamelioides is recognized by the
usually puberulent young stems, narrowly oblan-
ceolate leaves gradually tapering to the long-at-
tenuate base on slender petioles, small inflores-
cences with filamentous peduncles and pedicels,
and short corolla tubes. Compare this species with
H. decurrens (perhaps synonymous) and H. valerii
with cuneate to truncated leaf bases and lower
elevation habitat. Glabrous individuals resemble
a number of species, especially H. longepetiolata.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

173

Hoffmannia hammelii C. M. Taylor, sp. nov.

Species Hoffmanniae laxae similis, sed ab ea stipulis
brevioribus (0.7-1.5 mm longis) ac cymis secundis dif-
fert; etiam H. pallidiflorae similis, sed foliis basi acutis
ac lobulis corollinis brevioribus (3-4 mm longis) differt.

TYPUS Liesner2907 (holotypus CR, isotypus MO), from
near Sirena, Corocovado National Park, 0-200 m alt.,
5 July 1977, Puntarenas, Costa Rica.

Herbaceous subshrubs, 0.3-0.5(-1) m tall, stems
branched only near the ground, leafy stems 1.3-3 mm
thick, glabrous and terete, often drying dark; stipules 0.7-
1.5 mm long, ca. 1 mm broad at the base, glabrous,
deciduous. Leaves with petioles 6-27 mm long, 0.7-1.4
mm broad, glabrous; leaf blades (4-) 7- 18 cm long,
(1.5->4-7 cm broad, elliptic to elliptic-obovate or ellip-
tic-oblanceolate, apex acuminate, base acute (not de-
current), drying stiffly chartaceous, grayish above, usu-
ally paler or yellowish beneath, glabrous above and below,
2 veins 7-13/side. Inflorescences solitary and axillary
to leaves (1-2/node), 2-7 cm long but the flowering por-
tion only 1-2 cm long, with a single rachis or cymose
with a terminal flower and 2 lateral branches with 5-12
flowers each, peduncles 1 5-50 mm long, rachis with 2-
7(-l 1) flowers along 1 side, glabrous, pedicels 2-6 mm
long. Flowers with ovary/hypanthium 2 mm long and
1 .5 mm diam., calyx tube 0.5 mm long, calyx lobes 0.5-
0.8 mm long, ca. 1 mm broad at the base, triangular;
corolla rotate, white to yellow-green, tube 2-3 mm long,
lobes 3-4 mm long. Fruits ca. 5 mm diam., ellipsoid to
subglobose, red, glabrous.

Plants of the evergreen Pacific lowlands, from
near sea level to 200 m elevation. Flowering in
July-September and November; fruiting in Jan-
uary. This species is endemic to the Golfo Dulce
area and the Osa Peninsula.

Hoffmannia hammelii is recognized by its small
stature, leaves tapering equally to apex and base,
small flower groups on long slender peduncles,
leaves often drying grayish or reddish brown, and
restricted geographic range. This species appears
to be part of a species group including H. laxa, H.
pallidijlora, H. bullata, and H. discolor (Lemaire)
Hemsl. of Mexico. The species is named in honor
of Barry Hammel, who has made many important
contributions to our knowledge of Costa Rica's
Rubiaceae. Other collections seen were M. Cha-
varria et al. 254 CR, Kernan 813 & 1249 CR, MO,
and Knapp 2184 CR, MO.

Hoffmannia inamoena Standl., J. Wash. Acad. Sci.
18: 179. 1928. H. fimbrianthera Dwyer, Ann.
Missouri Bot. Gard. 56: 277. 1969. Figure 11.

Herbaceous subshrubs. 1-2 m tall, with few branches,
leafy stems 2.5-5 mm thick, sparsely to densely minutely
(0.1-0.2 mm) puberulent in early stages but soon gla-

brous and drying grayish, quadrangular; stipules 2-3 mm
long. Leaves with petioles 10-60 mm long, 0.8-2 mm
thick, minutely and sparsely puberulent or glabrous; leaf
blades 8-22 cm long, 3.5-10 cm broad, elliptic-oblong
to oblong or obovate, apex acuminate with tip 3-1 5 mm
long, base obtuse to acute and usually slightly decurrent
on petiole, drying membranaceous or thin-chartaceous
and grayish green above, glabrous above, very minutely
(0.05 mm) puberulent on the veins beneath or glabrous,
2 veins 9-14/side and loop-connected near the margin.
Inflorescences axillary or on older leafless nodes (1-2/
node), ca. 1 cm long and 1-5 flowers, glomerulate, sessile
or with peduncles 1-3 mm long, pedicels 1-3 mm long,
pubescent with hairs 0. 1-0.4 mm long or glabrous. Flow-
ers with crooked reddish hairs to 0.3 mm long (rarely
glabrous), hypanthium ca. 2 mm long and 1.7 mm diam.
distally, urceolate, calyx lobes 1.3-2.3 mm long, nar-
rowly oblong to narrowly triangular, apex obtuse; corolla
rotate, deep red, puberulent, tube ca. 1.5 mm long, lobes
ca. 4 mm long. Fruits 5-7 mm long, subglobose, becom-
ing white, puberulent.

Plants of evergreen forest formations from 400
to 1000 m elevation. Flowering in January and
May-July; fruiting in June and November-De-
cember. The species ranges from Bijagua, Alajue-
la, eastward along the Caribbean slope to Rio Re-
ventazon, and in the General Valley of the Pacific
slope.

Hoffmannia inamoena is recognized by the
minute puberulence on many parts, secondary
veins clearly loop-connected near the margin of
the leaf, very small sessile inflorescences, and rel-
atively large narrow calyx lobes. The material
placed here is rather poor and may not represent
a single species. Holm and Iltis (58 F) noted that
both flowers and fruit fall easily from the stems.

Hoffmannia laxa Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 22: 1 16. 1940. Figure 11.

Herbaceous subshrubs or slender shrublets 1-2 m tall,
branched only near the base, leafy stems 1-6 mm thick,
glabrous; stipules 1-3 mm long, triangular, glabrous, de-
ciduous. Leaves with petioles 1 .5-6(-9) cm long, 0.8-2.2
mm broad, glabrous; leaf blades 7-18(-21) cm long, 4-
9.5(-ll) cm broad, elliptic to elliptic-ovate or ovate,
apex acuminate, tapering gradually or abruptly to the
cuneate or acute base and slightly decurrent on petiole,
drying thin-chartaceous, grayish green to dark brownish
green above, glabrous above and below, 2 veins 7-1 1/
side. Inflorescences 1-3 in each leaf axil (2-6/node), 3-
8 cm long, ca. 2 cm broad, pendant, open paniculate or
racemose with few (8-15) distant flowers, reddish in life,
peduncles 10-45 mm long, 0.2-0.8 mm thick, glabrous,
rachis often elongate with alternate flowers and minute
(0.5 mm) bracteoles, pedicels 4-9 mm long (to 16 mm
in fruit), 0.2-0.3 mm thick (dried). Flowers glabrous,
hypanthium ca. 2 mm long and 1.8 mm diam., subglo-
bose, calyx lobes 0.2-0.7 mm long, triangular; corolla

174

FIELDIANA: BOTANY

funnelform, white to pale green, tube l-2(-3) mm long,
lobes 3-5 mm long and 1 mm broad at the base; anthers
ca. 2 mm long. Fruits to 8 mm long, 4-5 mm diam.,
globose to ellipsoid, greenish white when immature,
whitish purple or red to bluish black or purple-black at
maturity, with a small persisting calyx tube; seeds black.

Plants of evergreen lower montane forest for-
mation on both the Caribbean and Pacific slopes,
200-1600 m elevation. Flowering in February-
June and August-October; fruiting in January and
March-October. This species is known only from
Monteverde eastward to the slopes of Volcan Poas
and in western Panama.

Hoffmannia laxa is recognized by the often long-
petiolate somewhat larger leaves, the lack of pu-
bescence, the open few-flowered inflorescences with
long thin filamentous (when dried) peduncles and
pedicels, and the small flowers with very short
corolla tubes. The name H. capillacea Dwyer has
been applied incorrectly to material of this species.

Hoffmannia leucocarpa Stand!.. J. Wash. Acad.
Sci. 1 5: 9. 1925. //. carpinteraeStendl, N. Amer.
Fl. 32: 199. 1934, nom. nov. for//, macrophylla
Standl., J. Wash. Acad. Sci. 15: 9. 1925, non //.
macrophylla Hemsl. (?= H. trichocalyx Standl.,
J. Wash. Acad. Sci. 18: 181. 1928.) Figure 10.

Slender shrubs or small treelets, l-2.5(-4) m tall, with
few or no lateral branches, leafy stems 2.5-8 mm thick,
glabrous (rarely ferruginous-villous), usually hollow;
stipules 1.5-4 mm long, 2-4 mm broad at the base,
triangular, glabrous, deciduous. Leaves with petioles 1.5-
7(-10)cm long, 1.3-3.8 mm broad, glabrous: leaf blades
14-24(-35) cm long, 5-12(-15) cm broad, elliptic to
broadly elliptic, apex tapering abruptly and acuminate,
tapering more gradually to the cuneate base and slightly
decurrent on petiole, drying chartaceous to stiffly char-
taceous, pale greenish brown to dark grayish green above,
glabrous above and below, 2 veins 11-14/side. Inflo-
rescences 1-3 /axil below the leaves (2-6/node), 4-8(-l 5)
cm long, with 1 or 2 orders of branching (rarely with a
single rachis), often pendant, peduncles 2-5(-8) cm long,
0.5-2 mm thick, usually terminated by 3 subequal 1
branches, glabrous or less often minutely puberulent,
pedicels 2-8(-10) mm long, sparsely to densely puber-
ulent or villose. Flowers with hypanthium ca. 2 mm long
and 1.5 mm diam., obconic, puberulent with thin hairs
ca. 0.2 mm long, calyx green to pink, orange or red, calyx
tube 1-2 mm long, lobes 0.5-2 mm long, bluntly tri-
angular; corolla rotate to funnelform, pale pink to yellow
marked with orange or red, glabrous to villosulous, tube
4-8 mm long, 1.5-2 mm diam., lobes (2-)3-7 mm long,
2 mm broad at the base; anthers ca. 4 mm long. Fruits
8-10 mm long and 7-8 mm diam., globose to obovoid
(ca. 1 cm diam. in life), often bisulcate in development,
white to pink or red.

Plants of the wet Caribbean cloud forests and
nearby areas, from 1000 to 2300 m elevation.
Flowering in February-July; fruiting in February,
April, June-July, and October-December. The
species ranges from the eastern part of the Cor-
dillera de Tilaran (Monteverde) to the western
slopes of Volcan Irazu in central Costa Rica (but
see below).

Hoffmannia leucocarpa is recognized by the
larger firm-textured leaves on long petioles, pe-
dunculate and branched (usually pendant) inflo-
rescences, and the larger flowers with corolla tubes
and lobes usually equal in length. The fruit can be
white or pink to red. This species differs from
larger-leaved specimens of H. longepetiolata by
the more rigid leaf blades with short-acuminate
apices, larger inflorescences, and larger corollas.
This species may intergrade with H. arborescens,
forming a complex that includes H. pittieri. Dried
specimens can be similar to H. dotae, but that
species has smaller inflorescences and much
broader and more succulent corolla tubes and the
leaves have winged petioles. Most material placed
here has branched inflorescences, but a few have
monopodial circinnate inflorescences; this is an-
other example of the great variability within Hoff-
mannia species and why determining species in
the genus is so difficult.

The preceding circumscription excludes a num-
ber of specimens from the San Vito area formerly
placed here. These are treated as unusual elements
of//, pittieri or H. arborescens. Hoffmannia tricho-
calyx probably represents an unusually pubescent
representative of this species; it is based on Stan-
dley & Torres 47690 (holotype), 47480, 47819. &
47924, all us.

Hoffmannia liesneriana L. O. Williams, Fieldiana
Bot. 36: 54. 1973. Figure 8.

Herbs or slender subshrubs, 0.3-1(-1.5) m tall, stems
usually unbranched, quadrangular with opposing flat and
sulcate surfaces, leafy stems 4-12 mm thick, glabrous
(strigillose): stipules 1-3 mm long, 3 mm broad at the
base, triangular, glabrous. Leaves with petioles (2-)6-l 3
cm long, 1.5-3 mm thick, glabrous; leaf blades (1 1-)13-
24 cm long, (6-) 7- 13 cm broad, broadly ovate-elliptic
to broadly elliptic-oblong, apex abruptly narrowed and
short acuminate, abruptly narrowed to an obtuse or
rounded base, drying chartaceous, dark grayish or gray-
ish brown above, glabrous above, sparsely to densely
minutely (0. 1-0.2 mm) puberulent on the veins beneath,
2 veins (7-)10-14(-l 7)/side, usually loop-connected near
the margin. Inflorescences borne at older leafless nodes
or near the ground, 1-4/node, 2-11 cm long, with many
flowers in 1 -several dense clusters or on helicoid branch-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

175

es but rarely expanded, reddish to reddish purple, pe-
duncles 4-50 mm long, glabrous or sparsely puberulent,
pedicels 0-4 mm long, slender. Flowers subglabrous or
puberulent with hairs ca. 0.2 mm long, reddish, hypan-
thium 3-4 mm long, 2-2.6 mm diam., calyx lobes 2-5
mm long, ca. 0.4 mm broad; corolla salverform, rose or
pinkish red to red-brown, tube 1-3 mm long, lobes 3-7
mm long, ca. 1.3 mm broad; anthers ca. 4 mm long,
white. Fruits 4-6 mm long, 2-3 mm diam. (dried), red;
seeds ca. 0.2 mm long.

Plants of lowland rain forest formations on the
Caribbean slope, 50-600(-1 100) m elevation.
Flowering in March and May-November; fruiting
in the same months and in January. This species
apparently ranges from Rio San Lorenzo, Alajuela,
eastward to Guapiles, Limon, in Costa Rica.

Hoffmannia liesneriana is recognized by the large
leaves with long petioles, the short unbranched
stems, flowers in close clusters on cauliflorous in-
florescences, long calyx lobes, and lower-elevation
habitat. Unlike so many of our other Hoffmannia
species, this species has leaves that are often
abruptly narrowed at the base (but some collec-
tions do have long-decurrent leaf blades). This spe-
cies resembles Psychotria capacifolia with white
flowers. Compare H. davidsoniae and H. eliasii
Dwyer of Panama.

Hoffmannia longipetiolata Polak., Linnaea 4 1 : 567.
1877. H. tonduzii Standl., Contr. U.S. Natl.
Herb. 20: 205. 1919. H. woodsonii Standl., Ann.
Missouri Dot. Card. 28: 471. 1941. Figure 11.

Small shrubs or herbaceous subshrubs, 1-2 m tall,
usually few-branched and slender-stemmed, leafy stems
0.8-5 mm thick, glabrous or minutely puberulent along
narrow longitudinal ridges or with crooked hairs 0.5 mm
long in early stages; stipules 0.7-3 mm long, 2-3 mm
broad at the base, triangular, glabrous, deciduous. Leaves
with petioles 3-15(-40) mm long, 0.5-1 mm broad, gla-
brous or puberulent; leaf blades 4- 1 2(- 1 5) cm long, 1 . 5-
4(-6) cm broad, elliptic-obovate to elliptic or ovate-el-
liptic, apex short- to long-acuminate, tip to 1 5 mm long,
base acute to cuneate and somewhat decurrent, drying
membranaceous to thin-chartaceous, brownish green to
grayish green above, glabrous above, glabrous or mi-
nutely puberulent on the veins beneath, 2 veins 4-7 (-9)7
side. Inflorescences 1-3/axil (2-6/node), 1-2 cm long,
cymes or fascicles with (l-)3-6(-12) flowers, peduncles
2-5 mm long, pedicels 2-8(-10) mm long, ca. 0.3 mm
thick, glabrous or rarely sparsely and minutely (0. 1 mm)
puberulent. Flowers greenish yellow, glabrous, hypan-
thium 1.5-2 mm long, 1.3 mm diam. distally, obconic,
calyx lobes 0.5-1 (-2) mm long, ca. 1 mm broad at the
base, triangular and acute; corolla rostrate, yellow, pink
or cream white, tube 1.5-3 mm long, 1.3-1.5 mm diam.,
lobes 2-4 mm long, 1 .5-2 mm broad at the base, reflexed;
anthers 3-3.5 mm long. Fruits 5-6 mm diam., subglo-
bose, red to pink.

Plants of wet evergreen forest formations on both
the Caribbean and Pacific slopes, 600-1 800(-2000)
m elevation. Flowering in January-October, with
the majority in May-July; fruiting in April and
September-February. The species ranges from the
Cordillera de Guanacaste eastward to central Pan-
ama.

Hoffmannia longipetiolata is recognized by the
smaller or medium-sized leaves that taper grad-
ually to apex and base, little or no pubescence,
terete stems, very small few-flowered inflores-
cences, and small glabrous yellow flowers. The thin
peduncles and pedicels, short corolla tubes, and
larger corolla lobes are also helpful in recognizing
this species. This is one of the most commonly
collected species of Hoffmannia in Costa Rica.
Originally it was thought that the smaller-leaved
specimens determined as H. tonduzii were dis-
tinct, but there are too many intermediate collec-
tions to be able to segregate the larger-leaved spec-
imens (some of which have long petioles).
Collections from lower (600-900 m) elevations
and from western Panama often have larger leaves
than collections from central highland Costa Rica.
Two species that are very similar to H. longipe-
tiolata are H. decurrens (with densely puberulent
young stems) and H. psychotriifolia (with larger
fasciculate flowers and narrowly ellipsoid fruits).

Hoffmannia nesiota J. D. Smith, Bot. Gaz. 61: 374.
1916.

Shrublets, branchlets subterete, glabrous; stipules not
seen. Leaves with petioles 7-8 cm long, 2-3 mm thick,
pubescent; mature leaf blades 18-26 cm long, 10-13 cm
broad, elliptic to broadly elliptic-oblong, acuminate, base
obtuse and slightly decurrent, stiffly chartaceous and pu-
bescent or glabrescent, 2 veins 12-16/side. Inflores-
cences cymose with peduncles 1-5 cm long, pubescent
or glabrescent, lacking bracts and bracteoles, pedicles 6-
10 mm, slender. Flowers 15-16 mm long, with calyx
lobes 1.5-2 mm long, triangular; corolla 11 mm long,
rotate, puberulent, tube 5-6 mm long, lobes 5-6 mm
long, linear; ovary trilocular, style 12 mm long. Fruits
ca. 1 cm diam.; seeds ellipsoid ca. 0.5 mm long.

A poorly known species endemic to Cocos Is-
land. The species is only known from the type
collection: Pittier 12387 us, collected at Wafer Bay,
June 1898, and Barclay 2178 us, April 1838. The
Barclay collection is much more pubescent than
the type but appears to represent the same species.
Fruits of the Barclay collection are two-locular,
not three-locular as described for the type.

Hoffmannia nicotianifolia (Mart. & Gal.) L. O.

176

FIELDIANA: BOTANY

Williams is a species of Mexico and northern Cen-
tral America; the name has been incorrectly ap-
plied to Costa Rican collections.

Hoffmannia pallidiflora Standl., J. Wash. Acad.
Sci. 15:9. 1 925. H. rexmontis Dwyer, Ann. Mis-
souri Hot. Gard. 67: 241. 1980. Figure 11.

Herbs or subshrubs, 0.5-1.5 m tall, erect or leaning
on others, usually unbranched, leafy stems 2-6 mm thick,
glabrous, apparently succulent in life and distorted on
drying; stipules 2-3 mm long, 2 mm broad at the base,
glabrous, caducous. Leaves with petioles (1.2-)2-6 cm
long, 0.8-1 .8 mm broad, petioles of the same node often
unequal in length, glabrous; leaf blades 10-25 cm long,
4-10 cm broad, broadly elliptic to elliptic-oblong or el-
liptic-obovate, apex acuminate with a narrowed tip 5-
15 mm long, tapering gradually to a cuneate base (or
abruptly obtuse) and decurrent on petiole, drying mem-
branaceous, dark yellowish green or grayish green, gla-
brous above and below, 2 veins 8- 11 /side. Inflores-
cences solitary and axillary to basal leaves or cauliflorus
(2/node), 3-8 cm long, with a single unbranched rachis
or with 2 lateral helicoid branches 1-3 cm long, pedun-
cles 15-35 mm long, 0.3-1 mm thick, flowers 3-30,
pedicels 0-3 (-6) mm long, ca. 0.5 mm thick. Flowers
glabrous, hypanthium 2-3 mm long and 1 mm diam. at
apex, narrowly obconic, calyx lobes 1-2 m long, trian-
gular to ligulate, acute; corolla funnelform, white or pale
yellowish green, tube 2-4 mm long, ca. 1.3 mm diam.,
lobes 4.5-8 mm long, 0.8-1.7 broad. Fruits white be-
coming red or dark reddish purple, 4-6 mm diam. (to 1
cm in life), globose or ellipsoid-oblong during develop-
ment.

Plants of the wet cloud forests along the Carib-
bean escarpment, (400-)700-1400 m elevation.
Flowering in January-June; fruit in February-June,
August, and November-December. Collections
have been made from Bijagua, Alajuela, south-
eastward to the upper Rio Grande de Orosi, Car-
tago, and in western Panama.

Hoffmannia pallidiflora is recognized by the gla-
brous, slender, usually simple unbranched or two-
branched pedunculate inflorescences, very thin
leaves usually decurrent at the base and often long-
petiolate, narrowly obconic hypanthium, and re-
striction to the middle elevations of the Caribbean
slope. The thin foliage dries yellowish to very dark
green. The slender unbranched inflorescences have
flowers along only one side and usually dry yel-
lowish.

Hoffmannia piratarum Standl., J. Wash. Acad. Sci.
18: 180. 1928.

Shrublets, branchlets obtusely tetragonal, glabrous;

stipules caducous. Leaves with petioles 25-45 mm long;
leaf blades 10-20 cm long, 5-6 cm broad, elliptic-lan-
ceolate, long-acuminate, the tip often falcate, drying
membranaceous, dark green and glabrous above, sparse-
ly short villous and glabrescent beneath, with 8-9 2
veins on each side. Inflorescences in leaf axils, 1-3/nodc,
to 3 cm long, peduncles 1-2.5 cm long, short villous,
pedicels 2-5 mm long. Flowers puberulcnt proximally,
hypanthium 3 mm long, calyx lobes 2-3 mm long, tri-
angular or rounded; corolla glabrous, tube ca. 4 mm long.
2 mm diam. at base and 5 mm at apex, corolla lobes ca.
6 mm long. Fruits ca. 6 mm long.

Plants of Wafer Bay, Cocos Island. Flowering
in January. Standley knew it only from the type
collection Pittier 16259 us, and the preceding de-
scription is based on his description.

Hoffmannia pirtieri Standl., Contr. U.S. Natl. Herb.
18: 140. 1916. H. panamensis Standl., Contr.
U.S. Natl. Herb. 20: 204. 1919.

Shrubs or subshrubs, 1.5-3(-4) m tall, usually with
unbranched main stems, leafy stems 2-9 mm thick, be-
coming woody and 12 mm thick, glabrous or densely
pubescent on the young stems; stipules ca. 1 .5 mm long,
ca. 2.5 mm broad at the base, broadly triangular, gla-
brous. Leaves with petioles 1.5-7 cm long, 1.5-5 mm
broad, glabrous or rarely puberulent in early stages, with
narrow lateral margins continuous with the lamina mar-
gins; leaf blades 1 2-26(-35) cm long, 4-1 2(-l 6) cm broad,
elliptic-obovate to elliptic-oblong or oblanceolate, apex
acuminate with tip 8-18 mm long, gradually narrowed
to the cuneate base and long-decurrent on the petiole,
drying chartaccous and dark brown or dark grayish green
above, glabrous above and below (rarely puberulent in
early stages), 2 veins 9-14/side and weakly loop-con-
nected near the margin. Inflorescences 1-5/axil from
thickened reduced peduncles, 2-5(-9) cm long, with
(5-)10-15 flowers, peduncles 4-40 mm long, with 1
branches 3-30 mm long (often 2-branched with alternate
flowers along 1 side), glabrous to densely reddish pu-
berulent, pedicels to 9 mm long. Flowers glabrous to
reddish puberulent, hypanthium 2-3 mm long, 1.5-2
mm diam., oblong-obconic, calyx orange-red, calyx lobes
0.5-1 mm long; corolla yellowish tipped with red or
orange, tube 3-6 mm long, 0.7-1.6 mm thick, lobes 3-
5 mm long, 1.7 mm broad at the base. Fruits ca. 5 mm
diam., deep green becoming deep red.

Plants of evergreen montane forest formations,
1 100-1 700(-2200) m elevation. Flowering in Feb-
ruary-August; fruiting in January-February and
August. The species (as here interpreted, see be-
low) ranges from the San Vito de Coto Brus area
into the adjacent Chiriqui Highlands of Panama.

Hoffmannia pittieri is recognized by the taller,
more woody stems, larger leaves with cuneate long-
decurrent leaf bases, inflorescences with slender-
pedicellate flowers, short corolla lobes, and re-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

177

stricted latitudinal range. The inflorescences vary
from short to moderately long with few to many
branches. The interpretation presented here as-
sumes that the very similar H. leucocarpa does not
range eastward of Volcan Irazu. There is probably
intergradation with H. arborescens (which usually
has larger more branched inflorescences), and it
may be necessary to place all this material under
a more broadly denned H. arborescens.

Hoffmannia psychotriifolia (Benth. in Oerst.) Gri-
seb., Fl. Brit. W. Ind. 321. 1861. H igginsia psy-
chothaefolia Benth. in Oerst., Vidensk. Meddel.
Dansk. Naturhist. Foren. Kjobenhavn 1852: 50.
1853. Figure 11.

Slender shrubs or herbaceous subshrubs, l-2.5(-4) m
tall, with few lateral branches, leafy stems 0.8-4.5 mm
thick, glabrous, terete; stipules 1-1.5 mm long or reduced
to a ridge, 1-2 mm broad at the base, triangular, glabrous.
Leaves with petioles 8-35(-50) mm long, 0.7-1.5 mm
thick, glabrous; leaf blades 5-14(-20) cm long, 1 .5^.5(-6)
cm broad, narrowly elliptic to oblanceolate or narrowly
elliptic-obovate, apex acuminate with tip 1-2 cm long,
gradually narrowed to the acute or cuneate and decurrent
base, drying membranaceous to thin-chartaceous, dark
green or dark grayish brown above, distinctly paler be-
neath, glabrous above and below, 2 veins 5-9/side. In-
florescences usually fasciculate from a thickened solitary
short-shoot (brachyblasts) in the leaf axils, with 5-30
flowers per node, peduncles rarely present or 0-3(-8) mm
long, pedicels 2-8 mm long, ca. 0.3 mm thick, glabrous
or minutely puberulent. Flowers glabrous, to 18 mm
long, hypanthium 2-3 mm long, 0.7-1.7 mm diam. dis-
tally, oblong-obconic, with longitudinal ridges, calyx lobes
0.6-2(-3) mm long, triangular to ligulate; corolla fun-
nelform, pale yellow to white or pink, tube 4-9 mm long,
1-3 mm diam., lobes 3-6 mm long, 1.4-2 mm broad at
the base. Fruits 6-12 mm long, 5-8 mm diam., oblong-
ellipsoid, becoming red and translucent.

Plants of evergreen montane forest formations,
(900-)1 100-2100(-2400) m elevation. Flowering
in January-July and September-November; fruit-
ing in February-September and November. The
species ranges from southern Mexico and Guate-
mala to Panama and occurs in the West Indies.

Hoffmannia psychotriifolia is recognized by the
thin-glabrous leaves tapering gradually to both apex
and base, stems rounded and slender, small, usu-
ally fasciculate inflorescences, slender pedicels,
larger flowers with well-developed calyx lobes,
longer ridged ovary, and oblong fruit. This species
may be difficult to separate from H. longipetiolata
(including H. tonduzii), but that species has small-
er flowers, few-flowered, often pedunculate inflo-
rescences, and young stems that may have puber-
ulent longitudinal ridges.

Hoffmannia refulgens (Hooker) Hemsley was
based on cultivated material thought to have been
collected in South America but most probably from
northern Central America or southern Mexico (D.
Lorence, pers. comm.). This name has been used
for Costa Rican collections now placed under H.
bullata.

Hoffmannia subauriculata Stand! .. J. Wash. Acad.
Sci. 18: 179. 1928. H. haydenii Dwyer, Ann.
Missouri Bot. Gard. 56: 277. 1969. Figure 8.

Herbaceous subshrubs, 1-2 m tall few-branched, leafy
stems 1 .5-5(-7) mm thick, subglabrous; stipules 2-3 mm
long. Leaves subsessile with poorly denned petioles be-
cause of the long-attenuate lamina base and winged lat-
eral margins 2-4 mm broad (short petioles 1-3 mm long
sometimes present below the auricles); leaf blades 8-
2 5 (-40?) cm long, 3-10(-13) cm broad, obovate to ob-
lanceolate or broadly elliptic, apex short-acuminate, ta-
pering very gradually to the cuneate or long-attenuate
base (5-7 x l cm) and usually slightly auriculate near
the stem, drying dark, glabrous above, subglabrous or
minutely puberulent on the veins below, 2 veins 9- 137
side. Inflorescences 2-4 on lower leafless nodes, 3-9 cm
long, usually with few or no lateral branches (cymose to
cincinnoid), with ca. 6-12 flowers, peduncles 2-4 cm
long, glabrous or with reddish curved hairs to 0.2 mm
long, pedicels 2-6(-12) mm long. Flowers with minute
curved hairs or glabrous, with calyx lobes ca. 1.5 mm
long, triangular; corolla not seen at anthesis. Fruits ca.
8x6 mm, ellipsoid, bright red.

Plants of wet evergreen forests in the central
mountains and on the Caribbean slope, from 100
to 1 800 m elevation. Flowering in June-July and
November; fruiting in December-January. The
species ranges from near Monteverde to central
Panama.

Hoffmannia subauriculata is distinguished by
the long-decurrent leaves with broad (ca. 5 mm)
lateral margins (along what would otherwise be a
petiole) and their slight auriculate expansion near
the base. The inflorescences appear to develop only
at leafless nodes. This species is similar to H. am-
plexifolia, which has three leaves per node.

Hoffmannia valerii Standl., J. Wash. Acad. Sci.
18: 178. 1928. Figure 9.

Weak-stemmed shrubs or subshrubs, 0.5-1. 5(-3?) m
tall, usually without lateral branches, leafy stems 1-4
mm thick, with thin crooked multicellular hairs 0.3-1.5
mm long; stipules 1-4 mm long, rounded distally or with
an awn, caducous. Leaves with petioles 8-60 mm long,
0.8-1.6 mm thick, villous with hairs 0.3-1 mm long;
leaf blades 4-14(-19) cm long, 1.8-7(-9) cm broad,

178

FIELDIANA: BOTANY

broadly elliptic to narrowly elliptic-oblong or ovate-el-
liptic, apex acuminate, base obtuse to acute and some-
what decurrent on petiole, drying membranaceous to
thin-chartaceous, dark brown or grayish green above,
glabrous or with few scattered hairs above, with more
numerous crooked hairs 0.3-1.5 mm long beneath, 2
veins 6-10/side. Inflorescences axillary or at leafless
nodes, 2-4/node, l-2(-3) cm long, fasciculate or con-
tracted cymes with 3-9 flowers, peduncles 1-4 mm long,
pedicels 2-8(-12) mm long, filiform, villous or subgla-
brous. Flowers puberulent with crooked hairs that dry
reddish, hypanthium 2-3 mm long, 1.5-2 mm diam.
distally, calyx lobes l-3(-4) mm long, narrowly trian-
gular; corolla rotate, pale yellow, tube 1-3 mm long,
lobes 3-5(-7) mm long; anthers 2-3 mm long. Fruits 4-
6 mm long, 4-5 mm diam., ellipsoid, becoming orange
or bright red.

Plants of the wet evergreen Caribbean slope and
lowlands, often along streams and in wet areas,
50-950 m elevation. Flowering in February-Au-
gust and November; fruiting in January, July, and
November. The species ranges from near Arenal,
Alajuela, to western Panama.

Hoffmannia valerii is recognized by the long hairs
on many younger parts, slender unbranched stems,
small short-pedunculate or fasciculate inflores-
cences, short corolla tubes, and lower-elevation
habitats. The lype (Valeria 57 us) was collected at
600 m elevation near Arenal, Guanacaste Prov-
ince. This species resembles H. gesnerioides (Oer-
sted) Kuntze of Nicaragua at 800-1500 m eleva-
tion, but that species has longer (2-4 cm) peduncles,
attenuate leaf bases, and dense spreading villose
pubescence on all parts. Costa Rican material
identified as H. boraginoides Dwyer ined. appear
to be plants of//, valerii that have unusually long
(2.5-3 mm) calyx lobes. The excellent DUKE col-
lections from La Selva display considerable vari-
ation in sepal lobe length, and it seems best to
consider these plants all part of H. valerii.

Hoffmannia vesiculifera Standl., Publ. Field Co-
lumb. Mus., Bot. Ser. 4: 285. 1929. H. kirkbridei
Dwyer, Ann. Missouri Bot. Gard. 56: 278. 1969.
//. morn Dwyer, loc. cit. 67: 236. 1980. Figure 8.

Herbs or subshrubs, 0.3-1.5 m tall, main stems usually
unbranched, leafy stems 2.5-1 mm thick, usually densely
villous with multicellular crooked reddish brown hairs
to 1.5(-3) mm long; stipules to 2-5 mm long, little de-
veloped and difficult to see among the pubescence, ca-
ducous. Leaves with petioles 14-45 mm long, with a
lateral longitudinal inflated chamber (vessicle or pouch)
12-20 mm long and 4-9 mm broad on each side (2/
petiole), with conspicuous reddish brown hairs; leaf blades
12-27(-40) cm long, 7-15(-20) cm broad, elliptic to el-
liptic-obovate (less often obovate-oblong or oblanceo-

late), apex acuminate to short-acuminate, base obtuse to
cuneate and decurrent (to the inflated vessicles), drying
thin-chartaceous, dark brown or grayish above, paler
grayish or reddish brown beneath, with few scattered
crooked multicellular hairs above and below, more
densely hirsutulous on the midveins above and below,
the crooked reddish hairs 0.4-1 .3(-3) mm long, 2 veins
8-17/side. Inflorescences axillary or terminal on short
lateral axillary shoots and subtended by leaf-like bracts
ca. 2 cm long, solitary (2/node), sessile or with peduncles
(= lateral branches?) to 3 cm long, flowers 10-many. in
dense subsessile capitulae or on condensed helicoid cymes
(with the inflorescences occasionally expanding to 7 cm
and becoming branched in fruit, as in the type: Cooper
231 F), pedicels to 4 mm long. Flowers yellowish but
drying reddish brown, with conspicuous hairs 1-2 mm
long, hypanthium ca. 2.5 mm long, and 1.5 mm diam.,
calyx lobes 2.5-7 mm long, linear, with few to many
reddish hairs; corolla rotate, glabrous, cream yellow to
reddish, tube 2-4(-8) mm long, corolla lobes 4-17 mm
long, to 3 mm broad at the base, lanceolate; anthers 3.5-
4 mm long. Fruits 4-6 mm long, 3-5 mm diam., be-
coming bright red or white, thin-walled with longitudinal
ribs, often with long ( 1 mm) crooked hairs.

Plants of wet evergreen forests of the Caribbean
slope and lowlands, (20-)300-1500 m elevation.
(Note: All Costa Rican collections came from above
300 m elevation.) Flowering in February-June and
September; fruiting in April and October-Febru-
ary. The species ranges from central Costa Rica
to central Panama.

Hoffmannia vesiculifera is our most distinctive
species of Hoffmannia. The unusual inflated struc-
tures of the petiole are found in no other Costa
Rican species of Rubiaceae and resemble those
found on some Melastomaceae. The long crooked
multicellular hairs, condensed inflorescences, and
long-linear calyx lobes are additional distinctions.
The variation in leaf size and form and inflores-
cences (dense heads to helicoid cymes or, rarely,
branching infructescences) is considerable but does
not warrant segregation of species or subspecies.
The inflorescences may all be borne on lateral ax-
illary branches that are variously reduced, as ev-
idenced by the paired leaf-like bracts subtending
the inflorescences. Only one of our collections, Lent
91 1 F, cited small ants in the petiole chambers.

Holtonia Standley

Holtonia Standley is now considered part of
Elaeagia.

Isertia Schreber

REFERENCE B. Boom, A revision of the genus

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

179

Isertia (Isertieae: Rubiaceae). Brittonia 36: 425-
454. 1984.

Shrubs or trees, branchlets slender and subterete or
thick and quadrangular, glabrous or puberulent; stipules
interpetiolar or intrapetiolar, sometimes deeply divided
and apparently 2/node, persisting. Leaves opposite, pet-
iolate, often large; leaf blades drying thin-chartaceous to
coriaceous, venation pinnate, domatia absent. Inflores-
cences terminal and solitary, paniculate-thrysiform or
racemose-thyrsiform, often large, the secondary branch-
es terminating in dichasia or scorpioid cymes, flowers
sessile or pedicellate. Flowers bisexual, radially sym-
metrical, hypanthium ovoid to subglobose, calyx lobes
4-6, small, equal or unequal, persisting; corolla tubular-
funnelform to salverform, corolla tube short or elongate,
villous in the throat, corolla lobes 4-6(7), short and
spreading, valvate or imbricate in bud; stamens 4-7,
filaments inserted near the mouth of the tube, anthers
dorsifixed, loculate with the interior of the thecae divided
into small chambers, included or exserted; ovary 2-6(-7)-

locular, ovules numerous on axile placentas, style linear,
stigma with 2-6(7) oblong lobes. Fruits berry-like with
fleshy exocarp and 2-6 pyrenes, each pyrene (nutlet) with
a bony endocarp and 2-many seeds, globose; seeds min-
ute, angular, brownish, the testa deeply foveolate.

A genus of 14 species, ranging from Central
America through northern South America and oc-
curring naturally only in western Cuba and Gua-
deloupe in the Caribbean. Boom has divided the
genus into two sections: section Cassupa with 2-
3( 4) locules and stigmas and fleshy fruit and sec-
tion Isertia with (4-)5-7 locules and stigmas and
hard fruit. Our two species represent both sections
of the genus. The large leaves, large many-branched
solitary terminal inflorescences, flowers with long
narrow corolla tubes, and loculate anthers distin-
guish this genus.

Key to the Species of Isertia

la. Leaf blades usually dull greenish or grayish beneath, usually acute at the base and decurrent on the
petiole; corolla yellow to orange or red, corolla tubes to 28 mm long; fruit oblate, ca. 7 mm diam.
I. haenkeana

1 b. Leaf blades usually whitish gray beneath, subtruncate to obtuse at the base and not conspicuously
decurrent on the petiole; corolla white, corolla tubes to 55 mm long; fruit ellipsoid, ca. 10 mm
diam. . /. laevis

Isertia haenkeana DC., Prodr. 4: 437. 1830. 7.
deamii Bartlett, Proc. Amer. Acad. Sci. 33: 59.
1907. 7. deamii var. stenophylla J. D. Smith,
Bot. Gaz. 61: 374. 1916. Figure 50.

Shrubs or small trees, 2-6(-20) m tall, branches thick
and often slightly narrower at the node, leafy branchlets
3-10 mm thick, quadrangular with rounded edges, be-
coming terete, densely puberulent with short (0.2-0.6
mm) grayish hairs; stipules 4/node (or interpreted as 2
with 2 almost separate lobes), 7-14(-45) mm long, 2-8
mm broad at base, narrowly triangular with long-acute
apex, glabrous abaxially or sericeous along the midvein
(strigulose), drying dark. Leaves often smaller beneath
the inflorescence, petioles 5-50 mm long, 1.8-4 mm thick,
with adaxial margins continuous with the lamina mar-
gins, densely puberulent; leaf blades (7-)14-45(-64) cm
long, (4-)7-16(-28) cm broad, elliptic to elliptic-oblong
or elliptic-obovate, apex short- or long-acuminate, base
gradually narrowed and acute to attenuate, decurrent on
petiole, drying chartaceous, dark brown above, glabrous
and often lustrous above, with thin erect or appressed
whitish hairs 0.2-0.5 mm long on the veins beneath, 2
veins 14-22/side and loop-connected near the margin.
Inflorescences 8-22 cm long, 6-12 cm broad, peduncle
a thick extension of the stem, to 5 cm long, with a thick
central rachis and many lateral branches, strigulose, the
lateral branches (dichasia) with a short (1-2 cm) reddish
orange peduncle and a terminal flower at the dichotomy

of 2 longer scorpioid branches with 4-9 flowers, the sec-
ondary branches subtended by narrow bracts 3-1 1 mm
long, pedicles 0-2 mm long, bracteoles 1-2 mm long.
Flowers with hypanthium and calyx tube 2-3 mm long
and equally broad, cupulate, subentire with 4 minute
lobes, sparsely strigulose externally; corolla bright yellow
turning orange or reddish, tube 17-25(-28) mm long, ca.
1.5 mm diam. at base and 2-3 mm distally, lobes 5-6,
5-7 mm long, with prominent dense yellowish hairs ca.
2 mm long on the inner face basally; stamens 5 or 6,
filaments and anthers 3.5-6 mm long; ovary 4-6(-7) loc-
ular. Fruits berry-like, 4-5 mm long and 6-8 mm diam.,
oblate, smooth and strigulose, usually with 5 or 6 car-
tilaginous lobes and 5-6 multiseeded pyrenes; seeds 0.6-
0.9 mm long.

Plants of evergreen lowland wet forest forma-
tions, from near sea level to 600 m elevation in
southern Central America. Probably flowering
throughout the year; in central Panama flowering
is primarily in the early rainy season (May-July),
with fruit maturing in the late wet season and early
dry season (Croat, 1978). This species ranges from
Mexico to Panama, Columbia, and Venezuela and
is found in westernmost Cuba.

Isertia haenkeana is recognized by its large op-
posite leaves with many secondary veins and de-

180

FIELDIANA: BOTANY

current lamina base, four persisting stipules at each
node, cupular calyx with minute lobes, narrow yel-
low to red corolla tubes, short corolla lobes with
bearded hairs within, and rounded berry-like fruit
with hardened pyrenes within. This species resem-
bles Palicourea guianensis.

Isertia laevis (Triana) B. M. Boom, Brittonia 36:
433. 1984. Cassupa laevis Triana, Ann. Sci. Nat.
Paris, Ser. IV 9: 44. 1858. C. panamensis Standl.,
Contr. U.S. Natl. Herb. 18: 135. 1916. /. pana-
mensis (Standl.) Standl., Publ. Field Columb.
Mus., Bot. Ser. 8: 346. 1931. Figure 30.

Small trees to 10(-1 5) m tall, often with several trunks
ca. 10 cm diam., leafy branchlets 4-12 mm thick, qua-
drangular with rounded edges, densely appressed-pu-
berulent or strigulose with short (ca. 0.3 mm) yellowish
or brownish hairs; stipules apparently 4/node but united
at the base (for 1-2 mm) and leaving a scar around the
stem above the leaf bases, 6-12 (20-40) mm long, 3-6
mm broad, triangular-subulate, coriaceous, glabrous or
sparsely puberulent on the abaxial surface. Leaves op-
posite, petioles 1.6-7.5(-12) cm long, 2-5 mm thick,
minutely (0. 1-0.2 mm) puberulent; leaf blades smaller
beneath the inflorescences, 15-60 cm long, 7-22 cm
broad, oblong to ovate-oblong or elliptic-oblong, apex
acuminate or acute apex, tip to 2 cm long, base obtuse
to rounded and truncate (rarely slightly decurrent on
petiole), drying chartaceous to subcoriaceous, usually dark
brown above, glabrous or subglabrous above and with
the minor venation often impressed, minutely puberu-
lent on the veins beneath, whitish canescent between the
veins and the major and minor venation clearly outlined,
2 veins 1 5-22/side and loop-connected near the margin,
proximal 2 veins arising at 90 to the midvein, 3 veins
subparallel. Inflorescences (7-)14-35 cm long, (4-)6-15
cm broad, ovoid to ellipsoid in outline and with many
short (4 cm) lateral branches, peduncle 2-6 cm long, to
6 mm thick, lateral branches usually a 5-flowered di-
chasium (the distal branches with 2 flowers on the upper
side), with short (1-3.5 cm) secondary peduncles, sparse-
ly and minutely puberulent, flowers sessile or short (2-
4 mm) pedicellate, bracteoles 1-3 mm long, broadly ovate.
Flowers sweet smelling and apparently opening in the
evening, hypanthium and calyx tube 5-7 mm long, 3-5
mm diam., an elongate cup glabrous externally, the calyx
lobes minute and 3-6 or obscure; corolla salverform,
white, tube 32-55 mm long, 2-5 mm diam., sparsely
and minutely puberulent externally, corolla lobes 6-7,
1 0- 1 4 mm long, with long (1mm) yellowish hairs inside
near the mouth; stamens 6-7, filaments ca. 3 mm long
and laminar, anthers 7-9 mm long, included; ovary 2-
3-locular, style 32-55 mm long, stigma with 2 oblong
lobes 3-5 mm long. Fruits berries to 12 mm long, 8-1 1
mm diam., ellipsoid to subglobose, smooth and usually
glabrous, becoming black; seeds 0.7-1 mm long.

Trees of open secondary vegetation in evergreen
rain forest formations in both the Caribbean and
southern Pacific lowlands, from near sea level to

about 800 m elevation in Costa Rica. Probably
flowering throughout the year but with flowers and
fruit collected mostly in May-September. This
species ranges from Costa Rica and Panama along
the Andes to Bolivia and in the adjacent Amazon
Basin.

Isertia laevis is recognized by its large opposite
leaves with whitish surfaces beneath, four persist-
ing stipules at each node, solitary terminal inflo-
rescences with long-tubed white flowers, subentire
calyx tube, and subglobose berries. Panamanian
material of this species was called Isertia hypoleu-
ca by Croat (1978) and Dwyer (1980), but I. hy-
poleuca Bentham is a different species occurring
in Venezuela, the Guianas, and the Amazon basin.

Ixora Linnaeus

Shrubs or small trees, branchlets terete or angular,
mostly glabrous; stipules interpetiolar (also slightly unit-
ed above the petioles to form a very short intrapetiolar
ridge), simple, usually broad at the base and acuminate,
persisting. Leaves opposite and decussate or in whorls
of 3, sessile or petiolate, laminae often coriaceous, ve-
nation pinnate, domatia absent. Inflorescences terminal
or less often axillary, cymose, thyrsoid or paniculate,
branching often trichotomous, often with a rounded or
flattened top (corymbose), flowers pedicellate and sub-
tended by 2 bracteoles. Flowers bisexual and mono-
morphic, hypanthium ovoid, calyx tube short, calyx lobes
4 (5), short or extended, corolla salverform, white to
pinkish, red, scarlet, or yellow, corolla tube narrow, gla-
brous or puberulent at the throat, corolla lobes 4 (5-8),
linear-lanceolate to ovate, imbricate or convolute in bud
and rotate at anthesis; stamens 4 (5-8), inserted in the
throat or mouth of the tube, filaments short or none,
anthers oblong to linear, dorsifixed, exserted or partly
exserted; ovary 2-locular with 1 ovule in each locule,
peltately attached to the middle of the septum, style
filiform, stigmas 2. Fruits baccate, red becoming black,
the pericarp fleshy or leathery, with 1-2 pyrenes; seeds
concave-convex or plano-convex to subglobose.

A pantropical genus of ca. 400 species with the
largest number of species in Africa and the Indo-
Pacific area. The colorful inflorescences, narrow
corolla tubes, stamens borne near the apex of the
corolla tube, and two-seeded fruit help to char-
acterize the genus. The flowers are protandrous in
a distinctive manner: pollen is transferred from
the clustered anthers into a concave area of the
stigmatic head; elongation of the stigma disperses
the pollen. Later, the stigma lobes open to expose
their receptive surfaces for pollination. Four spe-
cies are known to occur in Central America; two
are native and two are widely cultivated as or-
namentals.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

181

Key to the Species of Ixora

la. Flowers closely crowded together in capitate inflorescences; corolla tubes 25-45 mm long; plants
cultivated for ornament 2

Ib. Flowers crowded in distal cymose groups in an open paniculate inflorescence; corolla tubes 3-8 mm

long; native wild species 3

2a. Flowers red or orange; leaves sessile or subsessile /. coccinea

2b. Flowers white; leaves petiolate /. finlaysoniana

3a. Fruits ca. 10 mm diam.; flowers sessile in small glomerules, peduncles 0.7-3 mm thick; leaf blades
1 2-26 cm long /. floribunda

3b. Fruits ca. 4 mm diam.; flowers in panicles and pedicellate, peduncles 0.3-1 mm thick; leaf blades
7-16 cm long /. nicaraguensis

Ixora coccinea L., Sp. PI. 110. 1753. Figure 31.

Shrubs 0.5-3(-5) m tall, much branched, leafy branch-
lets 1-4 mm thick, glabrous, terete; stipules 0.5-1.5 mm
long at the broad base and with a narrow tip 2-14 mm
long, glabrous. Leaves opposite, sessile or subsessile with
petioles to 2 mm long; leaf blades (2-)3-l 0(-1 6) cm long,
( 1-) 1 .5-4.5(-6) cm broad, oblong to elliptic-oblong, ovate-
oblong or oblong-obovate, apex rounded to obtuse or
acute and often with minute (0.5-1 mm) slender tip, base
rounded and subtruncate or subcordate to obtuse, drying
stiffly chartaceous or subcoriaceous, glabrous above and
below. Inflorescences terminal, to 10 cm long, flowers
closely clustered (subcapitate), peduncles 1-3 cm long,
flowers usually sessile at the apex of short-stipitate triads
(cymes). Flowers with hypanthium 1-1.5 mm long, tur-
binate, calyx lobes 4, ca. 1 mm long; corolla red or or-
ange-red, tube 25-45 mm long, only 0.5-1 mm diam.,
glabrous or subglabrous, corolla lobes 4, 10-15 mm long,
4-8 mm broad; stamens 4, anthers exserted and soon
deciduous; style usually exserted. Fruits 8-1 5 mm diam.
(rarely developed in cultivars).

Ixora coccinea is a widely cultivated species
throughout Central America, from near sea level
to about 1500 m elevation. The rounded clusters
of brilliant reddish flowers with long slender tubes
and the evergreen leaves make it an attractive or-
namental. The species originated in India and is
now grown throughout the tropics and subtropics.
Several varieties and forms have been recognized;
see F. R. Fosberg and H.-H. Sachet, Three culti-
vated Ixoras. Baileya 23: 74-85. 1989. This spe-
cies is called cruz de Malta, flor defuego, jazmin,
jazmin rojo, and jazmin de coral.

Ixora finlaysoniana Wallich ex G. Don, Gen. hist.
3: 572. 1834. Figure 31.

Shrubs or small trees to 5 m tall, leafy branchlets 2-
6 mm thick, glabrous, becoming terete with age; stipules
2-7 mm long, triangular and acuminate or broad with
1 or 2 awns. Leaves opposite, petioles 4-20 mm long,

1-2 mm thick, glabrous; leaf blades 6-18 cm long, 2-
6.5 cm broad, narrowly oblong to narrowly oblong-ob-
ovate, apex bluntly obtuse or rounded (acute), base grad-
ually narrowed and obtuse or acute, drying subcoria-
ceous, glabrous above and below. Inflorescences terminal,
5-10 cm long, to 10 cm broad and rounded, of densely
congested cymose branches forming a capitate cluster,
peduncles 0-3 cm long. Flowers glabrous, hypanthium
1-2 mm long, calyx lobes 3-4 mm long 1 .5-2 mm broad,
petaloid in texture; corolla white, tube 20-30 mm long,
0.3-1 mm diam., lobes 4-6(-8) mm long, l-2(-3) mm
wide; stamens 5, exserted, anthers narrow; stigma ca. 2.2
mm long, slender and narrowly 2-lobed.

Ixora finlaysoniana is widely cultivated in the
tropics and planted at lower (0-1 000 m) elevations
in Central America. The rounded inflorescences
of many white flowers with long slender tubes, the
lack of pubescence, the foliaceous sepal lobes, and
the short-petiolate coriaceous oblong leaves char-
acterize this species. Native of southeast Asia; this
species is referred to as corona de la reina and
mono de reina.

Ixora floribunda (A. Rich.) Griseb., Cat. PI. Cub.
134. 1866. Siderodendronfloribundum A. Rich,
in Sagra, Hist. Cuba 1 1: 24. 1850. Figure 43.

Small trees to 1 5 m tall, short-shoots frequently pres-
ent, leafy branchlets 2.2-6 mm thick, smooth and gla-
brous, terete; stipules 4-10 mm long, united above the
leaf bases to form a short (1-4 mm) sheath, triangular-
subulate and often shifted to the side above the petiole
(not strictly interpetiolar), glabrous abaxially. Leaves op-
posite, petioles 10-25 mm long, 1.5-2.5 mm thick, gla-
brous; leaf blades 12-26 cm long, 4-10 cm broad, oblong
to elliptic-oblong, or obovate, apex obtuse to short-acu-
minate, base obtuse to attenuate and decurrent on pet-
iole, drying stiff-chartaceous to subcoriaceous, concol-
orous, glabrous above and below, 2 veins 6-1 0/side, not
loop-connected distally. Inflorescences terminal, 1-3, 5-
1 2 cm long, panicles with 2-3 pairs of opposite branches,
primary peduncles 1-3 cm long, secondary peduncles
equally long and densely puberulent with short (0.2-0.3

182

FIELDIANA: BOTANY

mm) grayish hairs, flowers sessile and 3-12 in distal
cymose clusters. Flowers with hypanthium ca. 1.5 mm
long and 1.2 mm diam., turbinate, covered with short
(0.2 mm) thin erect whitish hairs (and with some smaller
glandular hairs); calyx tube 0.5-1 mm long, calyx lobes
minute; corolla white, tube ca. 4 mm long, 0.3-0.6 mm
diam. in proximal half, glabrous externally, white-villous
within, lobes 4, 3-4 mm long, ca. 1 .5 mm wide, glabrous;
filaments 0.6-1.5 mm long, anthers ca. 3.3 mm long, ca.
1.5 mm broad; stigma lobes ca. 1.5 mm long and un-
equal. Fruits 6-10 mm long, subglobose, sessile, sparsely
puberulent, drying red and smooth, persistent calyx tube
0.4-0.7 mm high and ca. 1.2 mm broad.

Trees of both lowland rain forest formations and
seasonally deciduous formations, from 1 to 800
m elevation in Central America. Flowering in Jan-
uary-February; fruiting in March-May in Central
America. This species, apparently uncommon in
Central America, ranges from Honduras and El
Salvador to Colombia and some of the West In-
dies.

Ixora floribunda is recognized by its stiff-gla-
brous elliptic-oblong leaves, terminal inflores-
cences of white salverform flowers, puberulent ca-
lyx, and sessile red berries. Our collections come
from the area between Canas and Bagaces and
Monte Aguacate on the seasonally deciduous Pa-
cific slope of central Costa Rica. The species has
also been collected in the Caribbean rain forest
formations of Honduras and Nicaragua. It has been
called pah de Maria.

Ixora nicaraguensis Wernham, J. Hot. 50: 243.
1912. 7. rauwolfioides Standl., Trop. Woods 1 1:
25. 1927. Figure 43.

Shrubs or small trees to 10 m tall, often with many
branches, leafy branchlets 1.5-3 mm thick, glabrous, te-
rete, grayish; stipules 3-6(-8) mm long, 2-3 mm broad
at the base, broadly triangular and acuminate to cuspi-
date with a short awn to 3 mm long. Leaves opposite,
petioles 3-9 mm long, 0.7-2 mm broad, glabrous, with
lateral (or adaxial) ridges; leaf blades 6-1 3(-l 6) cm long,
2-5(-7) cm broad, elliptic-oblong, oblong or ovate-ob-
long, apex abruptly narrowed and acute or short-acu-
minate, base obtuse to somewhat attenuate and slightly
decurrent on petiole, drying chartaceous, glabrous on
both surfaces, 2 veins 6-12/side, often obscure and
weakly loop-connected distally. Inflorescences 1-3 and
terminal, 3-9 cm long and equally broad, open pyra-
midal panicles with 2-3 pairs of opposite branches, pe-
duncles 1-25 mm long, 0.3-1 mm thick, minutely (0.1-
0.2 mm) puberulent, flowers on slender pedicels 0-10
mm long in distal triads, distal bracts ca. 0.5 mm long.
Flowers with hypanthium and calyx tube 1-1.7 mm long,
minutely puberulent or glabrous, calyx lobes obsolete or
minute (0. 1-0.5 mm); corolla white, glabrous externally,
tube 3-6 mm long, ca. 1 mm diam., lobes 4, 3-4 mm

long, 1-2 mm broad, oblong and rounded distally; fila-
ments 0.5-1 mm long, anthers 2-3 mm long, 0.3 mm
thick; stigma ca. 1.5 mm long, exserted. Fruits 5-6 mm
long, 4-5 mm diam., subglobose to ovoid, glabrous and
red; pyrenes 5 mm long.

Plants of evergreen rain forest formations on
both the Caribbean and southern Pacific slope of
Costa Rica, from near sea level to 300 m elevation.
Flowering in February-March and June-October;
fruiting in March and July-November. This spe-
cies ranges from Belize to eastern Panama.

Ixora nicaraguensis is recognized by its small
white flowers on open thin-branched inflores-
cences, slender corolla tubes, and small globose
two-seeded fruit, borne on thin-branched infruc-
tescences. This species may be mistaken for a Psy-
chotria, but the corolla lobes are valvate in bud in
Psychotria (and related genera).

