ILLINOIS LIBRARY

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BIOLOGY

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no. 33

FLORA COSTARICENSIS

t\ iiliam Burger, Kditor

Family #202 Rubiaceae

vVilliam Burger

Charlotte M. Tavlor

Kurcniber 30. 1993

'ublication 1454

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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

in

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

IV

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

16

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

7

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

2

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]

3

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

6

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

12

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

30

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

30

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

32

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) . .

34

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

36

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"

37

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

40

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 . .

70

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 . .

75

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

15

Coccocypselum hirsutum

C. cordifolium

Coccocypselum lanceolatum

Geophila repens

G. cordifola

Geophila macropoda

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

16

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

17

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.

18

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

19

D. serrulata

cJi

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

20

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

21

Hoffmannia

subauriculata

H. vesiculifera

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

22

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.

24

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

25

P. macrophylla

sycnotria aggregata

Psychotria cartagoensis

Psychotria uliginosa

10 cm

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

26

FIELDIANA: BOTANY

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

27

Pentagonia

donnell-smithii

cm

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

28

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

29

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.

30

FIELDIANA: BOTANY

Psychotria poeppigiana

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

31

Psychotria glomerulata

c-8

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

32

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

33

Gonzalagunia panamensis

ovatifolia

U9

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

34

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

35

Randia brenesii

Randia

loniceroides

Randia

thurberi

Randia aculeata

R. armata

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

36

FIELDIANA: BOTANY

Randia

altiscandens

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

37

R. genipoides

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

38

FIELDIANA: BOTANY

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

39

Genipa americana

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

40

FIELDIANA: BOTANY

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

41

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

43

Tocoyena

pittieri

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

44

FIELDIANA: BOTANY

Exostemma

caribaeum

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

45

Guettarda crispiflora

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

46

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

47

Chomelia microloba

Hamelia rovirosae

Chomelia recordn

Chomelia spinosa

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

48

FIELDIANA: BOTANY

Sabicea panamensis

Sommera donnell-smithii

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

49

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

51

Rustia costaricensis

A

Simira maxonii / \ Rustia occidentalis

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

52

FIELDIANA: BOTANY

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

53

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.

54

FIELDIANA: BOTANY

R. monteverdensis

FIG. 4 1 . Rondeletia spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

55

Rondeletia

povedae

Rondeletia

chaconii

R. hamelifolia

FIG. 41 A. Rondeletia spp.

56

FIELDIANA: BOTANY

axillaris

FIG. 42. Hamelia spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

57

Ixora nicaraguensis

Raritebe palicoureoides

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

58

FIELDIANA: BOTANY

FIG. 44. Faramea: species with larger leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

59

parvibractea

6J3

FIG. 45. Faramea: species with smaller leaves.

60

FIELDIANA: BOTAN\

R. cornifolia

Rudgea reducticalyx

C. impetiolaris

R. trifurcata

FIG. 46. Coussarea and Rudgea spp.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

61

Psychotria

umbelliformis

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

62

FIELDIANA: BOTANY

Coussarea talamancana

FIG. 48. Coussarea: species with larger leaves.

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

63

tilaranensis

FIG. 49. Palicourea: species with conspicuous bracts.

64

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

65

Ufl

P. lasiorrhachis

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

66

FIELDIANA: BOTANY

Palicourea

purpurea

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

67

FIG. 53. Palicourea: species with smaller leaves.

68

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

69

Psychotria goldmanii , ^

rt.

P. steyermarkii

70

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

71

Psychotria

tapantiensis

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

72

FIELDIANA: BOTANY

berteriana

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

73

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.

74

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

75

marginata

Psychotria

monteverdensis

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

76

FIELDIANA: BOTAN\

Psychotria

mexiae

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

77

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

78

FIELDIANA: BOTANY

panamensis var.

compressicaulis

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

79

Psychotria

rosulatifolia

viridis

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

80

FIELDIANA: BOTAW

Psycnotna lamarinensis

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

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE

81

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-

82

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

83

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-

84

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

85

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 3—4 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,

86

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 0 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

87

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, 2—4 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-

88

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

89

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-

90

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

91

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

92

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

93

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 4—5 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

94

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

95

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 2—4 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-

96

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

97

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.

98

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

99

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 (84°43'W), and

the southern Cordillera de Talamanca, Limon

(82°59'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 0 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

(3—4 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 (1—4 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 0—40 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 Cm; 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

11

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 0 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 1.8—4 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, 4—7 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

3

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 5—45 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 3—4 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 0 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 2—4 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 2—4 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 0 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