Ladenbergia Klotzsch

Small to large trees, the bark with bitter substances;
stipules usually large, interpetiolar or also united distally
(intrapetiolar) and forming a ring around the stem above
the leaf bases after falling, triangular to obovate, with
colleters at the adaxial base, caducous. Leaves opposite
(occasionally whorled), petiolate, often subcoriaceous,
pinnately veined, some species with domatia. Inflores-
cences terminal, paniculate with opposite branching or
cymose, ebracteolate. Flowers bisexual, monomorphic,
medium to large, fragrant, hypanthium turbinate to cy-
lindrical, calyx tube cupular, calyx lobes short or long;
corolla funnelform or salverform, sericeous externally,
corolla lobes 5-6, valvate in bud, minutely papillose
within and on the margins; stamens 5-6, filaments very
short, inserted near the center of the tube, anthers linear,
dorsi fixed, included; ovary 2-locular, placentas elongate,
spongy and borne on the septum, ovules numerous in
each locule and vertically imbricate, style slender, stig-
matic lobes 2. Fruits elongated capsules, cylindrical or
flattened, with septicidal dehiscence (but the septum very
thin and dehiscence often appearing to be loculicidal).
splitting from the top into 2 woody or coriaceous valves;
seeds numerous, longitudinally imbricate and peltate,
flattened and elongate, body oblong and surrounded with
a thin flattened dentate or laciniate wing.

A genus of about 30 species ranging from Costa
Rica to Bolivia; most of the species are Andean.
The genus is similar to Cinchona and Condami-
nea. The large salverform and sericeous flowers
with long valvate corolla lobes, large stiff usually
broad leaves, large broad stipules, and 2-valved
capsules with winged seeds help to characterize
the genus.

Paul Standley's original separation of Costa Ri-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

183

can material into three species appears to represent
three valid morphological entities that are not
sympatric, and intermediate collections are not

apparent. Nevertheless, the species are very sim-
ilar, and they might prove to be three subspecies
of a single species.

Key to the Species of Ladenbergia

la. Tufts of stiff hairs (0.5-1 mm long) usually present at the base of the petiole in young leaves; leaf

blades often acute at the base [corolla tubes 10-23 mm long, corolla lobes 11-17 mm long];

(1200-)! 500-2 100 m elevation L. valerii

Ib. Base of the petiole glabrous or with minute (0.2 mm) appressed hairs; leaf blades rarely acute at

the base; 0-1 500 m elevation 2

2a. Corolla tubes 15-20 mm long, corolla lobes 8-12 mm long; fruit 2-5.5 cm long; central highlands

1000-1400 m elevation L. brenesii

2b. Corolla tubes 25-50 mm long, corolla lobes 16-20 mm long; fruit 6-9 cm long; evergreen lowlands,

0-1000 m elevation L. sericophylla

Ladenbergia brenesii Standl., Pub. Field Mus. Nat.
Hist., Bot. Ser. 18: 1323. 1938. Figure 29.

Trees, 5-25 m tall, leafy branchlets 4-12 mm thick,
glabrous or sparsely puberulent at the nodes with short
(0.1-0.3 mm) hairs, quadrangular, becoming terete and
pale grayish; stipules 16-35 mm long, 4-16 mm broad,
ovate to obovate, obtuse to acute at the apex, glabrous
or with few slender ascending hairs ca. 0.5 mm long.
Leaves with petioles 10-35 mm long, 1.8-3.5 mm thick,
glabrous and drying dark; leaf blades 12-25 cm long, 5-
1 3 cm broad, broadly elliptic to elliptic-oblong or ellip-
tic-obovate, apex abruptly narrowed and obtuse or bluntly
acute, base obtuse or slightly rounded, drying subcor-
iaceous, glabrous above, glabrous or with a few scattered
hairs below, 2 veins (6-)7-10/side. Inflorescences 6-18
cm long, pyramidal, peduncles 2-7 cm long, glabrous,
with opposite branches and bracts to 5 mm long, pedicels
0-3 mm long. Flowers with hypanthium 3-6 mm long,
2.5-5 mm diam., clavate-tubular, densely sericeous with
yellowish brown ascending hairs, calyx tube ca. 1 mm
long, calyx lobes l-2(-4) mm long, ca. 2 mm broad,
broadly rounded, glabrous or sparsely puberulent distally
and with a minutely ciliate edge; corolla white, tube 1 5-
20 mm long, 5-6 mm diam., densely sericeous, lobes 8-
12 mm long, 2-3.5 mm broad. Fruits (2-)3-6 cm long,
5-10 mm broad (to 16 mm when fully flattened), sub-
terete and narrowly oblong before dehiscing, valves pu-
berulent externally and lustrous within; seeds 12-15 mm
long, 34 mm broad, surrounded by a thin translucent
erose wing, body of the seed 3-4 mm long and ca. 2 mm
broad.

Trees of wet evergreen cloud forest formations
of the central highlands, 1000-1400 m elevation.
Flowering in March-July; fruiting in July-Novem-
ber. The species is endemic and ranges from the
Cordillera de Tilaran to the western parts of the
Cordillera de Talamanca.

Ladenbergia brenesii is recognized by its cloud
forest habitat, large puberulent flowers, long nar-

row capsules splitting into valves that become flat,
and broad glabrous leaves. Compare this species
to L. valerii with which it can easily be confused.
Also known as aquijilla and quina.

Ladenbergia sericophylla Standl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 1324. 1938. Figure 29.

Trees, 8-35 m tall, trunks to 60 cm. dbh, leafy inter-
nodes 5-12 mm thick, young stems quadrangular and
drying dark, quickly becoming pale gray and terete; stip-
ules 15-45 mm long, 10-22 mm broad, oblong or ob-
ovate-oblong, rounded at the apex, minutely puberulent
with thin appressed-ascending hairs 0.1-0.3 mm long.
Leaves opposite, petioles 15-4 5 (-60) mm long, 2-3.5
mm thick, glabrous and drying very dark, reddish in life;
leaf blades 14-24(-30) cm long, 8-16(-18) cm broad,
very broadly elliptic to broadly elliptic-oblong or slightly
obovate, apex abruptly narrowed and rounded or bluntly
obtuse, base abruptly narrowed and obtuse or slightly
attenuate, decurrent, drying stiffly chartaceous to sub-
coriaceous, usually dark brown above, glabrous above,
minutely (0. 1-0.3 mm) appressed-puberulent on the veins
beneath and with larger (1 mm) hairs in the vein axils
(domatia), 2 veins 5-8/side. Inflorescences 10-20 cm
long, to 1 8 cm broad, pyramidal, peduncles 1-5 cm long,
4-8 mm thick, glabrous or minutely appressed-puber-
ulent, lateral branches opposite and subtended by bracts
3-6 mm long, bracteoles subtending the flowers 1-2 mm
long, rounded apically, pedicels 1-6 mm long. Flowers
with hypanthium 5-7 mm long, 2-3 mm diam., tubular,
densely sericeous with ascending yellowish hairs, calyx
tube 2-4 mm long, cupulate, sparsely puberulent or gla-
brous, calyx lobes 2-3 mm long, 2.5-3 mm broad, round-
ed at the apex, glabrous and drying dark; corolla white
or yellowish white, tube 25-50 mm long, 3-5 mm diam.,
densely sericeous, lobes 16-30 mm long, ca. 4 mm wide
and narrowly oblong to lanceolate. Fruits 6-1 1 cm long,
10-14 mm broad, oblong-cylindrical and rounded-rect-
angular in cross-section, straight or curved (falcate), mi-

184

FIELDIANA: BOTANY

nutely appressed-puberulent, persisting calyx 3-6 mm
long and 6-8 mm broad, narrowed at the base to form
a pedicel 5-1 5 mm long; seeds 1 1-20 mm long, 4-5 mm
broad with thin wing, body of the seed ca. 3 x 1.5 mm.

Tall trees of evergreen rain forest formations of
the Caribbean lowlands and on the southern Pa-
cific slope, 20- 1 000 m elevation. Flowering in Jan-
uary-February, July, and September; fruiting in
January-May. This species is only known from La
Selva, the western parts of the General Valley, the
mountains bordering the Pacific near Canas Gor-
das, and the Osa Peninsula. While endemic to Cos-
ta Rica, it may also occur in westernmost Panama.

Ladenbergia sericophylla is characterized by its
large broad leaves with appressed hairs on the veins
beneath, large broad stipules, long salverform se-
riceous corollas, and large capsular fruit. The taller
height of these trees may explain why our collec-
tions of this species are so few.

Ladenbergia valerii Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 18: 1324. 1938. Figure 29.

Small trees, 3-20 m tall, leafy branchlets 2-10 mm
thick, quadrangular or terete in early stages, essentially
glabrous but with stiff retrose or erect hairs ca. 0.5 mm
long at the base of the petiole attachment; stipules 8-16
mm long, 5-8 mm broad, broadly elliptic or ovate, apex
rounded, sparsely puberulent with appressed hairs or
glabrous and with a few hairs at the base and along the
midrib abaxially. Leaves opposite, petioles 6-25 mm
long, 1.5-2.5 mm thick, glabrous or very sparsely ap-
pressed-puberulent (except for tufts of longer hairs below
the base), drying dark; leaf blades 6-17(-28) cm long,
3-8(-13) cm broad, broadly elliptic to elliptic-obovate
or narrowly elliptic, apex abruptly narrowed and short-
acuminate or bluntly acute, base obtuse to acute, drying
stiffly chartaceous to subcoriaceous. glabrous or with a
few thin hairs above, with thin straight hairs 0.3-0.7 mm
long on the major veins beneath, 2 veins 5-9/side. In-
florescences 7-20 cm long, pyramidal with opposite
branching, peduncles 2-7 cm long, sparsely puberulent
or glabrous, bracts ca. 3 mm long, distal flowers in triads
and subtended by bracteoles 0.5-1 mm long, pedicels 0-
3 mm long and merging with the hypanthium. Flowers
with hypanthium and calyx tube 5-8 mm long, 2-3 mm
diam., densely appressed yellowish sericeous, calyx lobes
3-6 mm long, 2.5-3.5 mm broad at the base, broadly
rounded to obtuse distally and becoming reflexed, gla-
brous; corolla white or white with longitudinal pink
stripes, salverform, tube 10-23 mm long, 1.5-4.5 mm
diam., densely sericeous with ascending hairs, lobes 5,
(11-)12-17 mm long, 2.5-4.5 mm broad, lanceolate to
narrowly oblong, papillate-puberulent within, becoming
recurved. Fruits 3-8(-l 1) cm long, 8-14 mm broad; seeds
10-14 m long, 2-4 mm wide, body of the seed 3 m long
and 1.5 mm wide.

Trees of montane cloud forest formations from

(1200-)! 600 to 2100 m elevation. Flowering in
January-September and November; fruiting in
May-August. The species ranges from the Cor-
dillera de Tilaran to the Cordillera de Talamanca
(as far east as 8304' W) and will probably be found
in nearby Panama.

Ladenbergia valerii is recognized by its higher-
elevation habitat, unusual tufts of stiff short hairs
beneath the petioles on otherwise glabrous or gla-
brescent stems, sericeous corolla with relatively
long corolla lobes, and capsular fruit. This species
is very closely related to L. brenesii of similar for-
ests at somewhat lower elevations.

Lasianthus Jack

Herbaceous subshrubs (in Central America), shrubs or
rarely small trees, glabrous or pubescent; stipules inter-
petiolar, broadly triangular to lanceolate, persisting or
deciduous. Leaves opposite and decussate, petiolate,
usually acuminate at the apex, pinnately veined and of-
ten with many arching secondary veins, without doma-
tia. Inflorescences axillary, mostly sessile fascicles or cy-
mose glomerules, (sometimes pedunculate and simple or
branched), flowers sessile or subsessile, bracts small.
Flowers bisexual (rarely unisexual and monoecious), ra-
dially symmetrical, mostly small, sometimes heterosty-
lous, hypanthium urceolate to ovoid or subglobose, calyx
lobes 3-6, acute or rounded, persisting; corolla salver-
form to funnelform, white, corolla tube densely hairy in
the throat, corolla lobes 4-6, spreading or erect; stamens
4-6, filaments very short and borne in the throat, corolla
lobes 4-6, anthers dorsifixed near the base, included or
slightly exserted; ovary 4-12-locular, ovules solitary in
each locule, erect from the base of the locule or septum,
style short or long, stigmas 4-10, linear or lobed. Fruits
usually succulent, blue to purple, black, or red, with 4-
1 2 pyrenes, the pyrenes 1 -seeded and 3-angled with flat
sides, the dorsal side grooved, keeled or winged; seeds
narrowly oblong.

A genus of about 1 50 species, of southeastern
Asia, Malaysia, tropical Australia, and Africa. Two
species are found in the West Indies, and one spe-
cies in our area and South America. Our species
with eight-locular ovary and style with eight stig-
mas is unique among Central American Rubi-
aceae. Our species was originally described as a
new genus, Dressleriopsis Dwyer.

Lasianthus panamensis (Dwyer) Robbrecht, Ann.
Missouri Bot. Gard. 69: 427. 1982. Dressleriop-
sis panamensis Dwyer, Ann. Missouri Bot. Gard.
67: 154. 1980. Figure 7.

Herbaceous rhizomatous subshrubs to 0.6(-1) m tall,
leafy stems 2-6 mm thick, hirsute with slender erect hairs

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

185

to 2 mm long; stipules 4-7 mm long, 3-6 mm broad at
the base, broadly triangular and hirsute along the midrib.
Leaves with petioles 3-9 cm long, 1.5-2.5 mm thick,
hirsute; leaf blades 1 0-1 9 cm long, 5-9 cm broad, oblong
to elliptic-oblong, apex narrowed abruptly and short-
acuminate, tip 0-8 mm long, base obtuse and rounded
to subcordate-auriculate, often unequal at the petiole,
drying thin-chartaceous, dark grayish brown, with scat-
tered thin straight or crooked hairs 0.5-2 mm long on
upper and lower surfaces, 2 veins 9-14/side and loop-
connected near the margin to form an arcuate submar-
ginal vein, often with a shorter (parallel) minor 2 vein
between the major. Inflorescences axillary, dense fasci-
cles of sessile flowers 1-2 cm broad, glomerulate or ver-
ticellate, bracts ca. 2 mm long and difficult to see among
the long hairs. Flowers monomorphic, with hypanthium
ca. 2 mm long, subglobose, hirsute, calyx lobes 4, 3-4
mm long, 2-3 mm broad, with hairs to 1 mm long;
corolla white, tube 3-4 mm long, lobes 5 (4, 6), 1.5-4
mm long, oblong; stamens 5 (4, 6), anthers ca. 1 mm
long: ovary with 8 locules, style to 5 mm long, stigmas
8, oblong and radiate. Fruits berry-like, to 1 1 mm diam.,
globose, purple-black, pyrenes 8 or fewer, 3 mm long,
with an oblique scar on the concave side.

Plants of wet evergreen forest formations on the
Caribbean slope (at ca. 100 m elevation) and in
the central highlands of Panama (to 1 000 m ele-
vation). Flowering and fruiting in July. This spe-
cies is known from the La Selva area, Heredia,
and in Panama and Colombia.

Lasianthns panamensis is recognized by its slen-
der hirsute stems to 1 m tall, long-petiolate oblong
leaves with many secondary veins forming an ar-
cuate submarginal vein, sessile fasciculate inflo-
rescences, and fleshy berries with up to eight py-
renes. No other Central American species of
Rubiaceae has ovaries with eight locules. Super-
ficially, these plants resemble some species of
Hqffmannia and a few species of Psychotria with
axillary inflorescences.

Lindenia Bentham

REFERENCE S. Darwin, The genus Lindenia
(Rubiacea). J. Arnold Arbor. 57: 426-449. 1976.

Small shrubs growing along rivers and streams,
branchlets terete, glabrous or puberulent; stipules inter-
petiolar, borne above the petiole bases, short, usually
persisting. Leaves opposite short-petiolate; leaf blades
narrow, often drying dark, chartaceous, pinnately veined,
domatia absent. Inflorescences terminal, short and few-
flowered, flowers often in triads or solitary, bracts and
bracteoles present, pedicels short. Flowers bisexual, ra-
dially symmetrical, large and showy, hyanthium elon-
gate-turbinate, with 5 longitudinal ribs or angles, calyx
with 5 elongate lobes, lobes equal or unequal, persisting
in fruit; corolla narrowly salverform with a narrow elon-

gate tube, glabrous within, corolla lobes 5, convolute in
bud and spreading at anthesis; stamens 5, sessile on the
mouth of the tube, linear and exserted; ovary 2-locular,
ovules very many, vertical, placentas longitudinally ad-
nate to the septum, style slender and bifid. Fruits woody
capsules, clavate to pyriform or obovoid, with persisting
calyx lobes distally, 2-locular, splitting septicidally into
2 valves from the top; seeds numerous, angulate/rhom-
boidal.

A genus of three species, with the other two
species endemic in the western Pacific islands of
Fiji and New Caledonia, respectively. The stream-
side habitat, narrow leaves on thick stems, few
terminal flowers, and very long corolla tubes make
this genus quite distinctive.

Lindenia rivalis Benth., PL Hartw. 84. 1841. Fig-
ure 15.

Small shrubs, 0.4-1. 2(-2) m tall, leafy branchlets 1.5-
6 mm thick, glabrous or minutely puberulent with thin
ascending hairs ca. 0.2 mm long, older stems drying
black; stipules 3-5(-10) mm long, 1.5-4 mm broad, the
broad basal portion 2-3 mm high and with a narrow
awn to 2(-5) mm long, glabrous, thin and brown. Leaves
clustered near the ends of branchlets, petioles 2-10(-16)
mm long, 0.4-1 .5(-2) mm broad, little differentiated from
the leaf base; leaf blades 3-12(-17) cm long, 0.8-3(-4)
cm broad, oblanceolate to very narrowly elliptic-oblong
or narrowly elliptic, apex tapering gradually and acute,
sometimes with a short (0.5-1 mm) spine-like tip, base
tapering gradually and acute, decurrent on petiole, drying
chartaceous to coriaceous, the margin often revolute,
glabrous above, glabrous or puberulent beneath with thin
erect hairs 0.2-0.3 mm long, 2 veins 6-8/side, not loop-
connected. Inflorescences of solitary flowers or short (2-
1 mm) pedunculate triads (or clusters of up to 7 flowers),
bracts difficult to see among the distal leaves 4-9 mm
long, pedicels to 10 mm long, poorly differentiated from
the flower base. Flowers nocturnal, hypanthium 5-9 mm
long, 2-3.5 mm diam., puberulent, calyx tube minute,
calyx lobes 10-1 7(-22) mm long, 0.7-2(-2.8) mm broad,
narrowly oblong and acute, green; corolla white or white
tinged with pink, tube 10-17 cm long, 2-3.5 mm diam..
puberulent externally with hairs 0.2-0.5 mm long, lobes
1 5-27(-35) mm long, 5-14(-16) mm broad, narrowly to
broadly elliptic, acute at the apex; stamens sessile, an-
thers ca. 10 mm long, 1-1.4 mm broad; style slender
equalling or slightly exceeding the tube, stigmatic area
ca. 10m long. Fruits to 4 cm long, body of the fruit 15-
25 mm long, 9-14 mm broad, with long persisting calyx
lobes before dehiscence, broadly obovoid or pyriform,
the woody valves twisting; seeds 1.5-2 mm long.

Small woody shrubs of stream sides and often
growing on rocks next to the water, in deciduous
forest areas of Guanacaste Province, 0-700 m el-
evation (to 1300 m in Honduras). Flowering in
February-October in southern Central America
(primarily April-July); probably fruiting through-

186

FIELDIANA: BOTANY

out the year (mostly August-March). The species
ranges from northeastern Mexico and along the
Pacific slope of Central America to central Pana-
ma.

Lindenia rivalis is recognized by its small stiff
narrow leaves on thick dark branches, short stat-
ure and riverside habitat in seasonally deciduous
areas, few terminal flowers, and extremely long
narrow corolla tube. Opler's observation (Opler
945 F) that the flower is nocturnal would be con-
sistent with the long narrow tube being an adap-
tation for Sphingid moth pollination. Our only
other Rubiaceae with such long/narrow corolla
tubes have much larger leaves. Also known asjaz-
mincillo and lirio de aqua.

Machaonia Humboldt & Bonpland

Shrubs or small trees, stems often armed with spines
(leafless short-shoots), glabrous or puberulent, terete;
stipules interpetiolar, triangular or subulate and often
with a distal awn, persisting. Leaves opposite (rarely ter-
nate or verticillate), sessile or petiolate, thin-textured,
pinnately veined. Inflorescences terminal, solitary, pa-

niculate with opposite branching (or rarely umbeliform),
bracteate, flowers in cymose or crowded distal clusters.
Flowers bisexual and radially symmetrical, small, 4-5-
parted, hypanthium turbinate or obovoid, slightly com-
pressed laterally, calyx tube minute or cupulate, calyx
lobes 4-5(-6), equal or unequal, persisting; corolla short-
funnelform or short-salverform, white, corolla tube with
long hairs in the throat, lobes imbricate in bud. rounded
distally; stamens 4-5, filaments short or long, borne in
the throat of the tube, anthers oblong, dorsi fixed and
versatile, included or exserted; ovary 2-locular, with 1
ovule pendulous from the apex of each locule. style slen-
der, with 2 stigmas. Fruits small and dry, capsule-like
and splitting from the bottom into 2 elongate cocci (mer-
icarps), pendulous for a short period from the apex to
the persisting stipe-like central axis; seeds elongate and
cylindrical.

A tropical American genus of about 30 species,
found in Mexico, Central America, the West In-
dies, and tropical South America. The genus is
recognized by its small flowers with short corolla
tubes, two-locular ovary with solitary pendulous
ovules, and unusual capsule-like fruit. The sub-
capitate clusters of distal flowers on an openly
branching panicle, small thin leaves, and occa-
sional presence of spines are also distinctive.

Key to the Species of Machaonia

la. Hypanthium/ovary of the flower with few small hairs; fruit 5-7 mm long (including calyx lobes),
brownish and with few hairs often in rows; lowland evergreen Costa Rica M. martinicensis

Ib. Hypanthium/ovary of the flower whitish with a dense covering of short whitish hairs; fruit 4-5 mm
long, yellow or whitish with many hairs; Belize and central Panama M. acuminata

Machaonia acuminata Humb. & Bonpl., PI. Ae-
quin. 1: 101. 1806.

Shrubs or small trees, occasionally with spines to 4
cm long at the nodes, leafy branchlets 1.2-6 mm thick,
glabrous to densely villose with hairs to 1 mm long,
terete; stipules 2-4 mm long, triangular-subulate with a
narrow awn apically, puberulent. Leaves often disti-
chous, petioles 3-10 mm long, 0.6-1 mm broad, puber-
ulent and often with the hairs restricted to adaxial side;
leaf blades (3-)4-7(-10) cm long, 1.5^(-6) cm long,
ovate to ovate-elliptic or broadly ovate, apex acute (ob-
tuse) to short-acuminate and often slightly curved, base
obtuse to rounded and subtruncate, drying thin-charta-
ceous, glabrate above, with thin whitish hairs 0.3-0.7
mm long on the lower surfaces, 2 veins 3-6/side, tufted
hairs rarely present in the leaf axils. Inflorescences 4-20
cm long, 3-10 cm broad, open panicles, peduncles 2-4
cm long and 1.5-3 mm thick, puberulent, bracts to 7
mm long and liner, peduncles of the lateral branches 5-
40 mm long, flowers closely crowded in subcapitate dis-
tal groups of 7 or more, flowers sessile or subsessile.
bracteoles ca. 1 mm long. Flowers with hypanthium ca.
2 mm long, turbinate or oblong, minutely puberulent

with thin ascending whitish hairs, calyx lobes 0.5-1 mm
long, erect, rounded distally, puberulent; corolla white,
4-5 mm long, tube 1.5-3 mm long, sparsely puberulent
or glabrous, lobes 1.5-2.5 mm long, ca. 1.1 mm broad,
rounded at the tip; stamens exserted, anthers 0.3-0. 6 mm
long; styles 1-3.5 mm long, stigmas 0.3-0.6 mm long.
Fruits 4-5 mm long (including the persisting calyx), 1 .5-
2 mm broad, narrowly obovoid or turbinate, minutely
(0.1-0.2 mm) puberulent with ascending whitish hairs,
splitting at the top into 2 valve-like parts and later sep-
arating at the base, persisting central axis 2.5-3.2 mm
long.

Plants of lowland evergreen and partly decid-
uous forest formations. Flowering in May-No-
vember; fruiting in August-September and No-
vember. This species is known from southern
Mexico, northeastern Guatemala, Belize, and cen-
tral Panama southward to Ecuador and Brazil.

Machaonia acuminata is recognized by its oc-
casional spines, thin small ovate leaves, small
flowers with whitish puberulent ovary, and small

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

187

fruit. This species has not been found in Honduras,
El Salvador, Nicaragua, or Costa Rica. Its close
similarity to M. martinicensis and peculiar distri-
bution makes one wonder if the two are really
different species.

Machaonia martinicensis (EXT.) Standl., Publ. Field
Mus. Nat. Hist., Bot. Ser. 22: 193. 1940. Tertrea
martinicensis DC, Prodr. 4: 481. 1830. M. ro-
tundata Griseb., Fl. Brit. W. Ind. 348. 1861. M
rotundata var. dodgei Standl., Publ. Field Mus.
Nat. Hist., Bot. Ser. 18: 1326. 1938. Figure 37.

Shrubs or (rarely small trees or vines), l-4(-8) m tall,
branches occasionally with spines to 35 mm long, leafy
branchlets 1.6-5 mm thick, smooth glabrate; stipules
2.5-4 mm long, rounded or triangular at the apex and
with a short narrow awn. Leaves opposite (rarely alter-
nate), petioles 4-12 mm long, 0.5-1.2 mm broad, gla-
brous or puberulent on the adaxial side; leaf blades 4-
9 cm long, 1.8-5 cm wide, ovate to ovate-oblong (ellip-
tic-oblong), apex obtuse to acute or short-acuminate,
base broadly obtuse or rounded and sometimes decur-
rent on petiole, drying thin-chartaceous, glabrous above,
glabrous below or with few short (0.2 mm) hairs beneath
or with tufts of hairs (domatia) in the vein axils, 2 veins
5-7/side. Inflorescences 6- 11 (-15) cm long, to 14 cm
broad in fruit, peduncles 2-6 cm long, ca. 2 mm thick,
with whitish hairs 0.2 mm long, bracts 3-5 mm long and
linear (or sometimes leaf-like), secondary branches with
(secondary) peduncles 1 0-20 mm long, with soft whitish
hairs 0.2-0.3 mm long, flowers sessile or subsessile in
distal subcapitate clusters of 5-1 5. Flowers with buds 5-
6 mm long, hypanthium 1.5-2.5 mm long, sparsely pu-
berulent, calyx lobes 0.7-1.2 mm long, broadly rounded
distally; corolla white or pale greenish, tube 1.5-3 mm
long, stiff, densely villous at the mouth within, sparsely
puberulent externally, lobes 1-2 mm long; stamens 5,
anthers ca. 0.6 mm long; style 3.5 mm long, stigmas 0.5-
1 .2 mm long. Fruits 5-7 mm long, 2-3 mm broad, nar-
rowly turbinate-oblong, with 2 longitudinal sulci and
separating at the base, surface reddish brown or brown,
with few hairs (often in lines), persisting axis 4-5 mm
long.

Shrubs usually growing near the ocean shore, in
mangroves, or along rivers near the ocean in ev-
ergreen forest areas of both the Caribbean and
Pacific coasts. Flowering in March-April and June-
August; fruiting in July-September. The species
ranges from southeastern Nicaragua southward to
Colombia and also in Jamaica.

Machaonia martinicensis is recognized by its
restriction to near-shore environments, small thin-
ovate leaves, occasional spines, small flowers in
close clusters in an open panicle, and the unusual
capsule-like fruit splitting from the bottom. The
species has been collected along the Caribbean
shore and on the Osa Peninsula in Costa Rica.

Macrocnemum P. Browne

Trees or shrubs, branchlets terete, glabrous or puber-
ulent; stipules interpetiolar, oblong to obovate, caducous
or persisting with the leaves. Leaves opposite, petiolate;
leaf blades with pinnate venation, domatia often present.
Inflorescences terminal or axillary, paniculate with op-
posite branches, bracteate, flowers in distal cymose
groupings, sessile or pedicellate. Flowers bisexual and
radially symmetrical, calyx tube very short and cupulate,
calyx lobes 5, small, persisting; corolla funnelform or
salverform, corolla lobes 5, broadly imbricate in bud
and spreading at anthesis, often wider than long, mi-
nutely puberulent within; stamens 5, filaments short and
villous, anthers oblong; ovary 2-locular, ovules many,
peltate and vertically imbricate on the central placenta.
Fruits capsular, bisulcate and dehiscing loculicidally into
2 valves; seeds flattened, narrowly elongate with thin
wings.

A small genus of about 20 species, ranging from
Central America to Colombia and in the West
Indies. The broad rounded stipules, broadly over-
lapping corolla lobes (in bud), bilocular capsules,
and many small imbricate winged seeds help to
distinguish this genus.

Macrocnemum glabrescens (Benth.) Wedd., Ann.
Sci. Nat. Paris, ser. 4, 1: 76. 1854. Lasionema
glabrescens Benth., Bot. voy. Sulph. 105. 1845.
Figure 40.

Trees (rarely shrubs) (4-)8-25 m tall, bark brown and
the trunk deeply fluted, leafy branchlets 2-7 mm thick,
glabrous or minutely (0.1 mm) appressed-puberulent,
angular in early stages but becoming terete; stipules
(8-) 10-20 mm long, 5-10 mm broad, apex oblong-ob-
ovate and rounded, with thin appressed hairs 0.1-0.2
mm long, leaving a scar around the stem above the node.
Leaves often with the petioles unequal at the same node,
petioles 5-20 mm long, 0.7-1 .8 mm broad, sulcate adax-
ially; leaf blades 7-17(-21) cm long, 4-9 cm broad, ob-
ovate to obovate-oblong, or ovate-oblong, apex abruptly
narrowed and obtuse or short-acuminate apex, tip to 1
cm long, base gradually narrowed and obtuse or acute,
drying thin- to stiff-chartaceous, very dark above, gla-
brous above, glabrous or minutely appressed-puberulent
on the veins beneath, usually with tufts of hairs ca. 1 m
long in the vein axils beneath (= domatia), 2 veins 5-
9/side. Inflorescences terminal or axillary, solitary or 3
when the first branching node is subtended by a pair of
leaves, 8-28 cm long, to 20 cm broad, paniculate with
few opposite lateral branches to 1 2 cm long, peduncles
to 1 8 cm long, bracts 3-6 mm long or leaf-like, lanceo-
late, flowers in distal cymes or clusters of more than 10,
sessile or with pedicels to 3 mm long, bracteoles 0.5-1 .5
mm long. Flowers with a hypanthium 3-4 mm long and
ca. 2.4 mm diam., glabrous or minutely puberulent, ca-
lyx tube minute, calyx lobes 0.3-1 mm long, broadly
triangular; corolla bright rose pink, magenta or the tube
becoming maroon, salverform, tube 6-10 mm long, 1 .3-

188

FIELDIANA: BOTANY

2 mm diam., with 5 longitudinal ribs, glabrous or rarely
sparsely and minutely puberulent, villous at the stamen
attachment within, lobes 3-4 mm long, 4-5 mm broad,
broadly ovate and rounded distally; stamens 5, filaments
2-4 mm long, villous below the middle, anthers 0.8-1
mm long; style 6-8 mm long, stigma lobes ca. 0.5 mm
long, green. Fruits 9-16(-20) mm long, 2-4 mm broad
(to 4 mm when split open), narrowly oblong-tubular,
with obscure longitudinal ribs, valves opening but re-
maining attached at both apex and base, persisting calyx
ca. 2 mm broad and 1 mm high (together with disc);
seeds 2-3 mm long, 0.5-1 mm broad, with a membra-
naceous wing at opposite ends, body of the seed ca. 0.6
mm long.

Trees of evergreen or partly deciduous forest
formations from near sea level to 400(-1000) m
elevation. Flowering in December-April; fruiting
in March-July in Costa Rica and Panama. The
species ranges from the evergreen lowlands of
southern Costa Rica, through Panama, to Colom-
bia.

Macrocnemum glabrescens is recognized by its
broadly rounded stipules, usually thin-obovate
leaves (often on petioles differing in length at the
same node), large open panicles with small bright
pink flowers, broadly overlapping corolla lobes,
and narrow capsules with opened valves remain-
ing attached at the base and apex. This species is
only known from near the Hitoy Cerere reserve,
Dominical, and the Golfo Dulce region in Costa
Rica, but the original description mentions Nicoya
and it is possible that it grows in moist forest on
the southern part of the Nicoya Peninsula. This
species appears to be common in central Panama;
see flower description in Croat ( 1 978, p. 8 1 1 ). This
species resembles Ferdinandusa panamensis.
Standley (1938) listed palo cuadrado as a common
name.

Malanea Aublet

Shrubs, woody lianas, or trees, stems glabrous or pu-
berulent, terete; stipules interpetiolar, simple, caducous.
Leaves opposite, petiolate, entire, pinnately veined, with
domatia. Inflorescences axillary, paniculate with short
opposite or subopposite lateral branches resembling spikes
(or with fasciculate flowers on the central rachis or lateral
branches), flowers sessile or subsessile, bracteolate. Flow-
ers bisexual and radially symmetrical, small, hypanthi-
um turbinate to campanulate, calyx lobes 4, short; co-
rolla funnelform to rotate, pale green to white, tube short,
throat villose within, corolla lobes valvate or slightly
imbricate, villous on the adaxial surface; stamens 4, fil-
ament borne on the throat of the corolla, anthers dor-
si fixed and partly exserted; ovary 2-locular, ovules sol-
itary in each locule and pendulous from the apex of the

locule. Fruits fleshy, oblong or ovoid, 2-locular, exocarp
thin-fleshy, endocarp woody or hard; seeds ellipsoid.

A genus of 20-30 species in tropical South
America and the West Indies. Malanea colom-
biana has been discovered along the Caribbean
coast in Belize, in northern Nicaragua, and in
southeastern Costa Rica. Malanea erecia Seem,
(photo F), said to have been collected by Seemann
in Panama, was not treated in the Flora of Pan-
ama.

Malanea colombiana Standl., Publ. Field Columb.
Mus., Bot. Ser. 7: 66. 1930. Chomelia coclensis
Dwyer, Ann. Missouri Bot. Gard. 67: 97. 1980.

Lianas or clambering shrubs with vining branches,
leafy stems 1.7-8 mm thick, with closely appressed-as-
cending straight hairs 0.4-0.8 mm long, glabrescent and
lenticellate; stipules 6-18 mm long, 2.5-8 mm broad,
oblong or obovate, with many ascending veins, ap-
pressed-puberulent at the base and midrib, deciduous.
Leaves with petioles (4-)7-22 mm long, 0.6-2 mm thick,
appressed-puberulent; leaf blades 5-13 cm long, 2.5-7
cm broad, ovate-elliptic to ovate or oblong-elliptic, apex
obtuse to short-acuminate or acute, base obtuse to slight-
ly rounded and subtruncate, drying stiffly chartaceous,
dark brown above, glabrous above or with appressed
hairs along the midvein and widely scattered elsewhere,
with appressed thin whitish hairs 0.3-1 mm long be-
neath, often with tufts of hairs .in the vein axils beneath.
2 veins 5-7/side, distal secondaries to arcuate-ascend-
ing, 3 veins numerous and closely parallel to form a
straight or sinuous pattern between the secondaries. In-
florescences 4-13 cm long, 3-8 cm broad, pyramidal
with progressively shorter opposite branches, peduncles
2-6 cm long, 0.6-1 .8 mm thick, sericeous, lateral branch-
es l-3(-4) cm long, bracts 1.5-4 mm long and linear,
bracteoles ca. 1 mm long, flowers sessile or with pedicels
to 1 .5 mm long. Flowers 3-4 mm long, hypanthium 0.5-
1 mm long, turbinate, calyx lobes 0.1-0.4 mm long,
broadly triangular or poorly developed; corolla white or
yellowish white, tube 1.5-2.5 mm long. ca. 1 mm diam.
at mouth, lobes 4, 1-2 mm long, 1 mm broad at the
base, villose within; anthers 0.5-0.8 mm long; style ex-
serted, stigma lobes ca. 0.4 mm long. Fruits 5-6 mm
long, 3-4 mm diam., oblong-cylindrical, becoming pur-
plish red and drying black, persisting calyx very short
(0.5 mm) and not elevated.

Plants of evergreen or partly deciduous vege-
tation, 2-200 m elevation. Flowering in June in
Costa Rica (Herrera 3081 CR, MO); fruiting in Au-
gust in northern Nicaragua (Molina 14926 &
151125 EAP, F). The species ranges from Belize to
Colombia.

Malanea colombiana is recognized by its very
small flowers with four-lobed perianth, corolla
lobes conspicuously villous within, fleshy oblong

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

189

fruit, unusual tertiary venation, and climbing hab-
it. The tertiary veins of the leaf are many and
parallel with inconspicuous transverse connec-
tions. The Costa Rican collection has minor ve-
nation that differs by having a more reticulate ar-
rangement of 3 veins with sublineolate 4 veins.
More material is necessary to confirm its place-
ment under this name. The flowers resemble those
of Elaeagia, Figure 39.

Manettia Mutis ex Linnaeus

Herbs or climbers, stems herbaceous or slightly woody,
glabrous or puberulent; stipules interpetiolar, small,
sometimes adnate to the petiole, triangular to laciniate,
persistent. Leaves opposite and decussate, petiolate or
subsessile; leaf blades often narrow, pinnately veined,
domatia absent. Inflorescences axillary (rarely terminal),
cymose to paniculate, fasciculate, umbellate or the flow-
ers occasionally solitary, bracts and bracteoles present,
flowers pedicellate. Flowers bisexual, monomorphic or
distylous, radially symmetrical, small (in Central Amer-
ica) to large, hypanthium turbinate, calyx lobes usually

4 (5, 8), short or long, with glands or teeth in the sinuses;
corolla funnelform or tubular, white, pink, red, lavender,
blue, or yellow, corolla lobes 4(-5), valvate in bud; sta-
mens 4, filaments attached in the throat or at the mouth,
anthers versatile, exserted or included; ovary 2-locular,
ovules numerous and imbricate, vertical on axile pla-
centas, style filiform, stigmas 2 clavate or bifid. Fruits
thin-walled capsules, obovoid or turbinate to subglo-
bose, biloculate and bisulcate, splitting septicidally from
the apex into 2 valves; seeds compressed -discoid, usually
with a thin wing surrounding the central seed.

A tropical American genus of about 100 species.
The slender twining stems, smaller narrow leaves,
small inflorescences, small flowers (in Central
American species), small obovoid capsules break-
ing into two valves, and winged or flattened seeds
with erose margin help to distinguish these plants.
This study has benefited from the herbarium an-
notations of In-Cho Chung and David Lorence.
Manettia luteo-rubra variety paraguariensis (Cho-
dat) Chung (= M. inflata Sprague) is an ornamen-
tal grown for its bright red flowers tipped with
yellow.

Key to the Species of Manettia

la. Leaf blades narrowly lanceolate and often subsessile; seeds ca. 1 mm diam., orbicular with an erose
margin (the wing minute or absent); flowers white, corolla tube 3-5 mm long M. barbata

Ib. Leaf blades rarely consistently lanceolate, petioles usually well developed; seeds 2-3 mm long,
oblong or obicular with a thin expanded marginal wing; flowers white, pink, red, or magenta, corolla
tubes 4-13 mm long 2

2a. Corollas white; capsules 4-6 mm long; leaf blades usually puberulent beneath; 1000-1600 m ele-
vation M. flexilis

2b. Corolla brilliant red to magenta or pink; capsules 6-10 mm long; leaf blades usually glabrous beneath;
30-1 100 m elevation . . M. reclinata

Manettia barbata Oersted, Vidensk. Meddel.
Kjobenhavn 1852: 47. 1853. M. stenophylla J.
D. Smith, Bot. Gaz. 56: 58. 1913. Figure 1.

Slender vines to 3 m high, leafy internodes 0.7-2.5
mm thick, glabrous or subglabrous; stipules 0.5-3 mm,
long, broadly triangular, glabrous. Leaves with petioles
1-5 mm long, 0.3-1.5 mm broad, glabrous; leaf blades
3-10 cm long, 0.4-3 cm broad, lanceolate to narrowly
elliptic-lanceolate or linear-lanceolate, apex tapering
gradually and long-acuminate or acute, base obtuse or
slightly rounded, drying chartaceous, pale grayish green
or grayish brown, glabrous above and below, 2 veins 4-
7/side. Inflorescences 10-15 mm long, axillary cymes of
usually 3 flowers, peduncles only 1-4 mm long, bracts
to 3 mm long, lanceolate, pedicels 2-5 mm long, brac-
teoles 0.5-l(-2) mm long. Flowers with hypanthium 2-
3 mm long, 1 .2-2 mm diam., calyx lobes 1 .2-2 mm long,
0.7-1.2 mm broad, ovate; corolla white, tube 3-6 mm
long, 2 mm diam., lobes 2-3 mm long, glabrous exter-

nally and villous within; anthers ca. 1 mm long; stigma
ca. 0.5 mm long, oblong to lanceolate. Fruits 4-8 mm
long, 3-7 mm broad, persisting calyx to 1.5 mm long
and usually recurved; seeds 0.7-1 mm diam., orbicular,
wing reduced to a dentate rim around the seed, body of
seed ca. 0.6 mm diam., discoid.

Uncommon plants of evergreen forest forma-
tions, from 500 to 1300(-2400?) m elevation.
Flowering in November-January; fruiting in Jan-
uary and March. This species is known only from
central and southern Costa Rica.

Manettia barbata is recogized by its narrowly
lanceolate leaves, twining habit, few-flowered and
short-pedunculate inflorescences, short corolla
tubes, smaller capsules, and circular-discoid seeds
with little evidence of a wing.

190

FIELDIANA: BOTANY

Manettia flexilis Brandegee, Univ. Calif. Publ.
Bot. 6: 196. 1915. M. estrellae Standl., J. Wash.
Acad. Sci. 15: 6. 1925. Figure 1.

Vines to 4 m high, leafy stems 0.7-2 mm long, pu-
berulent with curled whitish hairs ca. 0.2 mm long; stip-
ules 1-2 mm long, adnate to the petiole base, rounded
distally or with short (0.3 mm) thick spike-like projec-
tions. Leaves with petioles 2-10 mm long, 0.3-0.7 mm
thick, puberulent; leaf blades (2.5-)3-7 cm long, 1-3 cm
broad, oblong to ovate or lanceolate, apex acute to long-
attenuate, base obtuse to acute, drying membranaceous
or thin-chartaceous, dark green above, pale grayish green
beneath, glabrous or puberulent above, short pilose be-
neath with thin whitish hairs ca. 0.3 mm long, 2 veins
4-6/side, the upper secondaries strongly arcuate-ascend-
ing. Inflorescences axillary, cymose or umbelliform flow-
ers few (rarely 1), peduncles 1-7 mm long, bracts ca. 2
mm long, flowers closely crowded, pedicels 1-5 mm long.
Flowers with hypanthium 2-3 mm long, densely puber-
ulent with straight or curved multicellular hairs 0.2-0.5
mm long, calyx cup to 0.4 mm long, calyx lobes (2-)3-
4 mm long, 0.8-1.5 mm broad, lanceolate, with erect
glands ca. 0.3 mm long in the sinsues; corolla white or
rose red, tube 4-10 mm long, ca. 1.3 mm diam., short
villose externally, glabrous within, lobes 4, 2-5 mm long,
ca. 0.7 mm broad, ovate to triangular, densely villous
externally; stamens 4, anthers ca. 0.7 mm long. Fruits
5-7 mm long, 4-6 mm broad, broadly obovoid to subglo-
bose, rounded and truncated at the apex, with few scat-
tered hairs; seeds 2-3 mm long, oblong and with a thin
translucent wing, body of the seed ca. 1 mm long and
oblong.

Plants of evergreen and partly deciduous forest
formations in the central Cordilleras, from ca. 1 000
to 1 600 m elevation. Probably flowering and fruit-
ing throughout the year. The species ranges from
southern Mexico to western Panama.

Manettia flexilis is recognized by its smaller cap-
sules, winged seeds, puberulent stems and leaves,
and mid-elevation habitats. This species is infre-
quently collected.

Manettia reclinata Mutis in I ... Mant. PI. 2: 558.
1771. Nacibea coccinea Aubl., Hist. pi. Guiane
96, t. 37, f. 1. 1775. M. coccinea (Aubl.) Willd.
in L., Sp. PI. ed. 4, 1: 624. 1797. M. cuspidata
Bertero in Spreng., Syst. Veg. 1:415. 1825. M.
panamensis Duchass. & Walp., Linnaea 23: 753.
1850. M. costaricensis Wernham, J. Bot. 56,
suppl. 1: 38. 1919. M. orbifera Wernham, loc.
cit. 41. 1919. M, seleriana Loes., Verhand. Bot.
Vereins Brandenb. 65: 107. 1923. Figure 1.

Vines to 4 m high, stems often with 4 narrow longi-
tudinal ridges, leafy stems 0.7-2.5 mm thick, sparsely
puberulent (rarely glabrous), often with a row of minute
(0. 1-0.2 mm) hairs along the longitudinal ridges; stipules

1-2 mm long, broadly triangular, minutely puberulent.
Leaves with petioles 2-14(-20) mm long, 0.6-1.2 mm
broad, sparsely puberulent; leaf blades (2-)3- 10 cm long,
l-3(-5) cm broad, narrowly ovate to ovate-elliptic or
lanceolate, acute or acuminate at the apex, obtuse to
acute at the base, leaves drying thinly chartaceous and
dark green above, paler greenish gray beneath, usually
glabrous above and below, with 4-6 major secondary
veins strongly ascending on each side. Inflorescences ax-
illary or terminal, usually solitary with 2-4 flowers per
node, peduncles 0.5-2.5 cm long, puberulent, pedicels
10-25 mm long (to 35 mm in fruit). Flowers with hy-
panthium 3-5 mm long, oblong, calyx tube to 1 mm
long, calyx lobes 4-8, unequal. 4-7 mm long, 0.3-1.3
mm broad, linear to lanceolate; corolla red, rose red.
deep pink, scarlet, or magenta, tube 6-13 mm long, 2-
2.5 mm diam., glabrate to densely minutely puberulcnt
externally, with yellowish hairs in throat, corolla lobes
4 (5, 6), 2-4 mm long, 2.5-4.5 mm wide, ovate, puber-
ulent externally and glabrous within; stamens 4 (5. 6),
filaments ca. 1 mm long, anthers ca. 3 mm long; style
ca. 1 1 mm long. Fruits 6-10 mm long, 6-9 mm broad,
obovoid or subglobose, glabrous or pubescent with thin
curled hairs to 0.5 mm long, with 8 longitudinal ribs (2
in the sulci), calyx lobes persisting and recurved; seeds
2-2.5 mm diam., thin-discoid and orbicular, with an
erose translucent wing around the margin, body of the
fruit 0.7-1 mm diam.

Plants of wet evergreen forest formations, 0-
1300 m elevation. Flowering in October-May;
fruiting in December-May. The species ranges from
southern Mexico and the West Indies into north-
ern South America.

Manettia reclinata is recognized by its larger red
or pink flowers, few flowers per node, usually gla-
brous leaves, larger rounded capsules borne on
long pedicels, flat orbicular seeds with thin winged
margin, and vining habit. This species appears to
be much more common than its congeners in Cos-
ta Rica.

Mitracarpus Zuccarini

Annual or perennial herbs, erect or decumbent, stems
tctragonous in cross-section, glabrous or puberulent;
stipules adnate to the petiole base to form a sheath with
3-15 slender distal setae (awns), persistent. Leaves op-
posite and decussate, subsessilc to short-petiolate, leaf
blades usually narrow (linear to ovate), chartaceous,
domatia absent. Inflorescences axillary or terminal, cap-
itate or glomerulate with densely crowded flowers, the
heads sometimes subtended by 4 leaf-like bracts, flowers
sessile or subsessile. Flowers bisexual and monomor-
phic, very small, hypanthium turbinate to subglobose.
calyx tube short, calyx lobes 4(-5), unequal, persistent;
corolla white, salverform or funnelform. tube usually
with a ring of hairs within, glabrous or villous in the
throat, corolla lobes 4 (3), valvate in bud; stamens 4,
inserted on the throat, anthers oblong to linear, dorsi-
fixed, included or exserted; ovary 2-3-locular, with 1

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

191

ovule in each locule attached to a peltate placenta in the
center of the septum, style short or long, with 2 short
linear stigmas. Fruits thin-walled capsules, 2- or 3-locular
with circumscissile or transverse dehiscence from below
the middle (the upper portion breaking away with the
persisting calyx and exposing the seeds), septum per-
sisting with the basal part of the capsule; seeds ellipsoid
to oblate or globose, ventral (adaxial) surface with 4 sulci
radiating from a central area to give an X-like pattern
demarking 4 broadly rounded areas, abaxial surface
smooth.

A Neotropical genus of 30-45 species, with most
in Brazil. The genus is very similar to Crusea,
Diodia, and Spermacoce with its weedy-herba-
ceous habit, broad stipules with distal awns, nar-
rowly elliptic leaves, small flowers in verticillate
heads, and two-locular capsules with two seeds.
Mitracarpus is unusual in the circumscissile de-
hiscence of the capsule and the X-like (cruciform)
sulcus on the inner face of the seed.

Mitracarpus hirtus (L.) DC., Prodr. 4: 572. 1830.
Spermacoce hirta L., Sp. PI. ed. 2: 148. 1762.
S. villosa Sw., Prodr. 29. 1788. M. villosus (Sw.)
Cham. & Schlend., Linnaea 3: 363. 1828. M.
breviflorus Gray, PI. Wright. 2, 68. 1853. Fig-
ure 4.

persisting calyx and top of fruit 2-3.5 mm long, body of
the yellowish brown capsule ca. 1 mm long, 0.6-1 mm
broad; seeds 0.5-0.8 long and ca. 0.6 mm wide, oblong-
oblate, yellowish brown, with 4 sulci forming an im-
pressed X on the adaxial face, smooth or pitted on the
abaxial face.

Common weedy plants of open disturbed sites
in both deciduous and evergreen formations, 0-
1200(-1400) m elevation. Flowering primarily in
June-December in Costa Rica. The species is found
in the southwestern United States and throughout
tropical America; it has become established
through much of tropical Africa and has been found
in India, Burma, and the western Pacific.

Mitracarpus hirtus is recognized by its herba-
ceous weedy habit, setose sheathing stipules, nar-
row hispidulous leaves with few secondary veins,
axillary heads of small congested flowers, and un-
usual fruit and seed. It is found in both the Ca-
ribbean and Pacific lowlands and the central up-
lands. There has been considerable confusion
regarding the nomenclature of this species and es-
pecially the applicability of the names; we follow
the recent annotations of C. Dennis Adams. These
plants can be mistaken for species of Spermacoce,
Diodia, and Hyptis (Labiatae).

Herbs to 0.6(-1) m tall, erect or spreading, stems usu-
ally simple distally and with few branches mostly near
the base, often slightly woody at the base, leafy stems
0.7-2.5 mm thick, with 4 longitudinal ridges and some-
what quadrangular in cross-section, usually with curved
thin hairs 0.2- 1.5 mm long and longer (1.5-2 mm) trans-
lucent hairs at the nodes; stipules united to the petiole
base to form a broad sheath l-3(-4) mm long, the straight
distal margin of the sheath with 6-9(-l 3) linear setae 1-
5 mm long. Leaves subsessile, petioles 0-3 mm long, not
clearly differentiated from the base of the blade; leaf
blades 2-5(-8) cm long, 0.5-1. 5(-2) cm broad, narrowly
elliptic to lanceolate, or narrowly elliptic-oblong, apex
tapering gradually and acute, base cuneate, drying stiffly
chartaceous, glabrescent or sparsely hispidulous above,
sparsely hispidulous beneath with hairs ca. 0.3 mm long
and along the margin, 2 veins 2-3/side, strongly as-
cending. Inflorescences axillary or terminal, flowers
densely crowded in capitulae or verticils 4-10 mm high
and 8-15(-20) mm broad, bracts 2-4 mm long but dif-
ficult to distinguish among the sepal lobes in the tightly
congested heads, flowers sessile or subsessile. Flowers
with hypanthium 1-2 mm long, glabrous beneath and
hispidulous distally, calyx lobes 4, unequal, the 2 larger
1.5-2.5 mm long, lanceolate, thick centrally and with a
hyaline margin; corolla white, glabrous or minutely hairy
externally, tube 1.5-2 mm long, 0.3-0.9 mm diam., lobes
4, 0.6-1.1 mm long. 0.4-1 mm broad at the base, ovate
or triangular; stamens 4, filaments very short or absent,
anthers 0.4-0.7 mm long, usually exserted; style 1-1.6
mm long, stigma 0.3-0.5 mm long. Fruits with the top
part coming off to expose the 2 locules and 2 seeds,

Morinda Linnaeus

Small to large trees or shrubs (lianas), stems terete or
quadrangular in cross-section, glabrous or puberulent;
stipules interpetiolar and sometimes slightly connate to
form a short sheath above the petioles (intrapetiolar),
entire to cuspidate or bifid, glabrous. Leaves opposite or
3/node (sometimes only 1 at a flowering node), petiolate,
pinnately veined, often slightly succulent in life, usually
with tufts of hairs (domatia) in the vein axils beneath.
Inflorescences terminal or axillary, with 1 or 2 (rarely
more) capitate heads on a common peduncle (the heads
rarely sessile), large or small bracts sometimes present,
flowers sessile in the capitulum with the basal parts usu-
ally united. Flowers bisexual (rarely unisexual), radially
symmetrical, usually distylous, hypanthium free or unit-
ed with other flowers, calyx tube urceolate or hemi-
spheric, often entire distally, calyx lobes usually minute
or absent; corolla funnelform or salverform, corolla tube
glabrous or puberulent in the throat, puberulent at the
base within, corolla lobes 4-7, narrow and valvate in
bud; stamens 4-7, filaments short and inserted in the
throat of the corolla, anthers dorsifixed and versatile,
included or exserted, connective often prolonged distal-
ly; ovary 2- or 4-locular 1 ovule in each locule, ovules
erect and attached below the middle or at the base of
the septum, style slender, with 2 short or long style
branches. Fruits fused into a fleshy syncarp made up of
the united ovaries (or their bases), often large and turning
white, pyrenes 1 -seeded or part of a 2-4-locular woody
structure; seeds obovoid or reniform.

192

FIELDIANA: BOTANY

A pantropical genus of 50-80 species; most of
the species are native to the Old World. The flow-
ering capitula with the basal parts of the flowers
united, and the latter forming a fleshy largely syn-
carpous fruiting capitulum, make this genus very
distinctive among our Rubiaceae. Three species
have been recorded from southern Central Amer-

ica; two additional native species (M. asperula
Standl. and M. yucaianensis Greenm.) are found
in northern Central America. There is consider-
able variation within our species, and this can make
their identification difficult. Current holdings of
this genus are very limited; all the Costa Rican
material is from near the seashore.

Key to the Species of Morinda

1 a. Leaf blades usually less than 5 cm broad, usually drying grayish or yellowish; branches often scandent
M. royoc

Ib. Leaf blades more than 5 cm broad, usually drying very dark in color; branches never clambering
or scandent 2

2a. Inflorescences usually solitary, with only 1 capitulum per peduncle, fruiting syncarps 3-1 2 cm diam.;
largest leaf blades usually more than 20 cm long and 14 cm broad; only found near the ocean shore
M. citrifolia

2b. Inflorescences 1-3/node, with 1-3 capitula per primary peduncle, fruiting syncarp 0.5-3 cm diam.;

largest leaf blades usually less than 20 cm long and 14 cm broad; lowland rain forests

M. panamensis

Morinda citrifolia L., Sp. PI. 176. 1753. Figure 19.

Shrubs or small trees. (l-)2-8(-12) m tall, trunks to
1 5 cm thick, wood yellow, branchlets quadrangular or
terete, leafy stems 2.5-12 mm thick, glabrous; stipules
6-20 mm long. 5-14 mm broad, oblong to suborbicular-
triangular and rounded distally, glabrous. Leaves well
spaced along the stem, petioles 12-20 mm long, ca. 2
mm broad, glabrous, with slightly winged adaxial mar-
gins; leaf blades 12-28(-40)cm long, 7-16(-24)cm broad,
oblong to broadly elliptic-oblong or ovate-oblong, apex
obtuse to acute (very short acuminate), base broadly
obtuse to cuneate, drying chartaceous or thin-charta-
ceous, often dark in color, glabrous above and below but
with tufts of hairs in the vein axils beneath, 2 veins 6-
8/side. Inflorescences axillary and drying black, solitary
or 2-3/node, capitulae 9-20 mm long, to 20 mm diam.,
oblong to subglobose, peduncles 10-22(-30) mm long,
1.2-2 mm thick, glabrous, flowers sessile and united at
the base. Flowers united basally (hypanthia connate),
calyx tube minute and truncated with a scarious margin:
corolla white, slightly thickened, glabrous externally, tube
6-10 mm long, cylindrical, corolla lobes usually 5, 3-8
mm long, ca. 1.5 mm wide, obtuse, thick-fleshy; stamens
4-6, filaments slightly unequal, anthers to 5 mm long,
becoming twisted; stigma to 5 mm long and 0.8 mm
wide, erose. Fruits united into a syncarp. 4-1 2 cm diam.,
fleshy succulent and irregularly globose to oblate, white
with green "eyes" ca. 8 mm diam. formed by the calyx
and disc of individual flowers, pyrenes to 10 mm diam.;
seeds 3.5 mm long.

Plants usually found near ocean beaches and
lagoons along the Caribbean shore from Honduras
to Panama, 0-20 m elevation. Flowering and fruit-
ing throughout the year (Sanchez 1983). The orig-

inal range of this species was from India to the
East Indies and northern Australia. It has become
naturalized in a number of areas around the Ca-
ribbean.

Morinda citrifolia is recognized by its large white
Annona-\\ke syncarps, large opposite leaves on
thick stems, and seaside Caribbean habitat. Her-
barium specimens can be difficult to distinguish
from M. panamensis. Yema de huevo is a common
name; the fruits are edible.

Morinda panamensis Seem., Bot. voy. Herald 1 36.
1854. Figure 19.

Shrubs or trees. 3-25 m tall, branchlets quadrangular
in cross-section, leafy stems 1.5-6 mm thick, minutely
(0.1 mm) farinose puberulent. glabrescent; stipules ap-
parently with the distal part tearing off to leave a cupulate
base 1-3 mm long forming a shallow cup around the
stem, at first broadly oblong (to 10 mm long and 9 mm
broad) and covering the shoot apex. I^eaves with petioles
5-25 mm long. 1-2 m thick, glabrous or minutely (0.1
mm) puberulent; leaf blades 9-21 cm long, 4-13 cm
broad, broadly elliptic to elliptic-oblong or ovate-oblong,
apex bluntly short-acuminate with tip to 10 mm long,
base obtuse or cuneate, drying thin-chartaceous, dark,
glabrous above, glabrous or minutely (0.2-0.3 mm) pu-
berulent beneath and with dense tufts of hairs (domatia)
in the vein axils. 2 veins 4-7/side. central 2 veins 1-4
cm distant. Inflorescences 1-3/node, capitate, peduncles
4-35(-55) mm long, 0.7-1.5 mm thick, the primary pe-
duncle simple or with 3-4 equal or unequal branches to
3 cm long, usually glabrous, each capitulum 4-10 mm

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

193

long and 5-10 mm wide, oblong to globose, with 9 flow-
ers or more, flowers sessile and united. Flowers united
below, glabrous externally (rarely minutely puberulent)
and drying black, hypanthium partly free distally for 1-
2 mm, calyx tube ca. 0.3 mm long and subentire; corolla
white, tube 5-10 mm long, lobes 4, 3-6 mm long, 1-2
mm broad, narrowly oblong; stamens 4, anthers 2.5-3.5
mm long; stigmas 3.5-5 mm long. Fruit a syncarp, 15-
30 mm diam., subglobose, the calyx tube little (0-0.5
mm) elevated above the surface of the fruit and 2.5-4
mm diam.

Plants of wet evergreen lowland rain forest for-
mations, 0-600 m elevation. Flowering in March-
May in Central America; fruiting in July-Septem-
ber. This species ranges around the Gulf of Mexico
and the Caribbean from Florida (U.S.A.), Mexico,
Central America, and the West Indies to Panama.

Morinda panamensis is recognized by its un-
usual capitula of flowers with united ovaries, seeds
imbedded within a globose syncarp, thin leaves
drying very dark, and rain forest habitat. Some
herbarium specimens can be very similar to spec-
imens of M. citrifolia, and it is possible that there
are intermediates in nature. An unusual collection
from near Upala (Herrera 1783 CR, MO) is mi-
nutely puberulent on all parts.

Morinda royoc L., Sp. PI. 176. 1753. Figure 19.

Shrubs, vines, or small trees, l-3(-7) m tall, often with
scandent branches, leafy branchlets 1-5 mm thick, young
stems with obscure longitudinal ridges and quadrangular
or terete, minutely puberulent with thin hairs 0. 1 mm
long; stipules l-2(-4) mm long, 3-5 mm wide at the
base, triangular or with an awn 1-2 mm long, glabrous
and drying yellowish brown. Leaves with petioles 4-14
mm long, 0.5-1 mm broad, with adaxial margins, gla-
brous or minutely puberulent; leaf blades 4-1 1(-13) cm
long, 1-4.5 cm broad, narrowly elliptic-oblong to oblan-
ceolate or linear-oblanceolate, usually 4 times longer than
broad, apex acute to short-acuminate, base gradually
narrowed and acute or attenuate, drying stiffly charta-
ceous, grayish or yellowish, with the margin often in-
volute, glabrous above and below but often with barbate
hairs in the leaf axils and along the midvein, 2 veins 3-
6/side, arcuate and weakly loop-connected near the mar-
gin. Inflorescences solitary and axillary, the capitulae 4-
12 mm diam., usually oblong, sessile or with peduncles
0-7(-10) mm long, minutely puberulent, flowers sessile
and united. Flowers united together in the lower half of
their ovary, free portion of the hypanthium and calyx
1-3 mm long, calyx entire distally (or obscurely 5-lobed),
minutely puberulent; corolla white, 6-8 mm long, tube
ca. 5 mm long and 2 mm diam., cylindrical, minutely
(0. 1 mm) puberulent externally, lobes 1-2 mm long; sta-
mens 5-6, anthers ca. 2 mm long; style 3-5 mm long,
stigmas ca. 1.5 mm long. Fruit a syncarp, 8-25 mm
diam., irregularly globose, pyrenes 5 mm long.

Plants of the Caribbean lowlands (from pine
savannas to evergreen forest formations) in Cen-
tral America. Flowering in January-August. The
species ranges from Florida (U.S.A.), Mexico,
Central America, and the West Indies to northern
South America.

Morinda royoc is recognized by its vining
branches, smaller narrow leaves that rarely dry
very dark, small syncarpous capitulae, and small
stipules. We have seen no specimens from Costa
Rica, but the species has been collected near Blue-
fields, Nicaragua, and in central Panama. Most
Central American collections are described as
vines. Appunia guatemalensis can appear quite
similar, but the branches are not vining and the
fruit develop separately.

Mussaenda Linnaeus

Shrubs, erect or climbing; stipules interpetiolar, united
and solitary or separate and paired (4/node). Leaves op-
posite, petiolate, pinnately veined and often acuminate
at the apex, domatia absent. Inflorescences terminal, cy-
mose, bracts and bracteoles deciduous; a few flowers of
the inflorescence with a single greatly expanded colorful
and leaf-like expanded sepal lobe. Flowers bisexual and
radially symmetrical (except for those flowers where 1
sepal lobe is greatly enlarged), calyx tube turbinate or
ovoid, calyx lobes 5, 1 lobe greatly expanded in a few
flowers of most species; corolla narrowly funnelform or
salverform, glabrous or puberulent on the outer surface,
corolla tube puberulent in the throat, corolla lobes 5;
stamens 5, filaments very short, borne near the base or
the upper part of the tube, anthers sagittate at the base;
ovary 2-locular with many ovules, style 2-branched. Fruits
usually fleshy and indehiscent (rarely dry and loculici-
dally dehiscent); seeds small and ellipsoid.

A tropical Old World genus of about 200 spe-
cies. The greatly enlarged and colorful sepal lobes
on a few flowers of each inflorescence are found
in most species of the genus and account for the
ornamental appeal of the following species.

Mussaenda erythrophylla Schumach. & Thonn.,
Beskr. Guin. PI. 116. 1827. Figure 16.

Shrubs or woody climbers, 1.5-3(-8) m tall, leafy
branchlets 2-6 mm thick, densely velutinous with yel-
lowish hairs ca. 1 mm long, terete; stipules 4/node, 6-
1 1 mm long, ovate-triangular, glabrous on the inner face.
Leaves with petioles 4-18 mm long, with hairs to 1.5
mm long; leaf blades 4-12(-15) cm long, 3-7 cm long,
ovate to broadly ovate-elliptic or ovate-orbicular, apex
short-acuminate, base obtuse or rounded and subtrun-
cate, densely puberulent on both surfaces with slender

194

FIELDIANA: BOTANY

hairs 0.5-2 mm long, 2 veins 5-7/side, arcuate-ascend-
ing. Inflorescences ca. 10 cm long and 20 cm broad, at
the ends of distal un branched leafy stems, with opposite
or trichotomous branches, densely velutinous. Flowers
densely pubemlent externally, calyx lobes 6- 1 2 mm long,
1-3 mm broad, red; the enlarged sepal lobe leaf-like,
4.5-6.5 cm long, 3-6.5 cm broad, on petioles 2-8 mm
long, with ca. 7 palmate veins, bright red; corolla yellow
to pink, tube 15-30 mm long, lobes 3-9 mm long, 4-7
mm broad, broadly ovate, whitish and papillate puber-
ulent within.

Popular small shrubs or climbers grown for or-
nament in parks and gardens. The enlarged leaf-
like sepal lobes are bright crimson to deep red and
give a very colorful effect. They flower throughout
the year but usually do not produce fruits.

4 cm diam., peduncles 2-4 cm long, flowers many and
closely congested. Flowers not conivent basally; corolla
7-8 mm long, narrowly funnelform, lobes 5, ca. 1.5 mm
long; style long-exserted. Fruits tightly congested in the
spherical heads, splitting into 4 parts, seeds minute.

The solitary pedunculate globose heads and
larger drooping leaves distinguish this introduced
species. It is fast-growing in open sites until it
reaches about 10 m in height. Common names
used in southeast Asia are kadam, kedam. and
laran. The literature of this tree is to be found
under Anthocephalus, now a synonym ofBreonia.
See J. D. E. Fox, Anthocephalus chinensis, the La-
ran Tree of Sabah. Econ. Bot. 25: 221-233. 1971.

Neoiamarckia Bosser

Nertera Banks & Solander
Nomen conservandum

Trees; stipules interpetiolar, triangular, deciduous.
Leaves opposite, petiolate, entire, domatia absent. In-
florescences terminal, usually solitary, capitate and pe-
dunculate, flowers densely congested. Flowers bisexual
and radially symmetrical, calyx lobes small; corolla fun-
nelform, corolla lobes 45, imbricate in bud; ovary 2-
locular at the base, placentation axile with many vertical
ovules; stigma fusiform. Fruits capsules, loculicidally de-
hiscent from the apex, thin-walled; seeds 1-5 in each
locule, small and angular.

A genus of two species of southeast Asia. One
species is occasionally planted in Costa Rica.

Neoiamarckia cadamba (Roxb.) Bosser, Bull. Mus.
Hist. Nat. Paris 4 ser. sect. B. Adansonia 6: 247.
1984. Nauclea cadamba Roxb., Fl. Ind., ed. 1,
2: 121. 1824. Anthocephalus cadamba (Roxb.)
Miq., Fl. Ind. Bat. 2: 135. 1856. A. morindae-
folius Korth., Verh. Nat. Gesch. 1 54, t.. 48. 1 842.
A. indicus A. Rich., Mem. Soc. Hist. Nat. Paris
5: 238. 1834, nom. illeg. Cephalanthus chinensis
a uctt.. non Lamarck; A. chinensis auctt.

Trees, 5-15(-30) m tall, fast-growing in early stages,
leafy stems 2.3-10 mm thick, glabrous, quadrangular;
stipules oblong, covering the buds, caducous. Leaves
usually somewhat pendant, deciduous, petioles 22-55
mm long, 1-4 mm thick, glabrous or minutely puberu-
lent, drying dark; leaf blades (7-) 13-3 4 cm long, (5-)6.5-
18 cm broad, ovate to ovate-oblong or oblong, apex
abruptly narrowed and blunt or short-acuminate, base
obtuse to rounded and truncate or subcordate, drying
brownish, glabrous above, glabrous or minutely (0.05
mm) puberulent beneath, 2 veins (4-)8- 1 6/side, 3" veins
subparallel. Inflorescences solitary, terminal on short lat-
eral branchlets, each with a single globose capitulum 3-

Perennial herbs, creeping and repent, glabrous or
sparsely puberulent, much-branched and often rooting
at the nodes (often forming mats); stipules interpetiolar,
small, partly united to the petiole bases, entire or with
2 teeth, persistent. Leaves opposite, very small, petiolate
or sessile; leaf blades ovate to rounded, glabrous or pu-
berulent, venation pinnate or subpalmate; domatia ab-
sent. Inflorescences of solitary flowers, axillary or ter-
minal, the flowers sessile or subsessile. Flowers bisexual,
radially symmetrical, hypanthium ovoid to turbinate,
calyx tube truncated and entire or slightly lobed; corolla
broadly funnelform to tubular, corolla lobes 4-5, valvate
in bud; stamens with thin filaments attached to the base
of the tube, anthers basifixed, the connective apiculate.
exserted; ovary 2-locular, with 1 ovule borne from the
base of each locule, style deeply 2-branched and slender.
Fruits fleshy drupes (rarely dry), rounded, with 2 plano-
convex pyrenes.

A genus of 6- 1 2 species found in Australia, New
Zealand, Malaya, Indonesia, South China, and
some Pacific islands; in addition, a single species
ranges from Mexico through the higher moist
mountains of Central and South America to Chile.

Nertera granadensis (Mutis ex L.f.) Druce, Bot.
Soc. Exch. Club Brit. Isles 1916: 637. 1917. Go-
mozia granadensis Mutis ex L.f., Suppl. PI. 1 29.
1781. Nertera depressa Banks & Solander ex
Gaert.. Fruct. et Sem. PI. 1 : 1 24. 1 788. Figure 3.

Creeping prostrate or pendant herbs to 5 cm tall and
up to ca. 1 m long, terrestrial or low epiphytic, often
forming mats, stems 0.3-1 mm thick and rooting at many
nodes, often tetragonous in cross-section in life, glabrous
or with a few scattered hairs; stipules 0.3-1 mm long,
triangular, united at the base with the bases of the pet-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

195

ioles, persisting. Leaves quite variable on different plants,
petioles 0.7-6(-9) mm long, 0.2-0.4 mm broad, gla-
brous, sometimes sulcate above; leaf blades (1.5-)2.5-
8(-13) mm long, (1.2-)2-7(-12) mm broad, ovate-tri-
angular to ovate-orbicular or ovate-deltoid, apex obtuse
to rounded, sometimes with a minutely apiculate tip,
base rounded and truncate to obtuse, often decurrent on
petiole in larger leaves, drying thin-chartaceous but
semisucculent in life, glabrous above and below (rarely
puberulent), 2 veins 2-4/side, strongly ascending. Inflo-
rescences absent, the solitary sessile flowers axillary to
distal leaves, the flowers covered by surrounding leaves
and very difficult to see in pressed herbarium material.
Flowers minute (ca. 4 mm long), usually glabrous exter-
nally, hypanthium 0.5-1 mm long, calyx lobes absent;
corolla 1-3 mm long, greenish to yellow or white, tube
0.6-1 mm long, widely funnelform (subcampanulate),
lobes ca. 0.5 mm long, glabrous; anthers 0.3-0.5 mm
long. Fruits sessile, 4-7 mm diam., globose, bright or-
ange, orange-red, or deep red, fleshy part often translu-
cent, pyrenes 2-2.5 mm long and 1.2-2 mm wide, ob-
long.

Plants of wet evergreen montane forest forma-
tions, from (1000-)! 500 to 3400 m elevation in
Central America. Fruiting throughout the year in
Costa Rica. This species ranges from Mexico,
through the high mountains of Central America
and South America, as far south as Chile. This
species, when interpreted in a broad sense, is said
to be found throughout the range of the genus.

Nertera granadensis is a very distinctive species
with its low-creeping and mat-forming habit, very
small paired leaves, lack of pubescence, solitary
little flowers, bright orange-red berries, and re-
striction to higher-elevation wet forest habitats.
Collections with larger (10 mm) leaves can look
very different from those with smaller (4 mm)
leaves. It is common in some montane rain forests,
covering mossy banks and old logs. In Costa Rica
it seems to be restricted to the central Volcanic
Highlands and the Cordillera de Talamanca. The
brightly colored berries and mat-forming ability
have made this species valuable as a greenhouse
or moist-area ornamental. These plants resemble
small collections of Didymaea.

Oldenlandia Linnaeus

REFERENCES E. E. Terrell, Synopsis of Olden-
landia (Rubiaceae) in the United States. Phyto-
logia 68: 125-133. 1990. E. E. Terrell and W. H.

Lewis, Overview and annotated list of North
American species of Hedyotis, Houstonia, Olden-
landia (Rubiaceae) and related genera. Phytologia
71: 221-243. 1991. D. A. Halford, Review of the
genus Oldenlandia L. (Rubiaceae) and related gen-
era in Australia. Austrobaileya 3: 683-722. 1992.

Annual or perennial herbs (rarely subshrubs), stems
erect or decumbent, simple or branched, glabrous or
puberulent; stipules interpetiolar, small, acute to acu-
minate, often united to the base of the petioles to form
a short sheath, with 1 -several awns. Leaves opposite and
decussate, sessile or short-petiolate; leaf blades usually
narrow, often with a stiff mucronate tip, domatia absent.
Inflorescences axillary or terminal, open and branched
panicles and cymes or the flowers solitary or fasciculate
in the leaf axils, flowers sessile to long-pedicellate. Flow-
ers bisexual, monomorphic or distylous, usually small,
hypanthium turbinate to hemispheric, calyx lobes
(3-)4(-5-8), equal, narrowly to broadly triangular; co-
rolla rotate or salverform to funnelform, white to lav-
ender, pink, or purple, tube cylindrical, the throat often
puberulent, corolla lobes (3-)4(-5), valvate in bud; sta-
mens (3-)4(-5), anthers dorsifixed and sessile or on short
filaments inserted on the throat, included or exserted;
ovary 2-locular, usually with many horizontal ovules on
peltate placentas attached near the base of the septum,
style filiform, stigmas 2, linear to subglobose. Fruits cap-
sular, often papery, globose to oblong, usually with a
loculicidally dehiscent apex (beak), later also septicidally
dehiscent; seeds usually many (50-100), angular to
subglobose, smooth to reticulate or alveolate, often be-
coming viscid when moistened.

A pantropical and subtropical genus with prob-
ably ca. 100 species but with problems regarding
generic circumscription. Some authors have sug-
gested placing this genus under Hedyotis or Hous-
tonia, while others divide it into smaller genera;
see the references cited above. Our plants of this
genus are recognized by their delicate herbaceous
habit, the very small or linear-lanceolate leaves,
the axillary flowers or small few-flowered inflo-
rescences, the minute flowers on filiform pedicels,
and the broadly rounded thin-walled capsules with
small seeds.

Many Costa Rican collections formerly identi-
fied as O. corymbosa, O. herbacea, and O. land-
folia are probably the same species: O. corymbosa.
This conclusion is based on an overview of the
material and following the keys and descriptions
of Verdcourt in the Flora of Tropical East Africa
(1976).

Key to the Species of Oldenlandia

la. Leaf blades less than 5 mm long, usually broadly ovate; plants often forming small mats (not yet
collected in Costa Rica) O. callitrichoides

196

FIELDIANA: BOTANY

1 b. Leaf blades more than 5 mm long, narrowly elliptic-ovate to lanceolate or linear; plants diffuse or
foming loose mats 2

2a. Corolla 2-1 1 mm long; fruit with a beak 0.8-1 mm long (not yet known to occur in Costa Rica)
O. herbacea

2b. Corolla 0.5-2 mm long; fruit with a small or well-developed beak 3

3a. Flowers pink or white marked with lavender, flowers often 2-3/peduncle; fruit globose to globose-
oblate, rounded at the base but not saccate, with a beak 0.1-0.6 mm high O. corymbosa

3b. Flowers white, flowers usually solitary; fruit broader than long and distinctly rounded (saccate) at
the base, with a beak ca. 1 mm long O. lancifolia

Oldenlandia callitrichoides Griseb., Mem. Am.
Acad. 2, 8: 506. 1863. Oldenlandiopsis callitri-
choides (Griseb.) Terrell & W. H. Lewis, Brit-
tonia42: 185. 1990.

Prostrate herbs to 10 cm tall, much-branched and of-
ten forming mats, leafy stems 0.1-0.5 mm thick (when
dried), glabrous; stipules adnate to petiole base and form-
ing a short sheath to 0.5 mm long, sheath entire distally
or with a small appendage, glabrous. Leaves very small,
petiole 0.5-2 mm long; leaf blades 1-3.5 mm long, 1-3
mm broad, rounded-triangular to ovate, apex bluntly
obtuse or rounded, base broadly obtuse to rounded and
subtruncate, somewhat decurrent on petiole, drying
membranaceous, greenish, glabrous above and below (or
with a few broad-based hairs ca. 0.2 mm long), 2 veins
3-4/side, strongly ascending. Inflorescences of solitary
flowers in distal leaf axils, usually only 1 flower per node,
pedicels 1-9 mm long, filiform, glabrous. Flowers 2-3
mm long, hypanthium 1-1.5 mm long, calyx lobes 0.2-
0.6 mm long. Fruits 2-3 mm long (including the calyx
lobes), ca. 1 mm diam., distally truncated, with a few
whitish raphides on the greenish surface, calyx lobes ca.
0.5 mm long.

Distinctive little plants usually found on sandy
soil in open sunny situations, in tropical lowland
sites. This species has the smallest leaves found
among Central American Rubiaceae. In general
aspect, these plants resemble Pilea hernariarioides
(Sw.) Weddell and P. microphylla (L.) Liebm. This
species is known from the Yucatan peninsula and
from central Panama, but it has not been collected
elsewhere in Central America. Terell and Lewis
based a new genus on this distinctive species (see
synonomy).

Oldenlandia corymbosa L., Sp. PI. 1 19. 1753. Fig-
ure 3.

liptic-oblong, apex abruptly obtuse to acute and with a
minute (0.2 mm) apiculate tip, narrowed to the base,
drying chartaceous and with the margins usually revo-
lute, dark above, glabrous or minutely scabrid above and
below (or with the cystoliths appearing like minute tri-
chomes), 2 veins usually obscure. Inflorescences axil-
lary, with single flowers on slender pedicels or cymose-
umbellate with 3 (2-5) flowers on a common filiform
peduncle 2-8 mm long, both singular and umbellate in-
florescences often present on the same stem or node,
pedicels 2-6(-12) mm long, filiform and glabrous. Flow-
ers very small, hypanthium 0.5-1 mm long, obconical
to cupular, glabrous, calyx lobes 4, 0.7-1.5 mm long,
narrowly triangular, corolla rotate, white or white marked
with lavender, blue, or purple, tube 0.6-1 mm long, lobes
0.5-1.2 mm long; stamens 4, sessile on the distal half of
the tube, anthers ca. 0.4 mm long. Fruits 1.7-2.7 mm
diam., body of the fruit 1.2-2 mm long and truncated
at the apex, raphides prominent, calyx lobes ca. 0.5 mm
long, borne on slender pedicels 4-1 1 mm long; seeds ca.
0.3 mm long, ellipsoid to depressed conic, reticulate.

Plants of open weedy sites on sandy soils in
evergreen and in seasonally deciduous formations,
from near sea level to 1 500 m. Probably flowering
and fruiting throughout the year. The species has
been found in both the Caribbean and Pacific low-
lands and in the Meseta Central. The species ap-
pears to have originated in Africa but is now wide-
spread in the tropics.

Oldenlandia corymbosa is recognized by its short
weedy habit, almost linear subsessile leaves, mi-
nute flowers, and rounded capsules. The senior
author believes that this is probably the correct
name for all the Central American material for-
merly called O. corymbosa, O. herbacea, and O.
lancifolia. Separation of species on the basis of
inflorescence differences or whether or not the stip-
ules are bifid appears to be trivial.

Herbs to 15(-30) cm tall, prostrate or erect, with few
to many branches, leafy stems 0.2-1.3 mm thick, gla-
brous or with a few hairs at the nodes; stipules with a
sheath 0.3-2 mm long, with 2-5 slender unequal awns
to 1.5 mm long. Leaves sessile or with petioles to 1 mm
long; leaf blades (7-)15-30M5) mm long, 0.5-3.5(-6)
mm broad, linear to linear-oblong or very narrowly el-

Oldenlandia herbacea (L.) Roxb., Hort. Bengal.
11. 1814. Hedyotis herbacea L., Sp. PI. 102.
1753.

Annual or perennial herbs, 5-60 cm tall, erect to
spreading and decumbent, stems glabrous, with 4 lon-

BURGER & TAYLOR: FLORA COSTAR1CENSIS. RUBIACEAE

197

gitudinal ridges; stipules forming a very short (0.1-0.5
mm) sheath, truncate and with a few awns to 0.3 mm
long. Leaves sessile; leaf blades 6-55 mm long, 1-4 mm
wide, linear to linear-lanceolate, apex acute, base cu-
neate, drying chartaceous, glabrous or with a few short
hairs along the margins. Inflorescences axillary, flowers
1 or 2/node, pedicels 3-30 mm long, filiform, glabrous.
Flowers usually isostylous, hypanthium 0.5-1 mm long,
ovoid, glabrous to papillate or puberulent, calyx lobes
0.5-2.5 mm long, narrowly triangular, scabridulous on
the margins; corolla white or lavender, or the tube green
and the lobes with purple marks, corolla tube 2-1 1 mm
long, cylindrical, lobes 1-3 mm long, ovate. Fruits 2.2-
5 mm long, 1.5-2 mm diam.. subglobose to ovoid, drying
pale yellowish, glabrous to puberulent, crowned by the
dark calyx lobes, beak 0.8-1 mm long; seeds 0.2-0.4 mm
long, ovoid to ellipsoid, reticulate, brown.

Weeds of open sunny sites. Originally from Af-
rica but now naturalized in Asia and parts of the
Americas. The above description is based on
Verdcourt (1976). This species has not yet been
recorded from Costa Rica. Central American ma-
terial earlier placed under this name is likely to be
Oldenlandia corymbosa or O. lancifolia.

Oldenlandia lancifolia (Schumach.) DC., Prodr. 4:
425. 1830. Hedyotis lancifolia Schumach. in
Schumach. & Thonn., Beskr. Guin. PI. 72. 1 827.
Manettia bocaturensis Dwyer, Ann. Missouri
Hot. Gard. 67: 278. 1980.

Perennial (rarely annual) herbs to 90 cm long, pros-
trate or creeping, usually much-branched near the base
and with simple distal stems, leafy stems glabrous or
rarely minutely scabridulous; stipules forming a sheath
to 1 mm long, with 2-5 slender awns 0.5-1.5 mm long.
Leaves sessile or subsessile; leaf blades 1 0-60 mm long,
2-12 mm broad, linear or linear-lanceolate to narrowly
elliptic, apex acute, base cuneate, drying chartaceous,
dark green above, glabrous on both surfaces but often
minutely scabrid along the revolute margins, 2 veins 3-
5/side. strongly ascending but thin and difficult to see.
Inflorescences axillary, of solitary flowers or several flow-
ers on very short peduncles (or reduced lateral shoots),
pedicels 5-30 mm long, filiform, glabrous or scabridu-
lous. Flowers monomorphic, ca. 3 mm long, hypanthium
ca. 0.8 mm long and 1.5 mm diam., cupulate, glabrous
or with scattered short hairs, calyx lobes 1-1 .8 mm long,
triangular, glabrous or scabridulous; corolla white (some-
times tinged with pink or purple), tube ca. 1 mm long,
lobes 1-2 mm long; stigma lobes 0.7-1 .4 mm long. Fruits
2-3 mm long (including the 1 mm tall beak), 3.2-5 mm
in diam., depressed subglobose; seeds 0.3-0.4 mm long.

Herbs of both seasonally deciduous formations
and lowland rain forest areas. The species is wide-
spread in tropical Africa and has been introduced
to parts of the South America and the West Indies.
The preceding description follows that of Verd-

court (1976). Central American material earlier
ascribed to this species is probably O. corymbosa.
Likewise, Steyermark's use of this name in the
Flora de Venezuela (1974, pp. 408-41 1), may be
incorrect.

Osa Aiello

REFERENCE A. Aiello, A re-examination of
Portlandia and associated taxa. J. Arnold Arbor.
60: 38-126. 1979.

Small trees, stems slightly expanded at the nodes; stip-
ules united (interpetiolar), small, with an acute central
lobe. Leaves opposite, petiolate, attenuate at the apex,
drying thin-chartaceous and grayish green, entire, pin-
nately veined, domatia absent. Inflorescence of single
axillary flowers, borne on pedicels continuous with the
base of the hypanthium/ovary. Flowers large, radially
symmetrical, glabrous, calyx with 6 long narrow lobes;
corolla with a long tube and distally funnelform, corolla
lobes 5, broadly triangular; stamens 5, anthers linear;
ovary 2-locular, placentas borne on the septum, with ca.
1 ovules in each locule, style filiform. Fruits thin-walled
capsules, ellipsoid, with 6 longitudinal ribs, apparently
opening septicidally, the calyx lobes persisting distally;
seeds biseriate, slightly compressed, lacking wings, not
imbricate, with persisting funicle.

A monotypic genus known only from the Osa
Peninsula of Costa Rica. No other species of Costa
Rican Rubiaceae has such large flowers. This ge-
nus is related to Portlandia and to a lesser extent
to Hintonia.

Osa pulchra (D. Simpson) Aiello, J. Arnold Arbor.
60: 116. 1979. Hintonia pulchra D. Simpson,
Phytologia 29: 277. 1974. Figure 15.

A small tree or slender treelet, 2.5-15 m tall, leafy
stem 1 .5-6 mm thick, glabrous, slightly expanded below
the node; stipules 1-3 mm long, the broad base 1-2 mm
high with a narrowed acute tip 0.5-1.5 mm long, per-
sisting. Leaves with petioles 4-1 2 mm long, 0.7-1 .5 mm
thick, glabrous, poorly differentiated from the lamina
base; leaf blades 12-19 cm long, 3-6 cm broad, elliptic-
oblong or narrowly oblong, apex gradually narrowed and
acuminate, base acute to attenuate, drying thin-charta-
ceous and grayish green, glabrous above and below, 2
veins 6-8/side, 3 veins obscure, domatia lacking. Inflo-
rescences of solitary flowers in the axils of distal leaves,
pedicels ca. 1 5 mm long but merging imperceptibly into
the flower base, ca. 1 mm diam., glabrous, drying black.
Flowers glabrous externally, probably slightly pendu-
lous, apparently homostylous, hypanthium ca. 6 mm
long, obconic, calyx lobes 18-40 mm long, 0.5-2.5 mm
broad; corolla long-tubular and funnelform distally
(trumpet-shaped), white, tube 1 7-27 cm long, 4-5 mm

198

FIELDIANA: BOTANY

diam. and gradually flaring to 80 mm diam. at the mouth,
lobes ca. 1 5 mm long and 3040 mm broad at the base,
broadly obtuse; anthers more than 20 mm long, 0.7 mm
thick. Fruits ca. 3 cm long and 1.5 cm broad, oblong-
ellipsoid, dark brown, the sepal lobes persisting but
breaking off; seeds ca. 6 mm long, testa tuberculate.

This species is known from only a few collec-
tions in lowland rain forest near Rincon de Osa
at ca. 50 m elevation on the Osa Peninsula. Flow-
ering in January-February, with immature fruits
in June and mature fruits in January.

Osa pulchra is distinguished by its very large
trumpet-shaped flowers that are thin in texture.
No other Costa Rican member of the Rubiaceae
has so long a flower. The long narrow basal tube
suggests pollination by a long-tongued sphingid
moth. The flowers are reminiscent of those of or-
namental species ofBrugmansia (formerly includ-
ed in Datura, Solanaceae).

Palicourea Aublet

REFERENCE C. M. Taylor, Revision of Pali-
courea in Mexico and Central America. Syst. Bot.
Monogr. 26: 1-102. 1989.

Shrubs or small trees, glabrous or pubescent, stems
terete, trigonous or quadrangular; stipules interpetiolar
and often also intrapetiolar to form a sheathing tube with
2 triangular lobes or awns on each side (4/node) or some-
times elongated interpetiolarly into emarginate or bi-
lobed apices with the intrapetiolar sheath poorly devel-
oped, colleters present within at the base. Leaves opposite
and decussate or rarely 3-4/node, petiolate or rarely ses-
sile; leaf blades entire and often elliptic, apex acute to
acuminate, base rounded to acute, drying chartaceous to
subcoriaceous, pinnately veined, domatia present or ab-
sent. Inflorescences terminal, paniculate with opposite
or alternate branching, variable in form (from elongate
and racemose to thyrsoid or broadly corymbose) with
flowers usually in distal cymose groups, peduncles and
branches of the inflorescences often colored red, orange,
yellow, purple, or blue (rarely green), glabrous or pu-
bescent, bracts and bracteoles usually present (in Costa
Rica), flowers usually pedicellate. Flowers bisexual and
usually distylous, radially symmetric or slightly bilat-

erally symmetric when the corolla is curved or gibbous
on 1 side, glabrous or pubescent externally, hypanthium
usually turbinate, calyx tube with 5 calyx lobes (rarely
truncated or spathaceous); corolla yellow, orange, red,
purple, or blue (rarely white or cream), tubular to fun-
nelform or salverform, membranaceous to camose. the
corolla tube straight or curved, usually expanded (gib-
bous) near the base, glabrous or puberulent externally,
usually with a ring of hairs within the lower half of the
tube (more rarely with the hairs in the upper half), corolla
lobes 5, valvate in bud; stamens 5, inserted in the middle
or on the throat of the tube, anthers dorsifixed and bifid
at the base, included or exserted; ovary 2(-6)-Iocular,
with 1 erect ovule from a basal placenta in each cell,
stigmas 2-branched. Fruits fleshy, exocarp bluish to blu-
ish black or purplish black; pyrenes usually 2. hemi-
spheric with ca. 5 longitudinal ribs on the rounded back,
usually with a longitudinal sulcus on the flattened inner
face.

A Neotropical genus of about 200 species, rang-
ing throughout moist tropical vegetation from
Mexico and the West Indies to southern Brazil
and Paraguay. Most species are South American;
there are 3 1 in Costa Rica. As in Hoffmannia. this
genus has speciated profusely at middle and higher
(1000-2800 m) elevations. These plants are rec-
ognized by their colorful terminal inflorescences,
corolla tubes often slightly bent or expanded at the
base, ovaries with a single erect ovule in each loc-
ule, and the fleshy fruit usually with longitudinal
ribs when dried. The infructescences usually be-
come purple regardless of their color during an-
thesis. The stipules forming a short tube above the
node and with two distal awns or lobes, and the
leaves with many secondary veins are additional
characteristics distinguishing many Costa Rican
species.

Palicourea is separated from the closely related
genus Psychotria by the ring of hairs in the interior
of the lower half of the corolla tube, a tendency
for the corolla tube to be inflated or gibbous near
the base, the more colorful inflorescences, and the
generally larger more colorful corollas. Despite
these differences, dried fruiting material may be
very difficult to separate.

Key to the Species of Palicourea

la. Plants of lowland formations, rarely found from as high as 1000 m elevation 2a

Ib. Plants of montane forest formations, from (900-)1000 to 3200 m elevation 5a

2a. Flowers lavender or white marked with purple; stipule forming a sheath to 8 mm long [leaves

with 9-15 pairs of secondary veins; from wet Caribbean slopes at 500-1000 m elevation]

P. copensis

2b. Flowers red, orange, or yellow; stipular sheath 0-2 mm long; wide-ranging species 3a

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

199

3a. Leaves 3/node, stipular teeth 6/node, 8-1 2 mm long and persisting, leaf blades to 8 cm broad

P. triphylla

3b. Leaves 2/node, stipular teeth or lobes 4, 2-9 mm long, often deciduous, leaf blades to 16 cm

broad 4a

4a. Stipular lobes rounded; leaf blades 1 5-28 cm long, to 1 6 cm broad P. guianensis

4b. Stipular teeth acute; leaf blades 7-14 cm long, to 7 cm broad P. crocea

5a. Plants pilose with hairs 1.5-2 mm long; stipules united and bifid or emarginate on each side;

flowers yellow and pilose P. standleyana

5b. Plants glabrous to puberulent, the hairs not exceeding 1 mm in length; stipules 4-lobed, bifid or

emarginate; flowers white, yellow, blue, or purplish 6a

6a. Calyx teeth or calyx lobes regularly more than 1.5 mm long [often persisting in fruit and with
slightly smaller dimensions; flowers usually blue, purple, or white (red to yellowish only in P.

macrocalyx and P. orosiana)] 7a

6b. Calyx teeth or calyx lobes 0.2-1 .5 mm long 1 7a

7a. Calyx lobes more than 3.5 mm long 8a

7b. Calyx lobes less than 3.5 mm long 1 5a

8a. Calyx spathe-like and splitting down 1 side (to 20 mm long); corolla tube 3-4 cm long;

leaves with 1 2-22 pairs of 2 veins P. spathaceae

8b. Calyx 5-lobed (not spathe-like); corolla tubes 8-26 mm long; leaves with up to 19 pairs

of 2 veins 9a

9a. Corolla 20-26 mm long, ca. 4 mm diam. [calyx lobes 4-14 mm long, lanceolate; bracts
of the inflorescence to 25 mm long and lanceolate; secondary veins 12-18 pairs; 1400-

2200 m elevation] P. hammelii

9b. Corolla tubes less than 20 mm long and 4 mm diam lOa

lOa. Young stems minutely puberulent 1 la

lOb. Young stems glabrous 1 2a

1 la. Calyx lobes 10-14 mm long, corolla white; stipule lobes 7-20 mm long; leaves

with 15-19 pairs of 2 veins; western Panama P. bella

1 Ib. Calyx lobes 6-1 1 mm long, corolla yellow; stipule lobes 6-17 mm long; leaves

with 8-12 pairs of 2 veins; central Costa Rica P. orosiana

12a. Leaves with 17-20 pairs of 2 veins; stipule lobes 2-9 mm long [corolla white]; floral

bracts 12-16 mm long; central Costa Rica P. bellula

12b. Leaves with 7-14 pairs of 2 veins; stipule lobes 2-10 mm long 12a

1 3a. Floral bracts 10-20 mm long, persisting and enclosing the flowers; corollas white;

2600-2800 m elevation in the Talamanca mountains P. skotackii

13b. Floral bracts 6-10 mm long, deciduous; corollas white with yellow or blue; Cor-
dillera de Tilaran and central volcanic highlands, 1400-2200 m elevation . . 14a
14a. Corolla white and blue; stipule lobes 2-4 mm long; inflorescence to 22 cm long and

narrowly thyrsiform P . albocaerulea

14b. Corolla white and yellow; stipule lobes 4-10 mm long; inflorescences to 15 cm long

and paniculate P. macrocalyx

1 5a. Young stems usually puberulent [flowers white; inflorescences narrowly thyrsiform; leaves

2-7 cm broad; 800-1 600 m elevation] P. lancifera

1 5b. Young stems glabrous 1 6a

16a. Leaves l-2(-3) cm broad, blades to 8(-10) cm long; corolla blue and white (compare also

1 7a below) P. salicifolia

1 6b. Leaves becoming larger and broader; flowers variously colored 1 7a

17a. Flowers purple to violet, blue, or white (if reddish then reddish purple or lavender) 18a

1 7b. Flowers yellow to orange or red-orange 24a

1 8a. Leaf blades not more than 9 cm long [0.7-3.5 cm broad]; young stems often slightly puberulent

19a

1 8b. Leaf blades usually becoming more than 9 cm long; young stems glabrous or puberulent
beneath the inflorescence . . 20a

FIELDIANA: BOTANY

19a. Inflorescences blue, corolla white to violet or blue; leaves thick, petioles thick; 1800-

3200 m elevation p. adusta

19b. Inflorescences yellowish green, corolla white to yellow; leaves thin, petioles slender;

1 300-2300 m elevation p, montivaga

20a. Leaf blades l-3(-4) cm broad, to ca. 14 cm long 2 la

20b. Leaf blades usually becoming more than 4 cm broad, to 24 cm long 22a

2 la. Inflorescences thyrsiform and purple, to 16 cm long; calyx lobes ca. 0.4 mm long . . .

P. angustifolia

21b. Inflorescences paniculate and purple, to 8 cm long; calyx lobes ca. 1 m long

P. purpurea

22a. Corolla white, tube 3-6 mm long; calyx lobes ca. 0.3 mm long; leaves with 9-1 1 pairs of 2

veins P. tilaranensis

22b. Corolla blue to purple, tube 5-10 mm long; calyx lobes 1-3 mm long; leaves with 11-21

pairs of 2 veins 23a

23a. Inflorescence purple, paniculate; leaf blades to 26 x 13 cm, with 11-19 pairs of 2 veins

P. discolor

23b. Inflorescence blue, thyrsiform; leaf blades to 18 x 8 cm, with 15-21 pairs of 2 veins ....

P. brenesii

24a. Stems and veins on the lower leaf surfaces densely puberulent with short stiff scurfy or slender

hairs; flowers puberulent P. vestita

24b. Stems glabrous to sparsely puberulent; flowers glabrous or puberulent 25a

25a. Stipule lobes/teeth 0.5-2 mm long; leaf blades 3-12 cm long and 1-3.5 cm broad 26a

25b. Stipule lobes/teeth 2-8 mm long; leaf blades usually more than 7 cm long and 3 cm broad . . 27a
26a. Flowers yellow or white with yellow and purple; leaves with 12-18 pairs of 2 veins; 1200-

1 600 m elevation P. garciae

26b. Flowers white and yellow; leaves with 6-9 pairs of 2 veins; wide ranging in Costa Rica;

1 1 00-2000 m elevation P. montivaga

27a. Inflorescence branches red to orange (to purple in fruit); corolla tube 6-10 mm long, orange;

(800-) 1200- 1700 m elevation P. padifolia

27b. Inflorescence branches yellow to greenish yellow; corolla tube 7-1 3 mm long, yellow; (800-)1000-

2300 m elevation 28a

28a. Corolla tube 7-1 1 mm long; stems glabrous; leaves with 8-13 pairs of 2 veins .... P. padifolia
28b. Corolla tube 1 1-13 mm long; stems puberulent or glabrous; leaves with 1 1-19 pairs of 2 veins

. P. lasiorhachis

Palicourea adusta Standl., J. Wash. Acad. Sci. 18:
279. 1928. Figure 53.

Small shrubs or rarely little treelets, 0.5-3(-6) m tall,
leafy branchlets 0.7-6 mm thick, glabrous or with minute
(0.1-0.5 mm) appressed hairs, becoming terete; stipules
2-6 mm long, united above the petiole and forming a
very short (1-3 mm) tubular sheath, with 2 distal teeth
1-3 mm long. Leaves opposite, petioles 4-12(-22) mm
long, 0.5-1 mm thick, glabrous (rarely minutely puber-
ulent); leaf blades 3-7(-10) cm long. 1-3.5 cm broad,
narrowly to broadly elliptic, elliptic-oblong, apex acu-
minate with tip 5-10(-l 5) mm long, base broadly obtuse
to acute and slightly decurrent on the petiole, drying thin-
chartaceous to stiffly chartaceous, brown to greenish,
glabrous above and usually with linear cystoliths visible,
glabrous or with a few minute appressed hairs on the
veins beneath, 2 veins 6-1 I/side and usually joining a
vein at the edge of the leaf, intersecondaries present.
Inflorescences 6-12 cm long, 4-9 cm broad, open py-

ramidal with opposite branching, yellowish to blue-gray,
peduncles 1.5-5 cm long, 0.7-1.4 mm thick, puberulent
with ascending hairs 0.2-0.4 mm long, bracts 1-4 mm
long, linear-lanceolate, bracteoles 1-2.5 mm long, ped-
icels 1-5 mm long. Flowers usually glabrous externally,
hypanthium 1-1.5 mm long, turbinate, glabrous or
sparsely puberulent (hairs to 1 mm long), calyx lobes
0.5-0.8(-1.5) mm long. ca. 1 mm broad at the base;
corolla tubular, bluish purple, violet, or white marked
with purple or violet, usually glabrous, tube 6-10.5 mm
long. 1-2 mm diam. and slightly expanded at the base,
corolla lobes 5, 1.5-3 mm long, minutely whitish pa-
pillate along the inner margin; anthers 1.5-2 mm long.
Fruits 3-4.5(-6) mm long and 4 mm diam., obovoid,
becoming dark purple to deep blue and translucent, with
longitudinal ridges when dry, persisting calyx ca. 0.7 mm
high.

Plants of ridges and wet open areas in evergreen
montane rain forest formations, ( 1 800-)2200-3 1 00

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

201

m elevation. Probably flowering and fruiting
throughout the year. These plants appear to prefer
ridges and wet open areas. The species ranges from
the Cordillera de Tilaran to western Panama.

Palicourea adusta is recognized by its high-al-
titude habitat, small leaves, small calyx lobes, usu-
ally bluish flowers (less often yellowish or purple).
This species is similar to P. montivaga with white
flowers and P. salicifolia with yellowish inflores-
cences.

Palicourea albocaerulea C. M. Taylor, Syst. Bot.
Monogr. 26: 15. 1989. Figure 53.

Shrubs or small trees, 2-7 m tall, leafy stems 1.2-4
mm thick, glabrous, quadrangular or rounded; stipules
to 8 mm long, united at the base to form a sheath 1-4
mm long, apex truncated with 2 teeth 2-4 mm long.
Leaves opposite, petioles 8-22 mm long, 0.6-1.2 mm
thick, glabrous, often drying reddish brown; leaf blades
5-15 cm long, 2-5 cm broad, narrowly elliptic to nar-
rowly oblong or elliptic-oblong, apex acuminate with tip
6-15 mm long, base acute (obtuse) to attenuate and
slightly decurrent on petiole, drying chartaceous, gla-
brous above, glabrous beneath or with a few short (0. 1-
0.3 mm) hairs along the midvein beneath, linear cys-
toliths sometimes visible on the lower surface, 2 veins
7-1 I/side and merging with a marginal vein at the leaf
edge. Inflorescences 1 1-22 cm long, 3-8 cm broad, nar-
rowly pyramidal (thyrsoid) with opposite or alternate
lateral branches, peduncles 1.5-5 cm long, 1.6-2.2 mm
thick, glabrous and drying dark brown, bracts 6-10 mm
long, linear and acute, pedicels 3.5-10 mm long, brac-
teoles 2-6(-8) mm long, glabrous. Flowers glabrous ex-
ternally, hypanthium ca. 1 .5 mm long, ca. 0.5 mm diam.
and funnelform, calyx lobes 5, 3.5-9 mm long, 1-1.2
mm broad, narrowly oblong, often unequal; corolla tu-
bular, white or white with blue, tube 8-1 1 mm long, 2-
3 mm diam., slightly expanded near the base, corolla
lobes 1-2 mm long and 1 .5 mm broad at the base; anthers
ca. 2 mm long. Fruits 5-8 mm long (not including the
calyx lobes) and 5-10 mm diam., ovoid or globose, blu-
ish black, persisting calyx lobes ca. 7 mm long.

Plants of lower montane cloud forest forma-
tions, from 1450 to 1650 m elevation. Flowering
in September-December; fruiting in June. This
species is only known from the Cordillera de Ti-
laran and adjacent areas in Costa Rica.

Palicourea albocaerulea is recognized by its re-
stricted range, generally glabrous parts, long inflo-
rescences, long calyx lobes, and white corollas that
turn blue with age. The narrow leaves and long
slender petioles are also distinctive. This is our
only species in which the flowers change from white
to blue. Compare this species with P. lancifera
with shorter calyx lobes and flowers that do not
turn blue.

Palicourea angustifolia H.B.K., Nov. gen. sp. 3:
367. 1819. P. lanceolata Oersted ex Polakows-
ky., Vidensk. Meddel. Dansk. Naturhist. Foren.
Kjobenhavn 17. 1852.

Shrubs or small trees, 2-5(-10) m tall, leafy branchlets
1.3-4 mm thick, glabrous or rarely sparsely puberulent,
terete; stipules to 10 mm long, the tubular sheath 2-4
mm long and truncate but with slender awns 3-6(-8)
mm long, glabrous, persisting. Leaves opposite, petioles
2-9(-13) mm long, 0.6-1.2 mm thick, glabrous; leaf
blades 5-15(-23) cm long, l-4(-5) cm broad, lanceolate
to narrowly elliptic or very narrowly oblong, apex grad-
ually narrowed and acute or acuminate, tip 4-14(-25)
mm long, base acute and slightly decurrent on petiole,
drying chartaceous, glabrous above, glabrous beneath or
with short (0.4 mm) thin whitish hairs along the veins,
2 veins 10-14/side. Inflorescences 3-14(-16) cm long,
3-6(-8) cm broad, a narrowly pyramidal (thyrsoid) pan-
icle, peduncles 1.5-3(-6) cm long, 1-2 mm, thick, gla-
brous or minutely appressed-puberulent, lateral branch-
es opposite, subopposite or alternate, purple to magenta,
bracts 2-5 mm long, pedicels 0-3 mm long, bracteoles
ca. 1 mm long. Flowers glabrous or minutely (0. 1 mm)
papillate puberulent externally, hypanthium ca. 1 mm
long, calyx lobes ca. 0.4 mm long, triangular; corolla
slender funnelform, rose red to purple or fuchsia, tube
9-14 mm long, 1.5-2.5 mm diam., expanded near the
base, corolla lobes 5, 1.2-2.5 mm long; anthers 1.7-2.5
mm long, included. Fruits 4-6 mm long and 5 mm diam.,
globose to ovoid, becoming purple and drying black,
puberulent or glabrate, persisting calyx ca. 0.5 mm long.

Plants of evergreen montane forest formations
on both the Caribbean and Pacific slopes, from
(1400-)! 600 to 2000(-2300) m elevation. Flow-
ering and fruiting throughout the year but with
most flowering collections made in January-Sep-
tember. The species ranges from central Costa Rica
to Peru.

Palicourea angustifolia is recognized by the nar-
row leaves, truncated stipule sheath with long awns,
narrow inflorescences, and slender funnelform
purple corollas with relatively short lobes. This
species resembles P. padifolia and P. purpurea but
differs in the relatively narrower inflorescences,
shorter pedicels, and thinner, more funnelform co-
rollas that are usually minutely puberulent at the
base. In addition, P. padifolia has yellow corollas
and red or orange inflorescence branches.

Palicourea bella (Standl.) Dwyer, Ann. Missouri
Bot. Gard. 67: 299. 1980. Psychotria bella
Standl., J. Wash. Acad. Sci. 18: 185. 1928.

Shrubs or small trees to 5 m tall, leafy stems ca. 4 mm
thick and quadrangular, glabrous or with small (0. 1-0.3
mm) hairs; stipules 10-20 mm long, to 15 mm broad.

202

FIELDIANA: BOTANY

with thin leaf-like texture, united above the petioles,
oblong, rounded and bilobed to emarginate. Leaves clus-
tered distally, petioles 8-20 mm long, 1.2-2 mm thick,
puberulent; leaf blades 9-22 cm long, 3-8.5 cm broad,
elliptic to elliptic-obovate or oblong-elliptic, apex acu-
minate, base gradually narrowed and acute, drying char-
taceous, dark greenish brown, glabrous or with short (0.3
mm) hairs on the veins and smaller hairs between the
veins on both surfaces, 2 veins 12-19/side. Inflores-
cences solitary, 8-20 cm long, 7-10 cm broad, broadly
corymbiform, peduncles 5-13 cm long, to 2 mm thick
and sparsely strigulose, bracts 10-26 mm long, ovate-
oblong, translucent brown and sparsely hirtellous, sec-
ondary peduncles 1 5-25 mm long, bright pink to purple,
pedicels 2-8 mm long, obscured by the large bracts and
bracteoles to 8 mm long. Flowers with hypanthium ca.
2 mm long, glabrous or with conspicuous thin yellowish
hairs ca. 0.7 mm long, the calyx lobes 6-14 mm long,
3-6 mm broad, ovate to ovate-elliptic, drying thin trans-
lucent with scattered short (0.2 mm) hairs; corolla tu-
bular, white to pink, carnose, tube 14-20 mm long and
2-3 mm diam., dilated at the base, corolla lobes 4-5 mm
long, ovate; anthers ca. 3.5 mm long. Fruits ca. 6 mm
long and 6 mm diam., ellipsoid, glabrous or sparsely
puberulent.

Plants of evergreen montane forest formations
at (1500-)2000-2700 m elevation. Flowering in
April. This species is only known from Chiriqui
Province in Panama and a collection from the
Caribbean slope of central Costa Rica at 1500 m.

Palicourea bella is recognized by the broadly
two-lobed stipules, corymbiform inflorescences
with broad thin bracts, and the large thin calyx
lobes. In life, both inflorescences and calyx are red-
violet to bright pink in color. The similarity in
morphology, texture, and vestiture of bracts and
calyx lobes is an interesting example of heterotopy.
This species is similar to P. bellula (q.v.) as well
as to P. hammelii C. M. Taylor and P. ochnoides
Dwyer of western Panama. Grayum 7046 (CR, MO)
from Volcan Barva is tentatively placed here.

Palicourea bellula C. M. Taylor, Syst. Bot. Mono-
gr. 26: 24. 1989. Figure 49.

Shrubs or slender treelets, to 2.5 m tall, leafy stems
2.5-5 mm thick, glabrous, quadrangular and drying dark,
grayish in age; stipules 5-1 1 mm long, 7-10 mm broad,
united above the petioles for 2-3 mm to form a broad
tube, with 2 rounded lobes 2-4 mm high, thin in texture
and glabrous. Leaves opposite, petioles 4-18 mm long,
1.5-2.5 mm thick, glabrous, sulcate adaxially near the
base; leaf blades 7.5-14 cm long, 3.5-7 cm broad, broad-
ly elliptic to slightly elliptic-obovate, apex acuminate,
tip ca. 5- 1 0(- 1 5) mm long, base obtuse (acute) and slight-
ly decurrent on petiole, drying stiffly chartaceous, green-
ish, glabrous above, with thin whitish hairs 0.3-1 mm
long along the sides of the midvein and on some 2 veins
beneath, 2 veins ( 1 0-) 1 7-20/side. Inflorescences 1 or 3,

9-14 cm long, 9-11 cm broad, rounded panicles, red-
violet to fuchsia, peduncles 3-8 cm long, 1-3 mm thick,
glabrous, bracts 12-16 mm long, 3-4 mm broad, lan-
ceolate, bracteoles 5-12 mm long and 3-5 mm broad,
narrowly ovate, pedicels 4-12 mm long. Flowers gla-
brous externally, hypanthium ca. 1 mm long and 1.2
mm diam.. drying dark and glabrous, calyx tube 2-3 mm
long and broadly cupulate, calyx lobes 5. 4-7 mm long,
2-3 mm broad, ovate, red-violet to fuchsia; corolla white,
tubular, carnose, tube 12-15 mm long and 2.5 mm diam..
corolla lobes 2-3 mm long; anthers ca. 3 mm long. Fruits
not seen at maturity.

Plants of open sites in montane rain forest for-
mations near the continental divide at 1 900-2200
m elevation. Flowering in January-February, May,
and November. This species is only known from
between Volcan Viejo (Alajuela) and the south-
eastern slope of Volcan Barva near the upper part
of Rio Patria (Heredia Province) Costa Rica.

Palicourea bellula is distinguished by its colorful
glabrous inflorescences with conspicuous pink to
purple bracts and calyx lobes, tubular white co-
rollas, leaves with many closely parallel secondary
veins, broadly rounded stipule lobes, and restrict-
ed cloud forest habitat. This species is similar to
P. bella, but the calyx and corolla are usually small-
er and stiffer than in P. bella. Palicourea hammelii
and P. ochnoides are similar but lack the broad
imbricate calyx lobes.

Palicourea brenesii Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 18: 1333. 1938. P. talamancana
Standl. & L. O. Williams, Ceiba 1: 250. 1951.
Figure 52.

Shrubs, 1-4 m tall, erect or clambering, often with
only a few distal branches 2.5-5 mm thick, glabrous or
sparsely strigillose with hairs 0.5-1 mm long, drying dark,
quadrangular; stipules united to form a tubular sheath
3-6 mm long, truncated distally and with 2 prominent
teeth (on each side) 4-7(-10) mm long and 1-2 mm
broad at the base, usually glabrous, persisting. Leaves
opposite, petioles 1.5-3 cm long, 1.2-2.2 mm thick, gla-
brous; leaf blades 9-1 8 cm long, 3.5-8 cm broad, broadly
elliptic to elliptic-obovate or elliptic-oblong, apex abruptly
narrowed and short-acuminate, base obtuse to acute,
drying stiffly chartaceous, brownish or yellowish, gla-
brous above, glabrous or sparsely puberulent beneath,
2 veins 15-21 /side. Inflorescences 8-15 cm long, 4-9
cm broad (at broadest part), pyramidal or elongate thyr-
soid panicles, often blue throughout, peduncles 2-5 cm
long, 1.5-3 mm thick, sparsely to densely puberulent
with thin hairs ca. 0.2 mm long, bracts 5-8 mm long,
ca. 2 mm broad, lanceolate, pedicels 0-6(-8) mm long,
bracteoles 1-5 mm long. Flowers minutely puberulent
and glabrescent distally, hypanthium ca. 1 mm long and
0.8 mm diam., calyx tube broadly cupulate, calyx lobes
1-2.5 mm long, broadly triangular and obtuse; corolla

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

203

blue, tubular, glabrous near the base and densely ap-
pressed-puberulent distally or glabrous throughout, tube
8-1 1 mm long, ca. 2 mm diam.. narrowed near the mid-
dle, corolla lobes 5, 3-5 mm long; anthers ca. 2.5 mm
long. Fruits 4-6 mm long, 4-6 mm diam., globose, gla-
brate.

Plants of evergreen montane rain forest for-
mations, from 1800 to 2600(-3200) m elevation.
Flowering and fruiting in February-June. The spe-
cies has only been collected in the northern part
of the Meseta Central (near Zarcero and Palmira)
and along the Interamerican Highway in the west-
ern part of the Cordillera de Talamanca in Costa
Rica.

Palicourea brenesii is distinguished by the larger
leaves with many secondary veins, bright "china
blue" thyrsoid panicels, and small blue to violet
flowers with conspicuous calyx lobes. The usually
glabrous stems contrast with the puberulent pe-
duncle and inflorescence branches. The striking
bright blue color of the inflorescence is similar to
that found in species of Faramea. The disjunct
collections from the Cordillera de Talamanca have
smaller corollas and stipule lobes to 10 mm long.
This species is very similar to P. discolor after the
flowers have faded.

Palicourea copensis (Dwyer) C. M. Taylor, comb,
nov. Psychotria copensis Dwyer, Ann. Missouri
Bot. Gard. 67: 365. 1980. Figure 52.

Shrubs or small treelets, 2-5 m tall, leafy stems 2-5
mm thick, with curved yellowish or whitish hairs 0.4-
0.9 mm long, glabrescent; stipules 6-12 mm long, united
to form a broad tube to 10 mm long, 4-8 mm broad (to
14 mm broad below the inflorescences), bilobed with a
distal sinus 1-6 mm deep, glabrous but with a ciliolate
margin, persisting. Leaves with petioles 10-55 mm long,
1.3-2.3 mm thick, glabrous or sparsely puberulent be-
neath; leaf blades 9-23 cm long, 3.5-9 cm broad, elliptic-
obovate to elliptic-oblong, apex narrowly acuminate and
5-13 mm long, base cuneate and slightly decurrent on
petiole, drying chartaceous, dark green above (pale green
beneath), glabrous above or with a few hairs along the
major veins, sparsely pubescent beneath with crooked
whitish hairs 0.4-1.5 mm long, 2 veins 9-14/side and
arcuate-ascending distally. Inflorescences terminal or
pseudoaxillary, 1 or several per node, 2-7 (-1 1) cm long,
3-6(-l 2) cm broad, pyramidal with few opposite branch-
es, peduncles 8-26 mm long, ca. 1 .5 mm thick, pubescent
or glabrous, bracts 3-7 mm long, flowers in distal cymose
or irregular clusters. Flowers with hypanthium ca. 1 mm
long, obconic, glabrous or pubescent, calyx yellowish,
calyx lobes quite variable, 1-3 mm long, oblong and
obtuse; corollas lavender or white marked with purple,
salverform with a distinctly gibbous base, tube 7-12 mm
long, 2-4 mm diam., puberulent, corolla lobes 3-5 mm

long; anthers ca. 3 mm long. Fruits 6-7 mm long, 4-5
mm diam., obovoid, becoming blue or violet.

Plants of the very wet Caribbean escarpment,
between 400 and 1 000 m elevation. Flowering and
fruiting in March-May and November. The spe-
cies is known from the Caribbean slopes of Volcan
Barva and the P. N. Braulio Carrillo area in central
Costa Rica. It was originally described from El
Cope in Code Province, Panama.

Palicourea copensis is recognized by its white or
lavender flowers, variable sepal lobes, smaller
thick-branched inflorescences, large persisting
stipules, puberulence of usually curved hairs, and
lower montane habitat. This is a rarely collected
species that appears to prefer creek margins.

Palicourea crocea (Sw.) Roem. & Schult., Syst. Veg.
5: 193. 1819. Psychotria crocea Sw., Prodr. 44.
1788. Figure 50.

Shrubs or small trees, 1-5 m tall, leafy stems 1-4 mm
thick, glabrous or minutely puberulent, terete; stipules
united to form a short (0.5-2 mm) truncated tube with
narrow distal teeth 1-4 mm long, glabrous, deciduous.
Leaves opposite, petioles 6-12(-20) mm long, 1-1.5 mm
thick, usually glabrous; leaf blades 7-14(-19) cm long,
2.5-6(-7) cm broad, elliptic, narrowly elliptic, lanceolate,
to elliptic-ovate or elliptic-oblong, apex acute to acu-
minate, base obtuse to acute and slightly decurrent on
the petiole, drying membranaceous to thin-chartaceous,
greenish, usually glabrous above, with short (0.3 mm)
thin hairs along the sides of the midvein beneath, 2
veins 8-12/side. Inflorescences 6-15 cm long, 2.5-7 cm
broad, nararow pyramidal to rounded with opposite or
subopposite (alternate distally) branching, reddish to or-
ange or pink, peduncles 1.5-1 1 cm long, 1-2 mm thick,
glabrous or sparsely and minutely puberulent, bracts 1-
10 mm long, linear-lanceolate and deciduous, pedicels
4-8(-12) mm long, slender, bracteoles ca. 1 mm long
and deciduous. Flowers distylous, glabrous or minutely
puberulent externally, hypanthium ca. 1 mm long and
0.7 mm diam., calyx lobes 0.3-0.5 mm long, broadly
ovate; corolla tubular, dark red to yellow-orange or pink
(rarely bluish green), tube 5-9 mm long, 1 .5-3 mm diam.,
inflated near the base, corolla lobes 1-2.5 mm long; an-
thers ca. 2.7 mm long. Fruits 4-6 mm long and 4 mm
diam., subglobose or ovoid, becoming dark blue or black,
longitudinally ribbed when dry, persisting calyx minute.

Plants of secondary growth in evergreen rain
forest areas often along streams and paths in the
Caribbean lowlands, from near sea level to 600 m
elevation. Flowering and fruiting throughout the
year (flowering mostly in April-August in Central
America). The species ranges from central Mexico
and the West Indies to Paraguay.

Palicourea crocea is recognized by its smaller

204

FIELDIANA: BOTANY

stature and lowland habitat, short stipular tube,
colorful thyrsoid inflorescences, minute calyx lobes,
and orange or red corolla tubes. This species is
often found in swampy areas and other sites with
poor drainage. This species is sometimes confused
with P. padifolia, of higher elevations and with
longer stipule sheaths, with P. guianensis, with
longer stipule lobes, and with species ofHamelia.

Palicourea discolor K. Krause, Bot. Jahrb. Syst.
54: Beibl. 119: 40. 1916. P. macrosepala K.
Krause, loc. cit. 41. 1916. P. panamaensis
Standl., Ann. Missouri Bot. Gard. 25: 839. 1938.
Figure 52.

Shrubs or small trees, 1-4(1-6) m tall, leafy stems 3-
6 mm thick, glabrous and quadrangular; stipules united
to form a broadly tubular sheath 2-6 mm long, truncated
apically and with 2 narrow teeth 4-7 mm long, glabrous
and persisting. Leaves opposite, petioles l-3(-5) cm long,
1.7-2.7 mm thick, glabrous; leaf blades 10-26 cm long,
5-13 cm broad, elliptic to broadly elliptic-oblong, acute
to short-acuminate, the tip 5-10(-15) mm long, obtuse
or acute at the base, drying stiffly chartaceous, dark brown
or greenish above, glabrous above, sparsely puberulent
with inconspicuous (0.05-0.2 mm) thin whitish hairs on
the veins beneath, 2 veins 11-19/side, united distally
with a vein at the leaf edge. Inflorescences solitary or
the basal branches subtended by leaves and apparently
3, 12-22(-30) cm long, 10-1 5(-24) cm broad, pyramidal
with opposite or subopposite branches, bright purple in
life, peduncles 2-14 cm long, 2.5-4 mm thick, glabrous,
bracts 5-8 mm long, 1 mm wide, linear-lanceolate, ped-
icels 1-8 long, glabrous, bracteoles absent. Flowers gla-
brous externally, hypanthium 0.5-1 .3 mm long and 0.3-
1 mm diam.. calyx lobes 1.5-5 mm long, acute to Un-
gulate, very variable in some collections; corolla purple
or white flushed with purple, carnose, tube 7-13 mm
long, 1-2 mm diam., corolla lobes 1-3.5 mm long. Fruits
4-5 mm long and 4-5 mm diam., globose to globose-
ellipsoid, with longitudinal ribs (dried), calyx persisting
or deciduous.

Plants of forest interiors and shaded thickets in
evergreen montane forest formations, from 1 1 00
to 1900(-2600) m elevation. Flowering and fruit-
ing in February-August. The species ranges from
the upper Rio Grande de Orosi and Moravia de
Chirripo in the Cordillera de Talamanca of Costa
Rica southward into the Chiriqui Highlands of
western Panama.

Palicourea discolor is recognized by its relatively
large leaves and inflorescences, purple flowers and
inflorescence branches, and variable calyx lobes
to 5 mm long. This species is similar to P. brenesii
with bright blue inflorescences and smaller corol-
las and P. purpurea with smaller calyx lobes and
smaller leaves.

Palicourea garciae Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 22: 195. 1940, non P. garciae
Steryerm. 1971. Psychotria copeyana Standl. &
L. O. Williams, Ceiba 1: 251. 1940. Figure 53.

Shrubs or small trees, 1.5-5 m tall, leafy branchlets
1.5-3.5 mm thick, glabrous or with short (0.3 mm) as-
cending yellowish hairs, quickly becoming glabrescent
and terete, drying dark; stipules united to form a tubular
sheath 2-4 mm long, each side with 2 short ( 1 mm) distal
lobes and a small ( 1 mm) central sinus, glabrous or mi-
nutely appressed-puberulent. Leaves opposite, petioles
5-15 mm long, 0.6-1.2 mm thick, glabrous; leaf blades
5-12(-15) cm long, 2-4(-4.5) cm broad, narrowly ellip-
tic, narrowly elliptic-ovate to narrowly oblong or lan-
ceolate, apex acuminate with tip 7-15 mm long, base
acute to obtuse and slightly decurrent on petiole, drying
stiffly chartaceous, much darker above than beneath,
glabrous above or with hairs along the midvein, with
few or many minute (0.05-0.2 mm) hairs along the major
veins beneath and usually glabrescent, 2 veins 12-18/
side, arising from the midvein at near 90. Inflorescences
solitary, 4-12 cm long, 6-12 cm broad, open pyramidal
or rounded corymbiform with opposite (subopposite)
branching, green or flushed with purple, peduncles
(l-)2.5-4 cm long, very sparsely to densely puberulent
with short (0.2 mm) thin hairs, bracts 2-5 mm long,
linear-subulate, pedicels 1-5 mm long, bracteoles ca. 1
mm long. Flowers glabrous externally, hypanthium 1-
1.5 mm long, 0.8-1 mm diam., calyx lobes 5, 0.7-1.5
mm long, acute; corolla funnelform and gibbous near the
base, white or flushed with yellow or purple distally
(grayish to purple near the base), tube 6-8 mm long,
constricted in the middle (1-1..5 mm diam.) and gibbous
near the base, corolla lobes 2-3 mm long and 1 mm
broad at the base; anthers 2-2.5 mm long. Fruits 4-5
mm long, 3-4 mm diam., globose to ellipsoid, becoming
pale blue at maturity, with longitudinal ribs, persisting
calyx ca. 1 mm long.

Plants of montane rain forest formations, from
1200 to 1600(-2000) m elevation. Flowering in
March and May-August; fruiting in May-June and
August. The species ranges from central Costa Rica
to Colombia.

Palicourea garciae is recognized by the small
narrow leaves with many secondary veins, the small
corolla tubes strongly bent below the middle and
inflated at the base, and the small pale blue fruit.
The very short tubular sheath and bifid interpetio-
lar part of the stipule is noteworthy. Psychotria
copeyana was based on a specimen ( Williams <&.
Allen 16482 F) with immature inflorescences from
El Copey. Vegetatively this species resembles sev-
eral smaller-leaved species of Psychotria, but the
strongly gibbous flowers are very different.

Palicourea guianensis Aubl., Hist. pi. Guiane 1:
173, t. 66. 1775. Psychotria palicourea Sw., Fl.
Ind. Occ. 1797. Figure 50.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

205

Shrubs or small trees, 2-6(-10) m tall, leafy stems 1.5-
8 mm thick, glabrous or minutely puberulent, quadran-
gular, often drying dark and contracted beneath the node;
stipules free or united for 1-2 mm at the base, lobes, 5-
8(-12) mm long, lobes 2-4 mm broad, blunt, glabrous
and drying dark. Leaves opposite, petioles 8-24(-38) mm
long, 1.3-3 mm thick, glabrous; leaf blades (12-) 15-28
cm long, (5-) 7-1 8 cm broad, broadly ovate-elliptic to
ovate-oblong, elliptic-oblong or ovate, apex acute or acu-
minate, base obtuse to rounded and subtruncate, drying
thin-chartaceous, concolorous, glabrous or minutely (0. 1
mm) papillate-puberulent above and below, 2 veins 10-
15/side. Inflorescences usually solitary, 7-18 cm long,
4-12 cm broad, pyramidal or thyrsiform with many
closely crowded alternate or subopposite branches, red-
orange when flowering and later turning purple, pedun-
cles 3-12 cm long, 1.8-3.2 mm thick, minutely puber-
ulent with thin hairs ca. 0. 1 mm long, bracts absent or
0.5-7 mm long and often adnate to the primary branches,
pedicels 2-5(-8) mm long, minutely puberulent. Flowers
monomorphic, minutely puberulent externally, hypan-
thium ca. 1.5 mm long and 0.8 mm diam., tubular, calyx
lobes 0.2-0.6 mm long, broadly obtuse; corolla yellow
or orange, tube 8-25 mm long and 2-3 mm diam. at
base, contracted in the middle and a slightly inflated
distally, corolla lobes 1-2 mm long, triangular; ovary 2-
6-locular. Fruits 4-7 mm long, 4-5 mm diam., ovoid to
ellipsoid, becoming purple violet and drying black with
prominent longitudinal ribs, persisting calyx and disc ca.
1 mm high.

mm thick, glabrous or minutely puberulent, drying dark;
leaf blades 8.5-20 cm long, 2.5-8.5 cm broad, elliptic
to elliptic-obovate, apex acuminate with tip 5-15 mm
long, base gradually narrowed and acute or cuneate,
slightly decurrent on petiole, drying stiffly chartaceous,
dark olive green above, glabrous above or puberulent on
the midvein, puberulent beneath with crooked hairs 0.05-
0.3 mm long, 2 veins 12-18/side. Inflorescences 4-14
cm long, to 1 3 cm broad, open pyramidal, pink to purple
but drying dark, peduncle 15-35 mm long, 1.5-3 mm
thick, glabrous or very sparsely minutely puberulent,
proximal bracts 18-35 mm long, ca. 5 mm broad, lan-
ceolate, distal bracteoles 6-20 mm long, pedicels 4-15
mm long. Flowers glabrous externally, hypanthium ca.
2 mm long and 1 mm diam., calyx 1 3-20 mm long, pink
to purple, calyx lobes 4-1 2 mm long, lanceolate, unequal;
corolla white, tubular, tube 20-27 mm long, ca. 4 mm
diam., corolla lobes 3-6 mm long, triangular. Fruits 8-
10 mm long, 7-8 diam., ellipsoid.

Palicourea hammelii was originally described
from collections made between 1 400 and 2200 m
elevation in Chiriqui, Panama. It has been recently
collected on the Fila Matama at 1 600 m elevation
on the Caribbean slope of the Cordillera de Tala-
manca (Herrera & Chacon 2781 CR, MO). This spe-
cies resembles P. bella with ovate calyx lobes and
P. bellula with a shorter corolla tube.

Plants of open secondary formations in ever-
green and partly deciduous forest areas, from near
sea level to 900 m elevation. Flowering and fruit-
ing occur throughout the year; flowering mostly in
March-July in Costa Rica. The species ranges from
southern Mexico and the West Indies southward
to southern Brazil and Bolivia.

Palicourea guianensis is recognized by its low-
land habitat, the wide glabrous or very minutely
puberulent thin-textured leaves, the large rounded
stipule lobes, the conspicuous densely flowered
yellow or orange inflorescences, and yellow or or-
ange puberulent corollas. Most of our collections
come from the Caribbean lowlands and from the
Golfo Dulce area. Central American material has
two-locular ovaries, but specimens from eastern
South American may have three-or four-locular
ovaries. Palicourea guianensis resembles P. padi-
folia (q.v.).

Palicourea hammelii Taylor, Syst. Bot. Monogr.
26:43. 1989.

Small trees to 5 m tall, leafy stems 3-7 mm thick,
glabrous or minutely (0. 1 mm) appressed-puberulent,
drying dark; stipules 6-12 mm long, with a sheathing
base 3-5 mm long with 2 lobes (4/node) 3-7 mm long,
persisting. Leaves with petioles 13-35 mm long, 1.3-2

Palicourea lancifera Standl. & L. O. Williams, Cei-
ba 1: 249. 1951. Figure 53.

Shrubs, 1.5-3(-5) m tall, leafy stems 1.3-4 mm thick,
glabrous to (less often) densely puberulent with crooked
yellowish hairs 0.1-0.3 mm long, quadrangular at first,
glabrescent; stipules united to form a tubular sheath 3-
5 mm long, apex truncated with 2 narrow teeth 2-5 mm
long on each side. Leaves opposite, petioles 7-25(-30)
mm long, 0.4-1.2 mm thick, glabrous or minutely pu-
berulent; leaf blades 5-1 5(-l 8) cm long, 2-5(-7) cm broad,
elliptic to narrowly elliptic-oblong, apex acuminate with
tip 6-20 mm long, base acute and decurrent on petiole,
drying thin-chartaceous or chartaceous, dark greenish or
brownish above, glabrous above, minutely (0. 1-0.4 mm)
strigulose on the veins beneath, 2 veins 8-12/side and
connecting with a vein along the leaf edge. Inflorescences
solitary, 7-17(-21)cm long, 2-6(-9)cm broad, narrowly
pyramidal with many short branches, green or marked
with yellow, peduncles 1-4 cm long, 1-2 mm thick, gla-
brous to densely puberulent, bracts 5-8 mm long, nar-
rowly lanceolate, flowers often alternate along the lateral
branches, bracteoles 1.5-4 mm long, pedicels 2-7(-17)
mm long. Flowers glabrous or sparsely puberulent ex-
ternally, hypanthium 1-1.5 mm long, ca. 1 mm diam.,
calyx lobes 1.5-3 mm long, often unequal, acute or
rounded distally; corolla white or flushed with green or
yellow, tubular, tube 7-1 1 mm long, ca. 2 mm diam.
and expanded at the base, corolla lobes 1-2 mm long;
anthers 2-2.5 mm long. Fruits 4-5 mm long and 4-5
mm diam., ovoid or globose-oblate, persisting calyx ca.
1.5 mm high.

206

FIELDIANA: BOTANY

Plants of very wet lower montane cloud forest
formations, from 800 to 1 700 m elevation. Flow-
ering June-November; apparently fruiting
throughout the year. This species ranges from the
Reserva Forestal de San Ramon and Zapote de
San Carlos (Alajuela) eastward to Moravia de
Chirripo (Limon) and the western part of the Gen-
eral Valley (San Jose).

Palicourea lancifera is recognized by its narrow
long-petiolate leaves, narrowly thyrsoid greenish
inflorescences, slightly larger calyx lobes rounded
distally, and white or pale yellow corolla tubes.
This species resembles P. albocaerulea with longer
calyx lobes and P. leucantha J. D. Smith of Mexico
and northern Central America.

Palicourea lasiorrhachis Oersted, Vidensk. Med-
del. Dansk. Naturhist. Foren. Kjobenhavn 39.
1 852. P. veraguensis Dwyer, Ann. Missouri Bot.
Card. 67: 318. 1980. Figure 51.

Shrubs or small treelets, l-5(-7) m tall, leafy stems
1.8-5 mm thick, glabrous to pubescent with stiff yellow-
ish curved hairs to 0.5 mm long; stipules united and
forming a broad tube 2-9 mm long, truncated and with
narrow teeth 2-5(-8) mm long, glabrous or minutely
puberulent, persisting. Leaves opposite, petioles 7-24(-32)
mm long, 1-2 mm thick, glabrous or puberulent; leaf
blades 6-17(-26) cm long, 2-8(-10) cm broad, elliptic
to narrowly ovate-elliptic or elliptic-oblong, apex taper-
ing gradually and acuminate, tip 4-13(-20) mm long,
base acute to obtuse, drying chartaceous, dark grayish
green to brownish above, glabrous above, sparsely to
densely puberulent along the major veins beneath with
short (0.3 mm) yellowish or longer (1 mm) whitish as-
cending hairs (rarely glabrous), 2 veins (8-)l 1-14(-18)/
side and joining a vein along the leaf edge distally. In-
florescences 5-1 5(-20) cm long. 4-1 3 cm broad, narrow-
ly pyramidal with 4-8 pairs of opposite or subopposite
lateral branches from the central rachis, greenish to bright
yellow, peduncles 1.5-7(-10)cm long, 0.7-2.8 mm thick,
sparsely to densely puberulent with stiff multicellular
hairs 0.3-1 mm long, bracts 2-9 mm long, linear-su-
bulate, pedicels 1-6 mm long, minutely puberulent, yel-
low to bright green, bracteoles 0.5-2 mm long. Flowers
sparsely to densely pubescent externally (often glabres-
cent), hypanthium 0.5-1.5 mm long, ca. 1 mm diam.
distally, obconic, calyx lobes 0.3-1.2 mm long, obtuse:
corolla yellow to greenish, tubular-funnelform, tube 6-
15 mm long, narrowed in the middle, ca. 2 mm diam.
at the base and 3 mm diam. distally, corolla lobes 1-2.5
mm long, ca. 1.5 mm broad at the base, triangular; an-
thers 1.5-2 mm long or 2.5-3 mm long in short-styled
form. Fruits 4-5 mm long and 3-4 mm diam., obovoid,
with prominent longitudinal ribs, puberulent or glabres-
cent, blue or black.

Understory plants of very wet montane rain for-
est formations, from (1000-)! 300 to 2600 m el-

evation. Flowering throughout the year (mostly in
May-June); probably fruiting throughout the year.
This species ranges from the Cordillera de Tilaran
along the continental divide and Cordillera de Ta-
lamanca to western Panama.

Palicourea lasiorrhachis is recognized by the of-
ten narrow leaves, usually puberulent beneath, the
minutely puberulent inflorescences (rarely gla-
brous) with yellow pedicels, the small calyx lobes,
narrowly funnelform yellow corolla tubes, and
small obovoid fruit. This common species is quite
variable and includes individual collections that
differ greatly from each other in regard to leaf size,
leaf shape, puberulence, and aspects of flowers and
inflorescences. A group of collections from near
El Empalme have thicker stems; stifler, more ob-
long leaves; secondary veins arising at almost 90
angles; and dense puberulence. This species may
be difficult to distinguish from P. vestita (q.v.). It
is often confused with P. padifolia, which has red-
orange inflorescence branches and more tubular
corollas. Also compare P. montivaga (smaller
plants with white flowers) and P. adusta (with blue
flowers).

Palicourea macrocalyx Stand!.. J. Wash. Acad. Sci.
18: 278. 1928. Figure 49.

Small shrub, subshrubs, or little treelets. 0.5-2(-6) m
tall, leafy branchlets 1 .5^4(-10) mm thick, glabrous, flat-
tened or quadrangular but quickly becoming terete: stip-
ules with a broad thick sheath 2-6 mm long, truncated
and with narrow acute teeth 4-10 mm long and 1-3 mm
broad, glabrous and persisting. leaves opposite, petioles
7-30(-45) mm long, 0.9-2.2 mm thick, glabrous; leaf
blades 5-15(-18) cm long, 2-6(-9.5) cm broad, elliptic-
obovate to obovate-oblong or elliptic (rarely broadly ob-
long), apex acuminate with tip 4-13 mm long, base ob-
tuse to acute (subtruncate). drying stiffly chartaceous to
subcoriaceous. olive green or brownish above, glabrous
above and below or with small (0.3-0.5 mm) hairs along
the midvein, 2 veins 9-1 3/side. Inflorescences 3-8(-l 5)
cm long. 2-10 cm broad, pyramidal, yellow or marked
with purple, peduncles 1-8.5 cm long, 1-1.8 mm thick,
glabrous, bracts ca. 6 mm long, pedicels 1-8 mm long,
glabrous, bracteoles 1-4 mm long. Flowers glabrous ex-
ternally, hypanthium ca. 1 mm long, narrowly obconic.
calyx lobes 3-6 mm long, 1-2 mm broad, narrowly tri-
angular to ovate, green or yellow and sometimes marked
with purple: corolla tubular, yellow or greenish yellow,
tube 8-13 mm long, ca. 2 mm diam.. slightly narrower
in the center, corolla lobes 2-3 mm long, obtuse; anthers
1-2 mm long. Fruits 5-6 mm long, 4-5 mm diam., el-
lipsoid, becoming blue, longitudinally ribbed (when
dried), with persisting calyx to 6 mm long.

Plants of the very wet cloud forests along the
continental divide, from 1300 to 2200 m eleva-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

207

tion. Probably flowering throughout the year;
fruiting in April-May and August-October. En-
demic to Costa Rica, and collected primarily in
the Cordillera de Tilaran, but ranging to near San
Isidro de Heredia on the western slopes of Volcan
Irazu.

Palicourea macrocalyx is distinguished by the
stiff slightly obovate leaves with many secondary
veins, lack of pubescence, yellow inflorescences
sometimes marked with purple, the larger imbri-
cate greenish yellow calyx lobes, and yellow co-
rollas. The name of this species should not be
confused with P. macrosepala Krause, a synonym
of P. discolor. This species is very similar to P.
bellula, which has longer corolla tubes.

Palicourea montivaga Standl.. J. Wash. Acad. Sci.
18:279. 1928. Figure 53.

Shrubs, 1-3 m tall (rarely slender treelets to 6 m), leafy
stems 0.9-6 mm thick, glabrous or with a few thin hairs
ca. 0.3 m long distally, terete; stipules with a tubular
sheath 0.5-4 mm long, truncated distally and with tri-
angular or linear teeth 0.5-2.5 mm long, glabrous and
persisting. Leaves opposite, petioles 6-14(-25) mm long,
0.4-1.1 mm thick, glabrous, slightly sulcate above; leaf
blades 3-7(-9) cm long, l-2.5(-3) cm broad, lanceolate
or elliptic-lanceolate to narrowly elliptic-oblong, apex
tapering gradually and acute or acuminate, tip to 1 5 mm
long, base acute (obtuse), drying chartaceous, dark green
or brown above, glabrous above, glabrous or with a few
thin hairs 0. 1 -0. 5 mm long on the veins beneath, 2 veins
6-9/side. Inflorescences 2-10 cm long, 2-8 cm broad,
pyramidal with open opposite branching, peduncles 8-
45 mm long, 0.3-1.2 mm thick, glabrous or minutely
puberulent, inflorescence branches usually bright yellow
(green), bracts 2-3.5 mm long, linear, pedicels 1-6 mm
long. Flowers usually glabrous externally, hypanthium
0.5-1 mm long, ca. 1 mm diam. distally, obconic, calyx
lobes 0.4-1 mm long, acute or obtuse; corolla white or
yellow, tube 5-9 mm long, 1-1 .3 mm diam., corolla lobes
1-2 mm long; anthers ca. 1.5 mm long. Fruits 4-5 mm
long, 3-4 mm diam., obovoid, becoming longitudinally
ribbed.

Plants of montane rain forest formations, from
(1 1 00-) 1300 to 2300(-2900?) m elevation. Prob-
ably flowering throughout the year, but mostly in
April-August; fruiting April-November. The spe-
cies is endemic and ranges from Volcan Tenorio
southward along the continental divide to the
western slopes of Volcan Irazu.

Palicourea montivaga is recognized by its small
narrow leaves with long tips, glabrous parts, small
yellowish inflorescences, small white or yellowish
corollas, and montane habitat. This species is
closely related to P. adusta with smaller bluish
flowers and P. lasiorrhachis with relatively larger

yellowish floral parts and shorter leaf tips. The
plant attributed to this species from Panama
(Dwyer, 1 980) is P. lasiorrhachis. Material of P.
padifolia can also be confused with this species.

Palicourea orosiana C. M. Taylor, Syst. Bot.
Monogr. 26: 61. 1989. Figure 51.

Shrubs, 2-8 m tall, leafy stems 1 .8-4 mm thick, stri-
gulose with stiff straight hairs to 1 mm long; stipules with
a short (1.5-3 mm) sheath (difficult to see under the
strigulose hairs) and narrow distal teeth 6-17 mm long,
covered with long (1-2 mm) thin hairs. Leaves opposite,
petioles 4-12 mm long, 1-1.9 mm thick, puberulent in
early stages with straight or curved ascending hairs; leaf
blades 7-14 cm long, 2.5-4.5 cm broad, elliptic-oblong
to narrowly elliptic, apex acuminate with tip 6-10(-13)
mm long, base acute, drying stiffly chartaceous, yellow-
ish, upper surface glabrous, glabrous on 2 veins and
interveinal areas beneath but with straight stiffhairs 0.5-
1.5 mm long on the sides of the midvein, 2 veins 8-1 3/
side. Inflorescences solitary, 7-1 1 cm long, 7-8 cm broad,
pyramidal, yellowish, peduncles 3.5-6.5 cm long and 1 .5
mm thick, strigulose, bracts and bracteoles 3-7 (-1 3) mm
long, ca. 1 mm broad, yellowish and resembling the calyx
lobes, pedicels 2-6 mm long. Flowers greenish yellow,
hypanthium ca. 1.3 mm long and 1 mm diam., densely
pubescent, calyx lobes 5-1 1 mm long, 0.8-1 . 1 mm broad
near the base, narrowly triangular, acute, sparsely pu-
bescent with thin whitish hairs ca. 0.5 mm long, drying
yellow; corolla tubular, yellow, tube ca. 10 mm long,
with whitish ascending hairs, corolla lobes ca. 2 mm
long; anthers ca. 2.5 mm long. Fruits not seen.

Plants of the wet cloud forests of the Caribbean
slopes and continental divide, at 1200-2100 m
elevation. Flowering in May-June. This species
has only been collected above the upper Rio Gran-
de de Orosi near Tapanti, Cartago, in Costa Rica;
it is also known from western Panama.

Palicourea orosiana is distinguished by its pu-
berulent yellowish inflorescence and flowers, larg-
er yellowish bracts and calyx lobes, and the long
stipule lobes. The pubescence along the midvein
on the underside of the leaf is also unusual. This
species is similar to P. lasiorrhachis, which has
much smaller calyx lobes, and to P. macrocalyx,
which has glabrous inflorescences and broadly im-
bricate calyx lobes.

Palicourea padifolia (Willd. ex Roem. & Schult.)
C. M. Taylor & Lorence, Taxon 34: 669. 1985.
Psychotria padifolia Willd. ex Roem. & Schult.,
Syst. Veg. 5: 189. 1819. Psychotria mexicana
Willd. ex Roem. & Schult., loc. cit. 192. 1819.
Palicourea costaricensis Benth. ex Oerst., Vi-
densk. Meddel. Dansk. Naturhist. Foren. Kjo-

208

FIELDIANA: BOTANY

benhavn 1852: 38. 1853. Palicourea subrubra
Polak., Linnaea 41: 571. 1877. P. galeottiana
sensu Standley, not M. Martens (see Taylor &
Lorence, 1985, cited above). Figure 51.

Shrubs or small trees, (l-)2-7(-10) m tall, leafy stems
1.5-5 mm thick, glabrous (rarely hirsutulous), quadran-
gular but becoming terete; stipules with a tubular sheath
1-4 mm long, truncated distally and with narrow awns
2-10 mm long, glabrous. Leaves opposite (rarely 4/node),
petioles 4-1 6(-22) mm long, 0.6-1 .8 mm thick, glabrous;
leaf blades 6-16(-24) cm long, 2-6(-8.5) cm broad, nar-
rowly elliptic-oblong to narrowly oblong, elliptic-ovate
or lanceolate (rarely elliptic-obovate), apex acute to acu-
minate with tip 4- 1 2(-20) mm long, base acute to obtuse,
drying stiffly chartaceous, glabrous and with short (0.2
mm) cystoliths often visible above, glabrous beneath
except for thin whitish hairs 0.3-0.9 mm long on the
major veins beneath or only along the midvein, 2 veins
8-14/side. Inflorescences (5-)7-18 cm long, 4-14 cm
broad, broadly pyramidal, reddish purple to salmon red
or orange (yellow), peduncle 1.5-5 cm long, 1.2-2 mm
thick, glabrous or sparsely puberulent, bracts ( l-)2-6(-8)
mm long and linear-subulate, bracteoles 0.5-2 mm long,
pedicels l-6(-10) mm long. Flowers glabrous or sparsely
puberulent externally, hypanthium 0.7-1 mm long, 0.5-
1 mm diam., calyx lobes 0.5-1 mm long, triangular,
obtuse; corolla tubular to slightly funnelform, carnose,
orange or yellow, tube 8-15(-18) mm long and 1.5-2.5
mm diam., narrower in the middle, corolla lobes 2-3
mm long, ca. 1.5 mm broad at the base, obtuse; anthers
2.5-4 mm long. Fruits 4-6(-10) mm long, 4-6(-10) mm
diam., ovoid to ellipsoid or globose, longitudinally ribbed,
persisting calyx ca. 1 mm high.

Plants of evergreen lower montane forest for-
mations and in moist sites in deciduous forest for-
mations, (800-)1000-2000(-2400) m elevation.
Flowering and fruiting throughout the year (mostly
flowering in December-August and fruiting in Jan-
uary-August). The species is quite common in
Costa Rica and ranges from the Cordilleras de
Guanacaste and Tilaran, around the Caribbean
side of the central volcanic chain, through the Cor-
dillera de Talamanca. The species ranges from
eastern Mexico to Panama.

Palicourea padifolia is recognized by its usually
narrow leaves, short-tubular stipule sheaths, col-
orful inflorescences with (usually) reddish pedi-
cels, minute calyx lobes, narrow yellowish (or or-
ange) corolla tubes, and montane habitats with
high rainfall. This is the most commonly collected
species of Palicourea in Central America. This spe-
cies is easily mistaken for P. angustifolia. P. crocea,
and P. purpurea, but those species have purplish
flowers. Palicourea lasiorrhachis is similar but has
yellow inflorescence branches and corollas. Pali-
courea padifolia is very closely related to P. thyr-
siflora (Ruiz & Pavon) Roem. & Schult. of Ecuador

and Peru and may be conspecific, but neither of
the two species has been collected in Colombia or
the eastern half of Panama. Fruiting plants of this
species may be confused with species of Psy-
chotria, but the inflorescences are more colorful
in P. padifolia.

Palicourea purpurea C. M. Taylor, Syst. Bot.
Monogr. 26: 71. 1989. Figure 52.

Shrubs or small trees, (l-)3-6(- 10) m tall, leafy stems
1-4 mm thick, glabrous, stipules with sheaths 1-4 mm
long, truncated distally and with slender teeth 2-5 mm
long, glabrous and persisting. Leaves opposite, petioles
6-25(-35) mm long, 0.6-1.3 mm thick, glabrous; leaf
blades 7-19 cm long, 2-7 cm broad, narrowly elliptic to
elliptic-oblong, apex acuminate with tip 5-15 mm long,
base acute and slightly decurrent on petiole, drying char-
taceous, dark brown above, glabrous above and below
or with a few thin white hairs along the side of the mid-
vein beneath, 2 veins 6-10/side. Inflorescences 5-1 2 cm
long, 4-1 1 cm broad, pyramidal, purplish to deep lav-
ender in color, peduncles 2-6 cm long, 1 .2-1 .8 mm thick,
glabrous, bracts 2-7 (-10) mm long, linear-subulate,
bracteoles 1-3 mm long, pedicels 0-1 1 mm long. Flowers
glabrous externally, hypanthium 0.6-1 mm long and 1
mm diam. at apex, obconic, calyx lobes 0.3-1 long and
1 mm broad at the base, triangular; corolla tubular and
carnose, purple to lavender, pink, or white, tube 11-18
mm long and 2-4 mm diam.. slightly expanded at the
base, corolla lobes 2-4 mm long; anthers 3.5-4 mm long.
Fruits 4-6 mm long and 4-5 mm diam., globose, drying
black and with longitudinal ridges.

Plants of montane evergreen forest formations,
(750-)1 200-2800 m elevation. Flowering in March,
May-July, and December; fruiting in January-
September and December. The species is found
on the Caribbean slope and continental divide of
the Central Volcanic chain and in the area above
Rio Grande de Orosi and in western and central
Panama.

Palicourea purpurea is distinguished by its lon-
ger purplish corollas, purple inflorescence branch-
es, and globose fruit. It is very similar to P. padi-
folia, but that species has yellow or orange corollas
and red or orange inflorescence branches, and ovoid
or slightly flattened fruit. Palicourea discolor is
also similar but has larger leaves, inflorescences,
and calyx lobes. Compare P. angustifolia also.

Palicourea salicifolia Standl.. J. Wash. Acad. Sci.
18: 280. 1928. P. austinsmithii Standl., Publ.
Field Mus. Nat. Hist., Bot. Ser. 18: 1333. 1938.
P. caerulescens Suesseng., Bot. Jahrb. Syst. 72:
286. 1942. Figure 53.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

209

Shrubs, 0.5-4 tall, leafy stems 1-4 mm thick, glabrous
and quadrangular, young stems drying black, pale gray
in age; stipules with short (1-2 mm) tubes truncated and
with narrow teeth 1-2 mm long, glabrous. Leaves op-
posite, petioles 3-9(-15) mm long, 0.4-0.8 mm thick,
glabrous; leaf blades 4-8(-10) cm long, l-2(-3) cm broad,
narrowly elliptic to lanceolate or lanceolate-oblong, apex
usually tapering gradually and acute or acuminate, base
acute and slightly decurrent on petiole, drying stiffly
chartaceous, glabrous above and below (rarely minutely
puberulent along the mid vein), 2 veins 9-13/side. In-
florescences 2-6 cm long, 3-6(-8) cm broad, pyramidal,
green or marked with blue or purple, peduncles 3-18
mm long, glabrous and usually drying black, bracts 4-7
mm long, 1 mm broad at the base, pedicels 0-8 mm
long, bracteoles ca. 3 mm long. Flowers glabrous exter-
nally, hypanthium ca. 1.3 mm long and 1 mm diam.
distally, calyx lobes 2-3(-4) mm long, ca. 1 mm broad,
oblong and rounded distally; corolla funnelform, car-
nose, white and often flushed with blue or purple, tube
6-13 mm long, ca. 2 mm diam., narrowed in the middle,
corolla lobes 3-6 mm long, to 2 mm broad at the base;
anthers 2-4 mm long. Fruits 6-8 mm long and 5-7 mm
diam., obovoid or ellipsoid, flattened, becoming deep
blue (pedicels also blue), with strong longitudinal ribs
(dried).

Plants of evergreen montane forest formations,
from 1 500 to 2700 m elevation. Flowering in Jan-
uary-September; fruiting in December-March and
May-August. The species is known from Monte-
verde and the northern part of the Meseta Central
(near Zarcero), from the Cerro de Carpintera, and
in the Cordillera de Talamanca.

Palicourea salicifolia is recognized by the small
narrow leaves tapering at both apex and base, small
inflorescences that often dry black, slightly en-
larged calyx lobes, and funnelform white corollas
marked with blue or purple. The fruit have sharply
denned longitudinal ribs. Two collections from be-
low 2000 m (Dryer 768 & 965 from Monteverde
and Stork 1 175 from Carpintera) have dried pale
greenish and the inflorescences are little-branched.
Because of its small leaves, this species resembles
P. adusta and P. montivaga with shorter corolla
tubes and calyx lobes.

Palicourea skotakii C. M. Taylor, Syst. Bot.
Monogr. 26: 76. 1989. Figure 49.

Shrubs, 1.5-2 m tall, leafy stems 1.5-5 mm thick,
glabrous, quadrangular or flattened; stipules 6-8 mm
long, tubular sheath 3-8 mm long, broadly bilobed dis-
tally with obtuse lobes ca. 2 mm long and 3 mm broad,
glabrous, becoming coriaceous. Leaves opposite, petioles
10-24 mm long, 1-1.6 mm thick, glabrous; leaf blades
7-13(-17) cm long, 3-5-5. 5(-6.5) cm broad, elliptic to
elliptic-oblong or elliptic-obovate, apex abruptly nar-

rowed and acuminate, base acute and slightly decurrent
on petiole, drying stiffly chartaceous to subcoriaceous,
dark greenish above, glabrous above and below, 2 veins
9-14/side and joining a vein along the leaf edge distally,
parallel minor 2 veins usually present. Inflorescences
4-10 cm long, 5-1 1 cm broad, usually 3-branched,
rounded to pyramidal, peduncle 2.5-4 cm long and 3
mm thick, glabrous, bracts 1 5-30 mm long, pale green
(dried), bracteoles to 1 5 mm long, ovate and imbricate
around the flowers, persisting. Flowers glabrous exter-
nally, calyx lobes 2-6 mm long, lanceolate to ligulate,
usually unequal; corolla funnelform, carnose, white, tube
ca. 10 mm long, corolla lobes ca. 6 mm long; anthers
ca. 2.5 mm long. Fruits 6-7 mm long, 5 mm diam.,
ellipsoid to slightly obovoid.

Plants of high montane rain forest formations
at 2700-2800 m elevation. Flowering in March
and June. Endemic; like the type (Taylor & Skotak
4756 DUKE), all collections are from along the In-
teramerican Highway in the western part of the
Cordillera de Talamanca.

Palicourea skotakii is distinguished by its high-
altitude habitat, stipules with broad lobes, lack of
pubescence, large greenish ovate persisting bracts
and bracteoles, and white corollas. This species
resembles P. bella\ it also resembles Psychotria
chlorochlamys and its allies.

Palicourea spathacea C. M. Taylor, Syst. Bot. 9:
226. 1984. Figure 49.

Shrubs or small trees, 2-8 m tall, leafy stems 2-8 mm
thick, quadrangular, with thin brownish hairs to 0.5 mm
long but soon glabrescent; stipules 5-12 mm long, 4-8
mm broad, bilobed, puberulent near the base and along
the midrib abaxially. Leaves opposite, petioles 8-20 mm
long, 1-2.2 mm thick, appressed pubescent with thin
ascending hairs or glabrescent; leaf blades 12-28 cm
long, 4.5-14 cm broad, elliptic-obovate to broadly ellip-
tic, obovate or elliptic-oblong, apex short-acuminate with
tip 8-15 mm long, base narrowed gradually and acute
or cuneate, slightly decurrent on petiole, drying stiffly
chartaceous, dark green or brown above (much paler
beneath), glabrous above, with thin brownish ascending
hairs to 1 mm long on the midvein beneath and shorter
(0.1-0.3 mm) hairs on the minor venation and surfaces,
2 veins 12-22/side (and usually with a weaker 2 be-
tween them). Inflorescences solitary, drooping, 10-23
cm long, 8-16(-20) cm broad, open broadly pyramidal
or hemispheric, peduncles 3-9 cm long, 1.5-3.5 mm
thick, puberulent near the base, bracts 1 5-20 mm long
and 5 mm broad, bracteoles 8-15 mm long, ca. 4 mm
broad, pink to purple (white), glabrous pedicels 12-26
mm long, ca. 0.5 mm thick and black when dried. Flow-
ers glabrous externally, hypanthium ca. 1.5 mm long and
1.2 mm diam., cylindrical and drying black, calyx tube
spathe-like and 1 5-20 mm long by 5-8 mm diam., split-
ting open along 1 side, pink to purple; corolla tubular,
camose, white, tube 3-4 cm long, 4-5 mm diam., straight

210

FIELDIANA: BOTANY

or slightly curved, corolla lobes 4-7 mm long; anthers
4.5-5 mm long. Fruits ca. 8 mm long (not including the
persisting calyx), ca. 7 mm in diam., dark purple but
drying black and with longitudinal ribs.

Plants of the very wet lower montane rain forest
formations on the Caribbean slope of the Cordil-
lera de Talamanca, at 1300-1700 m elevation.
Flowering in March, April, and November-De-
cember; fruiting in April-May, July-August, and
December. This species is only known from the
upper drainage area of Rio Grande de Orosi near
Tapanti, Cartago Province.

Palicourea spathacea is a striking species; the
bright pink or purplish bracts, bracteoles, and large
spathaceous calyx make the inflorescences es-
pecially conspicuous. The long white corollas, large
leaves with many secondary veins and cuneate
bases, and stipules with a single obtuse apex on
each side are further distinctions. The calyx is in-
flated before anthesis and splits for half to three-
quarters of its length as the corolla emerges; there
are no calyx lobes.

Palicourea standleyana C. M. Taylor, Syst. Hot.
Monogr. 26: 81. 1989. Psychotria brenesii
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
18: 1347. 1938, not Palicourea brenesii ; Standl.,
1938. Figure 54.

Shrubs or small trees, 2-7 m tall, leafy branchlets 3-
6 mm thick, densely pilose with thin yellowish hairs 1-

3 mm long; stipules (6-)8-18 mm long, 3-8 mm broad,
with a tubular sheath 4-8 mm long, acute or with 2 short
(1-2 mm) lobes, sparsely pilose. Leaves with petioles 1-

4 cm long, 0.7-1.7 mm thick, pilose with straight or
slightly curved hairs to 2 mm long; leaf blades 9-23 cm
long, 3-9 cm broad, broadly elliptic, elliptic-ovate or
elliptic-obovate, apex acuminate to caudate-acuminate
with 8-15(-20) mm long, base obtuse to acute, drying
stiffly chartaceous, dark brown above, densely pilose
above and below with hairs 1-1.5 mm long, 2 veins
(8-) 10-1 7/side, connected distally to make a slightly ar-
cuate submarginal vein. Inflorescences terminal, 3 or 1
with 3 main branches from a short (0-1 5 mm) common
peduncle, 7-12 cm long, to 14 cm broad, open trichot-
omous panicles, primary (basal) lateral branches 3-7 cm
long, pilose, bracts 4-1 1 mm long, ca. 1.5 mm broad,
lanceolate, flowers in close distal clusters subtended by
ovate pilose bracts, pedicels 0-5 long. Flowers puberu-
lent externally, hypanthium ca. 2 mm long, calyx lobes
2-4 mm long but difficult to see among the hairs; corolla
tubular-funnelform, camose, deep lemon yellow to
greenish yellow, tube 10-12 mm long, 1.5-2 mm diam.,
expanded at the base and with hairs to 2 mm long, corolla
lobes 3-5 mm long; anthers 2.5-3 mm long. Fruits 6-8
mm long (not including the 3-mm-long calyx), 5 mm
diam., ellipsoid, pilose, becoming blue.

Plants of very wet cloud forest formations along
the Caribbean slope and continental divide, 1 100-
1600 m elevation. Flowering in December-July;
fruiting in January-June. The species has been col-
lected in the Cordilleras de Guanacaste and Ti-
laran, north of San Ramon (Alajuela), near Ta-
panti (Cartago), in Chiriqui, Panama, and Narino
Province, Colombia.

Palicourea standleyana is recognized by the
prominent pubescence on all parts, the large tu-
bular stipules, stiff leaves with many secondary
veins and submarginal veins, open three-branched
inflorescences, and bright yellow corollas. This
species is easily confused with Psychotria pilosa,
but that species has much smaller corollas and
smaller fruit and the corolla tubes lack the internal
ring of the hairs that define Palicourea.

Palicourea tilaranensis C. M. Taylor, Syst. Bot.
Monogr. 26: 84. 1989. Figure 49.

Small shrubs, ca. 1.5 m tall, leafy stems ca. 3 mm
thick, quadrangular and glabrous, young stems drying
black; stipules 5-8 mm long, with a very short (0.5-1.2
mm) tubular sheath, stipule lobes to 4 mm long, broadly
obtuse to deeply bilobed, persisting. Leaves opposite,
petioles 13-40(-65) mm long, 0.7-1.7 mm thick, gla-
brous; leaf blades 8-16 cm long, 3.5-6 cm thick, ovate
to ovate-elliptic or ovate-oblong, apex acuminate with
tip 7-14 mm long, base obtuse, drying stiffly chartaceous,
dark greenish brown, glabrous above and below, 2 veins
8-1 I/side. Inflorescences 12-15 cm long. 6-9 cm broad,
pyramidal panicles with opposite branching, peduncles
4-6 cm long, 1 .5-2 mm thick, glabrous, bracts 8-1 3 mm
long, 2-4 mm broad, lanceolate, bracteoles ca. 5 mm
long and 2 mm broad, flowers sessile or on pedicels to
4 mm long within the persisting bracteoles. Flowers gla-
brous externally, hypanthium 0.5-1 mm long, calyx lobes
0.3-1.5 mm long, triangular; corolla tubular, carnose,
white, tube 3-7 mm long, ca. 1.8 mm diam., slightly
narrowed in the middle, corolla lobes ca. 2 mm long.
Fruits not seen.

Plants of evergreen lower montane rain forest
formations, at about 15500-1800 m elevation.
Flowering in February and June. This species is
known only from the Monteverde Nature Reserve,
in the Cordillera de Tilaran.

Palicourea tilaranensis is recognized by its re-
stricted range, large and persisting bracts and brac-
teoles, small calyx lobes, and white corollas. It
resembles Psychotria palicoureoides and its allies.

Palicourea triphylla DC., Prodr. 4: 526. 1 830. Psy-
chotria triphylla (DC.) Muell.-Arg. in Mart., Fl.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

Bras. 6(5): 233. 1&&1. Palicourea longibracteata
Bartlingex DC., Prodr. 4: 527. 1830. Palicourea
parviflora Benth., Bot. voy. Sulph. 107. 1844.
Figure 50.

Shrubs or herbaceous subshrubs, l-3(-5) m tall, erect
and usually unbranched (rarely clambering), leafy inter-
nodes 2.3-7 mm thick, glabrous or sparsely pubescent
on the new growth and nodes with short (0.1-0.3 mm)
thin hairs, terete; stipules united for only 1-2 mm at the
base or free, stipule lobes 6/node. 6-12(-15) mm long
and ca. 2 mm broad at the base, narrowly triangular and
acute, glabrous abaxially and ciliolate along the edge.
Leaves 3/node (rarely 2 or 4), petioles 3-7(-l 5) mm long,
1.4-2 mm broad, minutely pubescent but glabrescent;
leaf blades 7-20 cm long, 3-8 cm broad, narrowly ellip-
tic-oblong to lanceolate, narrowly elliptic or oblanceo-
late, apex tapering gradually and acuminate with tip 5-
1 5 mm long, base cuneate to acute (obtuse) and slightly
decurrent on petiole, drying thinly to stiffly chartaceous,
minute (0.05-0.2 mm) thin hairs along the major veins
above, with thin whitish hairs 0.1-0.3 mm long on the
veins and surfaces beneath, 2 veins 7-1 l(-15)/side. In-
florescences 6-20 cm long (to 26 cm in fruit), 5-10 cm
broad near the base, pyramidal to narrowly thyrsoid-
cylindrical, red to orange or orange-yellow, peduncles 5-
12(-18) cm long, 1.5-3 mm thick, puberulent with thin
whitish hairs, bracts 5-20(-30) mm long linear-subulate,
pedicels 1-5 mm long, puberulent. Flowers distylous,
minutely puberulent, hypanthium ca. 1 mm long and
0.5 mm diam., tubular, calyx lobes 0.2-1 mm long,
broadly triangular; corolla tubular or slightly funnelform,
yellow or reddish distally, tube 8-14 mm long, 1.5-3
mm diam., inflated at the base, corolla lobes 1-2 mm
long; anthers 2.5-3.5 mm long. Fruits 4-5 mm long, 3-
5 mm diam., ovoid to globose with prominent longi-
tudinal ridges (dried), becoming blue, purple, or black
at maturity, glabrous or puberulent, calyx ca. 0.7 mm
high.

Plants of open wet or poorly drained sties in
evergreen or partly deciduous forest formations in
the Caribbean lowlands and on the Pacific slope
of central and southern Costa Rica and in Panama,
from near sea level to 700 m elevation, but to 1 300
m on the semideciduous Pacific slope. Flowering
and fruiting throughout the year (but flowering
mostly in April-July). The species ranges from the
Caribbean side of central Mexico and Cuba to
Brazil and Bolivia.

Palicourea triphylla is recognized by its ternate
leaves, puberulence of short thin hairs, long per-
sisting stipules with six teeth per node, orange-
yellow inflorescences with persisting bracts, small
corolla lobes, and gibbous corollas with multicel-
lular hairs. The species is often found in open
swampy sites.

Palicourea vestita Standl., J. Wash. Acad. Sci. 18:
277. 1928. Figure 51.

Shrubs or small treelets, 1-4 m tall, leafy stems 1.5-
5 mm thick, densely hirsutulous with yellowish hairs
0.3-0.9 mm long; stipules with a tube 3-6 mm long, with
narrow teeth 3-8 mm long, puberulent, persisting. Leaves
opposite, petioles 6-1 8(-30) mm long, 0.8-1 .6(-2.2) thick,
densely puberulent with yellowish hairs; leaf blades
(6-)8.5-19 cm long, (1.7-)2.5-7 cm broad, elliptic to
elliptic-oblong, elliptic-obovate or lanceolate, apex acu-
minate (acute) with tip to 1 cm long, base acute to cuneate
(obtuse), drying stiffly chartaceous, often yellowish green,
glabrous above or with thin hairs along the midvein,
pubescent along the major veins beneath with thin yel-
lowish hairs 0.4-1.2 mm long, 2 veins 12-20/side and
with minor short parallel 2 veins between them. Inflo-
rescences 5-12 cm long, 5-8 cm broad near the base,
thyrsiform pyramidal, yellowish, peduncles 2-8 cm long,
1-3 mm thick, densely pubescent, bracts 4-10 mm long
and 1.5 mm broad, bracteoles 1-4 mm long and per-
sisting, pedicels 2-5 mm long. Flowers sparsely and mi-
nutely puberulent externally, hypanthium 1-1.5 mm long,
obconic, calyx lobes 0.7-1.5 mm long, acute; corolla
bright yellow, tube 7-13 mm long, ca. 2 mm diam.,
slightly inflated at the base, corolla lobes 1.5-3 mm long;
anthers 2.1-2.5 mm long. Fruits 5-8 mm long, 3.5-6
mm diam., obovoid, persisting calyx ca. 0.7 mm high.

Plants of evergreen montane wet forest forma-
tions of the Cordillera de Talamanca, from 1 200
to 2600 m elevation. Flowering in April-August;
fruiting in June-August, October, and December.
The species is known only from Costa Rica and
western Panama.

Palicourea vestita is recognized by the short yel-
lowish hairs on most plant parts, the leaves with
many secondary veins, the elongate thyrsiform in-
florescences, and the bright yellow corolla tube.
This species is distinguished from the very similar
but more common P. lasiorrhachis by the more
numerous secondary veins and longer calyx lobes.

Pentagonia Bentham
Nomen conservandum

Shrubs or small trees (rarely large trees), main stems
often unbranched or with few lateral branches, stems
thick, flattened, quadrangular or terete in early stages,
glabrous or puberulent, often with conspicuous rounded
lenticels; stipules interpetiolar, free, large, triangular,
usually early deciduous. Leaves opposite and decussate,
usually very large, sessile or petiolate, the petioles some-
times with auriculate (leafy) developments at the base;
leaf blades entire or pinnatind, usually drying coriaceous,
the minor venation lineolate-parallel and often with 2
distinct orientations in the same leaf area (parallel with
the tertiary veins and at right angles to the tertiary veins),

212

FIELDIANA: BOTANY

domatia absent. Inflorescences axillary to distal leaves,
usually short-pedunculate, cymose or corymbose to glo-
merulate, generally with fewer than 25 flowers, bracts
developed or minute, flowers subsessile or pedicellate.
Flowers bisexual and monomorphic, radially symmet-
rical, often large, usually densely pubescent externally,
hypanthium conical to turbinate or campanulate, calyx
tube well developed, calyx lobes 5-6 or spathaceous,
lobes equal or unequal, often with glands inside at the
base; corolla tubular to funnelform, carnose. white to
yellow or red, glabrous within the throat, villous at the
stamen attachment, corolla lobes 5-6, valvate in bud,
short; stamens 5-6, filaments borne on the middle of the
tube, equal or unequal, usually villous at the base, an-
thers dorsifixed, included; ovary 2-locular, ovules many
on expanded elongate placentas borne on the septum,
stigmas subcapitate or branched. Fruits baccate, fleshy
or becoming hard and nut-like when dry, usually globose
and the surface often lenticellate-muricate, 2-locular, the
large calyx often persisting on the mature fruit; seeds
many, angular.

A genus of about 20 species, ranging from Gua-
temala into northern and western South America.
The very large leaves with distinctive minor ve-
nation, the small axillary inflorescences with larger
crowded flowers, the stiff pubescent tubular co-
rollas, and the globose indehiscent fruit with large
persisting calyx distinguish this genus. The very
large leaves are correlated with a lack of lateral
branches on a solitary vertical trunk in many spe-
cies. The leaves are often reddish or purplish be-.
neath, making the plants even more striking. The
genus is seriously in need of revision. The large
leaves often very variable in form and size, the
compact inflorescences resulting in crushed or hid-
den flowers, and a paucity of collections account
for the fact that the species are still poorly under-
stood.

Key to the Species of Pentagonia

la. Leaf blades entire, never lobed distally; Caribbean lowlands and central cordilleras 2a

Ib. Leaf blades conspicuously pinnatind or pinnately lobed; Pacific evergreen lowlands 6a

2a. Leaves and stems with hairs ca. 1 .4 mm long, leaves sessile, the leaf blade slightly auriculate
near the stem P. hirsula

2b. Leaves and stems with hairs less than 0.5mm long, leaf blades petiolate and never with expanded
auriculate tissue near the base 3a

3a. Calyx lobes usually less than 3 mm long; inflorescences usually open and branched, with the
branches easily seen at anthesis P. costaricensis

3b. Calyx lobes usually more than 4 mm long; inflorescences compact, the branches of the inflo-
rescence short and usually difficult to see (except in fruit) 4a

4a. Bracts of the inflorescence minute or undeveloped; calyx often absent at the apex of the mature

fruit; leaves obtuse to rounded/subtruncate at the base; Guatemala to Costa Rica

P. donnell-smithii

4b. Bracts of the inflorescence conspicuous in early stages, more than 10 m long and 4 mm broad;
calyx persistent on the mature fruit; leaves rarely rounded or subtruncate at the base; south-
ernmost Costa Rica and Panama 5a

5a. Leaves subsessile with a thick petiole less than 2 cm long, lamina long-cuneate at the base;
floral bracts 5-10 mm long P. wendlandii

5b. Leaves with well-developed petioles more than 3 cm long, lamina obtuse to cuneate at the base;

bracts 10-18 mm long P. macrophylla

6a. Leaf blade long-decurrent (at its base) on the petiole . . P. linajita

6b. Leaf blade not at all decurrent on the petiole P. gymnopoda

Pentagonia costaricensis (Standley) W. Burger &
C. M. Taylor, comb. nov. Nothophlebia costari-
censis Standl., Contr. U.S. Natl. Herb. 17: 438.
1914. Figure 14.

Trees, 5-18 m tall, leaf stems 6-12(-22) mm thick,
glabrous or subglabrous, flattened-quadrangular in early
stages, often with dark round lenticels ca. 1 mm diam.;

stipules 2-6(-8) cm long, lanceolate, glabrous or mi-
nutely appressed-sericeous. Leaves with petioles (2.5-)4-
7 cm long. 2.5-7 mm thick, glabrous or subglabrous and
drying brown; leaf blades (2 l-)27-l 00 cm long, ( 1 2-)l 6-
50 cm broad, broadly elliptic-obovate to very broadly
obovate, apex abruptly narrowed and broadly obtuse,
base obtuse and only slightly decurrent (rarely slightly
auriculate at the petiole as in Gomez- Laurito 9321 CR,
F), margin entire, drying coriaceous, glabrous above and

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

213

below or with minute sericeous hairs on the veins be-
neath, 2 veins 8- 11 /side, the minor venation closely
parallel in 2 different directions (but both systems not
always apparent). Inflorescences 3-12 cm long, 3-15 cm
broad, often dichotomously branched, the primary pe-
duncle 5-20 mm long, secondary branches 5-20 mm
long, major branches lacking subtending bracts, ultimate
branches usually with 3-flowered cymes, striate and gla-
brous with elongate (0.4-2 mm) lenticels, pedicels 0-7
mm long. Flowers with hypanthium 2-4 mm long and
2-3 mm diam., obconic, minutely puberulent or gla-
brous externally, calyx tube 47(-12) mm long, 4.5-6
mm diam., cupular to short-tubular, with parallel ve-
nation, rose-colored, calyx lobes 0.5-2 mm high, 2-4
mm broad, broadly rounded to obtuse or obscure, vari-
able and often unequal; corolla cream white or yellow,
tube 12-20 mm long, 3-5 mm diam., minutely yellowish
puberulent externally, corolla lobes 3-6 mm long; an-
thers white. Fruits (based on Hammel& Grayum 14292)
10-20 mm diam., globose with a persistent calyx tube
4-5 mm high and 4 mm diam. at the top, calyx lobes
often obscure.

Trees of evergreen lowland rain forest forma-
tions, 5-600(-900) m elevation. Flowering in
April-May and November; fruiting in May, July-
September, and November. The species is known
only from the Caribbean lowlands, from northern
Costa Rica to western Panama.

Pentagonia costaricensis is recognized by the very
large leaves, unusual minor venation, branched
inflorescences, short calyx lobes, and smaller fruit.
Standley erected the genus Nothophlebia for this
unusual species but admitted (Standley 1938, p.
1329) that it might be referred to Pentagonia. A
collection from 1300-1400 m near Las Alturas
(Almeda et al. 6699 F) with larger calyx lobes and
glabrous corolla tube is tentatively placed here.

Pentagonia donnell-smithii (Standl.) Standl., J.
Wash. Acad. Sci. 17: 170. 1927. Watsonamra
donnell-smithii Standl., Contr. U.S. Natl. Herb.
17: 442. 1914. Figure 14.

Small trees or treelets, 2-7(-12) m tall, often with a
single stem, leafy stems 5-22 mm thick, at first ap-
pressed-sericeous but quickly becoming glabrous; stip-
ules 2.5-9 cm long, 6-25 mm broad, lanceolate to nar-
rowly ovate, with appressed-ascending sericeous hairs
0.2-0.5 mm long externally. Leaves often reddish be-
neath when young, petioles (2-)3-l 2 cm long, 1 .5-6 mm
thick, appressed-puberulent with thin ascending hairs ca.
0.2 mm long; leaf blades (15-)26-90 cm long, (9-) 13-
50 cm broad, broadly elliptic to broadly elliptic-oblong
or broadly ovate, apex bluntly obtuse or rounded, base
rounded and subtruncate to cuneate-obtuse, margin en-
tire and without lobes, drying chartaceous to subcoria-
ceous, pale to dark grayish green, glabrous above, sparse-
ly to moderately puberulent with thin ascending hairs
on the veins beneath, 2 veins 10-14/side, minor ve-

nation closely parallel. Inflorescences 2.5-5 cm long, to
8 cm broad (including both inflorescences of the node),
primary peduncle 1-2 cm long and 2-3 mm thick branches
of the inflorescence usually short and obscured by the
flowers, bracts 0-2 mm long, pedicels 0-6 mm long.
Flowers with hypanthium 3-7 mm long, ca. 3 mm diam.
distally, calyx tube 6-10 mm long, ca. 6 mm diam.,
minutely ascending sericeous, calyx lobes 4-8 mm long,
34 mm broad, broadly rounded distally, corolla yellow
or white, corolla tube 1 5-30 mm long, 3-5(-6) mm diam.
and broadest near the base, with short thin hairs ca. 0.2
mm long, corolla lobes 5, 5-10 mm long, acute. Fruits
16-40 mm diam., globose or globose-ovoid, outer wall
hard and 2-3 mm thick, with small (0.5 mm) tuberculate
lenticels, calyx deciduous or less often persisting on the
mature fruit; seeds 3-5 mm long, ellipsoid, orange.

Trees of evergreen rain forest formations on the
Caribbean slope and lowlands of Costa Rica, from
near sea level to 900 m elevation. Flowering in
March-July and October-November in Costa Rica;
probably fruiting throughout the year. The species
ranges from Guatemala southward along the Ca-
ribbean to central Costa Rica.

Pentagonia donnell-smithii is recognized by the
very large leaves with unusual minor venation, the
small compact inflorescences lacking developed
bracts, the prominent calyx lobes, stifFcorolla tubes,
and usually globose fruit. The mature fruit in most
Costa Rican collections are lacking the calyx, but
the calyx persists regularly in Guatemala and Hon-
duras. Standley (1938) suggested that this species
be submerged in P. macrophylla. But while the
two species appear to be closely related, P. macro-
phylla has a red calyx, white corolla, and large
distinctive bracts subtending the branches of the
inflorescence in early stages. No such bracts are
seen in P. donnell-smithii. Also, this species does
not appear to occur in eastern Costa Rica, where
intergradation might be expected.

Sterile material of these species, and of P. cos-
taricensis and P. wendlandii, may not be separable,
since there is considerable variation within each
species. For example, a dwarf treelet 1 m tall with
very short (2 cm) petioles and slightly obovate
leaves resembling the leaves off. wendlandii was
collected near Puerto Viejo de Sarapiqui (A. Ji-
menez 3424 CR). This collection lacks the bracts
that are found in both P. macrophylla and P. wend-
landii and is therefore placed here under P. don-
nell-smithii.

Pentagonia gymnopoda (Standl.) Standl., J. Wash.
Acad. Sci. 17: 171. 1927. Watsonamra gym-
nopoda Standl., Contr. U.S. Natl. Herb. 1 7: 444.
1914.

214

FIELDIANA: BOTANY

Small treelets, 1-6 m tall, usually with a vertical un-
branched stem, leafy stems 6-16 mm thick, somewhat
flattened in early stages but becoming terete, glabrous,
grayish; stipules 4-6 cm long, 2-3.5 cm broad, broadly
triangular to lanceolate, apparently glabrous or minutely
appressed-puberulent. Leaves with petioles 5-12 cm long,
2.7-5 mm thick, glabrous or minutely (0. 1 mm) puber-
ulent; leaf blades 28-70(-100) cm long, 30-45(-70) cm
broad, ovate to oblong in general outline but deeply
pinnatitid to the distal part of the blade, with 5-7 major
oblong lobes, the lobes 5-25(-35) cm long and 2-7(-8)
cm broad, apex acute to acuminate, base truncate to
acute, drying chartaceous, dull grayish green above, gla-
brous above, minutely (0.1-0.2 mm) papillate-puberu-
lent on the veins beneath, 2 veins 5-9/side. Inflores-
cences closely crowded in the leaf axils, 2-5 cm long,
subsessile and partly covered by the broad stipules or
with a peduncle up to 1 cm long, bracts to 3 cm long,
flowers closely crowded and apparently sessile. Flowers
with hypanthium ca. 10 mm long, densely appressed-
puberulent, calyx tube 6-10 mm long and ca. 6 mm
diam., calyx lobes 5, 18-15 mm long, 5-8 mm broad,
stiff and parallel veined; corolla white to yellow or pale
green, tube ca. 20 mm long, puberulent externally, co-
rolla lobes ca. 10 mm long; stamens 5, filaments 8-10
mm long and unequal, anthers ca. 3 mm long; style ca.
9 mm long. Fruits 15-20 mm diam., globose to ovate
or pyriform, the persisting calyx to 22 mm long and 6
mm diam.

Poorly known plants of evergreen lowland rain
forest formations. Flowering in January-Febru-
ary; fruiting in January-February. The species
ranges from the Carara Biological Reserve
(8435'W) southeastward along the Pacific low-
lands to central Panama.

Pentagonia gymnopoda is recognized by the very
large deeply pinnatifid leaves, small axillary inflo-
rescences, congested flowers, and sessile fruit. The
broad stipules often cover part of the inflorescence.
Currently known from only about four collections
in Costa Rica. Compare the closely similar P. ti-
najita.

Pentagonia hirsute Standl., J. Wash. Acad. Sci.
17: 170. 1927.

Probably small treelets, young branches ca. 10 mm
thick, hirsute; stipules not seen. Leaves sessile and entire;
leaf blades ca. 60 cm long and 27 cm broad, broadly
obovate or broadly elliptic-obovate, apex short-acumi-
nate, lamina gradually narrowed below the middle and
cuneate but merging with a 3-cm-broad basal region that
is slightly expanded (auriculate) and cordate-clasping at
the stem, the auricles ca. 2 cm wide on each side, drying
chartaceous, the upper and lower surfaces with numerous
thin straight hairs 0.7-1 .8 mm long, 2 veins 14-1 7/side,
with 1-3 branches distally and obscurely loop-connected
near the margin. Inflorescences poorly known, small,
with crowded sessile flowers in the leaf axils. Flowers

poorly known, hypanthium densely hirsute, calyx ca. 24
mm long (including both tube and lobes), membranous,
hirsute with whitish hairs. Fruits unknown.

Pentagonia hirsuta is a poorly known species
from about 500 m elevation near Tsaki in the
Talamanca valley, Limon Province, on the Carib-
bean slope of easternmost Costa Rica. The type
and only collection (Tonduz 9415 F, us) was made
in March 1 895. The long hairs on stems and leaves
are very unusual within Pentagonia. The leaf shape
resembles that of P. wendlandii in the long base
but not in its shape; P. wendlandii seems to have
more narrowly obovate leaves not as auriculate at
the base (and without the long hairs).

Pentagonia macrophylla Ik-nth.. Hot. voy. Sulph.
105, t. 39. 1845. Watsonamra macrophylla
(Benth.) Kuntze, Rev. gen. pi. 302. 1891. W.
pubescens Standl., Contr. U.S. Natl. Herb. 17:
441. 1914.

Trees or shrubs to 5(-10) m tall, often unbranched.
leafy twigs 6-12 mm thick, minutely (0.1-0.3 mm) pu-
berulent, quadrangular or becoming terete; stipules 3-7
cm long, 12-15 mm broad at the base, triangular and
acuminate, glabrous or puberulent. Leaves with petioles
3-9(-12) cm long, 2.5-5 mm thick, minutely puberulent
or glabrescent; leaf blades 22^45(-65) cm long 1 1-28(-37)
cm broad, elliptic to elliptic-obovate or very broadly
elliptic, apex gradually narrowed and acute, base grad-
ually narrowed and obtuse or acute, drying subcoria-
ceous, grayish, glabrous above, minutely puberulent be-
neath, 2 veins 9-13/side, minor venation parallel.
Inflorescences axillary, ca. 5 cm long (including the flow-
ers), dense corymbs of 3-7 flowers, sessile or with short
(ca. 1 cm) peduncles, bracts 10-18 mm long. 6-9 mm
broad, oblong-ovate, apex obtuse, red, with many par-
allel veins, minutely sericeous or glabrous with a ciliolate
margin, pedicels 0-4 mm long. Flowers ca. 3 cm long,
hypanthium 6-8 mm long, glabrous or minutely seri-
ceous, calyx lobes 5, 6-12 mm long (becoming 20 mm
long in fruit), to 7 mm broad, usually red; corollas yellow,
sparsely to densely sericeous externally, tube 14-40 mm
long, 3-10 mm diam., corolla lobes 5, 4-7 mm long,
ovate with acute apices; stamens differing in size, anthers
3.5-8 mm long; style 20-25 mm long. Fruits 1 5-28 mm
diam., globose beneath the persisting calyx, red or or-
ange; seeds ca. 4 mm long.

Plants of lowland evergreen forest vegetation,
at 10-900 m elevation in Panama. Flowering
mostly in late April-September in central Panama
(Croat, 1978). The species ranges from near the
Panama border in Limon Province to Colombia.

Pentagonia macrophylla is distinguished from
its congeners by the larger bright red bracts and
red calyx lobes. We have not seen material of this

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

215

species collected in Costa Rica, but the species was
studied near BriBri (Limon) by Lucinda McDade
(see Oecologia 68: 218-223. 1986).

Pentagonia tinajita Seem., Bot. voy. Herald. 134.
1854. P. alfaroana Standl., J. Wash. Acad. Sci.
17: 171. 1927. Figure 14.

Small unbranched treelets, l-3(-4) m tall, stems at
first with 4 prominent ridges or flattened soon becoming
terete, leafy stems 5-12 mm thick, glabrous; stipules 3-
5.5 long, 1.2-2 cm broad, ovate-oblong, obtuse or bifid
with striate parallel venation, puberulent along the mid-
rib. Leaves with petioles 0-1 1(-20) cm long (variable in
length due to the decurrent lamina base), 2.5-4.5 mm
thick; leaf blades 30-80(-100) cm long, 22^10 cm broad,
pinnately lobed and ovate in general outline (from a
cuneate base), the sinuses 5-15 cm deep near the base
and becoming more shallow distally, pinnatifid in the
lower VB of the lamina, sinuses usually rounded, usually
obtuse at the apices, abruptly narrowed and cuneate at
the base with long-decurrent lamina margins running
down to the petiole or to the leaf base, leaves drying
thin-chartaceous and dark green above (paler green be-
neath), with few short (0.2 mm) hairs on the upper sur-
faces, with more frequent short (0.3-0.4 mm) thin hairs
on the lower surfaces, 2 veins 6-12/side, minor veins
0.2-0.4 mm apart. Inflorescences sessile or subsessile,
densely fasciculate, bracts to 2 cm long and 3-4 mm
broad, with parallel longitudinal veins and difficult to
distinguish from the calyx lobes. Flowers with hypan-
thium ca. 5 mm long, calyx tube 10-15 mm long, calyx
lobes 4-8 mm long, 3-4 mm broad, with the same stiff
texture brown color and parallel venation as the bracts;
corolla tube 2-2.5 cm long, 4 mm diam. at the base and
8 mm distally, stiff, corolla lobes 6-8 mm long. Fruits
10-20 mm long and 1 2-1 5 mm diam., globose or ovoid,
the persisting calyx 10-20 mm long (tube and lobes).

Poorly known plants of lowland rain forest for-
mations, 0-200 m elevation. Flowering in Janu-
ary; fruiting in February and April. The species
ranges along the Pacific lowlands of central Costa
Rica to western Panama.

Pentagonia tinajita is recognized by the large
thin pinnatifid leaves with decurrent lamina base,
congested axillary inflorescences, and smaller fruit.
Pentagonia alfaroana (based on Standley 40194
F, us) was said to differ because the larger leaf lobes
are also pinnatifid, but leaf lobing appears to vary
greatly and does not appear to be a sound basis
for separating a species. Compare P. gymnopoda,
which may be conspecific.

Pentagonia wendlandii Hook., Bot. Mag. pi. 5230.
1861. Watsonamra wendlandii (Hook.) Kuntze,
Rev.gen.pl. 320. 1891.

Shrubs or small treelets to 3 m tall, main stems usually
unbranched and with a cluster of leaves at the top, leafy
stems 8-20 mm thick, with 4 prominent ridges, minutely
appressed-puberulent or glabrous; stipules 2.5-6.5 cm
long, lanceolate to ovate-oblong, minutely appressed-
puberulent. Leaves entire and subsessile, petioles 5-15
mm long, 6-10 m thick; leaf blades 50-100 cm long,
25-50 cm broad, narrowly obovate to obovate-oblong,
apex broadly obtuse or rounded, gradually narrowed be-
low the middle, base cuneate and slightly auriculate, leaves
drying stiffly chartaceous or subcoriaceous and brown,
glabrous above, minutely (0.1 mm) puberulent on the
veins beneath or glabrescent, 2 veins 14-16/side. Inflo-
rescences to 5 cm long, with ca. 6-15 flowers, peduncles
ca. 5 mm long, bracts 6-10 mm long, ca. 5 mm broad,
oblong and brown, with whitish sericeous hairs along
the midrib, flowers subsessile (or with pedicels to 1 mm
long, fide Dwyer). Flowers (from Dwyer, 1980) with a
calyx tube ca. 10 mm long, campanulate, purplish red,
stiff, puberulent, with numerous glands at the base with-
in, calyx lobes 5, slightly unequal, 5-10 mm long, 1 calyx
lobe usually short and acute; corolla yellow, tube ca. 25
mm long, narrowly cylindrical, glabrous to puberulent
externally, corolla lobes 4-7 mm long, 2.2-2.6 mm wide,
oblong to triangular; stamens 5, filaments to 1 6 mm long,
unequal, anthers ca. 4 mm long, oblong; style ca. 18 mm
long, stigmas 3.5 mm long. Fruits to 45 mm long, oblong
rotund, drying black, the persisting calyx to 1 5 mm long.

Pentagonia wendlandii occurs in Caribbean
lowland rain forest formations, from near sea level
to ca. 500 m elevation. The species ranges from
Bocas del Toro Province to central Panama, but
we have seen no flowering or fruiting material from
Costa Rica. (A leaf associated with Gomez- Laurito
9321 and collected near BriBri may be this species,
but the flowering sheet appears to be P. costari-
censis.) The species appears to be closely related
to P. macrophylla Benth. of central Panama but
differs in the long-cuneate leaf base with a very
short petiole. See the discussion under P. donnell-
smithii.

Pentas Bentham

Herbs or shrubs, stems erect or clambering, puberulent
in most species, from a fibrous or woody rootstock; stip-
ules interpetiolar with 2-many awns or narrow lobes.
Leaves opposite or in whorls of 3-5, petiolate; leaf blades
entire and usually narrowly ovate to lanceolate, pin-
nately veined, without domatia. Inflorescences terminal
or axillary to the distal leaf pair, usually much-branched
and cymose. Flowers bisexual, radially symmetrical,
monomorphic, dimorphic, or trimorphic, calyx lobes 5,
equal or unequal with 1-3 longer than the others; corolla
usually salverform with a narrow tube, throat pilose
within, corolla lobes 5, ovate to oblong or lanceolate;
stamens usually included, anthers usually linear and with
short filaments; ovary 2- or 3-locular, with many ovules
in each locule. Fruits capsules, usually ovoid and lon-
gitudinally ribbed, beaked, splitting open at apex into 2

216

FIELDIANA: BOTANY

or 4 valves; seeds minute, tetrahedral or subglobose. with
reticulate testa.

A genus of ca. 40 species of Africa, Arabia, and
Madagascar. The following species is a popular
garden ornamental with bright red, rose, lilac, or
white flowers.

Pentas lanceoiata (Forssk.) Deflers, Voy. Yemen,
142. 1889. Ophiorhiza lanceoiata Forssk., F.
Aegypt.-Arab. 42. 1775. Figure 31.

Herbs or subshrubs with erect or clambering stems,
0.5-2 m tall, stems 1.5-5 mm thick, puberulent with
crooked whitish hairs ca. 0.5 mm long; stipules 2-9 mm
long, with a short base and 3-9 slender setae, persisting.
Leaves opposite, petioles 0-5(-l 5) mm long, puberulent;
leaf blades (3-)4-13(-18) cm long, (l-)2-6 cm broad,
ovate-lanceolate to lanceolate or elliptic, apex acute, base
acute and slightly decurrent on petiole, drying stiffly
chartaceous, sparsely to densely puberulent above and
below, 2 veins 6-9/side and strongly ascending. Inflo-
rescences terminal or with the distal leaf pair subtending
the lateral branches, 3-9 cm long and equally broad,
subcapitate or corymbose and hemispheric, puberulent.
Flowers often trimorphic (style exserted and anthers in-
cluded, anthers exserted and style included, both anthers
and style included), calyx tube 1-3 mm long, calyx lobes
5, usually unequal with the longer 4-12 mm long; corolla
brilliant rose red to lilac or white, tube 14-25(-40) mm
long, ca. 0.5 mm diam. but enlarged in the distal 2-6
mm and 3-6 mm diam., throat filled with erect whitish
hairs, corolla lobes 3-10 mm long, acute, ovate-lanceo-
late to ovate-oblong, acute.

A popular ornamental grown in gardens around
the world. In Central America plants with deep
red or pinkish red corollas are most common. It
has become naturalized in Colombia. Compare
Ixora coccinea, another garden favorite.

Pentodon Hochsteter

Annual or short-lived herbs, glabrous; stipules united
to the petiole bases and forming a short truncated sheath
with minute distal lobes. Leaves opposite, sessile or short-
petiolate, entire with obscure pinnate venation, domatia
absent. Inflorescences terminal or axillary and I/node,
racemose with 1-4 distal nodes and 1-4 long- pedicellate
flowers at each node or few flowered and irregular (as in
ours). Flowers bisexual, distylous, glabrous externally,
calyx with 5 small equal lobes; corolla funnelform, tube
glabrous or with hairs at the throat within, corolla lobes
5; stamens 5, included or exserted, subsessile from near
the base of the tube or between the petal lobes; ovary
2-locular, with many ovules in each locule borne on
peltate placentas from the septum. Fruits capsules, lo-
culicidally dehiscent; seeds many, small, black.

A genus of probably two African species, with
one now introduced into pans of the Americas.
These plants are distinguished from the closely
related Oldenlandia by the five-parted flowers.

Pentodon pentandrus (Schumach. & Thonn.)
Vatke, Oest. Bot. Zeitschr. 25: 231. 1875. Hed-
yotis pentandra Schumach. & Thonn.. Kongel.
Dansk. Vidensk. Selsk. Naturvidenske, Math.
Am. 3: 71. 1827. Oldenlandia pentandra (Schu-
mach. & Thonn.) DC., Prodr. 4: 427. 1830. H.
halei Torr. & Gray, Fl. N. Amer. 2: 42. 1841.
P. halei (Torr. & Gray) Gray, Syn. Fl. N. Amer.
l.pt. 2: 28. 1884.

Herbs with weakly erect or decumbent stems 10-30
cm high, stems 0.8-1.8 mm thick, glabrous and slightly
succulent; stipule sheath 0.5-1.5(-3) mm long, entire or
with 1-3 short (0.3-2 mm) subulate lobes. Leaves with
poorly defined petioles 0.5-10 mm long (in ours); leaf
blades 1.5-4(-8) cm long, 0.4-2.5 cm broad, narrowly
ovate-elliptic to elliptic (elliptic-lanceolate to linear-lan-
ceolate), apex obtuse to acute, base obtuse to cuneate
and decurrent on the petiole, drying membranaceous or
thin-chartaceous (often translucent), greenish and gla-
brous on both surfaces, 2 veins 3-4/side but difficult to
see. Inflorescences 2-5 cm long, of (l-)3-5 irregularly
arranged flowers in leaf axils or at apex (elongate race-
mose inflorescences not seen in North American mate-
rial), peduncles 0-8 mm long, pedicels 2-5 mm long.
Flowers glabrous externally, hypanthium 2-3 mm long,
calyx tube 0.3-1 mm long, calyx lobes 0.5-1 .5 mm long,
acute; corolla white in ours, tube 1.5-4 mm long, 2-3
mm diam., corolla lobes 5, 1-3 mm long, 0.7-2 mm
broad, ovate-triangular. Fruits 2.5-4 mm long, 2-3 mm
diam., obconic to oblong; seeds 0.3 mm long.

Plants of moist depressions in open sunny sites,
often along streams. Collected in central Nicara-
gua as early as 1869 and recently found along Rio
San Juan, and in Belize. Originally in tropical Af-
rica, Texas to Florida, and Cuba.

Pentodon pentandrus is recognized by its deli-
cate habit with small thin ovate-elliptic leaves,
small few-flowered and irregularly arranged inflo-
rescences, small five-parted flowers, and small
capsules with many seeds. The North American
material (as represented by P. halei from the
southern U.S.A. and illustrated in Correll & Cor-
rell, 1982, fig. 615) differs greatly from typical Af-
rican collections (Verdcourt, 1976, p. 264, fig. 38).
Measurements in parenthesis in the description
represent African material. Nevertheless, the spe-
cies varies greatly in Africa, and the North Amer-
ican material probably represents an early atypical
introduction (Verdcourt, 1976, p. 263). Compare
Sipanea biflora.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

217

Pogonopus Klotzsch

Trees or shrubs; stipules interpetiolar, free, small, del-
toid to narrow, deciduous with the leaves. Leaves op-
posite, petiolate; leaf blades usually thin, entire, pin-
nately veined, domatia absent. Inflorescences terminal,
paniculate and subcorymbose-cymose, pedunculate,
bracteolate, 1 or 2 flowers of the inflorescence usually
with a greatly enlarged sepal lobe forming a broad pet-
iolate leaf-like and colorful "petal." Flowers bisexual,
radially symmetrical (except for those with a single great-
ly enlarged sepal lobe or where the tube is curved), hy-
panthium turbinate, calyx tube short, calyx lobes 5, den-
tate and deciduous (each inflorescence usually with 1 or
2 flowers with a greatly expanded calyx lobe); corolla
cylindrical, straight or curved, barbate in the throat, co-
rolla lobes 5, short, valvate in bud; stamens 5, inserted
near the base of the corolla tube, filaments slender and
glabrous, anthers versatile dorsifixed above the middle,
exserted; ovary 2-locular, ovules many on placentas lon-
gitudinally adnate to the placenta, style slender with 2
linear or oblong branches. Fruits capsular, subglobose
to oblong-ovoid, 2-locular, areolate at apex, loculicidally
bivalvate; seeds many, horizontal, crowded.

A neotropical genus of three species; one species
ranges from Mexico to South America and is grown
as an ornamental. The greatly enlarged and bright-
ly colored laminar sepal lobe of one or two flowers
in each inflorescence is distinctive (but compare
Mussaenda erythrophylld).

Pogonopus speciosus (Jacq.) Schum. in Mart. Fl.
Bras. 6, pt. 6: 265. 1889. Macrocnemum spe-
ciousum Jacq., Hoit. Schoenbr. 1: 19,1.43. 1797.
Macrocnemum exsertum Oersted, Vidensk.
Meddel. Kjobenhavn 45. 1852. P. exsertusOer-
sted, Amer. Centr. 17. 1863. Figure 16.

Shrubs or small trees, 2-6(-l 2) m tall, leafy stems 1.5-
5 mm thick, minutely (0.2-0.4 mm) appressed-pubes-
cent and glabrescent in age, the older stems with con-
spicuous rounded to linear whitish lenticels 0.5-1 (-2. 3)
mm long; stipules 1-3.5 mm long, 3-4 mm broad and
with a rounded or cuspidate tip to 1 m long. Leaves with
petioles 2-15 mm long, 1-2 mm broad, appressed-pu-
bescent, often with lateral wings continuous with the
lamina margins; leaf blades 7-18(-25) cm long, 3-
7.5(-9.5) cm broad, elliptic-obovate or elliptic-oblong
(smaller leaves near the inflorescence usually ovate), apex
acuminate with tip 5-15 mm long, base tapering grad-
ually and cuneate-acute, decurrent on the petiole, drying
thin-chartaceous, glabrous above, sparsely pubescent on
the veins beneath with thin whitish hairs 0.2-0.4 mm
long, 2 veins 4-10/side, the minor venation forming a
small reticulum on both surfaces. Inflorescences termi-
nal or axillary, 4-18 cm long, peduncles to 5 cm long,
main axis usually with opposite branches, minutely ap-
pressed-pubescent, flowers usually in compact distal cy-
mose gruoups, bracteoles 2-4 mm long, lanceolate to
linear, pedicels 0-3(-5) mm long. Flowers with hypan-

thium 3 mm long, 2-3 mm diam., conical-turbinate,
sparsely and minutely puberulent, drying dark, calyx lobes
0.6-2 mm long, 0.2-0.3 mm broad, expanded calyx lobe
with a petiole 8-20 mm long and lamina 2-4(-6) cm
long and 1.5-3.5(-5) cm broad, broadly ovate or ovate-
orbicular, bluntly obtuse to rounded, dark red to bright
red or rose red, palmately veined; corolla tubular, rose
red to dark red, tube 1 2-28 mm long, straight or curved,
3-6 mm diam. distally, densely and minutely pubescent
with ascending hairs, corolla lobes 3-5 mm long, 2-3
mm broad at the base; stamens with filaments 20-23
mm long, anthers ca. 2.5 mm long; style 30-35 mm long,
stigmas 2-2.5 mm long. Fruits 6-9 mm long, 5-7 mm
diam., oblong-urceolate, minutely (0. 1 mm) puberulent,
with 2 longitudinal sulci, abruptly truncated at apex, the
linear calyx lobes often persisting.

Ornamental plants grown in both evergreen and
deciduous environments in Central America, from
near sea level to about 500 m elevation. Probably
flowering principally in the rainy season: August-
December. The species grows wild in southern
Mexico, the Guanacaste lowlands, and central
Panama and in Colombia and Venezuela.

Pogonopus speciosus is recognized by the bright
red or rose petaloid expansions of the calyx in a
few flowers of each inflorescence. This species is
not known from the wild in Nicaragua, El Sal-
vador, or Honduras; it is possible that it was
brought to Mexico as an ornamental in pre-Co-
lumbian times. Compare Mussaenda erythrophyl-
la, another ornamental with bright red petaloid
sepal lobes.

Posoqueria Aublet

Trees, treelets, or shrubs, branches soon becoming
terete, glabrous or pubescent, usually thick; stipules in-
terpetiolar, free, large, triangular, usually early decidu-
ous. Leaves opposite and decussate, petiolate; leaf blades
usually large, entire, often coriaceous, usually with
domatia. Inflorescences terminal, usually few-flowered
and short (but the flowers very long), corymbose to cy-
mose or umbellate, pedunculate, bracts absent or min-
ute, flowers pedicellate. Flowers bisexual and mono-
morphic, (4-)5(-6)-parted, radially symmetrical or
bilaterally symmetrical because of the curved corolla tube,
hypanthium little differentiated from the pedicel, calyx
tube short, calyx lobes short-dentate, usually persisting,
often auriculate at the base, with glands within; corolla
long-salverform or long-tubular, white, corolla tube long
and narrow, glabrous externally, glabrous or glandular-
papillate within, corolla lobes imbricate or contorted in
bud, rotate or reflexed; stamens unequal or subequal,
short and inserted on the mouth of the tube, glabrous or
pilose, anthers basifixed and sagitate, linear-oblong, ex-
serted, (the pollen is said to be released explosively in
some spp.); ovary 2-locular or incompletely 1-locular,
ovules many in each locule and erect on stipitate bila-
mellar placentas borne on the septum, style filiform, as

218

FIELDIANA: BOTANY

long as the tube, stigmas short and bind. Fruits baccate
or drying hard and indehiscent. globose to ovoid, 1 - or
2-locular, often large; seeds large, hard, rounded and
obtusely angled to flattened, imbedded in a gelatinous
pulp.

A Neotropical genus of 1 2- 1 6 species, with most
of the species in South America; a single species
ranges as far north as southern Mexico. The genus
is distinctive because of the larger stiff leaves, large
stipules, the long white flowers, and the globose

indehiscent fruit with large seeds imbedded in a
fleshy pulp. The corolla tubes are usually more
than 10 cm long and less than 1 cm diam. The
flowers are strongly fragrant at night when they
open but yellowish and odorless by morning; these
are probably pollinated by long-tongued moths. In
some species the stamens are held asymmetrically
and under tension over the mouth of the corolla
tube until they are disturbed, when they snap for-
ward. Compare Borojoa and Tocoyena.

Key to the Species of Posoqueria

la. Corolla tubes (20-)30-38 cm long; leaf blades (10-) 18-46 cm long leaf blades usually puberulent
with very thin hairs beneath (but difficult to see, except along the larger veins), usually slightly rough
and minor venation obscure beneath (uncommon) P. grandiflora

Ib. Corolla tubes 8-20 cm long; leaf blades 10-20(-24) cm long, essentially glabrous beneath .... 2a

2a. Leaf blades drying dull above with the tertiary veins not easily seen, the minor venation not raised;

fruit dull yellow to orange, ellipsoid to ovoid, with pericarp leathery and 410 mm thick

P. coriacea

2b. Leaf blades lustrous above with tertiary veins easily seen and sometimes slightly elevated; fruit
orange and globose, with pericarp hard and brittle, 1-3 mm thick P. latifolia

Posoqueria coriacea M. Martens & Galeotti, Bull.
Acad. Brux. 11: 240. 1844.

Small trees or shrubs, 3-8(-20) m tall, branchlets 2-
7 mm thick, glabrous; stipules 8-12(-28) mm long, ovate,
ovate-oblong to ovate-lanceolate or suborbicular, acute,
glabrous. Leaves with petioles 6-22 mm long, glabrous;
leaf blades (6-)10-22(-27) cm long, (3-)5-12(-18) cm
broad, narrowly elliptic to ovate or narrowly obovate,
apex obtuse to abruptly acute, base cuneate to rounded
and subtruncate, drying coriaceous, 2 veins 4-8/side.
Inflorescences 2-4 cm long and 2-3 cm broad (not in-
cluding the corolla lengths), cymose-corymbose, with 7-
25 flowers, peduncles 1-2 cm long, glabrous or minutely
puberulent, pedicels 5-10(-23) mm long, glabrate. Flow-
ers with hypanthium + calyx 6-8 mm long, subulate
glands present within the calyx, calyx lobes 5, unequal,
0.5-1.2 mm long, rounded; corolla greenish white, gla-
brous externally, long-tubular, tube 9-20 cm long, 1.5-
3 mm diam. in the lower half, pilose within the throat
and at the base of the lobes, corolla lobes 5, 15-25 mm
long, unequal, rounded distally; anthers 7-10 mm long.
Fruits 7-10 cm diam., ellipsoid to ovoid, brown, peri-
carp leathery, 4-10 mm thick; seeds 10-12 mm diam.,
angular, aril white.

Plants of lowland rain forest formations, from
near sea level to 1 100 m elevation (to 1500 m in
South America). Flowering January-March; fruit-
ing in March-April, June-August, and October-
November. The species ranges from Mexico to
Brazil.

Posoqueria coriacea is characterized by the long-

tubular flowers, dull leaf surfaces, and fruit with
thick leathery rind. The 2 veins are often some-
what impressed in thick mature leaves when dried.
The larger leaf dimensions noted above in paren-
theses are from Steyermark's Flora of Venezuela
treatment ( 1 974) and may not occur in Costa Rica.
The breeding system of this species was studied
by Bawa and Beach (1983).

Posoqueria grandiflora Standl., J. Wash. Acad. Sci.
18: 166. 1928. P. maxima Standl., Publ. Field
Columb. Mus., Bot. Ser. 7: 57. 1930. Figure 15.

Small trees, treelets, or shrubs, 2-7(-17) m tall, trunk
to 18 cm dbh, leafy stems 2.5-12 mm thick, glabrous or
minutely puberulent with thin erect hairs 0.1-0.2 mm
long; stipules 8-18 mm long, 4-1 1 mm broad, triangular,
glabrescent, deciduous. Leaves well spaced along the stem,
petioles 7-20 mm long, 2-4 mm thick, glabrous or mi-
nutely puberulent; leaf blades ( 1 3-)l 8-34(-46) cm long,
6-1 7(-23) cm broad, elliptic to elliptic-obovate or ellip-
tic-oblong, apex acuminate to short-acuminate or blunt-
ly rounded, base tapering gradually to slightly rounded
and obtuse or acute, drying subcoriaceous to coriaceous,
pale grayish green beneath, glabrous above, with thin
erect hairs 0.1-0.5 mm beneath or the hairs difficult to
see and apparently glabrous, slightly rough to the touch
beneath, 2 veins 4-8/side and prominent on both sur-
faces, minor venation obscure on both surfaces. Inflo-
rescences only 3-6 cm long (not including the corolla
tubes), corymbose with ca. 10-20 flowers, peduncles 1-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

219

2 cm long and 3-8 mm thick, pedicels 2-12 mm long,
ca. 2 mm thick, glabrous or minutely puberulent. Flow-
ers glabrous, hypanthium 3-6 mm long, little differen-
tiated from the pedicel, calyx tube 3-6 mm long, 3-6
mm diam., calyx lobes 5, 0.5-1.5 mm long, broadly
rounded distally; corolla long-tubular with rotate lobes,
white, tube ( 1 2.5-) 1 5-36 cm long 2.5-4 mm diam., lobes
5, 20-38 mm long, 7-1 3 mm broad, oblong-obovate and
rounded; stamens with unequal filaments to 1 5 mm long,
0.5-1 mm thick, anthers 9-12 mm long, to 2 mm broad,
linear-lanceolate. Fruits 7-12 cm long, 4-7 cm diam.,
ovoid to ellipsoid, with a slightly roughened bark-like
(minutely lenticellate) brownish surface, pericarp 5-12
mm thick; seeds 6-10 mm long, orange to white.

Trees and treelets of evergreen lowland Carib-
bean rain forest formations, from 40 to 300 m
elevation. Flowering in February, April, July, and
September-October; probably fruiting throughout
the year. The species ranges from northeastern
Costa Rica to Colombia.

Posoqueria grandiflora is recognized by its very
long flowers on small terminal inflorescences, the
relatively large corolla lobes, and the large leaves
with obscure minor venation and pubescence (when
present), which makes the lower leaf surfaces
slightly rough to the touch. This species appears
to be less common than its congeners in Costa
Rica. The species has been called "wild coffee" in
the Caribbean lowlands. An unusual collection
(Wilbur 207 '11 DUKE, F) from southern Costa Rica
has large leaves with conspicuous long hairs, and
flowers with very long corolla tubes, but is prob-
ably no more than an extreme form of this species.
The Colombian material ascribed to P. maxima
appears to be conspecific; Steyermark erred in
making it a subspecies of P. coriacea Mart. & Gal.

Posoqueria latifolia (Rudge) Roem. & Schult., Syst.
Veg. 5: 227. 1819. Solena latifolia Rudge, PI.
Guian. 1: 26, t. 40. 1806. Stannia panamensis
Walp. & Duchass., Linnaea 23: 755. 1850. P.
panamensis (Walp. & Duchass.) Walp., Ann.
Bot. Syst. 2: 797. 1852. Figure 15.

Small trees to 9(-20) m tall, trunks to 25(-40) cm dbh,
wood hard and reddish, leafy stems 2-6 mm thick, gla-
brous; stipules 7-1 8 mm long, 3-8 mm broad at the base,
triangular to ovate-oblong, sometimes slightly (1 mm)
united above the petioles, apex obtuse to acuminate, stiff,
glabrous. Leaves distant along the stem, petioles (4-) 7-
20 mm long, 1 . 5-4 mm thick, glabrous; leaf blades (7-) 1 0-
20(-24) cm long, (3-)4-10(-14) cm broad, elliptic-ob-
long to elliptic-ovate or ovate, apex obtuse or short-
acuminate, base abruptly narrowed and obtuse or round-
ed, drying stiffly chartaceous to subcoriaceous, glabrous

above and below, 2 veins 5-7/side, the minor venation
visible on both surfaces. Inflorescences 2-5 cm long (not
measuring the corolla tubes), corymbose with 7-1 8 flow-
ers, peduncles 1-2 cm long, 2-3 mm thick, bracteoles
ca. 0.5 mm long, pedicels 3-9 mm long, glabrous. Flow-
ers glabrous externally, sweet aromatic, hypanthium 3-
6 mm long, calyx lobes poorly developed, 0-0.5 mm
long; corolla long-tubular with usually reflexed lobes,
white, tube 8-14(-16) cm long, 2-3.3 mm diam., lobes
5 (4), 12-20(-26) mm long, 4.5-5.5 mm broad, narrowly
oblong, rounded; stamens 5 (4), filaments to 6 mm long,
ca. 0.4 mm thick, anthers 6-7.5 mm long, ca. 1 mm
broad, linear-oblong, with basal lobes ca. 1 mm long;
style ca. 9 cm long. Fruits 4-6 cm diam., globose to
ovoid, yellow or orange at maturity, the pericarp only
1-3 mm thick, surface becoming wrinkled; seeds 6-12
mm long, often triangular, translucent in life, arils white
to yellow-orange, in a fleshy sweet pulp.

Trees and treelets found in rain forests, partly
deciduous forests, and moist sites in deciduous
forests, from 2 to 700(-1200) m elevation. Prob-
ably flowering and fruiting throughout the year
(but mostly flowering in March-October and fruit-
ing in October-April). This common species oc-
curs in all the lowland evergreen areas of Costa
Rica. The species ranges from southern Mexico to
the Amazon basin of Brazil and Bolivia.

Posoqueria latifolia is recognized by the larger
stiff glabrous leaves, the small terminal inflores-
cences with very long tubular flowers, and the glo-
bose fruit. The barely visible minor (3 and 4)
venation appears to be a consistent way of differ-
entiating the leaves of P. latifolia from those of P.
coriacea. Croat (1978, p. 814) remarks that the
anthers are united at anthesis along one side of the
tube and burst apart when contacted. He also notes
that the flowers open late in the day and do not
persist on the following day. Common names re-
corded in Costa Rica are boca de vieja, carica,fruta
de mono, guayaba de mono, guayaba mica, man-
zana de mico, picarito, and querica. The names
jicarillo and querica are used in southeastern Nic-
aragua. The fruit's pulp is edible and sweet.

Psychotria Linnaeus
Nomen conservandum

REFERENCES C. Hamilton, A revision of Me-
soamerican Psychotria subgenus Psychotria (Ru-
biaceae), part I: Introduction and species 1-16.
Ann. Missouri Bot. Gard. 76: 67-1 1 1 . Part II: Spe-
cies 17-47. Ann. Missouri Bot. Gard. 76: 386-
429. Part III: Species 48-6 1 and appendices. Ann.
Missouri Bot. Gard. 76: 886-916. 1989. A. Mo-

220

FIELDIANA: BOTANY

lina, Revision de las Especies de Cephaelis en
Mexico, Centre America y las Antilllas. Ceiba 4:
1-38. 1953. (Other references are listed under spe-
cific species below, under the family description,
and at the end of the text.)

Shrubs or small treelets, less often medium-sized trees
or herbaceous subshrubs (rarely lianas), terrestrial or rarely
epiphytic, stems often slightly thickened at the nodes,
glabrous or puberulent; stipules usually united and in-
terpetiolar with 1 or 2(-5) apices or lobes on each side,
sometimes also united around the stem above the pet-
ioles (intrapetiolar) and forming a short tube or sheath,
rarely separate to the base and appearing free, deciduous
or persisting, colleters often present on the stems at the
adaxial base of the stipules (and drying reddish in sub-
genus Psychotria). Leaves opposite and decussate (rarely
3 or 4/node), petiolate or rarely subsessile, usually acu-
minate at the apex; leaf blades entire and pinnately veined,
raphides (cystoliths) present and obscure or conspicuous,
some species with domatia in the vein axils beneath
(some African species with bacterial nodules in the leaves).
Inflorescences mostly terminal and occasionally becom-
ing pseudoaxillary by further growth of an axillary branch,
less often consistently axillary, usually solitary at a node,
usually pedunculate and paniculate with opposite
branching, bracts large to small or rarely undeveloped,
bracts forming an involucre in some species formerly
placed in Cephaelis, flowers often borne in distal brac-
teolate cymes, sessile or pedicellate. Flowers bisexual (in
Central America), radially symmetrical, usually small,
often distylous with long-styled (pin) and short-styled
(thrum) forms within the same species, calyx tube usually
short and cupulate, with 4 or 5(-6) short calyx lobes or
without lobes and entire; corolla tubular to funnelform
or campanulate, white to pink or yellowish, corolla tube
usually short or narrow, often with tufted hairs at the
throat within, glabrous or puberulent externally, corolla
lobes 4-5(-6), always valvate in bud, often thickened at
the tips; stamens 4-5(-6), borne from the middle or up-
per part of the tube, filaments slender, anthers narrow,
included or exserted, disc ring-shaped and encircling the
base of the style; ovary usually 2-locular (rarely with 4-
6 locules), 1 erect ovule borne from the base of each
locule or from the base of the thick septum, style long
or short, with 2 (rarely 4-6) linear stigmas. Fruits fleshy
drupes, red, yellow, blue, purple, black, or white when
ripe, often with spongy arenchymatous tissue, usually
with 2 (4-6) hard pyrenes, pyrenes plano-convex with a
flattened interior (adaxial) face and a rounded exterior
(abaxial) surface, often with longitudinal ridges on the
exterior surface (rarely with transverse ribs or projec-
tions), usually with a median longitudinal sulcus on the
inner face.

Psychotria is the largest genus of Rubiaceae and
one of the largest genera of Angiosperms, with an
estimated 1,500-1,600 species in the tropics and
subtropics of both hemispheres. Together with
Piper (Piperaceae) and Miconia (Melastomaceae),

Psychotria is one of Costa Rica's three most spe-
ciose woody genera, especially common in the un-
derstory of evergreen forests and forest edges. Some
species are difficult to separate from similar spe-
cies in Coussarea, Faramea. and Palicourea (q.v.).
While Cephaelis appeared to be a very distinctive
genus in Central America, a great number of in-
termediate species have necessitated the inclusion
of its species in Psychotria. Our treatment has ben-
efited greatly from Hammers work on Psychotria
at La Selva (in Taylor, 1991). Likewise, Hamil-
ton's recent revision (see references above) of the
species in subgenus Psychotria has been especially
helpful, as these species are often difficult to dif-
ferentiate; we have not deviated from his species
concepts in this treatment.

Psychotria is generally characterized by the sol-
itary terminal inflorescences with opposite
branching (in most species), the 1 - or 2-lobed in-
terpetiolar stipules, entire and pinnately veined
opposite leaves, smaller flowers with minute calyx
lobes, short corolla tubes, valvate corolla lobes,
basal solitary ovules, and fleshy fruit usually with
2 hard seed-like pyrenes. The short shrubby habit,
preferences for areas of high rainfall, fruit often
with longitudinal ridges, and white or yellowish
flowers are additional characteristics. Those spe-
cies with axillary inflorescences tend to have un-
branched succulent stems or few-branched woody
stems (see below). All the fruits appear to be bird-
dispersed; they are fleshy and bright red, blue to
purplish and black, or arenchymatous and whitish.
Two species of Psychotria found in Costa Rica, P.
emetica and P. ipecacuanha, are used medicinally.

Most species of Psychotria in Costa Rica are
readily recognizable but the differences between
closely related species can be subtle. There appear
to be real problems of intcrgradation in the epi-
phytic species, and the succulent-stemmed her-
baceous species with axillary inflorescences pre-
sent problems of species delimitation. Except for
the groups just mentioned, the majority of species
stand well apart and give little evidence of hy-
bridization or intergradation. The difficulty in
identifying a specimen to species often is due to
the large number of species in the genus, and the
wide range of variation found within some species.
Below we provide keys to four groups within the
genus; groups 2 and 3 appear to be largely mono-
phyietic. The illustrations are an additional ave-
nue for identification and are grouped by inflo-
rescence position, subgeneric placement, and
general similarity.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

221

Key to Four Species Groups of Psychotria in Costa Rica

1 a. Plants nearly always epiphytic and with subcoriaceous leaves drying grayish; ovary usually with 4
locules; fruits red and often with 4 seeds; or plants of Cocos Island Group 1

Ib. Plants not epiphytic, leaves various; ovary with 2 (rarely 5) locules; fruit usually with 2 (rarely 5)
seeds; and plants not known from Cocos Island 3

2a. Inflorescences regularly axillary; main stems usually short (ca. 1 m) and succulent in a majority of
species, both succulent and woody stems often with hollow sections within when dried . . . Group 2

2b. Inflorescences terminal, only occasionally axillary where lateral shoots have continued to develop;
main stems usually woody and rarely with hollow sections when dried 3

3a. Leaves drying greenish, gray-green, yellow-green, yellowish brown, or brown (note that leaves treated
with isopropyl alcohol may turn reddish brown and that some species have pinkish venation in
life), domatia rarely present; stipules often persisting, the stipules not subtending and enclosing a
ring of reddish hairs (colleters) in early stages; fruit usually becoming blue, purple, or black (red
only in P. haematocarpa with very small inflorescences, or orange in later stages in P. racemosa
with 5 locules and 5 -seeded fruit); subgenus Heteropsychotria Group 3

3b. Leaves drying grayish to black, pinkish brown or dark reddish brown, domatia present in a few
species; stipules usually falling off as the leaves expand (caducous), and usually enclosing a short
ring of reddish hairs (these often persisting just above the stipule scar); fruit always red at maturity
(never with more than 2 seeds); subgenus Psychotria Group 4

Key to Group 1: Epiphytic Species of Psychotria in Costa Rica and Three Species of Cocos Island

The epiphytic species appear to represent a very natural (probably monophyletic) group. However,
there is the strong likelihood that a large number of our collections belong to a single polymorphic entity,
for which P. guadalupensis is the earliest name. Psychotria maxonii appears to be quite distinct, but
there are a significant percentage of collections that appear to be intermediate between P. guadalupensis
and P. pithecobia. See the discussion under P. guadalupensis.

la. Epiphytic plants of continental Costa Rica 2

1 b. Trees and shrubs of Cocos Island 4

2a. Leaves to 5(-7) mm wide, 10-28 mm long, narrowly lanceolate to narrowly oblong; inflores-
cences to 3 cm long P. maxonii

2b. Leaves 8-40 mm wide, 20-120 mm long, more than 40 mm long if lanceolate or narrowly

oblong; inflorescences 3-14 cm long 3

3a. Leaves with 6 or more pairs of secondary veins, the veins clearly elevated and visible on the
dried leaf surfaces, leaves more than 4 cm long and 2 cm broad; inflorescences with peduncles

2-7 cm long P. pithecobia

3b. Leaves with fewer than 6 pairs of major secondary veins, the veins often obscure on the surfaces

of the dried leaves, leaves 3-10 cm long and 1-5 cm broad, very variable (on different plants)

as regards size, shape, and texture; peduncles 0.5-3 (rarely to 5) cm long . . .P. guadalupensis

4a. Leaves to 25 cm long, drying subcoriaceous and dark reddish brown; stipules forming an acute

F/CMs-like cap over the shoot apex and early deciduous P. cocosensis

4b. Leaves to 1 5 cm long, drying thin-chartaceous and greenish or grayish; stipules with 2 acute lobes

on each side and persisting 5a

5a. Inflorescences less than 3 cm long, with thick (1 mm) lateral branches less than 5 mm long; stipule

lobes 24 mm long P. brachybotrya

5b. Inflorescences usually becoming more than 3 cm long, with slender (0.3 mm) lateral branches 5-
10 mm long; stipule lobes ca. 1 mm long Psychotria sp. A

Key to Group 2: Species of Psychotria with Axillary Inflorescences

la. Inflorescences axillary to both leaves of the node, usually with 2 or more inflorescences at distal
nodes, inflorescences usually less than 5 cm long; fruits becoming blue, black, or purple 2

222 FIELDIANA: BOTANY

Ib. Inflorescences axillary to only 1 leaf at each node, usually with 1 inflorescence at distal nodes,

inflorescences often more than 5 cm long; fruits white to yellow or red .8

2a. Inflorescences subsessile and forming a dense verticillate cluster at the node 3

2b. Inflorescences short-pedunculate, not forming dense verticillate clusters around the node

3a. Inflorescences lacking an involucre of bracts, calyx lobes to 3 mm long (not Psychoiria

spp.) Hoffmannia spp.

3b. Inflorescences subtended by an involucre of broad bracts, calyx lobes to 2 mm long

4a. Bracts subtending the inflorescences pale green; stipules to 10 mm long and 5 mm

broad; widely distributed at 1 200-2800 m elevation P. aubletiana

4b. Bracts reddish; stipules 12-22 mm long and almost as broad; 700-1500 m near Orosi

and Muneco P. cartagoensis

5a. Leaves drying greenish to greenish brown, shrubs or small trees with many branches, young
stems glabrous; stipules forming a short sheath; inflorescences with many branches, often

with more than 1 fruits P. cooperi

5b. Leaves drying grayish or dark brown, subshrubs or few-branched shrubs, young stems mi-
nutely puberulent; stipules not forming a sheath; inflorescences few-branched, rarely with

more than 5 fruits 6

6a. Inflorescences usually 4/node; plants growing to 2(-5) m tall, with lateral branches; leaf blades

drying stiffly chartaceous [often obtuse at the base; peduncles to 1 5 mm long] . . . P. erecta

6b. Inflorescences 2/node; plants growing to 1 m tall, unbranched; leaf blades drying thin-char-

taceous 7

7a. Leaf blades drying brownish, often cuneate basally; peduncle to 5 mm long, inflorescence

less than 2.5 cm long P. emetica

7b. Leaf blades drying greenish, often decurrent basally; peduncle to 70 mm long, inflorescences

more than 3 cm long P. aggregata

8a. (from Ib) Stipules lobed or fimbriate distally, to 3 cm long, translucent to opaque 9

8b. Stipules entire distally, bluntly obtuse to rounded, to 1 cm long, thick and opaque 11

9a. Stipules to 20 mm broad, drying yellowish and translucent, fimbriate to bluntly lobed [in-
florescences neither capitate nor hirsute; fruit becoming white] P. cartagoensis

9b. Stipules to 7 mm broad, drying dark and opaque, with stiff narrow or filiform teeth; fruit

becoming red, purple, or blue 10

lOa. Leaves pilose with hairs to 2 mm long; inflorescences to 15 cm long, much-branched and

lacking an involucre; fruit becoming purple or blue P. pilosa

lOb. Leaves glabrous or with short (0.5 mm) hairs; inflorescences to 5 cm long, capitate and

involucrate; fruit red P- ipecacuanha

1 la. Leaves with more than 18 pairs of closely parallel secondary veins, veins becoming prominently
raised and the leaf corrugated in age, bluntly obtuse to rounded distally; plants rarely exceeding

40 cm in height [fruits red] - P- polyphlebia

1 Ib. Leaves with fewer than 18 pairs of secondary veins, not becoming corrugated, usually acuminate

at apex; plants usually more than 50 cm tall

1 2a. Fruits becoming orange or red

12b. Fruits becoming white or yellowish green

13a. Leaf blades drying dark above and much paler beneath, young leaves glabrous; pyrenes with

a longitudinal costa on the back; commonly collected . P. uliginosa

13b. Leaf blades drying greenish to dark brown above and only slightly paler beneath; young
leaves densely hirsute beneath with hairs to 1 mm long; pyrenes lacking dorsal costae; rare

P. siggersiana

14a. Young leaves villose beneath with hairs to 1 mm long, a definite arcuate submarginal vein present
2-3 mm from the leaf edge, with 16-22 pairs of major secondary veins; fruit becoming greenish

yellow P capacifolia

14b. Young leaves minutely puberulent or glabrous, a definite submarginal vein absent, with 10-16
pairs of secondary veins (and often loop-connected in the distal part of the leaf); fruit becoming
white and spongy. (Note: This is a variable group of plants that may intergrade.)

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 223

1 5a. Inflorescences with thin widely spreading distal branches, bracts subtending the major branches
less than 4 mm long, flowers borne separately along the distal branches P. macrophylla

15b. Inflorescences with thick (1 mm) distal branches or the inflorescences compact and short (5 cm),
bracts subtending the basal branches or basal flowers more than 4 mm long 16

1 6a. Peduncles 0-3 cm long, flowers loosely clustered to capitate; leaves usually elliptic (rarely consis-
tently narrow); widespread in Costa Rica P. aggregata

1 6b. Peduncles to 1 5 cm long, flowers usually capitate on the peduncle or on 3 short branches on the

peduncle; leaves usually elliptic-lanceolate; eastern part of the Cordillera de Talamanca

P. aggregata (sensu stricto)

Key to Group 3: Species of Subgenus Heteropsychotria and Similar Species in Costa Rica
(Including Species Formerly in Cephaelis)

la. Flowers borne in dense capitate or subcapitate inflorescences with closely clustered flowers, bracts
usually conspicuous and enclosing the flowers; inflorescences terminal and solitary or 3 closely

grouped together (species formerly placed in Cephaelis and others) 2

Ib. Flowers in open or congested inflorescences but not capitate or subcapitate (flowers sometimes in
small distal capitula or glomerules on secondary branches within branched inflorescences); bracts

small to conspicuous or absent, bracts rarely enclosing and obscuring the flowers 16

2a. Inflorescences subtended by 2(-3) large bracts forming a basal cupulate involucre, capitulum

solitary, 3-10 cm broad 3

2b. Inflorescences usually with more than 4 basal bracts, lacking a single cupulate involucre at

the apex of the peduncle; inflorescence of 1 or 3 capitula, 1-5 cm broad 6

3a. Bracts reddish (rarely yellow), inflorescences erect; very common shrubs in lowland

evergreen formations 4

3b. Bracts purple to lilac, inflorescences erect or pendant; rarely collected species 5

4a. Stems and leaves glabrous; stipules with 2 rounded lobes on each side . . P. elata
4b. Stems and leaves densely tomentulose; stipules with 2 sharp teeth on each side

P. poeppegiana

5a. Montane (800-1200 m) plants; bracts purple, inflorescences pendant; leaf blades with

ca. 1 4 pairs of 2 veins P. correae

5b. Lowland (0-500 m) plants; bracts lilac, inflorescences erect; leaf blades with ca. 7 pairs

of 2 veins P. borucana

6a. Stipules with more than 2 slender teeth on each side (more than 6 teeth per node); unbranched
subshrubs less than 1 m tall; fruit red [medicinal plants rarely collected in Costa Rica] . . .

P. ipecacuanha

6b. Stipules with 2 slender teeth on each side, or with 2 rounded lobes or unlobed (with 4 or
fewer stipule lobes per node); plants mostly shrubby and more than 1 m tall (except P.

guapilensis); fruit blue or purple 7

7a. Corollas more than 4 cm long; fruit becoming more than 1 2 mm long; bracts usually bluntly

obtuse and green P. chiapensis

7b. Corollas less than 3 cm long; fruit less than 1 2 mm long; bracts acute to rounded and green

to purple 8

8a. Basal bracts of the inflorescences rounded to bluntly obtuse at the apex 9

8b. Basal bracts acute to acuminate 14

9a. Bracts white to pale green or bluish, glabrous, broadly rounded distally forming a tight

cupulate involucre beneath the congested capitulum [0-600 m elevation]

P. glomerulata

9b. Bracts green to deep purple, glabrous to puberulent, bluntly obtuse distally (rounded

in P. platypoda), not forming a definite cup at the base of the inflorescence 10

1 Oa. Bracts deep purple or reddish purple, inflorescences often densely compacted and spher-
ical; plants usually less than 1 m tall P. guapilensis

1 Ob. Bracts greenish to purple, inflorescences never spherical, often loosely compacted; plants
usually more than 1 m tall 11

224 FIELDIANA: BOTANY

1 la. Inflorescences usually of 3 pedunculate capitula; bracts white to pink (purple in fruit);
stipular tube 2-5 mm long; 2 veins 8-12/side [1000-2500 m elevation] ...P. dichroa

lib. Inflorescences usually of a single pedunculate capitulum; bracts whitish to green or
purple; stipular tube ca. 1 mm long; 2 veins 1 1-22/side .12

12a. Bracts broadly ovate and rounded at the base, purple; stipule lobes short and bluntly
rounded; 1200-2300 m elevation p, molinianum

1 2b. Bracts usually narrowed at the base, whitish to green or purple; stipules lobes short to
long, acute; 0-800 m elevation 13

13a. Bracts elliptic-obovate to oblanceolate; flowers usually puberulent, corolla tube 9-13
mm long; commonly collected p. suerrensis

1 3b. Bracts broadly rounded; flowers glabrous, corolla tube 3-6 mm long; rare in Costa Rica

P. platypodd

14a. (from 8b) Bracts orange or reddish orange, ovate and broadly overlapping; plants confined

to the southern Pacific slope, 600-1300 m elevation P. aura nti bract ea

1 4b. Bracts green or greenish with white, blue, or purple, linear lanceolate to ovate but not broadly

overlapping; plants wide ranging 15

1 5a. Bracts narrowly ovate, often marked with blue; inflorescences ca. 3 cm long and 2 cm broad

with erect bracts; stipule lobes thin and translucent, to 6 mm long; Caribbean slope, 900-

1 600 m elevation P. hazenii

15b. Bracts often linear-lanceolate, usually marked with white or purple; inflorescences short (1

cm) with broadly spreading bracts; stipule lobes stiff and opaque, to 4 mm long; widely

ranging in moist evergreen formations, 0-800 m elevation P. hoffmannseggiana

1 6a. (from 1 b) Young stems densely and conspicuously puberulent with hairs 0.3-2 mm long; peduncles

and branches of the inflorescence usually densely puberulent 17

16b. Young stems glabrous or sparsely puberulent with inconspicuous hairs less than 0.3 mm long;

peduncles and branches of the inflorescences glabrous or puberulent 25

17a. Hairs usually becoming more than 0.8 mm long, leaf blades drying dark above and 12-30

cm long, often with more than 14 pairs of major 2 veins; flowers in dense distal clusters,

subtended by conspicuous bracts 18

17b. Hairs rarely exceeding 0.8 mm in length; leaf blades usually drying greenish (rarely dark

brownish above), usually less than 20 cm long, with less than 14 pairs of major 2 veins;

flowers in open or small distal clusters, bracts various 19

1 8a. Leaf blades with 9-18 pairs of major 2 veins; flowers subtended by densely puberulent
lanceolate bracts ca. 2 mm broad; wide-ranging P. pilosa

18b. Leaf blades with (14-) 17-23 pairs of major 2 veins; flowers subtended by sparsely

puberulent ovate bracts 2-3 mm broad; southwestern Costa Rica .... P. mortoniana

19a. Ovary with 5 locules, fruits often with 5 seeds; stipules with 2 long (6-14 mm) stiff persisting

awns on each side; leaf blades with conspicuous subparallel 3 veins [9-21 (-26) cm long]

P. racemosa

19b. Ovary with 2 locules, fruits never with more than 2 seeds; stipules rarely with 2 long stiff

awns (P. umbelliformis); leaf blades rarely with conspicuous 3 veins 20

20a. Inflorescence umbelliform, flowers in 3 small glomerules on equal primary branches at the

apex of a long peduncle; leaves glabrous above [rare] P. umbelliformis

20b. Inflorescences not as above; leaves puberulent or glabrescent above .21

2 la. Leaf blades cuneate at the base and long-decurrent on the petiole, thin in texture; flowering

portion of the inflorescences often broader than long; deciduous and evergreen formations

of the Pacific slope P pubescens

21b. Leaf blades not cuneate and long-decurrent at the base (sometimes decurrent in P. steyer-

markii), mostly stiff-chartaceous when dried; flowering portion of the inflorescence rarely

broader than long; plants of evergreen formations

22a. Pubescence of young stems usually in narrow longitudinal lines; leaf blades 3-12 cm long

and 1-3 cm broad; stipules with narrow awns 2-5 mm long [plants of lower montane (800-

1 800 m) cloud forests; inflorescences few-branched and racemose] P. steyermarkii

22b. Pubescence not in narrow longitudinal lines; leaf blades 6-20 cm long, 2-5 cm broad; lines;

plants rarely collected above 800 m elevation

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 225

23a. Plants of the Osa Peninsula; inflorescences with hairs to 1.5 mm long, branches and flower
clusters distant and subtended by linear bracts to 8 mm long [leaf blades 5-12 cm long and
with 10-12 pairs of major 2 veins] P. acicularis

23b. Plants not known from the Osa Peninsula or from the Pacific slope below 400 m elevation;
inflorescences with hairs usually less than 1 mm long, flower clusters congested or distant,
subtended by linear bracts usually less than 4 mm long 24

24a. Inflorescences with crowded flowers and branches, usually less than 4 cm broad, narrowly
pyramidal, erect, calyx lobes 0.3-1.5 mm long; leaf blades 7-20 cm long, with 8-14 pairs of
major 2 veins; awns of the stipules to 3 mm long P. hebeclada

24b. Inflorescences usually with open branching and separate flower clusters, usually more than
3 cm broad, broadly pyramidal, often pendant; calyx lobes 0.2-0.5 mm long; leaf blades 6-
1 5 cm long and with 8-1 1 pairs of major secondary veins; awns of the stipules to 7 mm long

P. pittieri

25a. (from 16b) Inflorescences large and many-branched, usually becoming more than 12 cm long and

1 cm broad, broadly paniculate; leaf blades usually large (often to more than 1 8 cm long); fruit

rarely more than 4 mm diam. when dried (to 5 mm in P. solitudinum) 26

25b. Inflorescences smaller, rarely more than 12 cm long (and if so narrowly racemiform) and usually

less than 10 cm broad, paniculate to subcapitate or racemiform; leaf blades often less than 15 cm

long; fruit 3-12 mm diam. when dried 31

26a. Stipule lobes to 15 mm long and 5 mm broad at the base, conspicuous; flowers distant in
small (1-2) alternate sessile groups along the slender distal branches of the inflorescence;
fruits grayish blue to white P. microbotrys

26b. Stipule lobes rarely more than 5 mm long, usually less than 2 mm broad at the base; flowers
distant in small sessile groups only in P. solitudinum; fruit blue to black 27

27a. Stipules at first acute distally but splitting and developing 2 acute teeth or awns separated
by a broad U-shaped sinus; leaf blades often drying dark brown above; rarely collected below
400 m elevation P. berteriana

27b. Stipules at first rounded or obtuse at the apex, splitting into 2 broad lobes separated by a
narrow V-shaped sinus; leaf blades drying dark greenish brown to yellowish green or grayish
green; plants growing from to 1 700 m elevation 28

2 8a. Flowers in small (1-2) separate groups along the (usually dichotomous) slender distal branches
of the inflorescence [corolla tube 4-5 mm long, narrowed at the base and urceolate distally];
southwest Pacific slope of Costa Rica P. solitudinum

28b. Flowers in distal small cymes on multiple-branched inflorescences with opposite and cymose

branching; corolla various; plants not collected from the southwest Pacific area of Costa Rica

29a. Plants only known from the upper Rio Grande de Orosi and Tapanti above 1 300 m elevation;
floral bracts ca. 3 mm long, often persisting with the flowers and obtuse distally; corolla tube
ca. 6 mm long and tubular P. tapantiensis

29b. Plants not collected above 1200 m elevation; floral bracts to 3 mm long, early deciduous;
corolla tube 2-7 mm long 30

30a. Corolla tube 2-3 mm long, funnelform or tubular; floral bracts usually rounded at the apex;
leaf blades with 8-14 pairs of major 2 veins; fruit blue P. luxurians

30b. Corolla tube 6-7 mm long, narrowed at the base and expanded distally; floral bracts usually
acute at the apex; leaf blades with ca. 8-10 pairs of major 2 veins; fruit becoming blue or

black P. angustiflora

3 la. (from 25b) Inflorescences 12-20 cm long and only 2-5 cm broad, racemiform or thyrse-like,

peduncles 6-12 cm long; leaf blades with a distinct marginal vein within ca. 0.3 mm of the leaf

edge and connecting the distal ends of the major secondary veins; stipules narrowly 2-lobed . . .

P. cincta

31b. Inflorescences rarely over 14 cm long, usually more than 4 cm broad when over 10 cm long, not

racemiform or narrowly thyrse-like; leaf blades lacking a distinct marginal vein along the leaf edge;

stipules various 32

32a. Inflorescences with conspicuous bracts and bracteoles to 14 mm long and 3 mm broad, the bracts

persisting with the flowers . 33

FIELDIANA: BOTANY

32b. Inflorescences without conspicuous bracts, bracts rarely more than 3 mm long or more than 1 mm

broad, bracts often deciduous before the flowers mature 37

33a. Inflorescence small and compact, less than 3 cm long but expanding in fruit; stipules with

narrow awns 2-6 mm long [0-900 m elevation] p. brachybotrya

33b. Inflorescences larger and more open, to 10 cm long; stipule lobes less than 2 mm long or

triangular (not narrowly awned) 34

34a. Stipules very shortly (1-3 mm) bilobed with a U-shaped sinus between them [lateral branches

of the inflorescences not subtended by bracts; stipules 2-5 mm long] P. officinalis

34b. Stipules bilobed distally with a short or long V-shaped sinus between the lobes 35

35a. Flowers subtended by ovate bracteoles 2-3 mm long, lateral branches of the inflorescences

subtended by adnate bracts to 8 mm long; stipules 3-7 mm long P. brachiata

35b. Flowers subtended by lanceolate to oblanceolate bracts 4-14 mm long, lateral branches of

the inflorescence usually without subtending bracts; stipules 8-20 mm long 36

36a. Flowers subtended by lanceolate bracts 4-9 mm long; inflorescences with 4 or more nodes

with lateral branches; corolla tube 3-7 mm long P. capitata

36b. Flowers subtended by oblanceolate bracts 5-14 mm long; inflorescences with 1-3 nodes with

lateral branches; corolla tube 7-10 mm long P. calochlamys

37a. (from 32b) Stipules with 1 narrow lobe (usually 2/node), the awn-like lobe acute (rarely bifid)

distally and with an inner (adaxial) tooth; 500-1600 m elevation P. valeriana

37b. Stipules with 2 lobes on each side (4/node), the lobes separate laterally and without a tooth-like

adaxial (inner) appendage, or the lobes/teeth not developed; 0-2000 m elevation 38

38a. Inflorescences less than 1.5 cm long, less than 12 mm wide; fruit becoming red at maturity ....

P. haematocarpa

38b. Inflorescences becoming more than 2 cm long and more than 2 cm broad; fruit blue to purple or

black at maturity (orange during development in P. racemosd) 39

39a. Leaf blades linear-lanceolate to narrowly elliptic-oblong, largest blades less than 3 cm broad; 1000-

2000 m elevation P. goldmanii

39b. Leaf blades rarely linear-lanceolate, larger leaves over 3 cm broad; 0-1500 m elevation 40

40a. Inflorescence branches 0.7-1.5 mm thick when dried, flowering part of the inflorescence (beyond

the peduncle) broader than long 41

40b. Inflorescence branches slender (ca. 0.4 mm thick) when dried, flowering portion of the inflorescences

usually longer than broad [fruit less than 8 mm long and 7 mm diam. when dried] 43

4 la. Corolla tubes less than 3 mm long; inflorescences consistently with 4 lateral branches at the

first node; rare in Costa Rica at 1 500 m elevation P. allenii

41b. Corolla tubes 10-15 mm long; inflorescences with 2 or 4 opposite lateral branches at the first

node; from below 1 200 m elevation in Costa Rica 42

42a. Fruit 8-12 mm long and 7-10 mm diam., without transverse projections; leaf blades stiffly

chartaceous; calyx lobes ca. 0.2 mm long P. eurycarpa

42b. Fruit 4-5 mm long and ca. 6 mm diam., with transverse projections when dried; leaf blades

thinly chartaceous; calyx lobes 0.2-1 mm long P. domingensis

43a. Leaf blades drying membranaceous or thin-chartaceous, usually cuneate at the base and decurrent

on the petiole; plants often found in deciduous forest formations .44

43b. Leaf blades usually drying chartaceous, rarely cuneate and conspicuously decurrent on the petiole;

never collected in deciduous areas

44a. Leaf blades with 3-7 pairs of major 2 veins, often obovate; corolla tube 6-10 mm long; fruit

7-10 mm long P microdon

44b. Leaf blades with 9-15 pairs of major 2 veins, elliptic to ovate; corolla tube 2.5-4 mm long;

fruit 5-6 mm long P pubescens

45a. Fruits often with 5 seeds, ovary with 5 locules; stipules with stiff sharp awns 6-14 mm long and

persisting; leaf blades with 3 veins subparallel [with 7-12 pairs of major 2 veins] P. racemosa

45b. Fruit never with more than 2 seeds, ovary with 2 locules; stipules with awns or lobes 2-8 mm

long; 3 8 veins not subparallel 46

46a. Stipules with lobes or teeth absent or rarely 1 mm long, corolla tube 5-8 mm long (rare in Costa

Rica) P. phanerandra

46b. Stipules with lobes or teeth 2-8 mm long; corolla tubes 0.7-5 mm long 47

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 227

47a. Stipules with narrow lobes 2-3 mm long; fruits ca. 4 mm long and 6 mm with smooth rounded
surfaces (dried); leaf blades with long (1-3 cm) drip tips; inflorescences often small (to 6 cm long)
P. acuminata

47b. Stipules with narrow awns 3-8 mm long; fruits ca. 3 mm long and 4 mm diam. with longitudinal
ribs and transverse depressions; leaf blades with narrowed tips 1-2 cm long; inflorescences to 12
cm long P. deflexa

Key to Group 4: Species of Subgenus Psychotria and Similar Species in Costa Rica

la. Leaf blades 3-9 cm long, very rarely more than 7 cm long; small shrubs of evergreen forests ... 2
Ib. Leaf blades 5-30 cm long, the largest leaf blades usually more than 9 cm long; plants of evergreen

and deciduous habitats 6

2a. Young stems densely puberulent with short (0. 1-0.2 mm) reddish brown hairs; stipules acute
at the apex and caducous; inflorescences less than 1 6 mm long [ 1 200-2200 m elevation] . .

P. parvifolia

2b. Young stems glabrous (except for a ring of reddish hairs above the stipular scar); stipules
bilobed or bifurcate distally; inflorescences usually more than 20 mm long (except in P.

chagrensis) 3

3a. Inflorescences capitate and sessile, less than 1 7 mm long and subtended by large bracts to 1
cm long, corolla tube 4-8 mm long; stipules bifurcate distally with a V-shaped sinus [wet

lowlands, 0-300 m elevation] P. chagrensis

3b. Inflorescences paniculate and pedunculate usually more than 20 mm long and subtended by
inconspicuous bracts, corolla tubes 2-4 mm long; stipules with 2 distal lobes separated by a

broad U-shaped sinus 4

4a. Stipule lobes glabrous or minutely (< 0.05 mm) papillate puberulent; inflorescence branches

ca. 0.4 mm thick when dried [0-500 m elevation in Nicaragua and Panama]

P. fruticetorum

4b. Stipule lobes minutely (ca. 0.1 mm) ciliolate; inflorescence branches 0.2-0.3 mm thick when

dried; plants known to occur in Costa Rica 5

5a. Leaf blades rarely more than 5 cm long; petioles 2-10 mm long (the leaves not sessile);

inflorescences not exceeding 5 cm in length [0-1800 m elevation] P. graciliflora

5b. Leaf blades (at least the larger blades) usually exceeding 1 cm in length, petioles 0-50 mm

long (leaves sessile in some spp.); inflorescences usually exceeding 5 cm in length 21

6a. (from Ib) Young stems conspicuously puberulent with reddish hairs 0.2-2 mm long, internodes

puberulent above the ring of reddish hairs (colleters) at the stipule scar 7

6b. Young stems glabrous or inconspicuously puberulent with minute (0.03-0. 1 mm) hairs, but a ring

of reddish hairs (colleters) often present just above the stipule scars 14

7a. Inflorescences small (1-5 cm), flowers usually congested distally; peduncles to 2 cm long . .

7b. Inflorescences larger (4-15 cm long), flowers separate or congested, peduncles 1.5-15 cm long

8a. Leaf blades with an arcuate submarginal vein; stipules with a tube 4-14 mm long and

narrow awns; fruits 5-6 mm long [0-200(-800) m elevation] P. psychotriifolia

8b. Leaf blades usually lacking an arcuate submarginal vein but the distal veins often loop-
connected; stipules with a tube 0-5 mm long, usually lacking narrow awns; fruits 6-14

mm long 9

9a. Leaf blades usually elliptic-obovate or elliptic; stipules 3-1 1 mm long; fruits 6-8 mm

long; widespread at 20-800(-1200) m elevation P. nervosa

9b. Leaf blades usually oblanceolate to narrowly elliptic-oblong; stipules 8-14 mm long;

fruits unknown; Chiriqui Highlands ca. 1 200 m elevation P. boquetensis

lOa. Leaf blades with 14-33 pairs of major 2 veins, leaf blades 12-30 cm long; peduncles 4-14

cm long 11

lOb. Leaf blades with 8-13 pairs of major 2 veins, leaf blades 6-30 cm long; peduncles 1.5-8 cm
long 12

228 FIELDIANA: BOTANY

1 la. Leaf blades with 14-22 pairs of major 2 veins; lacking a definite submarginal vein, 3

veins subparallel; floral bracts ca. 3 mm long, narrowly triangular P. micrantha

1 Ib. Leaf blades with 20-34 pairs of major 2 veins, with definite submarginal veins and

subparallel 3 veins; floral bracts ca. 7 mm long and ovate P. sixaolensis

12a. Leaf blades rounded at the base and subcordate at the petiole, to 30 cm long, petioles 6-12
cm long; stipules ca. 2 cm long; rarely collected plants of the Caribbean escarpment [200-

500 m elevation] p. insignis

1 2b. Leaf blades not rounded and subcordate at the base (or if so not regularly exceeding 1 8 cm

in length), petioles usually less than 6 cm long; widespread 13

1 3a. Corolla tubes 2-2.5 mm long; fruit 4-6 mm long; peduncles 1.5-5 cm long; leaf blades usually
drying reddish brown or brown, major veins not impressed on the upper surfaces; (200-)400r

1600 m elevation P. jimenezii

13b. Corolla tubes 1.5-2 mm long; fruit 5-7 mm long; peduncles 3-8 cm long; leaf blades usually
drying grayish, major veins impressed on the upper leaf surfaces; 20-400 m elevation ....

P. neillii

1 4a. (from 6b) Larger leaf blades usually more than 1 cm broad and usually more than 20 cm long

14b. largest leaf blades less than 10 cm broad, rarely more than 20 cm long 21

15a. Inflorescences short (3-1 1 cm long) and compact (2-5 cm broad), with lateral branches to 1

cm long and often difficult to see; plants 0.5-2 m tall 16

1 5b. Inflorescences to 30 cm long and 4-15 cm broad, with lateral branches more than 1 cm long

and clearly visible; plants 1-10 m tall 18

16a. Leaf blades with 16-19 pairs of major 2 veins, usually more than 25 cm long, rounded

to obtuse at the apex; bracts ca. 5 mm long P. chitarriana

16b. Leaf blades with 8-14 pairs of major 2 veins, usually less than 25 cm long, short-
acuminate to acute at the apex; bracts 0.2-2 mm long 17

17a. Stipules with a basal tubular sheath and 2 narrow distal lobes; leaf blades with 10-14
major 2 veins on each side, usually obovate; inflorescences to 7 cm long, compact

panicles of cymes : P. alfaroana

1 7b. Stipules not tubular at base, broadly ovate and bluntly obtuse at the apex; leaf blades
with 8-1 1 pairs of major 2 veins, usually broadly elliptic; inflorescences to 3 cm long,

with short lateral branches and appearing subumbellate P. lamarinensis

18a. Stipules united over the shoot apex (as in Ficus) and tearing off as the leaves begin to grow,

narrowly conical in early stages; fruit 5-9 mm long (var. compressicaulis of)

P. panamensis

18b. Stipules united only at their base, open distally and often with the base persisting, ovate-
triangular in form; fruit 4-6 mm long

19a. Leaf blades subsessile and subcordate at base, usually 3-4 times longer than wide, 2 veins

arising from the mid vein at angles of 80-100 P. rosulatifolia

19b. Leaf blades with prominent petioles, never rounded at the base, blades 2-3 times longer than

wide, 2 veins usually arising at angles of 60-70 .20

20a. Secondary veins not usually loop-connected; stipules 8-30 mm long and acute to acuminate

at the apex; inflorescences to 30 cm long .P. grandis

20b. Secondary veins loop-connected distally to form an arcuate submarginal vein; stipules 5-12

mm long and rounded to acute; inflorescences to 10 cm long . P. limonensis

2 la. (from 5b and 14b) Flowers in dense sessile verticils on a single elongated (4-9 cm) rachis or in

dense clusters in a compact often globose arrangement, inflorescences solitary and apical, spiciform

or capitate/subcapitate, primary branches short (< 1.5 cm) or absent

21b. Flowers usually in open cymes, or on open spreading branches of the inflorescences when closely
clustered, never spiciform or globose-capitate, primary branches of inflorescences usually more

than 1 cm long

22a. Flowers in dense sessile verticillate clusters along the slender elongated (3-9 cm) rachis,
inflorescences spiciform with a single pair of proximal opposite branches or none; leaf blades
usually with domatia in the distal vein axils P- viridis

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 229

22b. Flowers in dense subcapitate globose of ellipsoid inflorescences with a rachis less than 4 cm

long; leaf blades lacking large domatia P. alfaroana

23a. Stipules united over the shoot apex and forming a cap with a single acute apex as in Ficus (usually
not split or open distally during development), narrowly conical to cylindrical and elongate, usually
becoming more than 5 mm long before splitting, caducous as the leaves begin to expand (note that
new lateral branches beneath the inflorescences often are enclosed within F/cws-like stipules after
other stipules have fallen); inflorescences with opposite branches; leaves lacking a submarginal

vein 24

23b. Stipules united over the shoot apex only in very early stages (if present the united cap usually
splitting before reaching 5 mm in length and open distally in later development), caducous or
tardily deciduous, larger F/cs-like stipules not present; inflorescences with opposite or whorled

branches; leaves with or without a submarginal vein 27

24a. Leaf blades usually with pit domatia or tufted hairs in the vein axils beneath, drying grayish

or pinkish gray above, with 10-13 pairs of major 2 veins; ca. 600 m elevation in the Cordillera

de Tilaran P. mexiae

24b. Leaf blades lacking pit domatia or tufted hairs in the vein axils, drying grayish to dark reddish

brown, with 5-16 pairs of major 2 veins; 400-2100 m elevation 25

25a. Stipules 2-6 mm long, central 2 veins arising at angles of 60-90, leaf blades 4-12 x 1.5-

5 cm [drying dark grayish above, with 5-7 pairs of major 2 veins; fruits 4-7 mm long] . .

P. orosiana

25b. Stipules usually becoming more than 6 mm long; central 2 veins arising at angles of 50-70,

leaf blades 6-22 x 2-10 cm 26

26a. Leaf blades usually drying grayish above, with 6-8 pairs of major 2, usually narrowly elliptic

and with long petioles; stipules becoming 5-45 mm long; dried fruits 7-10 mm long

P. sarapiquensis

26b. Leaf blades usually drying reddish brown to dark brown (less often dark grayish), with 6-16

pairs of major 2 veins, very variable in shape and texture; stipules becoming 8-80 mm long;

fruits 5-9 mm long P. panamensis

27a. (from 23b) Inflorescences only 2-4 cm broad, with short (< 2 cm) lateral branches (sometimes
becoming larger as the fruit develop) [flowers often closely congested distally]; leaf blades with 2
veins often weakly loop-connected near the margin; plants usually found in deciduous and partly

deciduous formations 28

27b. Inflorescences usually 4-8 cm broad, the lateral branches usually more than 2 cm long; leaf blades

with or without loop-connected 2 veins; plants not found in deciduous formations 31

28a. Corolla tube 4-5 mm long; fruit 6-9 mm long; inflorescences 1-3 cm long, with opposite

branches or the flowers subsessile at the end of the peduncle [broad ovate stipules with an

obtuse apex often subtending the young inflorescence] P. quinqueradiata

28b. Corolla tube 1-3.5 mm long; fruit 4-8 mm long; inflorescences 1 .5-10 cm long, with opposite

or whorled branches 29

29a. Inflorescences with only 2 branches at each node, often umbelliform; corolla tube 1.5-2 mm

long; stipules 6-14 mm long, acute and becoming narrowly 2-lobed at the apex; leaf blades

narrowly elliptic, to 20 cm long P. tenuifolia

29b. Inflorescences usually with 4 branches at the first node (2 longer and 2 shorter), rarely

umbelliform; corolla tube 2.2-3.5 mm long; stipules 2-8 mm long and bluntly obtuse; leaf

blades usually obovate to elliptic, to 1 5 cm long 30

30a. Pit domatia rarely present in distal vein axils beneath; leaves usually drying gray to pinkish

gray; calyx not usually persisting on the fruit; deciduous formations (in Costa Rica)

P. carthagenensis

30b. Domatia or tufts of hairs often present along the midvein beneath; leaf blades usually drying

gray to greenish gray [calyx often persisting on the fruit with lobes 0.5-3 mm long]; in both

evergreen and deciduous areas P. horizontalis

3 la. (from 27b) Inflorescences usually with 4 lateral branches at the first node (2 smaller descending

and 2 larger ascending); leaf blades usually narrowly elliptic to oblanceolate and with an arcuate

submarginal vein formed by the loop-connected secondaries (compare dichotomy 28 also) . . 32

31b. Inflorescences with 2 opposite lateral branches at the first node; leaf blades various 33

230 FIELDIANA: BOTANY

32a. Corolla tube 2-2.5 mm long; leaves usually drying grayish; wet lowlands (0-800 m elevation)

P. clivorum

32b. Corolla tube 3-5 mm long; leaves usually drying reddish or grayish brown; montane (2000-

2600 m elevation) forests p. sylvivaga

33a. Leaf blades usually with large (1-3 mm) domatia in the vein axils beneath, often opening by
ellipsoid slits [blades 1 1-24 cm long, usually elliptic-oblong and drying dark reddish brown;
inflorescence axes winged; peduncles ca. 2 mm thick; fruit 7-9 mm long; rarely collected in Costa

Rica] P. remota

33b. Leaf blades without large webbed domatia in the vein axils, smaller domatia present or absent

34a. Calyx lobes well developed (0.5-1 mm long); peduncles 1.5-2.5 mm thick when dried, primary
branches of the flowering inflorescences 1-1.5 mm thick when dried; leaf blades lacking domatia'.

often oblanceolate, to 22 cm long; montane (1300-2600 m elevation) forest species 35

34b. Calyx lobes minute (0.3 mm) or absent; peduncles 0.5-1 .5 mm thick when dried, primary branches
of the flowering inflorescences usually less than 1 mm thick when dried; leaf blades often with
domatia on the lower surfaces, not usually oblanceolate or more than 1 5 cm long (except in P.

sarapiquensis); 0-2500 m elevation 37

35a. Primary branches of the inflorescence usually diverging at more than 100 from the more
distal rachis ( reflexed); leaf blades with 10-15 pairs of major 2 veins, with or without an
arcuate submarginal vein [flowers on open cymes on tertiary branches of a much-branched

inflorescence, peduncles 5-9 cm long; 1 700-2000 m elevation] P. stockwellii

35b. Primary branches of the inflorescence usually diverging at 90 or less from the rachis, per-
pendicular or ascending; leaf blades with 8-1 1 pairs of major 2 veins; usually with a definite

submarginal vein 36

36a. Flowers in dense glomerules at the ends of the few (ca. 5) 1 or (ca. 9) 1 and 2 branches;

peduncles 1.5-5.5 cm long; 900-1500 m elevation P. monteverdensis

36b. Flowers in open cymes or pedicellate on 2 or 3 branches of the inflorescences; peduncles

4-1 2 cm long; 1 300-2600 m elevation P. sylvivaga

37a. Stipules developing 2 small (1-3 mm) slender lobes at the apex (4 acute tips per node) and usually

united to form a short tube at the base; corolla tube usually more than 3 rtim long 38

37b. Stipules acute at the apex (without small lobes), a short tube not apparent at the base; corolla tubes

rarely more than 3 mm long (except in P. chiriquina)

38a. Leaves drying dark reddish brown, petioles 7-18 mm long; stipules 4-10 mm long; 1500-

2500 m elevation P. chiriquina

38b. Leaves drying dark to pale grayish, petioles 0-5 mm long; stipules 2-5 mm long; 400-1900

m elevation P- orosiana

39a. Fruits 7-10 mm long, obovid-oblong; stipules conical, united and early caducous; leaf blades

glabrous beneath and without domatia [600-1600 m elevation] P. sarapiquensis

39b. Fruits 4-8 mm long (unknown in P. laselvensis), subglobose; stipules free distally, not united into

a conical cap; leaf blades glabrous or minutely puberulent and often with domatia beneath 40

40a. Highland (1500-2500 m) plants of easternmost Costa Rica and Chiriqui, Panama; leaves usually

drying reddish brown; corolla tube 3-6 mm long P. chiriquina

40b. Lower (0-1000 m) elevation plants; leaves usually drying grayish; corolla tube 1-3 mm long 41
4 la. Leaf blades with domatia rarely present in distal vein axils, blades to 13(-16) cm long; stipules

triangular; La Selva area, 50-300 m elevation P laselvensis

41b. Leaf blades with domatia often present along the central and proximal part of the midvein, blades
to 17 cm long; stipules ovate-triangular; widespread, 0-1000 m elevation P. marginata

Psychotria acicularis C. M. Taylor, sp. nov. Figure TYPUS-/. Utley A K. Utley 1036 (holotypus CR. >-

typi F, MO!), from region to southwest of airstrip, 20-60

:>:> - m alt., 20 July 1974, Rincon de Osa, Puntarenas, Costa

Rica.

Species Psychotriae pittieri Standley affinis. sed ab ea

bracteismajoribus(3-10mmlongis)etangustoribusdif- Small shrubs, 0.5-2 m tall, leafy stems

f ert thick, densely hirsutulous with thin hairs 0.4-1.5 mm

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 231

long, older stems glabrescent; stipules 3-8 mm long, with
a short (1-3 mm) tube and 2 slender teeth 3-7 mm long
on each side, densely hirsutulous in early stages, per-
sisting or deciduous. Leaves with petioles 2-9(-13) mm
long, ca. 0.7 mm thick, densely hirsutulous, often some-
what unequal at the same node; leaf blades 5-12 cm
long, 2-4.5 cm broad, narrowly ovate-elliptic to elliptic
or elliptic-oblong, apex acuminate with a tip 4-12 mm
long, base usually obtuse, drying thinly chartaceous or
membranaceous, dark green above, with slender hairs
0.5-1.2 mm long on the upper surface, especially along
midvein, becoming glabrescent, more densely hirsutu-
lous beneath with slightly shorter more persistent hairs,
2 10-15/side. Inflorescences terminal and solitary, 5-
12 cm long, open or elongate panicles with usually 4
branches at apex of the peduncle and 2 additional pairs
of opposite or alternate branches, peduncles 3-8 cm long,
ca. 0.7 mm thick (dried), hirsutulous with hairs to 1.5
mm long, bracts 3-10 mm long and linear, flowers sub-
sessile in condensed distal cymes, bracteoles 2-4 m long,
linear. Flowers densely puberulent externally, hypanthi-
um ca. 1 mm long, sericeous with ascending thin whitish
hairs ca. 0.5 mm long, calyx lobes 0.5-1 mm long, lan-
ceolate; corolla white, narrowly salverform, tube ca. 3
mm long and 0.5 mm diam., corolla lobes ca. 1.2 mm
long; anthers ca. 1 mm long. Fruits 3-3.5 mm long, 2.5-
3 mm diam., broadly ellipsoid, bluish with thin ascend-
ing hairs ca. 0.5 mm long, persisting calyx lobes ca. 0.5
mm long.

Plants of the rain forest interior on the Osa Pen-
insula, at elevations of 20-80 m. Flowering in May
and July; fruiting in May-August. This species is
only known from near Rincon de Osa on the Osa
Peninsula in southwestern Costa Rica.

Psychotria acicularis is recognized by its hir-
sutulous indumentum on all parts, small stature,
small leaves, open inflorescences with conspicuous
linear bracts, sessile flowers borne in distal clus-
ters, small blue fruit, and limited geographic range.
This species, a member of subgenus Heteropsy-
chotria, resembles P. pittieri vegetatively but dif-
fers in its linear bracts and inflorescences with four
branches at the first (basal) node. Additional spec-
imens seen were Aguilar 6423 (CR), Burch 4413
(DUKE, MO), Burger & Stolze5449(cR, F, MO), Duke
16111 (MO), Kennedy 1 934 (MO), Raven 21532 (CR,
F), and Utley & Utley 1174 (CR, F).

Psychotria acuminate Benth., Bot. voy. Sulph. 107.
1 845. P. cuspidata sensu Standl. et auctores, non
Bredem. ex Roem. & Schult. fide Steyermark
1974. Figure 56.

Shrubs or subshrubs, l-2(-3) m tall, much-branched,
leafy stems 1 .5-5 mm thick, glabrous, quadrate to terete;
stipules with a short (0.5-2 mm) truncated tube with 2
narrow awns 1.5-3 mm long on each side, these decid-
uous. Leaves with petioles 5-15(-18) mm long, 0.7-1.4

mm thick, minutely (0.05 mm) papillate-puberulent; leaf
blades 7-15(-19) cm long, 3-8(-10) cm broad, elliptic
to oblong-elliptic or ovate-elliptic, apex short- to long-
acuminate or caudate-acuminate with tip 1-3 cm long,
base acute to obtuse, drying thin-chartaceous and green-
ish or brownish, glabrous above and below, or with tuft-
ed domatia in the vein axils beneath, 2 veins 7-12/side,
midveins sometimes reddish beneath. Inflorescences
solitary and terminal, 2-6 cm long (to 7 cm in fruit); 2-
4(-5) cm broad, paniculate, peduncles 1-4 cm long, 0.5-
1 mm thick, minutely papillate-puberulent (or appar-
ently glabrous), primary branches opposite or alternate,
bracts subtending the inflorescence branches absent or
small (1 mm) and adnate to branches, floral bracts ab-
sent, pedicels 0-2 mm long. Flowers distylous, minutely
(0.05 mm) papillate-puberulent externally, hypanthium
ca. 0.5 mm long and 0.5 mm diam., calyx 0.3-0.7 mm
long with weakly denned lobes; corolla white or pale
yellow, tube 2-4(1-5) mm long and 0.5-1 mm diam.,
lobes 5, 1.5-2.5 mm long, rounded; anthers 1-1.2 mm
long, linear. Fruits 4 mm long, 5-7 mm diam., 2-lobed
and rounded, smooth, becoming blue, bluish gray, or
purple; pyrenes 2.5 mm long and 3 mm broad, hemi-
spheric with a longitudinal sulcus on the inner face.

Plants of evergreen lowland rain forest forma-
tions on both the Caribbean and Pacific slopes,
from near sea level to 900(-1 100) m elevation in
Costa Rica. Flowering in January-August in
southern Central America (usually May-June at
La Selva). Fruiting is primarily in June-Decem-
ber. The species ranges from Mexico and the West
Indies to Venezuela and Peru.

Psychotria acuminata is characterized by its gla-
brous or minutely papillate-puberulent parts, lus-
trous acuminate leaves with pronounced narrow
drip tips, small flowers and inflorescences, 2-awned
stipules, and smooth rounded bilobed blue fruit.
In some specimens, the stipules appear to be un-
developed. The small stature, leaf size, and foliage
drying greenish are similar to many other species,
but the small yellowish papillate-puberulent inflo-
rescences and smooth bilobed fruit are distinctive.
Also, the veins on the undersides of the leaves are
sometimes pink. The presence of domatia-like tufts
of hairs in the vein axils is uncommon among our
species of Psychotria subgenus Heteropsychotria.
The breeding biology was studied by Bawa and
Beach (1983). Psychotria valeriana of higher ele-
vations is similar but has a single stipular awn
with an adaxial tooth on each side of the stem.
[The name P. cuspidata deserves more careful re-
view; it may be that Steyermark failed to recognize
a specific distinction between the Caribbean ma-
terial and his Venezuelan specimens. C.M.T.]

Psychotria aggregata Standl., Contr. U.S. Natl.
Herb. 18: 128. 19 16. P. tonduziiSiandl, J. Wash.

232

FIELDIANA: BOTANY

Acad. Sci. 15: 287. 1925. Montamans pana-
mensis Dwyer, Ann. Missouri Bot. Gard. 67:
286. 1980. Figure 12.

Herbacous subshrubs with succulent stems, 0.5-1.5(-2)
m tall, leafy stems 1 .6-6 mm thick (dried), quadrangular,
glabrous; stipules 2-9(-12) mm long, to 7 mm broad,
triangular with a short (2-3 mm) cupular base and bifid
conical appendages 4-7 mm long, base persisting. Leaves
with petioles 3-1 0(- 1 3) cm long, 1 -2 mm thick, glabrous;
leaf blades 12-30(-38) cm long, (3-)5-17 cm broad, el-
liptic-oblong to elliptic-obovate or elliptic-oblanceolate,
apex tapering gradually and acute or acuminate, tip 5-
1 5(-22) mm long, base obtuse to cuneate or acute and
decurrent on petiole, drying chartaceous and dark to pale
grayish green, glabrous above, glabrous or minutely pa-
pillate-puberulent beneath, 2 veins 10-19/side. Inflo-
rescences axillary, solitary, 4-1 5 cm long, 2-4 cm broad,
congested to capitate or with 1-2 pairs of lateral branch-
es, peduncles l-5(-8) cm long, 1-2 mm thick, glabrous
or minutely and sparsely puberulent, bracts at the lateral
branches 2-8 mm long, flowers subsessile in closely
crowded distal glomerules on the 1 or 2 branches, brac-
teoles 1-4 mm long. Flowers with hypanthium 1-2 mm
long, glabrous or sparsely and minutely puberulent, calyx
0.5-2 mm long, lobes 5, 0.5-4 mm long, obtuse to linear;
corolla white, funnelform glabrous or minutely puber-
ulent externally, tube 2-4 mm long, ca. 1 mm diam.,
corolla lobes 5, ca. 1 mm long, obtuse and galeate; an-
thers ca. 1.2 mm long. Fruits 5-7 mm long (dried), 4-6
mm diam. (to 10 mm in life), ovoid to subglobose, spongy,
white; pyrenes ca. 6 mm long, with margin and median
keel thickened dorsally.

Plants of evergreen montane rain forest and cloud
forest formations, from 10 to 2300 m elevation
(but most common in the 400-1 700-m range). Ap-
parently flowering and fruiting throughout the year.
The species ranges from northern Costa Rica into
western Panama.

Psychotria aggregata is recognized by its short
succulent unbranched main stems, large essen-
tially glabrous leaves, axillary inflorescences with
flowers in dense bracteolate clusters, and spongy
white fruits. This species is quite similar to P.
macrophylla but differs in the more condensed
inflorescences and the pyrenes. (This and related
species have recently been studied by Molly Ne-
pokroeff, wis, 1 992.) The type of P. aggregata (Pit-
tier 3264 us from Horqueta, Chiriqui, Panama) is
a very poor specimen, but it is an early name for
this variable taxon, commonly called P. tonduzii
by previous authors.

Psychotria alfaroana Standl., J. Wash. Acad. Sci.
18:273. 1928. Figure 66.

Herbaceous erect subshrubs, 0.2-0.6(-1) m tall, usu-
ally with a single erect unbranched stem, rhizomatous,

leafy stems 2-4 mm thick, glabrous, terete; stipules to
20 mm long, basal sheath 5-8 mm long and 4 mm broad,
with 2 narrow distal lobes 3-12 mm long, glabrous or
sparsely puberulent, caducous. Leaves with petioles (2-)6-
25(-30) mm long. 1.5-2 mm thick, sulcate above, gla-
brous; leaf blades 1 1-22(-30) cm long, 4-1 1 cm broad,
obovate to elliptic-obovate or elliptic-oblanceolate. apex
short-acuminate or subacutc. base gradually narrowed
and cuneate, decurrent on petiole, drying thin- to stiffly
chartaceous, grayish or grayish green, glabrous above.
with minute (0. 1-0.3 mm) thin hairs along major veins
beneath, 2 veins 10-14/side, weakly connected by an
arcuate submarginal vein. Inflorescences terminal, sol-
itary, 3-8 cm long, 2-4 cm broad, compact globose or
elongate dense panicles of cymes, peduncle 1-4 cm long,
1.2-2 mm thick, glabrous, bracts 0.5-1 mm long, ped-
icels 1-2 mm long. Flowers glabrous externally, hypan-
thium ca. 1.5 mm long, calyx tube ca. 1 mm long, cu-
pulate, calyx lobes 5, 0.5-2 mm long, narrowly oblong;
corolla white, tube 2.5-4 mm long, ca. 2 mm diam..
lobes 5, 1 .5-3 mm long, ca. 1 mm broad at base; stamens
5, anthers 0.8-1.3 mm long. Fruits 8-12 mm long, 4-6
mm diam., ellipsoid, red at maturity, becoming black
when dried, persisting calyx to 3 mm long; pyrenes ca.
7 mm long, usually with 5 dorsal ribs.

Plants of wet rain forest formations of the Ca-
ribbean slope and continental divide, 20-
900(-1100) m elevation. Flowering in January-
September; fruiting in January-February and June-
August. The species ranges from the Cordillera de
Guanacaste in Costa Rica to Bocas del Toro Prov-
ince in Panama.

Psychotria alfaroana is recognized by its very
short stature, usually obovate leaves, compact
subglobose inflorescences, and larger fruit. The
bright red fruit and leaves drying grayish are char-
acteristics of subg. Psychotria. Standley and Vale-
ric made a number of collections of this species
near Tilaran, including the holotype (41579 us).
A collection from along Rio Corobici (Opler 138
F) is provisionally placed here; this is the only
collection from the Pacific lowlands. Several col-
lections with exceptionally large (35 x 15 cm)
leaves attenuate almost to the base and then
abruptly subauriculate are tentatively placed here:
Gdmez-Laurito 11570, Hammelet al. 16895, and
Herrera & Chacdn 2355 (all at CR).

Psychotria allenii Standl., Ann. Missouri Bot.
Gard. 27: 342. 1940.

Small trees or shrubs, (2-)4-10 m tall, leafy stems 1-
4 mm thick, glabrous or minutely and sparsely puber-
ulent; stipules united around the stem for 0.5-3 mm,
with 2 acute lobes 1-5 mm long, separated by a V- or
U-shaped sinus, persisting. Leaves with petioles 6-18
mm long, 0.8-1.3 mm thick, glabrous; leaf blades 5-16
cm long, 3-7(-8) cm broad, ovate-elliptic or ovate-ob-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

233

long to narrowly elliptic-oblong, apex acuminate with
tip ca. 5 mm long, base obtuse to slightly rounded, drying
dark olive green or grayish green and chartaceous, gla-
brous above, glabrous or minutely puberulent beneath,
2 veins 5-7/side, distal veins strongly arcuate-ascend-
ing. Inflorescences 8-10 cm long, 5-7 cm broad, pyra-
midal paniculate with distal branches progressively
shorter, peduncles 3.5-6.5 cm long, 0.9-2.3 mm thick,
glabrous or puberulent, the first node with 4 lateral
branches 2-3.5 cm long subtended by linear bracts 1-3
mm long, flowers mostly sessile and crowded in distal
cymes. Flowers with hypanthium 0.4-0.8 mm long, gla-
brous or sparsely papillate-puberulent, calyx ca. 0.7 mm
long with glabrous lobes ca. 0.3 mm long; corolla white,
tube 0.7-2 mm long and 1 mm diam., lobes 5, ca. 1.7
mm long; stamens 5. Fruits 6-1 5 mm long and ca. 6 mm
diam. when dried, bright blue; pyrenes smooth, subglo-
bose.

Plants of moist evergreen lower montane for-
ests, 50-900(-1500) m elevation. Flowering and
fruiting in February-August in Panama. The spe-
cies is known only from Monteverde in Costa Rica
(Koptur 74 MO with flowers and early fruits in
October). The species ranges to central Panama.

Psychotria allenii is recognized by its distinctive
pyramidal inflorescences with relatively thick pe-
duncles, four stout lateral branches at the first few
nodes, and small sessile flowers. This species is
quite similar to P. officinalis, but that species has
floral bracts and larger flowers and lacks the robust
lateral branches of the inflorescence. The Costa
Rican collection has narrower elliptic-oblong leaves
than Panamanian material, and it was a 2-m shrub
rather than a tree.

Psychotria angustiflora K. Krause, Bot. Jahrb. 54,
Beibl. 119: 43. 1916. P. mima Standl., Publ.
Field Mus. Nat. Hist., Bot. Ser. 22: 386. 3 1 Oct.
1 940 (holotype: Skutch 4589 F from Costa Rica),
not P. mima Standl., loc. cit. 22: 204. 10 Sept.
1 940 (holotype: L. B. Smith 1878 F from Brazil).
Figure 57.

Shrubs or small trees to 5 m tall, leafy stems 1.5-5
mm thick, glabrous, terete; stipules 3-8 mm long, 2-6
mm broad, triangular to ovate and rounded at the nar-
rowed tip, with a short (1-3 mm) tube, obtuse or bilobed
with a small ( 1 mm) sinus, glabrous, persisting. Leaves
with petioles 15-35(-60) mm long, 1.2-3 mm thick, gla-
brous, with 2 adaxial ridges; leaf blades (9-) 1 5-24(-30)
cm long, (4-)7-14(-16) cm broad, oblong to ovate-ob-
long, broadly elliptic or broadly elliptic-obovate, apex
short-acuminate with a tip 5-10 mm long, base broadly
obtuse (rarely acute), drying thin-chartaceous and green-
ish gray or greenish brown above, glabrous above and
below, 2 veins 8-1 I/side, central 2 veins ca. 2 cm dis-
tant in larger leaves. Inflorescences terminal, solitary
(3-) 17-45 cm long, (8-) 15-25 cm broad, pyramidal pa-

niculate with distant opposite lateral branches, peduncles
6-15 cm long, 1.5-2 mm thick, glabrous or minutely
puberulent, bracts to 7 mm long and 1.5 mm broad,
flowers subsessile in distal cymes or along slender distal
branches, subtended by bracteoles ca. 1 mm long. Flow-
ers minutely papillate-puberulent externally, hypanthi-
um ca. 0.7 mm long, calyx ca. 0.4 mm long, calyx lobes
ca. 0.2 mm long, acute; corolla pale yellow or greenish
yellow (rarely white), tubular-obovoid or funnelform,
tube 5-7.5 mm long, 0.9-1 .3 mm diam. in the lower half
and 2-3 mm distally, lobes 5, 1-2.5 mm long, triangular
and acute, erect. Fruits 4-5 mm long and 3-5 mm diam.
(to 8 mm in life), subglobose with ca. 6 longitudinal ribs,
becoming blue or black; pyrenes 1.5-3.5 mm long.

Plants of evergreen rain forest formations, from
near sea level to 900 m elevation on the Caribbean
slope. Flowering in January-August; fruiting in
June and October-November. The species ranges
from near La Selva, Heredia, to Brazil.

Psychotria angustiflora is recognized by its large
open inflorescences with small distal bracteoles,
narrowly obovoid yellowish corolla, larger leaves,
and ovate stipules that are obtuse or minutely two-
lobed at apex. The stipules may become two-part-
ed as the expanding stem breaks them apart. The
flower buds are quite narrow before the corolla is
fully expanded. The isotype of P. angustiflora
(Tonduz 12996 us) fits in well with the material
placed here. This species should be compared to
P. berteriana (smaller flowers, different stipules,
narrower leaves) and P. luxurians (smaller flowers,
larger bracteoles).

Psychotria aubletiana Steyerm., Mem. New York
Bot. Card. 23: 694. 1972. Cephaelis axillaris
Sw., Prodr. 45. 1788, not P. axillaris Willd. P.
aubletiana var. centro-americana Steyerm., loc.
cit. 698. 1972.

Herbs or subshrubs, 0.5-1 .5(-4) m tall, erect and few-
branched, stems slightly succulent, leafy stems 1-5 mm
thick, glabrous or rarely densely puberulent with soft
yellowish hairs ca. 0.2 mm long, quadrangular, drying
dark; stipules 5-10 mm long, to 6 mm broad, basal sheath
3-5 mm long, ovate with a short (2-4 mm) sinus and 2
irregular rounded lobes per side. Leaves with petioles 6-
20(-40) mm long, 0.7-1.8 mm thick, glabrous or mi-
nutely puberulent; leaf blades 4-15(-18) cm long, 1-
4(-6.5) cm broad, elliptic-oblong, narrowly oblong-ob-
ovate to narrowly elliptic-obovate, oblong-lanceolate or
narrowly elliptic, apex acuminate (rarely acute) with tip
4-1 2 mm long, acute at the base (obtuse in larger leaves),
drying stiffly chartaceous, dark greenish above, much
paler beneath, glabrous above, glabrous beneath or with
short (0.1-0.2 mm) hairs on major veins, 2 veins 6-87
side (with lesser parallel intermediate 2 veins). Inflo-
rescences axillary and sessile, 1.3-2 cm diam., densely
compact, subglobose, with distal 2-lobed stipules resem-

234

FIELDIANA: BOTANY

bling bracts, bracts obovate, purple or green, 2-7 mm
long, flower 5-1 5/head, sessile. Flowers with 5 calyx lobes
1-2 mm long; corolla white, glabrous, tube ca. 10 mm
long, lobes 5, 3-3.5 mm long, triangular, stamens 5,
anthers 1-2 mm long. Fruits 5-7 mm diam., ellipsoid,
bright blue; pyrenes 3-5 mm long and 2-4 mm broad.

Understory plants of evergreen montane rain
forest formations, from (600-)1 200 to 2300(-2800)
m elevation in Costa Rica. Probably flowering and
fruiting throughout the year. The species ranges
from Guatemala through Central America to Co-
lombia, Venezuela, and the lesser Antilles.

Psychotria aubletiana is recognized by its small
stature, sessile rounded inflorescences often encir-
cling distal nodes and subtended by an involucre
of broad bracts, short white corolla tubes, and
bright blue fruit. This species is not easily confused
with other species of Psychotria.

Psychotria aurantibractea C. M. Taylor, nom. nov.
Cephaelis pittieri K. Krause, Bot. Jahrb. Syst.
54, Beibl. 1 19: 45. 1916, not Psychotria pittieri
Stand i. Figure 18.

Shrubs, 1-3 m tall, leafy stems 1.3-4.5 mm thick,
glabrous, often narrowed below the node when dried;
stipules united for only 0.5-2 mm around stem, with 2
narrowly triangular or linear teeth 2-8 mm long, sepa-
rated by 2-3 mm, persisting. Leaves with petioles 5-40
mm long, 0.8-1.5 mm thick, glabrous; leaf blades 8-
21 (-28) cm long, 2-6 cm broad, narrowly oblong to nar-
rowly elliptic-oblong or lanceolate, apex gradually ta-
pering and acute or acuminate, tip to 1 5 mm long, base
cuneate to acute, drying membranaceous or thin-char-
taceous and dark brown, glabrous above, glabrous or
sparsely and minutely (0. 1 mm) puberulent on the veins
beneath, 2 veins 10-16/side. Inflorescences terminal, 1
or 3, 2.5-6 cm long, 4-6 cm broad (to 8 cm when tri-
partite), usually with 3 capitate or cymose clusters of
flowers on short (1-4 mm) primary branches, peduncles
5-25 mm long, 1.3-2.2 mm thick, minutely puberulent,
bracts 8-15 mm long, 4-9 mm broad, ovate to lanceo-
late, acuminate, orange (rarely red), sparsely puberulent
and ciliolate along margins, flowers sessile. Flowers en-
closed by bracts and bracteoles, hypanthium ca. 1 .5 mm
long, densely pubescent with straight ascending hairs,
calyx lobes 5, ca. 1.3 mm long, dentate with narrrow
lobes; corolla yellow or yellow-rose, tube ca. 5 mm long,
lobes ca. 2 mm long. Fruits 5-7 mm long, 2.5-5 mm
diam., ovoid-ellipsoid, with prominent longitudinal cos-
tae and short thin hairs, color unknown, persisting calyx
1 .3-2 mm long, conical; pyrenes ridged.

Plants of continually wet or seasonally dry ev-
ergreen forest formations, from 200 to 1200 m
elevation. Flowering in May-July; fruiting in Au-
gust-September and January. This species is en-
demic to southern Costa Rica, from San Isidro del

General to San Vito and the Osa Peninsula, and
on the adjacent Caribbean slope (at 83W).

Psychotria aurantibractea is recognized by its
terminal inflorescences composed of large ovate,
orange or reddish bracts, narrow leaves, and re-
stricted geographical range. The colorful bracteate
inflorescences made this and other species for-
merly placed in Cephaelis so distinctive in Central
America.

Psychotria berteriana DC., Prodr. 4: 515. 1830.
Figure 58.

Shrubs or small trees, 1.5-6(-8) m tall, leafy stems
(1.5-)3-5 mm thick, glabrous or minutely puberulent
with hairs 0.1-0.2 mm long; stipules at first triangular
and acute with a small distal cleft, becoming 2-lobed or
2-awned (on each side), basal sheath 0.5-2 mm long with
acute lobes 1-3.5 mm long. Leaves with petioles 10-
30(-52) mm long, 0.6-1.8 mm thick, glabrous or mi-
nutely puberulent; leaf blades 8-20(-26) cm long, 3-
9(-ll) cm broad, narrowly ovate-elliptic to elliptic or
elliptic-oblong, apex acuminate with 5-15 mm long, base
acute and cuneate to obtuse (rarely rounded and sub-
truncate in large leaves), drying membranaceous to thin-
chartaceous, dark green or brown, glabrous or with min-
ute (0.1-0.2 mm) hairs along the veins above, glabrous
or with thin whitish hairs ca. 0.2 mm long beneath, 2
veins 7-15/side. Inflorescences terminal and solitary. 5-
16(-22) cm long, 5-15 cm broad, oblong but becoming
broadly pyramidal paniculate with opposite or alternate
branches, yellowish in life, peduncles 5-10 cm long. 1 .3-
2.2 mm thick, sparsely puberulent with thin hairs ca. 0.2
mm long, bracts 3-8 mm long, ca. 0.6 mm broad, flowers
sessile in distal cymes, bracteoles 1-2 mm long. Flowers
minutely puberulent externally (rarely glabrous), hypan-
thium ca. 0.5 mm long, calyx lobes ca. 0.2 mm long;
corolla tubular-salverform, yellow to white, tube 1.5-4
mm long, 1-2 mm diam., glabrous proximally, lobes 5.
1-1.5 mm long; anthers 1-1.4 mm long. Fruits 34 mm
long (5 mm including the persisting calyx), 3-4 mm diam..
subglobose, fleshy, lustrous black; pyrenes with 3-5 dor-
sal ridges.

Plants of evergreen rain forest formations, from
near sea level to 1 700 m elevation (to 1 200 m on
the Pacific slope of southern Costa Rica). Flow-
ering throughout the year; probably fruiting
throughout the year. The species ranges from Mex-
ico to South America.

Psychotria berteriana is characterized by its large,
many-branched, pyramidal inflorescences, thin
leaves often with long slender petioles and drying
dark, two-lobed stipules, small flowers, and small
rounded fruit. Compare P. luxurians and P. an-
gustiflora (with longer thinner corolla tubes). A
broader interpretation of P. berteriana might re-
quire the inclusion of P. luxurians. The greenish

BURGER & TAYLOR: FLORA COSTARICENSIS. RUB1ACEAE

235

yellow inflorescences resemble those found in Pal-
icourea tilarensis.

Psychotria boquetensis Dwyer, Ann. Missouri Bot.
Gard. 67: 349. 1980.

Shrubs, 0.5-2 m tall, leafy stems 1.2-3.5 mm thick,
tomentulous with reddish hairs 0.4-1 mm long; stipules
8-14 mm long, 2-6 mm broad, oblanceolate to ovate,
obtuse to acuminate at apex (not bilobed in the type as
described by Hamilton, 1989), caducous. Leaves with
petioles 1-6 mm long, 0.7-1.2 mm broad, with curved
thin reddish hairs ca. 0.3 mm long; leaf blades 4-16 cm
long, 1-4 cm broad, narrowly elliptic-oblong to oblan-
ceolate, apex acuminate with tip 4-15 mm long, base
gradually narrowed and acute or cuneate, decurrent on
petiole, drying thinly chartaceous and dark reddish brown,
puberulent along the primary and secondary veins above
and below, with hairs 0.2-0.7 mm long along the margin,
2 veins 10-13/side, with minute tufted domatia in the
axils beneath. Inflorescences terminal or pseudoaxillary,
several at a node, 2-4 cm long, small panicles, peduncles
ca. 20 mm long and 0.6 mm thick, bracts represented
by dense whitish hairs 0.5-1 mm long, flowers sessile or
subsessile in compact cymes. Flowers glabrous on the
distal surfaces, hypanthium ca. 1 mm long and puber-
ulent, calyx lobes 1-1.5 mm long, narrowly triangular,
corolla white, urceolate, tube 1.5-2.5 mm long and 1
mm diam., lobes 5, 1-1.5 mm long; stamens 5, anthers
0.5-0.7 mm long. Fruits unknown.

Plants of the Boquete region of the Chiriqui
Highlands at about 1 200 m elevation. Flowering
material was collected in May-June. The species
is only known from western Panama.

Psychotria boquetensis is recognized by the nar-
row leaves with conspicuous reddish hairs, mem-
branaceous broad caducous stipules, small inflo-
rescences, and narrow calyx lobes to 1.5 m long.
The species is closely related to P. nervosa but
differs in the large calyx lobes, narrower leaves,
and smaller inflorescences. This distinctive species
is a member of subgenus Psychotria.

Psychotria borucana (A. Molina) C. M. Taylor &
W. Burger, comb, nov., Cat. fl. Peru (in press).
Cephaelis affinis Standl., Publ. Field Mus. Nat.
Hist., Bot. Ser. 8: 1 84. 1 930, not P. affinis Baker.
C. borucana A. Molina, Ceiba 4: 31. 1951. Fig-
ure 17.

Shrubs or woody herbs, 1.5-4 m tall, leafy stems 2.5-
8 mm thick, glabrous and drying dark, quadrangular or
flattened; stipules 9-20 mm long and 5-10(-15) mm
broad, basal sheath 1-2 mm long, ovate-triangular, ob-
tuse, with parallel venation, glabrous or minutely brown-
ish puberulent. Leaves with petioles 12-50 mm long,

1 .7-3(-5) mm thick, glabrous (as in the type) or minutely
papillate-puberulent with erect brownish hairs 0. 1 mm
long; leaf blades 1 5-28(-32) cm long, 7- 14(- 16) cm broad,
obovate to elliptic obovate (as in the type) to broadly
elliptic, apex short-acuminate (acute) with tip 5-15 mm
long, base gradually narrowed and cuneate to obtuse,
drying stiffly chartaceous, dark grayish above and mark-
edly paler beneath, glabrous above, glabrous beneath (as
in the type) or with minute hairs 0.1-0.2 mm long
throughout, 2 veins 6-10/side. Inflorescences terminal,
capitulae solitary, 2-4 cm broad, peduncle 5-20 cm long,
2.5-4 mm diam., glabrous or minutely puberulent, basal
bracts mostly 2, 4-15 mm long, broadly ovate to reni-
form, interior bracteoles 4-8 mm long, spatulate, pale
purple, lavender, or green marked with purple, flowers
enclosed by bracts. Flowers with hypanthium ca. 2 mm
long, calyx 1-3 mm long and 3 mm broad; corolla fun-
nelform, white, glabrous, 20-22 mm long, tube 14-19
mm long and 2-4 mm diam., lobes 5(-6), 2.5-3 mm long
1.5- 2 mm broad at the base, acute; stamens 5(-6), an-
thers subsessile, ca. 2.8 mm long; style ca. 16 mm long.
Fruits ca. 8 mm long and 8 mm diam., rounded-oblong,
becoming blue or purple, 1 0-ribbed when dried; pyrenes
with 4-6 ribs.

Plants of evergreen forest formations of the Pa-
cific slope, from around 50 to 500 m elevation in
southern and south-central Costa Rica. Flowering
in May-July; fruiting in late August and Novem-
ber. The species is also known from Peru.

Psychotria borucana is recognized by its large
leaves, long-pedunculate purple heads, two broad
inflorescence bracts with reticulate venation, long
corolla tube with short lobes, unusual stipules, and
restricted range (in Costa Rica). The leaves are
quite variable in shape. Compare P. correae with
pendant heads and more secondary veins.

Psychotria brachiata Sw., Prodr. 45. 1788. Figure
58.

Shrubs or small trees, (0.5-)1.5-3(-5) m tall, leafy
branchlets 1.2-5 mm thick, glabrous, quadrangular be-
coming terete; stipules 3-7 mm long, 2-5 mm broad,
rounded and bilobed with a sinus 0.5-2 mm deep and
2 rounded lobes, glabrous and persisting. Leaves with
petioles (1-) 1.5-3. 5 cm long, 1-1.5 mm thick, glabrous;
leaf blades 9-17(-21) cm long, 3-7(-ll) cm broad, el-
liptic to elliptic-oblong or elliptic-obovate, apex usually
short-acuminate with tip 5-10(-l 5) mm long, base acute
to attenuate (obtuse), drying thin-chartaceous or char-
taceous and greenish or brownish, glabrous above, gla-
brous beneath except for a line of short (0.5 mm) hairs
along sides of midvein, 2 veins 7-10/side. Inflorescences
terminal and solitary or 3 (4, 5), 10-20 cm long, 3-10
cm broad, open pyramidal panicles with opposite sep-
arated (6-30 mm) lateral branches mostly arising at 90
angles, peduncles 1.5-6 cm long, with thin hairs ca. 0.4
mm long (mostly in 2 longitudinal rows), often with 3
2 branches bearing the distal flower clusters, bracts 3-
8 mm long and 1.5-3 mm broad, lanceolate, floral bracts

236

FIELDIANA: BOTANY

2-3 mm long, broadly ovate, ciliolate along the margins,
pedicels 0-2 mm long, flowers borne in capitulae 5-10
mm broad. Flowers with hypanthium 1-1.5 mm long,
calyx 1-1.5 mm long, yellowish, calyx truncate or slightly
dentate; corolla yellow or white (often with bluish tips
and yellow throat), funnelform, tube 3-5 mm long and
0.7-2 mm diam., usually glabrous externally on the low-
er half, lobes 5, 1-1.5 mm long. Fruits 4-5 mm long
(including calyx) and 3-6 mm diam., obovoid to ellip-
soid but becoming subglobose and spongy in life (to 10
mm diam.), bright blue to dark blue or purple, with
prominent longitudinal ribs, persisting calyx ca. 1 mm
long; pyrenes 3-4 mm long, with 3-5 ribs.

Plants of secondary vegetation in evergreen
tropical and premontane wet forest formations on
the Caribbean slope and in the General Valley,
from 5 to 800 m elevation in Costa Rica. This
species appears to flower and fruit throughout the
year in Central America. At La Selva flowering is
primarily in May-July and fruiting is mostly in
July-September. The species ranges from the West
Indies and Mexico through Central America to
Peru.

Psychotria brachiata is recognized by its open
inflorescences with well separated short opposite
branches bearing closely clustered yellowish flow-
ers in distal groups subtended by prominent per-
sisting bracts. The small stature, nearly glabrous
leaves usually drying greenish, spongy bright blue
fruit with longitudinal ribs, and persisting stipules
are additional features. Immature inflorescences
of P. brachiata are racemose with the congested
flowers in unexpanded capitula at the ends of the
opposite lateral branches.

Steyermark (1972, pp. 585-586) distinguishes
P. caerulea Ruiz & Pavon from P. brachiata by
its larger corollas (11-12 mm long), external co-
rolla surface that is densely puberulent through-
out, more prominent calyx lobes, larger clusters
of flowers, and larger floral bracts forming a more
definite involucre. Specimens that fit Steyermark's
definition of P. caerulea appear to occur in Bocas
del Toro, Panama (D. Simpson in herb. 1977),
and it is possible that they also occur in nearby
Costa Rica. The question of whether or not these
two species intergrade and are conspecific might
make a worthwhile study.

Psychotria brachybotrya Muell.-Arg. in Mart., Fl.
Bras. 6(5): 327. 1881. P. iquitosensis Standl.,
Publ. Field Columb. Mus., Bot. Ser. 8: 195. 1930.
Figure 56.

Shrubs or subshrubs, l-2(-3) m tall, leafy branchlets
0.8-4 mm thick, glabrous; stipules separate or with a

short (0.5-1 mm) sheath, lobes 2/side, 2-7 mm long,
0.5-1 mm broad at base, linear to narrowly triangular,
usually separated by a U-shaped sinus, glabrous, per-
sisting. Leaves with petioles 3-10 mm long (poorly dif-
ferentiated from the decurrent leaf base), glabrous; leaf
blades 9-16(-18) cm long, 3-7(-8) cm broad, ovate-el-
liptic to ovate or elliptic, apex usually tapering gradually
and short-acuminate or acute, tip 5-10 mm long, taper-
ing gradually or abruptly to the attenuate base (acute to
broadly obtuse above the narrowed base), drying thinly
chartaceous and greenish, glabrous above and below, 2
veins 4-8/side and arcuate ascending, distal 3 veins of-
ten subparallel. Inflorescences terminal and solitary, 1 .2-
3 cm long but enlarging in fruit, 1-2 cm wide, narrowly
paniculate with short opposite lateral branches or some-
times subcapitate, peduncle 3-18 mm long, ca. 1 mm
thick, with thin whitish hairs to 0.5 mm long, 2 branches
0-5 mm long, cymes congested, usually subtended by 3
involucrate narrowly ovate to lanceolate bracts, median
bracts 3-6 mm long and 2-3 mm broad, with 2 shorter
lateral bracts, all 3 united at base, flowers sessile. Flowers
with hypanthium ca. 0.5 mm long, puberulent, calyx
lobes 5, 0.2-0.3 mm long; corolla white, salverform, tube
1.5-4 mm long, ca. 1 mm diam. distally, glabrous or
puberulent, lobes 5, 0.8-1.5 mm long. Fruits ca. 4 mm
long and 4 mm diam., globose to oblong, becoming pur-
ple-black at maturity (blue-black on Cocos Island), with
prominent longitudinal ribs (transverse ridges some-
times visible when dried), calyx minute (0.3 mm high)
or obscure; pyrenes 3-4 mm long, with 4-5 prominent
ridges.

Plants of the lowland rain forest formations on
both the Caribbean and Pacific slopes, from near
sea level to 400(-900) m elevation and usually
found in shaded, poorly drained sites. Flowering
in July-August; fruiting in August-October and
December. In addition to the mainland plants, a
distinctive population of this species is found on
Cocos Island (see below). The species ranges from
Costa Rica and Panama to Brazil and Bolivia.

Psychotria brachybotrya is distinguished by its
well-separated stipule lobes, glabrous leaves dry-
ing green, small compact inflorescences, flowers
subtended by an involucre of bracts, small white
flowers, and purplish black fruit. Inflorescences
change in shape as they grow and expand. Collec-
tions from Cocos Island placed under this name
differ from the mainland material in having small-
er leaves with blades (4-)6-9 cm long and 1.5-3.5
cm broad; the stipules, flowers, and fruit are very
similar. Compare this species to P. hoffmannseg-
giana, P. platypoda, and P. officinalis. Hammel (in
Taylor, 1991) noted that P. brachybotrya is usually
a larger plant than P. officinalis and the latter has
a linear ventral sulcus on the pyrene.

Psychotria ca loch lam ys Standl., Publ. Field Co-
lumb. Mus., Bot. Ser. 8: 199. 1930.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

237

Small shrubs or treelets, 1.5-4.5 m tall, leafy stems
2.5-4.5 mm thick, glabrous or sparsely and minutely
puberulent; stipules 1 3-28 mm long and 5-9 mm broad
at the base, ovate-triangular with 2 attenuate-acute lobes
6-10 mm long, greenish to wine red or violet, with fine
parallel venation, chartaceous. Leaves with petioles 3-
16(-25) mm long, 0.9-2 mm thick, glabrous; leaf blades
8-23 cm long, 2-8.5 cm broad, elliptic-ovate to oblong-
lanceolate or lanceolate, apex gradually tapering and acu-
minate, tip 5-10 mm long, base acute to obtuse, drying
chartaceous and greenish, glabrous above and below, 2
veins 1 1-17/side, often with lesser 2 veins between the
major. Inflorescences solitary and terminal, 12-14 cm
long and 6-7 cm broad, hemispheric panicles with 2-
3(-4) branches at the first and second nodes, peduncles
3-8 cm long and 1.5-2 mm thick, white, minutely (0.1
mm) puberulent, bracts subtending the first whorl of
branches 1 .5-2 cm long, 3-4 mm wide, narrowly elliptic,
resembling the stipules in texture, distal bracteoles 5-14
mm long, 2-4 mm broad, lanceolate, persisting, white
to violet, enclosing the sessile flowers. Flowers ca. 15
mm long, hypanthium 0.6-1 mm long, calyx 0.6-0.8 mm
long, lobes 5, triangular; corolla white to purple or pale
violet, yellow in throat, tubular-salverform, tube 7-10
mm long, lobes 5, 2-3 mm long; anthers ca. 1.5 mm
long and curved. Fruits ca. 6 mm long and 4 mm diam.,
oblong, dark purple or black; pyrenes with 4-5 ribs.

In Costa Rica this disjunct species is only known
from the hills near Golfito, Puntarenas, at about
1 00 m elevation, flowering in December-January.
The species also occurs in Choco, Colombia, and
Amazonian Peru.

Psychotria calochlamys is distinguished by its
large deciduous stipules, long and persisting white
or lilac bracts and bracteoles, and large flowers.
The unusual form and size of the stipules is similar
in P. capitata, a close relative. The inflorescence
with large white (or purplish) bracts and flowers
is unlike any other Costa Rican species of Psy-
chotria and reminiscent of some species of Pali-
courea. The disjunct occurrence in Costa Rica is
similar to that of some other species in the Golfo
Dulce region. This species may prove to be con-
specific with P. stipulosa Muell. Arg. of Amazo-
nian South America, but that species (interpreted
in a broad sense) includes variation not seen in
our material.

The leaves drying grayish and the bright red fruit
are characteristics of the subgenus Psychotria. This
species has a disjunct distribution and is not known
from Costa Rica. It is found in southern Mexico
and Guatemala and again in Colon and Code
provinces in Panama.

Psychotria camponutans (Dwyer & Hayden) Ham-
mel, Selbyana 12: 139. 1991. Cephaelis cam-
ponutans Dwyer & Hayden, Ann. Missouri Bot.
Gard. 55: 35. 1968.

Semisucculent herbs or subshrubs 0.5-1.2 m tall, rhi-
zomatous, stems unbranched, leafy stems 24 mm thick,
quadrangular or terete, glabrous, drying black and con-
tracted below the nodes; stipules 6-8(-15) mm long, 5-
6(-10) mm broad, broadly ovate with a serrated margin
and 2 long lobes, glabrous and thin, distal parts decid-
uous but with a persisting truncated sheath 1-1.5 mm
long. Leaves with petioles 18-55 mm long, 1.2-2 mm
thick, glabrous, drying black; leaf blades (1 1-)18-25 cm
long, (4-)7-9 cm broad, narrowly elliptic-oblong to ob-
long or oblong-obovate, apex acuminate with tip 6-13
mm long, acute at base and slightly decurrent on petiole,
drying thinly chartaceous, dark or grayish above, gla-
brous above and below, 2 veins 8-15/side but these
difficult to see. Inflorescences axillary and I/node, 1-2
cm long, to 2 cm diam., capitate, peduncle 4-10 mm
long, with involucrate bracts 8-15 mm long, 5 mm broad,
ovate and acuminate, glabrous and green, flowers sessile
and congested, each enclosed by 2 lanceolate bracteoles
6-8 mm long. Flowers glabrous, calyx lobes 5-6, ca. 1
mm long; corolla white, funnelform, tube ca. 3 mm long,
corolla lobes 1-1.5 mm long. Fruits 7-8 mm long, 4-5
mm diam., ellipsoid to obovoid, crowned by a persisting
calyx, deep maroon red or pale rose red, bracts red to
purple; pyrenes 6-7 mm long, ribbed.

Plants of the wet evergreen forests of the Carib-
bean slope, from 300 to 900 m elevation. Flow-
ering in April and July; immature fruit were col-
lected in September. Costa Rica to central Panama.

Psychotria camponutans is distinguished by its
short semisucculent erect stems with small axillary
capitate inflorescences, reddish fruit, and restric-
tion to the Caribbean slope. Compare P. uliginosa
and P. cartagoensis.

Psychotria calophylla Standl., Contr. U.S. Natl.
Herb. 18: 129. 1916.

Small trees, 3-18 m tall, with larger (13-26 x
5-12 cm) elliptic to obovate leaves usually reddish
tomentulose beneath, with an arcuate submarginal
vein; the inflorescences paniculate with usually 3
ranks of branching, the flowers in distal glomer-
ules, and larger (10-14 x 6-9 mm) ellipsoid fruit.

Psychotria capacifolia Dwyer, Ann. Missouri. Bot.
Gard. 67: 353. 1980. Figure 54.

Herbs or subshrubs, 0.4-1 (-2?) m tall, usually suc-
culent and unbranched, leafy stems 3-12 mm thick, gla-
brous or with hairs to 1 mm long at nodes, terete, drying
dark; stipules 3-6 mm long, to 8 mm broad, broadly
obtuse to truncate and entire, with a conical caducous
lateral appendage to 3 mm long, densely hirtellous. Leaves
densely hirtellous in early stages but usually glabrescent

238

FIELDIANA: BOTANY

in age, petioles 3-10 cm long, 2-4 mm thick, with hairs
ca. 1 mm long in early stages; leaf blades 2040 cm long,
9-1 6(-20) cm broad, oblong to elliptic-oblong or elliptic-
obovate, apex abruptly narrowed and short-acuminate,
base cuneate to obtuse, drying thinly chartaceous and
grayish green, usually glabrous above, glabrescent or with
thin hairs to 1 .4 mm long beneath, 2 veins 1 7-22/side,
connected by a distinct slightly arcuate submarginal vein
2-3 mm from the margin. Inflorescences axillary, I/node,
(8-) 15-3 5 cm long, 5-12 cm wide, open panicles with
opposite branching, peduncles 7-22 cm long, 1.5-2.2
thick, with thin yellowish to reddish brown hairs 0.2-
0.7 mm long, bracts 1-2 mm long, flowers sessile on
distal dichotomous branches or distal clusters, bracteoles
ca. 0.5 mm long. Flowers puberulent externally, hypan-
thium ca. 0.7 mm long, calyx to 1 mm long, calyx lobes
5, 0.3-0.5 mm long, triangular; corolla white or pale
yellow, funnelform, tube 2.5-4 mm long, 0.5-1 mm diam.
basally and 2 mm distally, lobes 5, ca. 1 mm long. Fruits
5-6 mm long, 3-4 mm broad, ovoid, yellow-green or
whitish, with ca. 10 longitudinal ribs; pyrenes ca. 5 mm
long.

Plants of evergreen rain forest formations on the
Caribbean slope, ranging from 5 to 1300 m ele-
vation. Flowering in January-September; fruiting
in March-November. The species ranges from
southern Nicaragua to western Panama.

Psychotria capacifolia is recognized by its short
succulent stems, pubescence on younger parts, large
leaves, axillary often large puberulent inflores-
cences, well-developed (only slightly arcuate) sub-
marginal vein, and the yellowish green fruit. The
corolla interior is said to have a "ball" of hairs
within (Dwyer, 1980, p. 354). This species closely
resembles some material placed under P. macro-
phylla, P. siggersiana, and P. aggregate, but the
differences used in the keys appear to reflect a real
discontinuity of gene flow.

Psychotria capitata Ruiz & Pav., Fl. Peruv. 2: 59,
pi. 206. 1799.

Shrubs or small treelets l-3(-4) m tall, leafy stems 2-
4 mm thick, glabrous; stipules 8-18 mm long, 4-7 mm
broad, ovate-lanceolate, acute at apex, with 2 triangular
lobes 1-10 mm long, glabrous, deciduous. Leaves with
petioles 4-20 mm long, 1-2 mm thick, glabrous, often
drying yellowish green; leaf blades 6-16(-22) cm long,
2.5-6(-8) cm broad, elliptic-oblong to elliptic, ovate-
elliptic or elliptic-lanceolate, apex short-acuminate with
tip 5-12 mm long, base acute or obtuse and slightly
decurrent on petiole, drying stiffly chartaceous, greenish,
glabrous above and below (except for short hairs along
the sides of the midvein beneath), 2 veins 9-14/side,
arising at almost 90 from midvein. Inflorescences ter-
minal and solitary, 5-1 3 cm long, 3-6 cm broad, an open
racemose or congested panicle, white at anthesis, pe-
duncles (l-)2.5-7 cm long, ca. 1.5 mm thick, glabrous
(rarely puberulent), branches opposite or ternate (alter-

nate) and usually without subtending bracts, flowers ses-
sile in distal cymose groupings of 2-4 subtended by white
lanceolate bracteoles 4-9 mm long and 1-2 mm broad.
Flowers glabrous externally, hypanthium ca. 1 mm long,
calyx ca. 0.5 mm long, calyx lobes 4-5, ca. 0.3 mm long,
triangular; corolla white or cream (yellow in throat), tube
3-7 mm long, 1 mm diam. near base, lobes 4-5, 1.5-4
mm long; stamens 4 or 5 anthers 1-2 mm long. Fruits
5-7 mm long, ca. 5 mm diam., subglobose, dark blue-
black to purple-black or black; pyrenes ribbed.

Plants of lowland evergreen forest formations,
0-200 m elevation. Flowering and fruiting
throughout the year (mostly in December-July).
The species ranges from Belize (Croat, 1978), the
Caribbean lowlands of Nicaragua and Panama to
Peru and Brazil; it is not known from Costa Rica.

Psychotria capitata is recognized by its white
flowers and inflorescences, conspicuous bracts on
the distal parts of the inflorescences, leaves usually
glabrous and drying greenish, and the large bilobed
stipules. This species is similar to P. calochlamys,
which has larger bracts and stipules and less prom-
inent secondary venation. Compare also Psycho-
tria officinalis and Palicourea tilaranensis. This
species appears to be less variable in Central
America than in South America (cf. Steyermark,
1974).

Psychotria cartagoensis Nepokroeff, nom. nov.
Cephaelis latistipula Standl., J. Wash. Acad.
Sci.18: 281. 1928, not Psychotria latistipula
Benth. Figure 13.

Herbaceous subshrubs, 0.3-1 m tall, stems erect and
usually unbranched, leafy stems 2-6 mm thick, glabrous,
quadrangular or terete; stipules (5-) 1 2-22 mm long, 1 2-
18 mm broad, ovate in general outline, bilobed with a
narrow sinus 2-7 mm deep, glabrous, stiffand persisting.
Leaves well separated along the stem, petioles 2-6(-7)
cm long, 1.2-3.7 mm thick, glabrous, drying dark; leaf
blades 13-27 cm long, 5-9 cm broad, elliptic-oblong,
oblong-lanceolate, elliptic-obovate, or ovate-elliptic, apex
abruptly narrowed or rounded and shortly acute or short-
acuminate, base acute to cuneate, drying stiffly charta-
ceous, dark gray-green above, much paler beneath, 2
veins 9-13/side, arising at 60-80 and arcuate distally.
Infloresences axillary and sessile or subsessile, broadly
capitate, 1-2 cm long and 24 cm broad, becoming dark
magenta or purplish, bracts 7-8 mm long and 2-4 mm
broad, innermost bracteoles lanceolate, flowers enclosed
by many bracteoles. Flowers glabrous externally, hypan-
thium ca. 2 mm long, turbinate, calyx cup 0.3-1 mm
long, calyx lobes 4-5(-6), 1-4 mm long, lanceolate; co-
rolla 4-5 mm long, funnelform, white, lobes 5, 1.2-3
mm long, acute; anthers 2.5 mm long. Fruits ca. 6 mm
long and 4 mm broad (not including the 2 mm long
persisting calyx), reddish purple.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

239

Understory plants of evergreen wet forests of
the Caribbean slope, from 800 to 1 500 m eleva-
tion. Flowering in March (Standley 39695 us the
type of C. latistipuld) and May-June; fruiting in
June and November. The species has only been
collected in the valley of Rio Sarapiqui (Heredia
Province) and near Orosi and Mufieco (Cartago)
in central Costa Rica.

Psychotria cartagoensis is recognized by its short
unbranched habit, broad bilobed stipules, axillary
and subsessile heads, and absence of pubescence.
Among our species, Hoffmannia congesta is most
likely to be confused with this species, but that
species lacks the involucrate bracts and has many-
seeded fruits. This species and its allies have been
studied by Molly Nepokroeff (wis, 1992); P. \vil-
buriana Dwyer and P. dukei Dwyer of Panama are
closely related.

Psychotria carthagenensis Jacq., Enum. PL Carib.
16. 1760. Figure 65.

Shrubs, subshrubs, or small treelets, 0.3-3(-4) m tall,
leafy branches 1.5-4 mm thick, glabrous and terete, be-
coming gray; stipules 3-8 mm long, 1.5-5 mm broad,
ovate to oblong or slightly obovate, apex rounded or
bluntly acute, glabrous, drying dark reddish brown, ca-
ducous. Leaves with petioles 3-10 mm long, ca. 1 mm
thick, with decurrent lamina base; leaf blades (4-)6-
13(-15) cm long, (1.7-)2-5(-7) cm broad, oblanceolate
to narrowly elliptic-obovate or narrowly elliptic, apex
obtuse to very shortly acuminate (tip ca. 4 mm long),
base gradually narrowed and cuneate or acute, decurrent
on petiole, drying chartaceous, gray or reddish gray, gla-
brous on both surfaces, 2 veins 6-8/side, domatia rarely
present in distal vein axils. Inflorescences terminal and
solitary, 3-8(-10) cm long, 2.5-4 cm broad, a compact
panicle with usually 3 nodes of opposite branches di-
minishing in size toward apex (also verticillate or um-
bellate), peduncle to 4.5(-6) cm long, 0.6-1 .3 mm thick,
glabrous, bracts subtending the lateral branches 0.5-1.5
mm long, subulate to broadly ovate, flowers in distal
cymes, pedicels 0-2 mm long, bracteoles to 0.5 mm long.
Flowers glabrous or minutely (0.05 mm) papillate-pu-
berulent externally, hypanthium ca. 1 mm long, calyx
ca. 0.5 mm long, subentire or with 5 short triangular
lobes; corolla white, funnelform-salverfbrm, tube 2.5-3
mm long, ca. 1 mm diam. near base, lobes 5, 1.2-2 mm
long; anthers 1-1.5 mm long. Fruits ca. 5 mm long and
4 mm diam., oblong or ellipsoid, becoming red or or-
ange, with 10 vertical ribs, glabrous; pyrenes with 2 sulci
on the inner face and 5 ribs on the exterior surface.

Plants of shaded sites in seasonally dry decid-
uous forest formations of Guanacaste and north-
ernmost Alajuela Province, from 5 to 200(-600)
m elevation (rarely to 1400 m elsewhere). Flow-
ering in November-August (primarily February-

July); fruiting throughout the year. The species
ranges from Mexico and the West Indies to Bolivia
and Argentina.

Psychotria carthagenensis is recognized by its
deciduous forest habitat, smaller oblanceolate or
obovate leaves, lack of conspicuous pubescence,
small flowers and inflorescences drying reddish,
and inflorescences often with nodes bearing two
larger and two smaller lateral branches. The ten-
dency of the leaves to dry gray or reddish and the
red or orange fruit are characteristics of members
of subgenus Psychotria. The correct spelling of this
species is carthagenensis (not carthaginensis}. This
species is similar to P. tenuifolia of similar habitats
but with bilobed stipules.

Psychotria chagrensis Standl., J. Wash. Acad. Sci.
15: 105. 1925. Figure 60.

Shrubs, 1-3 m tall, many-branched and often flat-
topped, leafy branchlets 1-3 mm thick, glabrous, reddish
brown or dark; stipules 3-10 mm long, 1 -3 mm broad
near base (broader when subtending the inflorescence),
narrowly tubular to narrowly ovate and acute at apex
with 1 or 2 slender awns 1-3 mm long, glabrous and
drying dark reddish brown, caducous. Leaves usually
closely clustered distally, petioles 1-12(-17) mm long,
ca. 0.7 mm thick, glabrous; leaf blades 2.5-8(-10) cm
long, 0.9-3(-3.5) cm broad, obovate to oblanceo-
late, elliptic-obovate or elliptic, apex acuminate or cau-
date-acuminate to bluntly obtuse, tip ca. 5 mm long,
base cuneate to acute and decurrent on petiole, drying
stiffly chartaceous, gray or reddish brown, glabrous above
and below, 2 veins 6-9/side (but sometimes difficult to
see), joined by an arcuate submarginal vein near margin.
Inflorescences terminal (rarely pseudoaxillary), 6-16 mm
long, to 7 mm broad, sessile, subtended by broad stipule-
like dark reddish brown glabrous bracts to 1 cm long,
fasciculate or capitate, with few sessile flowers, bracteoles
4-5 mm long. Flowers distylous, glabrous externally, hy-
panthium ca. 1 mm long, calyx 2-4 mm long, lobes 1-
2 mm long, narrowly triangular, corolla funnelform, white,
tube 49 mm long and 1-1.3 m diam., lobes 5, 2-3 mm
long; stamens 5, anthers ca. 1 m long. Fruits becoming
8 mm long (not including persisting calyx), 3-6 mm
diam., oblong-ellipsoid to ovoid, persisting calyx 2-3
mm long, becoming red or purple pyrenes ca. 6 mm long,
weakly ribbed.

Understory plants often found in low wet de-
pressions in evergreen rain forest formations, from
near sea level to 700 m elevation on the Caribbean
slope and Osa Peninsula. Probably flowering and
fruiting throughout the year (flowering mostly in
March-August). At La Selva flowering is most
common in May-June and fruiting in November-
December. The species ranges from southern Nic-
aragua to Colombia and Peru, but with disjunct

240

FIELDIANA: BOTANY

populations in Veracruz, Mexico, and Izabal,
Guatemala.

Psychotria chagrensis is recognized by its short
(often flat-topped) habit, very small obovate leaves,
distinctive stipules, small sessile fasciculate or cap-
itate inflorescences, and unusual calyx lobes. The
leaves drying grayish or reddish and the red fruit
are characteristic of subgenus Psychotria. The
slender secondary veins often have point inden-
tations on the lower surface after the leaves have
been dried. The very small leaves are unusual
among the woody plants of the lowland rain forest
floor; compare P. graciliflora, P. parvifolia, and
Randia loniceroides.

from near sea level to 700 m elevation. Flowering
in May-September and January; fruiting in July-
March. The species ranges from southern Mexico
and Belize to central Panama.

Psychotria chiapensis has the longest corolla
tubes among Central American Psychotria species;
it is pollinated by long-tongued sphingid moths
(cf. Bawa & Beach, 1983). The terminal subcapi-
tate inflorescences (often in groups of three) with
broad bracts and the relatlively large four-edged
fruit are also distinctive. The paucity of puberu-
lence, and leaves usually broadest at or above the
middle are additional characters that make this
species stand apart. It is called cocobolito in Pan-
ama.

Psychotria chiapensis Standl., Contr. U.S. Natl.
Herb. 23: 1390. 1926. Cephaelis tetragona J. D.
Smith, Bot. Gaz. 61: 376. 1916, not Psychotria
tetragona Seem., 1867. Figure 18.

Psychotria chiriquiensis (Standl.) C. M. Taylor,
comb. nov. Cephaelis chiriquiensis Standl., Ann.
Missouri Bot. Gard. 28: 469. 1941. Figure 17.

Shrubs, treelets, or small trees, 2.5-8(-10) m tall, much-
branched with rounded crown, leafy stems 2-6(-8) mm
thick, terete and often conspicuously contracted for 5-
10 mm below the node after drying, glabrous or with
short (0.2 mm) hairs in new growth, center with white
spongy pith; stipules 2-5 mm long, 2 4(-8) mm broad,
broadly triangular, rounded with a small apical sinus,
with 2 triangular acute lobes to 2 mm long. Leaves with
petioles 7-35 mm long, 0.8-2.3 mm thick, glabrous
(sometimes puberulent near inflorescences); leaf blades
9-20(-23) cm long, 3-8(-9) cm broad, elliptic-obovate
to elliptic-oblong or narrowly oblong-obovate, apex short-
acuminate or acute with tip 5-10 mm long, base usually
gradually narrowed and cuneate base, slightly decurrent
on petiole, drying thin-chartaceous and grayish green to
dark brown above, distinctly paler beneath, glabrous
above and below except for short (0.2 mm hairs) along
sides of midvein beneath, 2 veins 9-12/side. Inflores-
cences terminal, 1 (or 3 when basal lateral branches are
axillary to distal leaves), 4-12 cm long, 5-8 cm broad,
subcapitate or corymbiform with 3-7 dense bracteate
flower clusters, peduncles l-8(-15) cm long, 1.3-3 mm
thick, minutely puberulent with hairs 0.2 mm long, bracts
6-10 mm long, 4-8 mm broad, ovate to ovate-elliptic,
obtuse to rounded, minutely ciliolate along margins, with
6-many flowers closely clustered and sessile. Flowers
distylous, calyx resembling the bracts in texture and col-
or, 3-5 mm long, calyx lobes 5, ca. 1.5 mm long, tri-
angular; corolla salverform, white, glabrous externally in
ours, tube 20-45 mm long, 1-3 mm diam. for most of
its length, lobes 5, 9-15 mm long and 3 mm broad;
anthers ca. 4 mm long. Fruits 12-16 mm long (not in-
cluding the calyx) and 9-1 3 mm diam., ellipsoid to ovoid,
purple-black, strongly 4-angled, persisting calyx 2-3 mm
long; pyrenes 10-13 mm long, 7 mm broad and 3 mm
thick, bony, with 3 dorsal ribs and concave areas be-
tween.

Plants of evergreen lowland rain forest forma-
tions on both the Caribbean and Pacific slopes,

Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla-
brous; stipules 3-5 mm long, with a short (1 mm) tube
and 4 rounded ovate lobes 2-4 mm long at each node,
these separate on the internode but overlapping above
petioles, persisting. Leaves well separated along stems,
petioles 15-40(-57) mm long, 0.5-1.5 mm thick, gla-
brous; leaf blades 8-16(-20) cm long, 3-7 (-9) cm broad,
elliptic to elliptic-oblong, apex acuminate with tip 5-10
mm long, base obtuse to acute, drying stiffly chartaceous,
dark green to yellowish brown or grayish brown, glabrous
above and beneath, 2 veins 1 1-22/side, loop-connected
distally to form an arcuate submarginal vein. Inflores-
cences terminal (pseudoaxillary), solitary or 3, 3-12 cm
long, 3-8 cm broad, capitulum often with 3 short (2-5
mm) primary branches (especially in fruit), involucral
bracts 12-20 mm long, 20-50 mm broad, broadly ovate,
purple to reddish violet, peduncle l-6(-8) cm long, 2-
3.5 mm thick, glabrous, flowers subsessile within many
imbricate bracteoles 8- 12 x 5-1 Omm, elliptic to oblong.
Flowers glabrous externally, hypanthium ca. 1 mm long,
0.5 mm diam., turbinate, calyx tube ca. 1 mm long, lobes
ca. 0.5 mm long; corolla tubular-funnelform, purplish,
tube 9-12 mm long and 1.7 mm diam., lobes 5-6, 1.5-
2 mm long; anthers to 2 mm long. Fruits elliptic, blue,
enclosed within the inflorescence; pyrenes with 4-5 ridg-

Plants of lower montane rain forest formations,
from 1 200 to 1 800 m elevation. Flowering in Jan-
uary-September; fruiting in October-December.
This species ranges from Volcan Tenorio to south-
ward along the Caribbean slope and continental
divide to the Chiriqui Highlands.

Psychotria chiriquiensis is recognized by its in-
volucrate capitate inflorescences with broad pur-
ple bracts, long-petiolate leaves with many prom-
inent secondary veins, lack of pubescence, and
rounded stipule lobes usually overlapping above

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

241

the petiole. This species is named in honor of An-
tonio Molina R., who monographed the Mexican
and Central American species ofCephaelis in 1953
and has made many important collections in Cen-
tral America. This species is very similar to P.
data, with larger red bracts, heads not three-
branched, and white corollas. It is also similar to
P. dichroa.

Psychatria chiriquina Standl., Contr. U.S. Natl.
Herb. 18: 129. 1916. Figure 61.

Shrubs or small trees, 2-6 m tall, leafy stems 1-4 mm
thick, glabrous, dark reddish brown; stipules 4-8(-10)
mm long, 3-5 mm broad, ovate to elliptic, apex acute,
ciliolate to erose or shortly bilobed, united around stem
at base, glabrous or puberulent, caducous. Leaves with
petioles 7-18(-35) mm long (sometimes variable on the
same stem), 0.4-1.3 mm broad, glabrous; leaf blade 5-
15 cm long, 2-6 cm broad, narrowly elliptic, elliptic-
oblanceolate to oblanceolate or oblong-obovate, apex
gradually tapering and acute or acuminate, tip ca. 10
mm long, gradually narrowed to an acute base and de-
current on petiole, drying stiffly chartaceous, dark red-
dish brown above, glabrous above and below except for
hairs along sides of midvein, occasional pit domatia in
vein axils beneath, 2 veins 6-12/side. Inflorescences
terminal, usually solitary or 3, 3-10(-l 7) cm long, a few-
branched panicle, often trichotomous, peduncles 1.5-5
cm long, 0.5-2 mm thick, essentially glabrous, pedicels
0-3 mm long. Flowers glabrous externally, calyx ca. 1
mm long, truncate or with 5 small (0.6 mm) lobes; corolla
white, salverform or funnelform, tube 3-6 mm long, 1.5-
2 mm diam., glabrous or puberulent externally, lobes 5,
1.5-2 mm long; stamens 5, anthers ca. 1 mm long. Fruits
5-8 mm long, 4-6 mm diam., oblong to subglobose,
becoming red, persisting calyx 0.5-1 mm long; pyrenes
with 45 rounded ribs.

Psychotria chitariana Dwyer & C. Hamilton, Phy-
tologia64: 221. 1988.

Subshrubs, 30-100 cm tall, leafy stems 49 mm thick,
glabrous, drying dark; stipules 15-18 mm long, ovate,
with 2 awns ca. 3 mm long, caducous. Leaves with pet-
ioles 0-27 mm long, ca. 3 mm thick, glabrous, with
lateral margins and flattened above; leaf blades 25-40
cm long, 10-17 cm broad, elliptic-obovate to obovate-
oblong or oblanceolate, apex obtuse to acute, base grad-
ually narrowed and cuneate or expanded and auriculate
at the petiole, drying thinly chartaceous, grayish brown
to reddish brown, glabrous above and below, 2 veins
12-17/side, loop-connected 2-3 mm from the margin.
Inflorescences terminal and solitary, dense panicles of
congested cymes, globose or hemispheric, 3-1 1 cm long,
3-5 cm broad, peduncles l-2.5(-^.5?) cm long, bracts
ca. 5-6 x 1-3 mm, glabrous, flowers closely congested,
pedicels 3-5 mm long, bracteoles to 0.5 mm long. Flow-
ers glabrous externally, calyx ca. 1 mm long, cupulate,
lobes 5, short; corolla funnelform, greenish white, tube
4. 5-5 mm long, 1-1.5 mm diam., lobes 1.5-2. 3 mm long,
triangular or oblong; anthers ca. 1.5 mm long. Fruits not
seen.

Known only from the Caribbean slope of the
Cordillera de Talamanca in Costa Rica, from 200
to 800 m elevation. Flowering in April-May; im-
mature fruits were collected in September. The
species ranges from the valley of Rio Chitaria east
of Turrialba (Liesner et al. 15400 CR, MO holotype)
to the valley of Rio Estrella.

Psychotria chitariana is distinguished by its short
stature, large obovate leaves with rounded and
obtuse apex, and unusual inflorescence. The leaves
drying dark grayish (or reddish brown) is a char-
acteristic of subgenus Psychotria. Compare P. al-
faroana with more open inflorescences and acute
leaf bases.

Plants of evergreen montane cloud forest for-
mations, from (900-)1500 to 2500 m elevation.
Flowering in January-May (July-August in Nic-
aragua); fruiting in August-April (November-May
in Nicaragua). The species is known from northern
and central Nicaragua and from the easternmost
part of the Cordillera de Talamanca in Costa Rica
and the Chiriqui Highlands in Panama.

Psychotria chiriquina is recognized by its higher
elevation habitats, stipules often with two small
distal lobes, narrow leaves tapering gradually to
apex and base, pit domatia, small flowers, and
poorly developed calyx lobes. The red fruit and
leaves/branchlets drying dark reddish brown are
characters of subgenus Psychotria. Compare P.
sarapiquensis, with a shorter calyx, and P. sylvi-
vaga. This species is also similar to P. panamensis
and with very different stipules.

Psychotria cincta Standl., Publ. Field Columb.
Mus., Hot. Ser. 7: 90. 1930. Figure 58.

Shrubs, l-2.5(-4) m tall, leafy branches 1.5-4 mm
thick, glabrous and drying greenish or dark; stipules
forming a small basal (1-3 mm) sheath, with 2 awns (4/
node), awns 6-14 mm long, ca. 0.5 mm broad, glabrous,
persisting. Leaves with petioles 8-20 mm long, 1-1.5
mm thick, glabrous; leaf blades 12. 5-21 (-28) cm long,
3. 5-7 (-9) cm broad, elliptic-oblong to oblong or ovate-
oblong, apex tapering gradually and acuminate, tip 8-
18 mm long, base acute to obtuse, drying stiffly char-
taceous, green or grayish green, glabrous above and be-
low, 2 veins 9-1 1 (- 1 4)/side, leaf margin with a distinctly
thickened vein along the edge. Inflorescences terminal
and solitary, usually pendant, 12-20 cm long, 2-4 cm
broad, racemiform thyrsoid panicles with short (1-2 cm)
usually alternate lateral cymose branches, peduncles 6-
14 cm long, 0.6-1.2 mm thick, sparsely and minutely
puberulent with short (0.1-0.2 mm) thin hairs, bracts

242

FIELDIANA: BOTANY

linear, adnate to lateral branches or absent, bracteole 0-
0.5 mm long, pedicels 0-2 mm long. Flowers minutely
puberulent externally, hypanthium 0.5 mm long, calyx
0.2-0.4 mm long, truncate or with minute lobes; corolla
funnelform or salverform, white or greenish white, tube
2-3 mm long, ca. 0.5 mm diam., lobes 5, 1-1.5 mm long.
Fruits 4-6 mm long, subglobose, obscurely ribbed, gla-
brous, red becoming black; pyrenes shallowly ribbed.

Plants of evergreen lowland Caribbean rain for-
est formations, from near sea level to 500 m ele-
vation. Flowering in February-July; fruiting in
August. The species ranges from Nicaragua to Co-
lombia.

Psychotria cincta is recognized by its persisting
stipules with long narrow awns, the often oblong
leaves usually drying yellowish green, the long nar-
row pendant inflorescences with unusually short
lateral branches, and the small puberulent flowers.
In addition, the thickened leaf margins are nota-
ble. This distinctive species is presently known
only from La Selva and nearby areas in Costa Rica.
Compare P. deflexa, which lacks the thickened leaf
margins and has pyrenes with transverse ribbing.

Psychotria clivorum Standl. & Steyerm., Publ. Field
Mus. Nat. Hist., Bot. Ser. 23: 87. 1944, based
on P. limonensis var. angustifolia Standl., loc.
cit. 17: 282. 1937. Figure 61.

Shrubs or small trees, 1.5-6 m tall, leafy stems 2-4
mm thick, glabrous; stipules 10-16 mm long, 2-5 mm
broad (broader beneath the inflorescences), with a sheath
1-4 mm long, ovate to elliptic, usually bilobed with 2
acute lobes and a short (1-3 mm) sinus at apex, glabrous,
dark reddish brown, caducous. Leaves variable in shape
(on different plants), petioles 6-35 mm long, 1-1.7 mm
thick, glabrous; leaf blades 10-22 cm long, 3.5-8 cm
broad, narrowly elliptic-oblong to elliptic or oblanceo-
late, apex acuminate, tip 7-15 mm long, base acute to
cuneate and often long-decurrent on petiole, drying
membranaceous to thinly chartaceous, grayish or gray
tinted with red, glabrous above and below (midvein rare-
ly sparsely puberulent beneath), 2 veins 8-12/side, loop
connected 1-3 mm from margin, small pit domatia usu-
ally present. Inflorescences terminal and solitary, 6-10
cm long, 3-7 cm broad, open panicles, usually with 4
lateral branches from 2 proximal nodes, peduncles 1-5
cm long and 2.2 mm thick, minutely puberulent or gla-
brous, reddish brown, bracts to 2(-3) mm long, pedicels
0-2 mm long. Flowers glabrous externally, hypanthium
ca. 0.6 mm long, conical, calyx tube ca. 0.5 mm long
with minute (0.2 mm) broadly triangular lobes; corolla
funnelform, white, tube 2-2.5 mm long, 1.2-1.5 mm
diam., lobes 1-2 mm long; anthers ca. 1 mm long. Fruits
5-6 mm long, 4-5 mm diam., ellipsoid or obovoid, lon-
gitudinally ribbed, red; pyrenes with 3-5 longitudinal
ridges.

Plants of evergreen forest formations of the Ca-
ribbean slope; from near sea level to 800 m ele-
vation (to 1400 m in Guatemala). Most flowering
collections have been made in March-May. A lit-
tle-collected species ranging from Veracruz, Mex-
ico, to central Panama.

Psychotria clivorum is recognized by its larger
narrow leaves gradually narrowed to the base with
numerous pit domatia and distinct submarginal
vein, stipules tubular at base and sometimes bi-
lobed distally, inflorescences with whorled lateral
branches at first two nodes (usually of two unequal
pairs), small flowers, and reddish fruit. The ten-
dency of the leaves to dry grayish and the stipules
to dry reddish is characteristic of subgenus Psy-
chotria. Specimens of this species may be difficult
to separate from material of P. orosiana and P.
sarapiquensis.

Psychotria cocosensis C. Hamilton, Phytologia 64:
222. 1988. Figure 62.

Shrubs or small trees to 5 m tall, leafy stems ca. 2-7
mm thick, glabrous; stipules (7-) 1 2-35 mm long, ( 1 .5-)2-
5 mm broad, lanceolate, glabrous, drying dark reddish
brown, caducous. Leaves with petioles 5-16(-23) mm
long, 1-2.5 mm thick, glabrous; leaf blades (8-) 12-1 7
cm long, (3-)5-7 cm broad, narrowly elliptic to elliptic-
oblong or broadly elliptic-obovate, apex acuminate or
caudate-acuminate with tip 4-12 mm long, base acute
to attenuate and decurrent on petiole, drying subcoria-
ceous, reddish brown to grayish brown, glabrous above
and beneath, 2 veins 9-1 2/side, small pit domatia often
present in the vein axils beneath. Inflorescences terminal
or pseudoaxillary, solitary or 3-parted, 1.3-3 cm long,
few-branched panicles with distal 3-flowered cymules,
peduncles 0-7 mm long, ca. 1 mm thick, glabrous, bracts
0.5-1.5 mm long, triangular, glabrous, pedicels 2-3 mm
long. Flowers puberulent externally, hypanthium ca. 0.5
mm long, glabrous, calyx tube ca. 1 mm long and 2 mm
broad, cupulate, slightly denatate; corolla funnelform,
white, tube ca. 3 mm long and 2 mm diam., lobes 5, ca.
4 mm long, 2 mm wide; stamens 5, anthers ca. 2 mm
long. Fruits ca. 6 mm long, 5-6 mm diam., ellipsoid to
globose, orange then red and finally purple, drying dark
reddish brown, persisting calyx 0.8-1.5 m long.

Plants known only from Cocos Island, from near
sea level to 50 m elevation. We have seen four
collections: Foster 4132 F, Holdridge 5169 us, Pit-
tier 12375 us (the holotype), and Pittier 16279 GH,
us. Flowering in April-June; fruiting in January,
April, and June.

Psychotria cocosensis is similar to P. panamen-
sis but differs in its subcoriaceous leaves and larger
fruits with persisting cupulate calyx. The very small
domatia, short inflorescences, and flowers with long

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

243

corolla lobes are distinctive. The red fruit and ten-
dency of the leaves to dry reddish brown are char-
acteristics of subgenus Psychotria. The Holdridge
collection is unusual in having broadly elliptic-
obovate leaves that have dried grayish and sig-
nificantly paler beneath.

Psychotria cooperi Standl., Publ. Field Columb.
Mus., Bot. Ser. 4: 296. 1929. Figure 13.

Small trees or shrubs, 3-8 m tall, to 10 cm dbh, leafy
stems 1 .4-5 thick, glabrous, terete or slightly tetragonal;
stipules 5-9(-l 1) mm long, to 8 mm broad, with a broad
tube, truncate to rounded or bilobed with a small ( 1 mm)
sinus at apex, glabrous, persisting or deciduous. Leaves
opposite or 3 at a node, petioles 7-25 mm long, 0.7-1.7
mm thick, glabrous; leaf blades 8-21 cm long, 3.5-7 cm
broad, elliptic-oblong to elliptic or elliptic-obovate, apex
acuminate with tip 3-10 mm long, base gradually nar-
rowed and acute or cuneate base, slightly decurrent on
petiole, drying chartaceous, dark yellowish green to dark
brown above, distinctly paler beneath, glabrous except
for short (0.2-0.3 mm) stiff hairs along the sides of the
midvein above and below (sometimes with thin hairs
along the secondary veins beneath), 2 veins 7-1 I/side,
arcuate-ascending near the margin. Inflorescences ter-
minal or pseudoaxillary and becoming verticilate, sessile
or subsessile, 1-3.5 cm long, to 3.5 cm broad, a compact
globose irregularly branched panicle, peduncle 1-7 mm
long, bracts to 10 mm long, bracteoles ca. 1 mm long
and digitate-glandular, pedicels to 6 mm long in fruit.
Flowers glabrous externally, distylous, hypanthium ca.
0.7 mm long, calyx lobes 5, ca. 1 mm long; corolla fun-
nelform, white, tube 4-5 mm long, 1.3 mm diam., lobes
reflexed at anthesis, ca. 2 mm long. Fruits 4-5(-10?) mm
long, 2.5-3.5 mm diam., obovoid or turbinate and trun-
cated distally, dark blue or purple, persisting calyx ca. 1
mm long; pyrenes 3-4 mm long, with 4-5 longitudinal
ribs.

Plants of lowland rain forest formations from
20 to 300(-600) m elevation, often found in light
gaps and stream sides. Flowering and fruiting
throughout the year with most collections from
January-March. The species ranges from north-
eastern Costa Rica to Colombia.

Psychotria cooperi is recognized by its small con-
gested axillary inflorescences (sometimes appear-
ing verticillate), the broad short-tubular stipules,
leaves often drying yellowish green, small white
flowers, and blue or purple obovoid fruit. The
flowers are mostly sessile in terminal inflores-
cences but become axillary as the stems continue
to grow. These small treelets are called cocobolito
in Bocas del Toro, Panama. Fruiting material may
resemble Palicourea copensis.

Psychotria correae (Dwyer & Hay den) C. M. Tay-
lor, comb. nov. Cephaelis correae Dwyer &
Hayden, Ann. Missouri Bot. Gard. 55: 36. 1 968.
Figure 17.

Shrubs or small trees, 2-3.5(-5) m tall, leafy stems 3-
6 mm thick, glabrous, quadrangular; stipules with a very
short (1-3 mm) sheath with 2 broadly rounded often
overlapping lobes ca. 4 mm long and 4 mm broad sep-
arated by a U-shaped sinus, glabrous. Leaves with pet-
ioles 1 5-80 mm long (opposing petioles often unequal),
1.2-2.5 mm thick, glabrous; leaf blades 1 1-28 cm long,
5-13 cm broad, broadly elliptic-oblong, ovate-oblong,
apex abruptly narrowed and short-acuminate with tip
ca. 4 mm long, base broadly obtuse, drying chartaceous,
green, glabrous above and below, 2 veins 13-17(-20)/
side. Inflorescences solitary and terminal or pseudoax-
illary, capitulae 1-2 cm long, 2-3 cm diam., involucre
5-10 cm wide, peduncles (3-)5-12(-22) cm long, ca. 2
mm thick, glabrous, erect and becoming pendant, the 2
large involucral bracts 2.5-5 cm long and 4-5 cm broad,
ovate to reniform, rose red to magenta or purple, gla-
brous, bracteoles 8-15 mm long, obovate. Flowers gla-
brous externally, calyx lobes 5, ca. 1 mm long; corolla
narrowly funnelform, white, tube to 10 mm long, cylin-
drical, lobes 5, ca. 3 mm long; anthers ca. 3 mm long.
Fruits 10-15 mm long, ca. 6 mm diam., obovoid-oblong,
blue; pyrenes with 4-5 slightly elevated ridges

Plants of evergreen cloud forest formations along
the central Cordilleras at 900-1100 m elevation.
Flowering in March-August; fruiting in October.
This species is only known from the Cordilleras
de Guanacaste and Tilaran and the Province of
Code in Panama.

Psychotria correae is recognized by the larger
leaves with many secondary veins, the unusual
stipules, the long-pedunculate inflorescences be-
coming pendant in later stages, and the large pur-
plish bracts subtending the capitulum. Specimens
of this species can be confused with P. elata, which
has erect capitulae with smaller involucral bracts
and shorter peduncles, and to P. chiriquiensis,
which lacks involucral bracts. This attractive spe-
cies was named in honor of our respected Pana-
manian colleague Mireya Correa.

Psychotria deflexa DC., Prodr. 4: 510. 1830. P.
patens auct. non Sw. fide Steyermark 1972. Fig-
ure 56.

Shrubs, (0.5-)l-3 m tall, leafy stems 0.5-5 mm thick,
glabrous or sparsely puberulent in early stages; stipules
united around stem for 0.5-2 mm and with 2 subulate
or linear awns 3-8 long on each side, glabrous, persisting.
Leaves with petioles 3-9(-12) mm long, 0.5-1.3 mm
thick, glabrous; leaf blades 7-15(-18) cm long, (1.5-)2-

244

FIELDIANA: BOTANY

5(-7) cm broad, narrowly ovate-elliptic to narrowly el-
liptic, elliptic-lanceolate or rarely lanceolate, apex ta-
pering gradually and acute or acuminate, tip 10-20 mm
long, base broadly obtuse to acute, drying chartaceous,
greenish or brown, glabrous above and below, 2 veins
4-8/side. Inflorescences terminal (rarely pseudoaxillary)
and solitary, 4-1 1 cm long and 2-4 cm broad, narrowly
pyramidal or thyrsiform panicles with slender central
rachis and short (5-14 mm) opposite or subopposite
lateral branches (expanding somewhat in fruit), pedun-
cles 2.5-5 cm long, 0.4-1.1 mm thick, glabrous or mi-
nutely puberulent, lateral branches without subtending
bracts, pedicels 1-2 mm long, bracteoles ca. 1 mm long.
Flowers glabrous externally, hypanthium ca. 1 mm long,
calyx ca. 0.5 mm long, dentate; corolla white, funnel-
form, tube 1.6-3 mm long, 0.3-1 mm diam. (in Costa
Rica), lobes 4-5, ca. 1 mm long; stamens 4, anthers ca.
1 mm long. Fruits 2-3 mm long and 3-4 mm diam. (8-
10 mm when aerenchymatous), oblate and somewhat
bilobed, longitudinally ridged, becoming purple, violet,
or blue (pulpy white within); pyrenes with 3-4 ridges and
transverse ribs.

sheath, with 2 short (1-2 mm) obtuse or rounded lobes
separated by a U-shaped sinus, glabrous. Leaves with
petioles 4-25 mm long, 0.7-1 .2 thick, glabrous; leaf blades
4-1 1(-13) cm long, 1.8-4.3 cm broad, narrowly elliptic
to lanceolate, apex tapering gradually and acuminate, tip
5-10 mm long, base acute and slightly decurrent on pet-
iole, drying subcoriaceous, dark brownish green above,
glabrous above and below, 2 veins (6-)8-12/side. Inflo-
rescences 1-3 and terminal, 3-10 cm long, 3-8 cm wide,
usually subtended by a pair of smaller (15 mm) leaves,
capitula ca. 1 5-25 mm long and equally broad (enlarging
in fruit), subtended by 2 involucrate bracts 12-20 mm
long and 8-15 mm broad, white becoming pinkish (dark
purple in fruit), peduncles 3-9 cm long and 1.5-3.5 mm
thick, glabrous, flowers sessile within bracteoles. Flowers
glabrous externally, hypanthium ca. 1.5 mm long, calyx
ca. 3 mm long, lobes ca. 1.5 mm long and triangular;
corolla funnelform, white to pink, tube 5-10 mm long,
ca. 2 mm diam., corolla lobes 2-3 mm long, bluntly
acute. Fruits 8-9 mm long (including the 1 mm high
calyx), 4 mm diam., narrowly obovoid, blue-black, dry-
ing dark reddish brown; pyrenes usually smooth.

Understory plants of wet evergreen or partly
deciduous forest formations from ca. 500 to 1200
m elevation on the Caribbean slope of the cordil-
leras and near sea level on the Osa Peninsula.
Flowering in June-August; fruiting in July-March.
This species ranges from Mexico to Peru and Bo-
livia.

Psychotria deflexa is recognized by its persisting
narrowly awned stipules, narrow leaves, general
lack of pubescence (only the slender narrow inflo-
rescence is minutely puberulent), small flowers,
and purple or blue fruit with transverse ribs. Flow-
ers in Costa Rican material appear to be much
smaller than those described by Steyermark (1974,
p. 1284). This species may resemble P. cincta and
P. microbotrys. There are collections (Kernan &
Phillips 661 & 789 CR) from Corcovado National
Park that have unusually thin pedicels and inflo-
rescences. However, these aberrant samples are
bridged by other collections (Liesner 2842 & 3198
CR, also from the Osa Peninsula) that appear to
be intermediate with the more normal inflores-
cences. These populations are worthy of further
study.

Psychotria dichroa (Standl.) C. M. Taylor, comb,
nov. Evea dichroa Standl., Contr. U.S. Natl.
Herb. 18: 124. 1916. Cephaelis dichroa (Standl.)
Standl. Publ. Field Columb. Mus., Bot. Ser. 4:
296. 1929.

Small shrubs, 0.6-2 m tall, leafy stems 1.3-4 mm thick,
glabrous; stipules 2.5-6 mm long, with a short (2-3 mm)

Plants of evergreen montane forest formations,
1 200-2600 m elevation. Flowering January-Sep-
tember. This species is only known from the Chi-
riqui Highlands of western Panama and the Ca-
ribbean slope of the Cordillera de Talamanca.

Psychotria dichroa is recognized by the leaves
with prominent 2 veins, one to three terminal
small involucrate heads, white flowers, and high-
land habitat. In Costa Rica, it is rarely collected
and appears to be restricted to the understory of
high-elevation (2000-2500 m) Quercus forests.
This species is similar to P. chiriquiensis with red-
dish purple capitula.

Psychotria domingensisJacq.,Enum. PI. 16. 1760.
Psychotria pavetta Sw., Prodr. 45. 1788. Pali-
courea domingensis (Jacq.) DC., Prodr. 4: 529.
1 830. Palicourea pavetta (Sw.) DC., loc. cit. 525.
1830. Psychotria mombachensis Standl., Publ.
Field Columb. Mus., Bot. Ser. 8: 188. 1930.

Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla-
brous, often with a shrunken area below the node after
drying; stipules 2-5 mm long, with 2 triangular lobes
separate almost to base by a broad (1-2 mm) U-shaped
sinus, acute or awned, glabrous, deciduous or persisting.
Leaves with petioles 5-20 mm long, 0.7-1.7 mm thick,
glabrous, often drying yellowish; leaf blades 8-19 cm
long, 3-7 cm broad, elliptic-oblong to narrowly elliptic-
obovate, apex acuminate or gradually tapering and acute,
base acute to obtuse and slightly decurrent on petiole,
drying membranaceous to thin-chartaceous, yellowish
green to greenish brown or brown, glabrous above and
below, 2 veins 8-1 I/side. Inflorescences solitary or 3,
terminal or pseudoaxillary, 4-9 cm long, 3-6 cm broad,

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

245

a rounded open panicle with opposite branching, pe-
duncle 3-20 mm long, ca. 1.5 mm thick and yellowish
when dried, glabrous or minutely papillate-puberulent,
bracts absent or 1-6 mm long and linear, flowers in distal
cymules of 3, pedicels 0-3 mm long, ca. 1 mm thick,
bracteoles minute or absent. Flowers glabrous or mi-
nutely puberulent externally, hypanthium ca. 1.2 mm
long, calyx tube ca. 0.5 mm long, lobes 0.2-1 mm long,
narrowly acute; corolla white, tubular-salverform, tube
ca. 12 mm long, 0.8-1.7 mm diam., lobes ca. 4-5 mm
long and 2 mm broad. Fruits 4-5 mm long, ca. 6 mm
diam., subglobose to ellipsoid, purple-black, with 4-5
longitudinal costae and smaller transverse ribs when
dried; pyrenes angled.

Plants of evergreen or seasonally deciduous for-
mations, from 20 to 900 m elevation. Flowering
in April and June (the Nicaraguan type of P. mom-
bachensis (Maxon et al. 7818 F) was flowering in
July); a Costa Rican collection from near Tilaran
(Standley & Valeria 45222 us) was fruiting in Jan-
uary. The species is frequent in the West Indies
and Guatemala-southern Mexico but is known
from only a few collections in southern Central
America.

Psychotria domingensis is recognized by the in-
florescences with thick branches and relatively few
flowers, larger fruit with longitudinal and trans-
verse ridges, thin glabrous leaves that often dry
yellowish green, two-lobed stipules, and dried stems
often conspicuously contracted beneath the node.
The few collections from Costa Rica come from
both evergreen and deciduous areas. This species
is rather similar to P. eurycarpa of wet forests, but
that species has stiffer leaves lustrous above, poor-
ly developed calyx lobes, and larger fruit without
transverse ribs. It is also similar to P. microdon
and some species of Coussarea (see fig. 47).

Psychotria dukei Dwyer, Ann. Missouri Bot. Gard.
67: 371. 1980.

Shrubs or herbaceous subshrubs, 0.5-1. 5(-2) m tall,
leafy stems succulent in life, 1.8-7 mm thick (dried),
glabrous; stipules 7-20 mm long, 8-1 8 mm broad, ovate-
triangular, bilobed or fimbriate, yellowish and membra-
naceous, lobes 1-4 mm long, deciduous. Leaves with
petioles 1-5 cm long, 1-2.3 mm broad, glabrous; leaf
blades (1 1-) 13-27 cm long, 4-1 1 cm broad, elliptic-ob-
long to oblong or elliptic-ovate, apex short-acuminate
with tip 5-10 mm long, base obtuse to cuneate and de-
current on petiole, drying chartaceous, dark green or
greenish brown, glabrous above, minutely (0. 1 mm) pu-
berulent along the major veins beneath, 2 veins 10-18/
side. Inflorescences axillary, solitary at each node, 2.5-
9 cm long, pyramidal, often with the flowers crowded in
3 capitula on 3 primary branches 3-20 mm long (rarely
with further secondary branches), peduncles 8-35(-60)

mm long, ca. 1.2 mm thick, minutely puberulent or gla-
brous, bracts 2-10 mm long, triangular to lanceolate,
bracteoles 2-5 mm long, often forming a small involucre
beneath the sessile flowers. Flowers glabrous externally,
hypanthium ca. 1 mm long, calyx tube ca. 0.5 mm long,
entire; corolla tubular-funnelform, white, tube 3-4 mm
long, ca. 0.5 mm diam. basally, lobes ca. 1.5 mm long.
Fruits 5-6 mm long, 3-4 mm diam., oblong, white; py-
renes longitudinally ridged.

Plants of lower montane rain forest formations
from (800-)1 100 to 1800 m elevation. Flowering
in March and June-August; fruiting in March-
November. The species ranges from the Cordillera
de Tilaran to Choco, Colombia.

Psychotria dukei is recognized by its usually un-
branched succulent stems, axillary inflorescences
with flowers in capitate clusters, white fruit, and
unusual stipules. This species is easily confused
with P. aggregata or P. macrophylla, but the large
thin stipules and inflorescence bracts are distinc-
tive. This species is rarely collected in Costa Rica.
Compare P. dukei with P. wilburiana Dwyer of
Panama.

Psychotria elata (Sw.) Hammel, Selbyana 1 2: 1 39.
1991. Cephaelis elata Sw., Prodr. 45. 1788. C.
punicea Vahl, Eclog. Amer. 1:19. 1796. C. cos-
taricensis Schlechtend., Linnaea 28: 546. 1856.
Evea elata (Sw.) Standl., Contr. U.S. Natl. Herb.
18: 123. 1916. C. elata forma lutea Standl., Publ.
Field Mus. Nat. Hist., Bot. Ser. 18: 1279. 1938.
Figure 17.

Shrubs, subshrubs, or small trees, 0.5-5(-8) m tall,
leafy stems 1.3-5 mm thick, glabrous, terete or qua-
drangular; stipules united around stem for 1-2 mm, with
2 ovate lobes on each side, 2-5 mm long and ca. 3 mm
broad, obtuse to broadly rounded and separated by a U-
or V-shaped sinus, glabrous, persisting. Leaves with pet-
ioles 4-22(-30) mm long, 1-2 mm wide, glabrous; leaf
blades 6-25 cm long, 2.5-7(-8.5) cm broad, oblong to
elliptic-oblong, oblong-obovate, or oblanceolate, apex
acute to short-acuminate, tip 4-10 mm long, base cu-
neate to acute or obtuse, drying thinly to stiffly charta-
ceous, green, glabrous above and below, 2 veins (9-)l 3-
20(-23)/side. Inflorescences terminal and solitary (rarely
2-3), capitate and globose-hemispherical, capitula 1.5-
4 cm long, involucre 2-7 cm broad, peduncles 2-13 cm
long, 1.5-3 mm thick, erect or becoming pendant, gla-
brous, the 2 involucrate basal bracts 15-55 mm long,
1 5-45 mm broad, deep red to orange-red (yellow), ovate
to reniform, rounded to acuminate, glabrous, bracteoles
5-10 mm long, flowers sessile. Flowers glabrous exter-
nally (rarely puberulent), calyx ca. 1 mm long, dentate;
corolla white (rarely pink or yellow), narrowly funnel-
form, tube ca. 16 mm long, 1.5-2.5 mm diam., lobes 5,
2.5-4 mm long, narrowly triangular; stamens 5, anthers
2-3 mm long. Fruits 5-10 mm long, 2-5 mm diam.,

246

FIELDIANA: BOTANY

ovoid, blue becoming black; pyrenes 4-7 mm long and
3-4 mm broad, 5-ribbed.

Common shrubs of evergreen forest edges and
forest interiors, from 30 to 1700 m elevation.
Flowering and fruiting throughout the year (pri-
marily flowering in January-August). Fruiting at
La Selva in July-September. The species ranges
from Mexico and the West Indies, through Central
America to Colombia.

Psychotria elata is recognized by its glabrous
parts, usually narrowly oblong leaves with many
secondary veins, solitary long-pedunculate heads,
large brilliant red (rarely yellow) bracts subtending
the capitulum, and white corollas. The yellow-
bracted form appears to be most common in the
General Valley and Golfo Dulce area. This is one
of Central America's most distinctive species of
Rubiaceae, common in light gaps and on the edges
of forests. The flowers are visited by butterflies
and hummingbirds (Freeman & Stiles, 1990);
breeding biology was studied by Bawa and Beach
(1983). This is one of the species that made Ce-
phaelis so distinctive a genus in Central America.
Compare P. poeppigiana with conspicuous pubes-
cence, P. chiriquiensis with purple capitula lacking
the large involucral bracts, P. correae with longer
peduncles and larger bracts, and P. dichroa with
several smaller heads.

Psychotria emetica L.f, Suppl. PI. 144. 1781. Fig-
ure 13.

Small herbs and subshrubs, 0.2-1 m tall, rhizomatous,
stems usually unbranched, terete, leafy internodes 1-4
mm thick, appressed-puberulent with stiff crooked
brownish hairs 0.2-0.4 mm long; stipules 2-4 mm long,
1-2 mm broad at base, narrowly triangular and ap-
pressed-puberulent, drying dark, usually caducous. Leaves
with petioles 3-12(-20) mm long, 1-2 mm broad, ap-
pressed-puberulent with stiff brownish hairs; leaf blades
(7-)8.5-13(-17) cm long, 2-5(-6.5) cm broad, elliptic to
narrowly elliptic-oblong, elliptic-obovate or oblanceo-
late, apex acute or short-acuminate, tip ca. 5 mm long,
base cuneate to acute and slightly decurrent on petiole,
drying thinly chartaceous, dark grayish green or dark
grayish brown, glabrous above, sparsely appressed-pu-
berulent with short (0. 1-0.3 mm) hairs beneath, 2 veins
5-9/side. Inflorescences axillary, usually solitary in each
axil (2/node), 1-2 cm long, condensed cymose or sub-
capitate, with 3-10 flowers, peduncles 3-12 mm long,
ca. 0.7 mm thick, puberulent, bracts 1-2 mm long, tri-
angular-acute, pedicels 0-5 mm long. Flowers with hy-
panthium ca. 1 mm long, conical and appressed-puber-
ulent, calyx lobes 1-1.5 mm long, narrowly triangular;
corolla white, funnelform, glabrous externally, tube 2-4
mm long, 1-1.5 mm diam. at mouth, lobes 5, 1.5-2 mm
long, acute. Fruits 8-10 mm long and 4-6 mm diam.,

oblong or ellipsoid, persisting calyx 1-1.5 mm long, be-
coming blue; pyrenes ca. 5 mm long, smooth.

Plants of shaded understory in lowland ever-
green rain forest formations, from 10 to 300 m
elevation. In Costa Rica and Panama flowering in
February-August; fruiting mostly in June-No-
vember. This species ranges from Guatemala to
Bolivia.

Psychotria emetica is recognized by its short,
usually unbranched stems, small condensed axil-
lary inflorescences, and bright blue fruit. These
plants are occasionally cultivated and have been
called raicilla and raicilla macho in Costa Rica,
Nicaragua, and Panama. This species differs from
the closely similar P. erecta by its smaller stature,
puberulent stems, and floral details. The roots con-
tain the drug ipecac but it is of inferior quality to
that found in P. ipecacuanha.

Psychotria erecta (Aubl.) Standl. & Steyerm., Publ.
Field Mus. Nat. Hist., Bot. Ser. 23: 24. 1943.
Ronabea erecta Aubl., Hist. pi. Guiane 1: 156.
1775. Figure 13.

Shrubs or treelets, (0.5-)l-3(-8) m tall, leafy stems
1 .5-7 mm thick, sparsely appressed-puberulent with thin
ascending hairs ca. 0.3 mm long, becoming glabrescent
and drying dark; stipules with a persistent short (0.5 mm)
base and single slender awn 2-6 mm long, ca. 0.5 mm
wide, triangular to subulate, appressed-puberulent. Leaves
with petioles 8-18(-35) mm long, 0.7-2(-2.5) mm thick,
appressed-puberulent with hairs 0.2-0.3 mm long; leaf
blades 8-20 cm long, 3-9 cm broad, elliptic-oblong to
oblong or slightly obovate, apex abruptly narrowed and
short-acuminate, tip 4-1 1 mm long, base obtuse to acute,
drying chartaceous to subcoriaceous, dark olive green or
brownish green, glabrous above, glabrous or sparsely pu-
berulent beneath with thin appressed hairs, 2 veins 5-
8/side. Inflorescences 1-3 in each axil (2-6/node), 1-3
cm long, 5-10 mm broad, subcapitate condensed cymose
with 3-7 closely crowded flowers, peduncles 0-8(-18)
mm long, ca. 0.7 mm thick, appressed-puberulent with
ascending yellowish hairs, flowers subsessile or sessile,
bracteoles to 1 mm long. Flowers with hypanthium 1-2
mm long, sparsely appressed puberulent near the base,
calyx ca. 1 mm long, glabrous, lobes ca. 0.2 mm high;
corolla white, tubular-salverform, glabrous, tube 3^ mm
long, 1 mm diam., glabrous externally, lobes 5(-6), 1.5-
3 mm long. Fruits 8-10 mm long, 5-8 mm diam., ellip-
soid or oblong, glabrous, becoming blue-black; pyrenes
5-7 mm long, smooth.

Plants of evergreen rain forest formations, from
near sea level to 700(-1000) m elevation on both
the Caribbean and Pacific coasts. At La Selva this
species is usually found on ridges near light gaps.
Flowering in May-October; probably fruiting

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

247

throughout the year. This species ranges from
Mexico and the West Indies to Bolivia.

Psychotria erecta is recognized by its axillary
small subcapitate inflorescences with few small
flowers, blue fruit, and oblong leaves often drying
dark. This species is similar to P. emetica but grows
to a larger size with broader and thicker leaves and
glabrescent stems. Both species have axillary, not
pseudoaxillary, inflorescences. Compare Appunia
guatemalensis.

Psychotria eurycarpa Standl., J. Wash. Acad. Sci.
18: 275. 1928. Figure 47.

Shrubs or small trees, 1.5-5(-7) m tall, leafy stems 1-
3 mm thick, glabrous or rarely minutely papillate-pu-
berulent; stipules 1-3 mm high and 3-4 mm broad, trun-
cate to broadly rounded with a small (0.2 mm deep)
sinus, persisting or deciduous. Leaves with petioles 12-
28(-35) mm long, 0.7-1.5 mm thick, glabrous and often
drying yellowish; leaf blades 6-14(-21) cm long, 2.7-
7(-l 1) cm broad, ovate-elliptic to elliptic or broadly el-
liptic, apex short-acuminate with a narrowed tip 3-10
mm long, base obtuse to acute, drying stiffly chartaceous,
yellowish green, glabrous and lustrous above, glabrous
beneath, 2 veins 4-7/side. Inflorescences terminal and
solitary, 5-1 1 cm long, 5-9 cm broad, broadly pyramidal
panicles with opposite branching or umbellate, the first
node often with 4 lateral branches, peduncles 1 5-55 mm
long, 1-2 mm thick, yellowish, glabrous, bracts ca. 1 mm
long, flowers sessile or subsessile in glomerules of 2-5.
Flowers nocturnal, glabrous or very minutely papillate-
puberulent externally, hypanthium ca. 1 mm long and
0.6 mm diam., obconic, calyx tube 0.3-0.7 mm high
with minute (0.2 mm) lobes; corolla narrowly funnel-
form, white, tube 6-1 5 mm long and 0.8 mm diam. (1.7
mm distally), lobes 5 (6, 7), 3-8 mm long, 0.5-1 mm
broad, narrowly oblong. Fruits 8-20 mm long (to 25 mm
when succulent), 7-1 5 mm diam., oblong to subglobose,
blue-black or dark purple (often drying pale yellowish
brown), longitudinal ridges present but not prominent
when dried; pyrenes 12-17 mm long, obscurely 3-5-
angled.

Plants of lowland rain forest formations and
lower montane cloud forest formations, from 20
to 900(-1100) m elevation. Flowering in Febru-
ary-May; fruiting in October-April and June. This
species is only known from the Caribbean coastal
plain, Cordillera de Guanacaste, Cordillera de Ti-
laran, and the Caribbean slopes of the Central
Highlands as far east as Rio Reventazon, and in
western Panama.

Psychotria eurycarpa is recognized by the usu-
ally glabrous parts often drying yellowish green,
the short stipules (truncated or with minute apical
sinus), few-branched inflorescences with thick pe-
duncles and thick lateral branches and inconspic-
uous bracts, longer corolla tubes, and the large

fruit. Breeding biology was studied by Bawa and
Beach ( 1 983). A collection (Taylor 3546) from Las
Cruces (San Vito) is tentatively placed here; it is
the highest elevation ( 1 1 00 m) and easternmost
collection in Costa Rica. Folsom 9188 DUKE, from
La Selva with corolla tubes to 18 mm long and
corolla lobes to 1 2 mm long, is provisionally placed
here. Individuals of this species may be difficult
to distinguish from P. domingensis (with bilobed
stipules and flowers in distal triads) and Coussarea
psychotrioides (q.v.).

Psychotria fruticetorum Standl., J. Arnold Arbor.
11:42. 1930. Figure 60.

Shrubs, subshrubs, or small treelets, 0.3-3(-5) m tall,
leafy stems 0.7-3.5 mm thick, glabrous; stipules 24 mm
long, 1-3.5 mm broad at base, triangular to ovate, trun-
cate to acute, with 2 awns 1-1.5 mm long, glabrous or
minutely (0.05 mm) papillate, drying reddish brown,
caducous. Leaves with petioles 0-6(-8) mm long, 0.7-
1.2 mm thick, glabrous; leaf blades 3-8.5(-l 1?) cm long,
1 4(-5?) cm broad, elliptic to obovate or oblanceolate,
apex acute to subacuminate, base cuneate to acute, dry-
ing stiffly chartaceous to subcoriaceous, grayish to dark
reddish brown, glabrous above and below, 2 veins 4
7/side, often with minute pit domatia or tufts of hairs
in the axils of 2 veins beneath. Inflorescences terminal
and solitary, 3-4.5 cm long, 3-4 cm wide, rounded pan-
icles with opposite or whorled branches, peduncles 5-
20 mm long, 0.5-0.8 mm thick, glabrous, bracts ca. 1
mm long, flowers subsessile in distal cymes, pedicels 0.5-
1 mm long. Flowers glabrous externally, hypanthium ca.
0.8 mm long, obconic, calyx lobes ca. 0.5 mm long;
corolla funnelform to salverform, white, tube ca. 2 mm
long, 1-1.2 mm diam., lobes 5, ca. 1.5 mm long and 0.7
mm broad at the base; stamens 5, anthers ca. 0.8 mm
long. Fruits 4-5 mm long, 3-4 mm diam., ellipsoid with
ca. 10 longitudinal ridges, bright red; pyrenes ridged.

Plants of wet evergreen lowland forest forma-
tions (and in open pine savannas in Nicaragua),
from near sea level to 200 m elevation (to over
1 300 m in Mexico and Guatemala). Flowering in
southern Central America in March-May and Sep-
tember; fruiting in July-December. The species
ranges from Mexico, mostly along the Caribbean,
to Panama but is disjunct from Nicaragua to cen-
tral Panama.

Psychotria fruticetorum is recognized by the
small, often cuneate and subsessile leaves, cadu-
cous stipules with two distal awns, general lack of
pubescence, small domatia, small inflorescences,
and small flowers and fruit. The tendency of the
leaves to dry grayish or reddish and the bright red
berries are characteristics of subgenus Psychotria.
Presently, this species is not known from Costa
Rica, though it grows near Bluefields, Nicaragua,

248

FIELDIANA: BOTANY

and in central Panama. It may be restricted to
limestone soils.

Psychotria glomerulata (J. D. Smith) Steyerm.,
Mem. New York Bot. Gard. 23: 670. 1972. Ce-
phaelis glomerulata J. D. Smith, Bot. Gaz. 16:
12, pi. 1. 1891. Figure 18.

Small shrubs or treelets, 0.6-2(-4?) m tall, much-
branched, leafy stems 1-5 mm thick, quadrangular, gla-
brous; stipules with a short (0.5-2 mm) sheath, truncated
and entire at apex, glabrous, persisting. Leaves with pet-
ioles 3-1 1(-15) mm long, 0.8-1.3 mm thick, glabrous,
yellowish green; leaf blades 5-14(-17) cm long, 1.5-
5(-6.5) cm broad, narrowly oblong to narrowly elliptic-
oblong, oblong or lanceolate, apex acuminate with tip
4-1 2 mm long, base obtuse to acute or cuneate and slightly
decurrent on petiole, drying chartaceous to stiffly char-
taceous, glabrous above and below, 2 veins 9-13/side
and loop-connected near the the margin. Inflorescences
terminal and solitary, capitula 1.2-2.5 cm long, 1.5-3.5
cm broad, oblate to subglobose, subtended by 4 decus-
sate involucrate bracts 10-16 mm long and 1 2 mm broad,
oblong to broadly obovate and rounded distally, greenish
yellow or whitish and sometimes edged with blue or
purple, glabrous, peduncle to 5(-15) mm long, flowers
tightly enclosed within the bracts and spatulate bracte-
oles. Flowers glabrous externally, hypanthium ca. 1 mm
long, calyx lobes 5, 0.5-1 .5 mm long; corolla funnelform,
white, tube 10-15 mm long, 1-2 mm diam.. lobes 5,
1.5-3 mm long; stamens 5, anthers 2-3 mm long. Fruits
6-13 mm long and 5-10 mm diam., ellipsoid, bright
blue; pyrenes 4-6 mm long and 3-4 mm diam., appar-
ently smooth.

Understory shrubs of evergreen rain forest for-
mations, from near sea level to 600 m elevation
along the Caribbean slope and coastal plain. Prob-
ably flowering and fruiting throughout the year but
flowering mostly In March-April and August-Oc-
tober. This species ranges from southern Mexico
and Belize along the Caribbean lowlands to Pan-
ama.

Psychotria glomerulata is recognized by its short
stature, narrow leaves, lack of pubescence, solitary
sessile terminal heads with four (to eight) white or
yellow-green involucrate bracts, and lowland Ca-
ribbean habitat. Our figure is based on the original
illustration. This species is similar to P. guapilensis
with long stipule lobes; P. apoda Steyerm. of South
America may be synonymous.

Shrubs or small treelets, 1.5-3(-5) m tall, leafy stems
0.6-4 mm thick, glabrous, flattened or quadrangular at
first, soon terete and with thickened nodes; stipules 1-3
mm long, 1.5-2.5 mm broad, with 2 acute lobes and
distal sinus 0.3-2 mm deep, glabrous, deciduous or per-
sisting. Leaves with petioles 3-22 mm long, 0.3-1 mm
thick, glabrous; leaf blades (2-)3-10(-13) cm long,
(0.8-)l-2.5(-3) cm broad, narrowly elliptic-oblong to
lanceolate or linear-lanceolate, apex long-acuminate with
tip 7-15 mm long, base gradually narrowed or abruptly
acute and slightly decurrent on petiole, drying charta-
ceous, dark green or brown above, glabrous above and
below, 2 vein 8-1 3/side, often with parallel intermediate
2 veins. Inflorescences terminal and solitary (pseu-
doaxillary), 4-14 cm long, to 1 1 cm broad, open pyra-
midal panicles with distant slender opposite or subop-
posite branches, peduncles 15-52 mm long, 0.7-1.2 mm
thick and glabrous, bracts 3-10 mm long and linear to
triangular (or leaf-like), distal branching cymose (often
minutely puberulent), flowers usually sessile in triads.
Flowers glabrous on the exterior, hypanthium ca. 1 mm
long, obconic, calyx tube ca. 1 mm long, subentire or
with 5 acute lobes to 0.5 mm long; corolla funnelform,
greenish white, tube 4-6 mm long, 0.6-1 mm diam.,
lobes 5, 1-2 mm long. Fruits 3-5 mm long, 2.5-4 mm
diam., ellipsoid-oblong, blue, purple, or white, persisting
calyx 0.3-0.7 mm long; pyrenes 3-5-angled or -ridged.

Plants of evergreen montane forest formations,
from 1000 to 2000 m elevation. Flowering in
March, April, and October; fruiting in January and
March-August. The species ranges from the Cor-
dillera de Guanacaste eastward along the high-
lands to Darien, Panama.

Psychotria goldmanii is recognized by its higher-
elevation habitat, smaller narrow long-acuminate
glabrous leaves, calyx with minute lobes, longer
corolla tube, and small blue fruit. The leaves often
dry with the lower surfaces markedly paler in color
and obscure 3 venation. The stipules with two
small lobes are also distinctive. Few other species
of Psychotria in southern Central America develop
such narrow linear-lanceolate leaves (an extreme
form of which is represented by the type of P.
torresiana, Standley 39769 us). Compare P. stey-
ermarkii with more numerous 2 veins and P. val-
eriana with one stipule appendage. Smaller-leaved
specimens of this species may be difficult to dis-
tinguish from Palicourea montivaga, but that spe-
cies has shorter leaves, very slender petioles, a
tendency to develop a short stipular tube, inflo-
rescences that dry yellowish, and corollas that are
swollen at the base.

Psychotria goldmanii Standl., Contr. U.S. Natl.
Herb. 18: 130. 1916. P. torresiana Standl., J.
Wash. Acad. Sci. 15: 288. 1925. P. eugenifolia
Dwyer, Ann. Missouri Bot. Gard. 67: 375. 1980.
Figure 55.

Psychotria graciliflora Benth. in Oerst, Vidensk.
Meddel. Dansk. Naturhist. Foren. Kjobenhavn
1852: 35. 1853. P. vallensis Dwyer, Ann. Mis-
souri Bot. Gard. 67: 438. 1980 (fide Hamilton
1989). Figure 60.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

249

Shrubs, 0.6-2(-3) mm tall, much-branched and often
flat-topped, leafy stems 0.7-2(-3) mm thick, glabrous
(except for reddish colleters at the node), usually with 2
(opposite) longitudinal ridges between nodes; stipules
with a narrowly triangular to ovate base 1-2 mm long
and 2 awns 1-2 mm long, with ascending hairs 0.1-0.2
mm long, reddish brown, deciduous. Leaves mostly clus-
tered near the ends of branchlets, petioles 2-10 mm long,
0.5-1 mm broad, glabrous; leaf blades 1 .5-6(-8) cm long,
0.7-3 cm broad, elliptic to ovate-elliptic, apex bluntly
obtuse to acute or subacuminate, base cuneate to acute
and decurrent on petiole, drying chartaceous, dark gray
or reddish gray above, glabrous above and below, 2
veins 3-6/side, pit domatia rarely present. Inflorescences
terminal and solitary, 2-6 cm long, 2-5 cm broad, py-
ramidal rounded panicles with 1-2 pairs of opposite
branches or umbellate, peduncles l-2(-3.5) cm long, ca.
0.5 mm thick, glabrous and drying dark, bracts and brac-
teoles to 1 mm long, flowers often sessile in distal cymes
or triads. Flowers glabrous externally, hypanthium ca.
0.7 mm long, obconic, calyx tube ca. 0.5 mm long, lobes
0.2-1 mm high; corolla white, salverform, tube 2.5-3.5
mm long, 0.5-0.8 mm diam., lobes 5, 1-2 mm long;
stamens 5, anthers ca. 0.9 mm long. Fruits 4-6 mm long,
3-5 mm diam., globose to ellipsoid, with ca. 10 longi-
tudinal ridges, bright red or orange; pyrenes ca. 4 mm
diam.

Understory plants of wet evergreen forest for-
mations on both the Caribbean and Pacific slopes,
from near sea level to 1700 elevation (to 2500 m
elsewhere, and not found below 1000 m on the
Pacific slope of Costa Rica). Flowering in Febru-
ary-August (mostly in March-May); fruiting
throughout the year. The species ranges from
southern Mexico to Colombia.

Psychotria graciliflora is recognized by its very
small leaves, usually horizontal glabrous branches,
caducous stipules with two slender pubescent awns,
small open inflorescences with slender peduncles,
and small flowers with minute calyx lobes. The
bright red fruit and tendency for leaves to dry
grayish or reddish are characteristics of subgenus
Psychotria. Costa Rican material of this species
has smaller leaves with fewer secondary veins than
material from Honduras and Nicaragua (Hamil-
ton, 1989, cited under the genus). This species
resembles P. carthagenensis, P. chagrensis, and P.
parvifolia (see fig. 60).

Psychotria grandis Sw., Prodr. 43. 1788. Figure
64.

Shrubs or small trees, (1.5-)4-8(-10) m tall, trunks to
20 cm diam., leafy stems 4-10 mm thick, terete, glabrous
or minutely papillate-puberulent in early stages; stipules
8-30 mm long, 5-20 mm broad, broadly ovate-trian-
gular with the margins often revolute, acute to acuminate

and often minutely (1 mm) 2-lobed, usually glabrous,
often inflated at the base, deciduous. Leaves with poorly
denned petioles 0.5-3.5 cm long, 1.3-3 mm broad, gla-
brous; leaf blades ( 1 1-) 1 8-40 cm long, (3-)6-l 6 cm broad,
obovate to elliptic-obovate or oblanceolate, apex obtuse
to short-acuminate with tip ca. 5 mm long, gradually
narrowed to the cuneate base and long-decurrent on pet-
iole, drying chartaceous to subcoriaceous, grayish green
to pinkish gray or dark grayish above, glabrous above
and below, 2 veins 12-16/side. Inflorescences terminal
or pseudoaxillary and solitary, 12-25(-30) cm long, 10-
18 cm broad, open pyramidal panicules with opposite
branching or with 4 branches (2 larger and 2 smaller) at
lower nodes, peduncles 11-18 cm long, 2-4 mm thick,
glabrous or minutely (0.05 mm) papillate-puberulent,
flowers 2-5 in distal cymules, bracts ca. 2 mm long and
triangular, pedicels 1-3 mm long, bracteoles ca. 0.5 mm
long. Flowers mostly glabrous externally, hypanthium
ca. 0.8 mm long, obconic, calyx tube ca. 0.6 mm long,
lobes 5, 0-0.3 mm high, triangular; corolla white, fun-
nelform, tube 2-4 mm long, 1.5-2 mm diam., lobes 5,
1.5-2 mm long and 1.1 mm broad at the base; stamens
5, anthers 0.8 mm long. Fruits 5-7 mm long and 5 mm
diam., broadly ellipsoid or subglobose, orange or red,
surfaces smooth and rounded; pyrenes ca. 5 mm long,
with 4-5 rounded dorsal ribs.

Trees of wet evergreen forest formations on both
the Caribbean and Pacific slopes of Costa Rica,
from near the seashore to 800 m elevation. Flow-
ering in February-July; fruiting mostly in July-
February. The species ranges from southern Mex-
ico and the Greater Antilles to northern South
America.

Psychotria grandis is recognized by its often larger
(tree) habit, large glabrous-obovate leaf blades de-
current on the petioles, large ovate-acuminate
stipules with reflexed lateral margins, large inflo-
rescences, and small flowers. The red fruit and
vegetative parts drying grayish or reddish are char-
acteristics of the subgenus Psychotria. Guatemalan
material is often puberulent and with larger inflo-
rescences; Nicaraguan collections have larger fruit.
This species is closely related to P. costavenia Gri-
seb., which ranges from Mexico and Cuba into
southern Nicaragua and is smaller in its parts.

Psychotria guadalupensis (DC.) Howard, J. Ar-
nold Arbor. 47: 139.1 966, sensu lato. Loranthus
guadalupensis DC., Prodr. 4: 294. 1830. Vis-
coides pendulum Jacq., Select. Stirp. Amer. 73,
pi. 51, f. 1. 1763. P. parasitica Sw., Prodr. 44.
1 788, nomen illeg. P. pendula (Jacq.) Urb., Symb.
Ant. 1: 445. 1900, non P. pendula Hooker f.,
1880. P. peperomiae Standl., Contr. U.S. Natl.
Herb. 18: 132. 1916. P. orchidearumSiandl.J.
Wash. Acad. Sci. 18: 276. 1928. Figure 60.

250

FIELDIANA: BOTANY

Small epiphytic shrubs, stems 0.2-1 m long, leafy stems
1-4 mm thick, glabrous, often slightly succulent; stipules
with a short (0.5-2.5 mm) usually membranaceous trans-
lucent sheath, breaking apart and becoming thickened
and whitish at the base. Leaves with petioles 1-6 mm
long, 0.7-1.2 mm thick, glabrous; leaf blades 12-40 mm
long, 7-22 mm broad, ovate or ovate-elliptic to obovate,
apex bluntly obtuse to rounded, base obtuse to acute and
slightly decurrent on petiole, drying stiffly chartaceous
to subcoriaceous, gray to dark gray above, paler beneath,
glabrous above and below, with 2-5 obscure 2 veins per
side arising at angles of 20-40 (veins rarely prominent).
Inflorescences terminal and solitary, 1.5-5 cm long,
equally broad, few-flowered cymes to pyramidal panicles
with opposite or 3 -partite branching, peduncles 3-30
mm long, 0.4-1 . 1 mm thick, glabrous and reddish in life
but often drying black, bracts 2-6 mm long, longer bracts
linear, flowers in distal cymes or triads, pedicels 0.5-5
mm long. Flowers glabrous externally, hypanthium 1-
1.2 mm long, 0.5-0.7 mm diam., calyx tube ca. 0.3 mm
long, calyx lobes 0.4-1 mm long, triangular and acute,
often reddish; corolla white or rose, salverform, tube
(4-)6-8 mm long, ca. 1 mm diam., lobes 4, 1.5-2 mm
long. Fruits 3-6 m long, 3-5 mm diam., globose to ob-
late, usually with 4 pyrenes, red or purple and finally
black.

Epiphytes of lower montane cloud forest for-
mations of the Caribbean slope and continental
divide, from (200-)1000 to 2300 m elevation.
Flowering in January-August; fruiting in Febru-
ary-September and December. The species ranges
from Mexico and the West Indies to the Guianas.

Psychotria guadalupensis is recognized by its
epiphytic habit, unusual stipular tube, thick leaves
often with obscure venation, variable inflores-
cences with slender peduncles, longer-tubed flow-
ers, and four-locular ovary. The leaves drying
grayish and red fruit are characters shared with
subgenus Psychotria. The differentiation between
this species and material placed under P. pithe-
cobia may be artificial, though it does separate a
large portion of the specimens effectively. Costa
Rican material placed here has longer corolla tubes,
and leaf secondary veins are less numerous and
less prominent when dried as in P. pithecobia, and
they arise from the midvein at a smaller angle. But
there do appear to be a few intermediates, and
paucity of collections makes it difficult to tell
whether or not the longer corolla tubes of P. guad-
alupensis are a consistent difference. (See Ho-
ward's discussion of variation in this species, J.
Arnold Arbor. 47: 139-142. 1966.)

The preceding description refers to Costa Rican
material (excluding specimens assigned to P. pi-
thecobia) and differs from that given by Steyer-
mark (1974). For example, corolla tubes are ca. 3
mm long in Venezuela (and in P. pithecobia), much

shorter than usually seen in this material. It seems
probable tbat the length of the corolla tube varies
as greatly (between different plants) as do so many
other characters of inflorescences and leaves in this
complex.

The type of P. orchidearum Standl. (Standley &
Valeria 50863 us) has rather small (2 cm long)
leaves and small inflorescences, but the leaves are
not as narrow as those in P. maxonii. The type of
P. peperomiae Standl. (Pittier 3235 us from Pan-
ama) has somewhat larger leaves and is interme-
diate between the larger-leaved forms of this spe-
cies and the type of P. orchidearum. The fact that
all these plants share the same habitats and alti-
tudinal range suggests that they may be part of a
single polymorphic complex.

Psychotria guapilensis (Standl.) Hammel, Selby-
ana 12: 139. 1991. Cephaelis discolor Polak.,
Linnaea 41: 572. 1877, not/*, discolor (Griseb.)
Rolfe. C. tonduzii Krause, Bot. Jahrb. Syst. 54,
Beibl. 1 19: 45. 1916, not P. tonduzii Standl. C.
nana Standl., J. Wash. Acad. Sci. 17: 171. 1917,
not P. nana K. Krause. Evea guapilensis Standl.,
J. Wash. Acad. Sci. 15: 104. 1925. C. nicara-
guensis Standl., Trop. Woods 16: 46. 1928. C.
guapilensis (Standl.) Standl., Publ. Field Co-
lumb. Mus., Bot. Ser. 4: 295. 1929. Figure 7.

Herbs or subshrubs, 25-90 cm tall, with only 1 or 2
unbranched terete stems 2-4 mm thick, glabrous; stip-
ules with basal portion 2-4 mm long, with 2 narrow
triangular lobes 3-8 mm long, glabrous, often deciduous.
Leaves with petioles 2-9.5 cm long, 0.8-2 mm thick,
glabrous; leaf blades 9-24 cm long 3.5-10 cm wide, el-
liptic-oblong to oblong or elliptic-obovate, apex abruptly
narrowed and acuminate, tip 6-1 5 mm long, base obtuse
to acute and slightly decurrent on petiole, drying thinly
chartaceous, dark green or brown, glabrous above and
below, 2 veins 10-14/side. Inflorescences terminal and
solitary, ca. 5-8 cm long, the dense capitula 2-3 cm long
and 2.5-5 cm broad, globose to oblate, peduncles 4-40
mm long, ca. 1.5 mm thick, glabrous, with many ovate-
involucrate bracts 6-10 x 6-10 mm, covering the outer
surface and becoming dark reddish purple in color, brac-
teoles spatulate and with purple hairs, flowers sessile.
Flowers glabrous externally, hypanthium ca. 1.5 mm
long, calyx short-cupulate, lobes 4-5(-6), 0.8-1.5 mm
long, triangular, corolla pale pink to purple, funnelform,
tube ca. 2.5 mm long, lobes 5(-6), 1.5-2 mm long. Fruits
ca. 10 mm long, pyriform to ellipsoid, bright blue; py-
renes 5-6 mm long and 4 mm broad, smooth to slightly
angled on the dorsal surface.

Understory plants in primary rain forest for-
mations of the Caribbean slope in Costa Rica,

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

251

from 30 to 900 m elevation. Probably flowering
and fruiting throughout the year but most collec-
tions have been made in July-September. The spe-
cies ranges from southeastern Nicaragua to the
Choco region of Colombia.

Psychotria guapilensis is recognized by its small
herbaceous habit, long and slender petioles, dense
rounded short-pedunculate terminal heads cov-
ered by many purplish to maroon bracts, and bright
blue fruit. This species resembles P. glomerulata.

Psychotria haematocarpa Standl., J. Wash. Acad.
Sci. 18: 274. 1928. Figure 56.

Shrubs, 1-2.5 m tall, leafy stems 0.8-2.5 mm thick,
glabrous and terete, with 2 opposite longitudinal ridges
in early stages; stipules with a short (ca. 1 mm) truncate
sheath with 2 filiform awns per side 2-5 mm long, early
deciduous or the base of the stipule becoming thickened.
Leaves with petioles 2-8(-14) mm long, ca. 1 mm thick,
glabrous, drying yellowish; leaf blades 6-15 cm long, 2-
5.5 cm broad, elliptic to oblong-elliptic, apex acuminate
with tip 5-13 mm long, base attenuate or acute and
decurrent on petiole, drying membranaceous or thin-
chartaceous, green or grayish green, glabrous above and
below, 2 veins 6- 1 0/side and loop-connected near (1.5-
3 mm) the margin. Inflorescences terminal (pseudoax-
illary), solitary, rounded to subcapitate 4-10 mm long,
to 10 mm broad, condensed cymes with 5-9 flowers,
peduncles 2-5 mm long, ca. 0.5 mm thick, glabrous (or
minutely puberulent), bracts 0.5-2 mm long, green, gla-
brous, pedicels 0-2 mm long. Flowers minutely papil-
late-puberulent externally, hypanthium ca. 1 mm long,
calyx 1-2 mm long, thin, lobes 5, 0.8-1.3 mm long,
triangular; corolla short-salverform, pale greenish or
white, tube 2-2.5 mm long, ca. 0.5 mm diam. at base,
lobes 0.5-1 mm long, acute. Fruits 5-6 mm long, 4-5
mm diam., elliptic-oblong to globose, orange and finally
bright red (dark purple?) at maturity; pyrenes with 45
rounded ridges.

Plants of evergreen forests on the Caribbean slope
from 50 to 1000 m elevation and on the Pacific
slope at 500-1000 m. Flowering in July-August
near La Selva; fruiting in October-February. This
species ranges from Nicaragua to Colombia.

Psychotria haematocarpa is recognized by its
small stipules with deciduous filiform awns, small-
er green dried leaves with loop-connected second-
ary veins, very small condensed inflorescences, and
red fruit. The calyx resembles the corolla in size
and texture. This is one of only a few species of
subgenus Heteropsychotria with red fruit, rather
than blue or black. These are the smallest inflo-
rescences of any of our species of Psychotria (or
any of our shrubby species of Rubiaceae), and this
small size may explain the paucity of collections,
as the flowers and fruit are very difficult to see

among the leaves. Compare P. brachybotrya, with
larger bracts and purple fruits, and smaller spec-
imens of P. hoffmannseggiana, with larger bracts.

Psychotria hazenii Standl., Publ. Field Columb.
Mus., Dot. Ser. 7: 96. 1930. P. ramonensis
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser.
18: 1360. l938.CephaelischlorochlamysSlandl,
loc. cit. 1278. 1938. Figure 55.

Small shrubs, 1-2 m tall, leafy stems 0.9-3 mm thick,
glabrous, drying yellowish green; stipules with a tubular
sheath 3-7 mm long, with 2 narrow lobes 4-1 1 mm long
and 0.5-1.5 mm broad, linear to narrowly lanceolate,
glabrous. Leaves with petioles 6-20(-40) mm long, 0.3-
0.8(-1.3) mm broad, glabrous; leaf blades (3.5-)6-l 1(-17)
cm long, (l-)1.8-3.5(-4) cm broad, narrowly elliptic to
elliptic-oblong, apex tapering gradually or abruptly and
acuminate, tip 5-12 mm long, base acute to obtuse and
slightly decurrent on petiole, drying chartaceous, dark
green or brownish, glabrous above and below (or sparsely
and minutely puberulent along the midvein beneath), 2
veins 11-17/side. Inflorescences terminal and solitary
(or 3), 2-4 cm long, 1 .5-2.5 cm broad, loosely condensed
bracteate heads, peduncles 10-25 mm long (to 30 mm
in fruit), ca. 1 mm thick and glabrous, bracts subtending
the inflorescence branches 1-2 cm long, 4-7 mm broad,
ovate and acute or acuminate, glabrous externally (pu-
berulent within), chartaceous, green with bluish tips,
flowers sessile within enclosing bracts. Flowers with hy-
panthium ca. 1.8 mm long and 1.4 mm diam., obovoid-
oblong, white, glabrous, calyx tube ca. 0.7 mm long,
lobes 5, 1-2 mm long, ca. 1.2 mm broad at the base,
triangular, bluish distally; corolla not seen. Fruits 4-8
mm long, oblong, becoming bright blue with a spongy
texture, persisting calyx 1-2 mm long.

Plants of lower montane rain forest formations
along the Caribbean slope and continental divide,
from 900 to 1 600 m elevation. Probably flowering
in May-July; fruiting in August-November. In
Costa Rica this species is only known from the
Caribbean slopes of the Cordillera de Tilaran and
central Volcanic Highlands; it is also found in Co-
lombia and Ecuador.

Psychotria hazenii is recognized by the tubular
stipules with two long narrow awns, the slender
petioles, the condensed inflorescences with large
broadly imbricate bracts, and bright blue fruit.
Our inability to find material with corollas sug-
gests that flowering is nocturnal. This species re-
sembles P. steyermarkii with smaller bracts and
P. pittieri; compare also Palicourea skotakii.

Psychotria hebeclada DC., Prodr. 4: 513. 1830.
Probably including P. molliramis (K. Schum. &
Krarse.) Steyerm., Mem. New York Bot. Gard.

252

FIELDIANA: BOTANY

23: 529. 1972. Palicourea molliramisK. Schum.
& Krause, Bot. Jahrb. Syst., 40: 331 1910. Fig-
ure 55.

Shrubs or subshrubs, 0.7-2(-3) m tall, leafy stems 1-
4 mm thick, terete, sparsely to densely puberulent or
hispidulous with curved whitish hairs 0.2-0.8 mm long,
often contracted beneath the node when dried; stipules
with a short truncate tube 0-2 mm long, with 2 distal
linear lobes or spines 1-3 mm long, puberulent, persist-
ing or breaking apart. Leaves with petioles 4-15(-30)
mm long, 0.8-1.2 mm thick, puberulent; leaf blades 7-
20 cm long, 3-8 cm broad, elliptic to ovate-elliptic or
elliptic-obovate, apex acuminate with tip 4-14 mm long,
base cuneate to acute (obtuse), drying membranaceous
to thin-chartaceous, dark green to brownish green above,
much paler beneath, sparsely puberulent or subglabrous
above, puberulent beneath (especially along the midvein)
with thin straight hairs 0. 1-0.4 mm long, 2 veins (6-)8-
14/side. Inflorescences terminal and solitary (or 3),
(2.5-)3-6(-12) cm long, 1.5-4(-8) cm broad, narrowly
pyramidal or subcylindrical open panicles of closely
spaced flowers (in early stages), expanding in fruit, pri-
mary branches opposite or alternate, peduncles to 45(-60)
mm long, 1-1.8 mm thick, with curved or crooked thin
hairs 0.2-0.6 mm long, bracts 2-4 mm long, linear (not
apparent beneath the proximal branches), pedicels 1-
2(-5) mm long, often purplish, bracteoles 1-2 mm long.
Flowers distylous, puberulent externally, hypanthium ca.
0.7 mm long and 0.5 mm diam., obconic, calyx lobes 5,
0.3-1.5 mm long, ca. 0.3 mm broad, ligulate; corolla
white or tinged with pink or purple near the mouth,
salverform, tube 2-4 mm long, 1-2 mm diam. and often
constricted in middle, lobes 5, 1.5-2 mm long; anthers
1.2-1.7 mm long. Fruits 4-5 long, 3-5 mm diam., sub-
globose, with 8 or 1 sharply denned longitudinal ridges,
glabrous, purple or blue-black; pyrenes 2.5-4.5 mm long,
ridged.

Plants of evergreen forest formations on the Ca-
ribbean slope (20-600 m elevation) and on the
Pacific slope of southern Costa Rica from 40 to
700(-1000?) m. Flowering in March-August in
Costa Rica and Panama; fruiting in May-January.
The species ranges from southern Mexico to Ec-
uador and Venezuela.

Psychotria hebeclada is recognized by its small
stature, distinctive pubescence on most parts, two-
awned stipules, relatively compact pyramidal to
subcylindrical conical inflorescences (in early
stages), flowers with well-developed calyx lobes,
and small fruit finally becoming black. These plants
are often found along rivers, streams, and forest
edges. This species was called P. pubescens in the
Flora of Barro Colorado Island, but P. pubescens
is found in deciduous forests and has smaller calyx
lobes. It appears that P. hebeclada is synonymous
with P. molliramis as used by Steyermark (1974).
Poor material of this species may resemble P. ra-
cemosa. Liesner 14126 CR with glabrous inflores-

cences, large calyx lobes, and minutely papillate-
puberulent corolla tube is tentatively placed here;
it may be a closely related South American species.

Psychotria hoffmannseggiana (Willd. ex Roem. &
Schult.) Muell. Arg. in Mart., Fl. Bras. 6(5): 356.
1881. P. furcata DC., Prodr. 4: 512. 1830. P.
involucrata sensu Standley and others, in part,
not Sw. (cf. Steyermark, 1974). Figure 56.

Shrubs, 0.7-2(-4) m tall, leafy stems 0.7^4.5 mm thick,
minutely (0.1-0.2 mm) glabrous to papillate-puberulent
(conspicuously pubescent with thin straight hairs in a
few collections); stipules 2-4 mm long, basal sheath 0.3-
2 mm long, with 2 awns 1-3 mm long (4/node) separated
by a U-shaped sinus, glabrous (pubescent), usually per-
sisting. Leaves with petioles 2-8 mm long, ca. 0.5 mm
thick, glabrous or with hairs 0. 1-0.2(-0.5) mm long; leaf
blades 5-14(-17) cm long, 1.5-6(-8) cm broad, ovate-
elliptic, narrowly ovate, elliptic or elliptic-oblong (lan-
ceolate), apex tapering gradually and acuminate with tip
3-14 mm long, base cuneate to acute and often decurrent
on petiole, drying thinly chartaceous and usually green-
ish, glabrous above or puberulent only on midvein, gla-
brous or puberulent beneath with thin hairs 0. 1-0.3 mm
long (more rarely with hairs ca. 0.5 mm long), 2 veins
5-9/side. Inflorescences terminal and solitary, 6-25 mm
long, involucre up to 6 cm broad, capitate or condensed-
cymose with (3-)7-30 congested flowers, peduncles 3-
20 mm long, 0.7-1 .8 mm thick, pubescent or glabrescent,
bracts 3-22(-35) mm long, 1-6 mm broad, linear-lan-
ceolate, conspicuous and persisting, green to purple,
flowers sessile. Flowers glabrous or puberulent exter-
nally, hypanthium 1-2 mm long, calyx lobes 0.2-0.8 mm
long, acute to rounded; corolla white, funnelform, tube
2-6 mm long, 0.8-2.5 mm diam., lobes 1.3-2 mm long;
stamens 4 or 5, anthers ca. 1 mm long. Fruits 3-6 mm
long, 4-5 mm diam., subglobose to slightly oblong, with
8 longitudinal ridges, becoming purple to dark maroon
or black, glabrous or puberulent; pyrenes ridged.

Plants of wet evergreen lowland rain forest for-
mations, from near sea level to 700 m elevation,
both on the Caribbean slope and in the General
Valley and Golfo Dulce area. Flowering primarily
in May-August; fruiting in July-March. The spe-
cies ranges from Veracruz, Mexico, into Venezuela
and Brazilian Amazonia.

Psychotria hoffmannseggiana is recognized by
its persisting two-awned stipules, small capitate
inflorescences with conspicuous narrow spreading
or recurved persisting bracts, small flowers, and
purple fruit. The subcapitate inflorescences can
expand after anthesis and become slightly
branched. The leaves can vary greatly in size in
different plants and, while most of our plants have
little pubescence, a few are densely pubescent.
Plants of southwestern Costa Rica often have con-

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

253

spicuously larger bracts than specimens from other
areas. This species is very similar to P. officinalis,
which has larger branched inflorescences. The ma-
terial placed here was called P. involucrata Sw. by
Standley (1938 and in herb.), but Steyermark (1972,
p. 603) showed that the Swartz name is a synonym
of P. officinalis. Some authors may prefer to retain
P. furcata as a distinct species, but the South
American material of P. hoffmannseggiana in-
cludes a wide range of variation, some of which
is similar to that seen in the type of P. furcata.

cences, and the small flowers often with large calyx
lobes. The leaves are often undulate along the edge
in life and may have an arcuate submarginal vein.
The red fruits and tendency for the leaves to dry
grayish are characteristics of subgenus Psychotria.
There seems to be great variation in flowers and
inflorescence, with thicker puberulent peduncles
being correlated with more congested flowers. The
calyx lobes also seem to vary greatly in their de-
velopment. Compare P. quinqueradiata.

Psychotria horizontalis Sw., Prodr. 44. 1788. P.
longicollis Benth. in Oerst., Vidensk. Meddel.
Dansk. Naturhist. Foren. Kjobenhavn 1852: 33.
1853. Figure 65.

Small shrubs to little treelets, 1-3 m tall, leafy stems
14 mm thick, glabrous (rarely minutely puberulent),
usually becoming pale grayish; stipules 2-7 mm long, 1-
4 mm broad at base, triangular to narrowly ovate, obtuse
to acute, usually glabrous, drying reddish brown, cadu-
cous. Leaves with petioles l-7(-ll) mm long, 0.5-1.3
mm thick, glabrous or minutely puberulent; leaf blades
3 9(-l 3) cm long, 1 .5-4.5(-6) cm broad, elliptic to ovate-
elliptic, obovate, lanceolate or oblanceolate, apex acute
to short-acuminate, base obtuse or slightly auriculate,
drying thin-chartaceous or chartaceous, dark gray to
pinkish gray or brown, glabrous above and below (rarely
with thin hairs ca. 0.2 mm long on the veins beneath),
2 veins 5-9/side, occasionally with tufted hairs or pit
domatia in vein axils beneath. Inflorescences terminal
and solitary, 2-6 cm long (to 13 cm in fruit), l.5-4(-7)
cm broad, rounded open panicles with opposite or
whorled branching (often globose-umbellate), peduncles
l-3(-7) cm long, 0.5-1.3 mm thick, glabrous or puber-
ulent, with 2-5 flowers separate or closely clustered in
distal cymes, pedicels 0-2 mm long, bracteoles 0.2-1
mm long. Flowers glabrous externally, hypanthium 0.7-
1 mm long, obconic, calyx tube ca. 0.6 mm long, lobes
5, 0.5-3 mm long, linear to ligulate; corolla funnelform,
white, tube 2.3-3.5 mm long and 0.7-1.4 mm diam.,
lobes 5, ca. 1.5 mm long; anthers 0.7-1 .2 mm long. Fruits
4-8 mm long, 3-6 mm diam., ellipsoid to ovoid, orange
becoming bright or dark red; pyrenes ca. 4 x 3 mm,
with 3-5 rounded ridges.

Plants of both evergreen and seasonally decid-
uous forest formations, most often collected from
the seasonally dry Pacific slope, 20-1600 m ele-
vation. Flowering most often in May-August;
fruiting throughout the year (June-December in
Costa Rica). This species ranges from Mexico to
Ecuador and Brazil; it is also found in Cuba and
Hispaniola.

Psychotria horizontalis is recognized by its
smaller often subsessile leaves often with domatia,
unlobed stipules, often umbelliform inflore

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