(navigation image)
Home American Libraries | Canadian Libraries | Universal Library | Community Texts | Project Gutenberg | Children's Library | Biodiversity Heritage Library | Additional Collections
Search: Advanced Search
Anonymous User (login or join us)
Upload
See other formats

Full text of "Flora Costaricensis"

ILLINOIS LIBRARY 

,M URBANA-CHAMPAIGN 

BIOLOGY 



o vu,^ 

re 

rt.xv, 
no. 33 





FLORA COSTARICENSIS 

t\ iiliam Burger, Kditor 

Family #202 Rubiaceae 

vVilliam Burger 
Charlotte M. Tavlor 



Kurcniber 30. 1993 
'ublication 1454 



>U BLUSHED BY ! 



TURA 






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



FTELDIANA 



Botany 

NEW SERIES, NO. 33 



FLORA COSTARICENSIS 

William Burger, Editor 

Family #202 Rubiaceae 

William Burger 

Curator 

Department of Botany 

Field Museum of Natural History 

Chicago, Illinois 60605-2496 

Charlotte M. Taylor 

Missouri Botanical Garden 

St. Louis, Missouri 63166-0299 



Accepted April 16, 1993 

Published December 30, 1993 BIOLOGY LIBRARY 

Publication 1454 101 " 




PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY 






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

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



Table of Contents 



INTRODUCTION v 

ACKNOWLEDGMENTS v 

RUBIACEAE 1 

KEYS TO THE RUBIACEAE OF COSTA RICA 2 

Key 1 : Technical Keys to the Traditional 

Tribes and Genera of Rubiaceae 2 

Key 2: Artificial Key to Genera and 

Illustrations 8 

Illustrations of Rubiaceae 15 

Description of Genera and Species (Aliber- 
tid) 82 

Psychotria 220 

LITERATURE CITED 324 

LIST OF ACCEPTED SPECIES 325 

INDEX . 328 



List of Illustrations 



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

ia, and Diodid) 15 

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

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

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

pus, Richardia, and Spermacoce 18 

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

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

of Spermacoce 20 

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

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

nia 22 

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

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



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

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

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

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

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

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

and Warszewiczia 30 

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

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

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

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

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

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

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

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

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

and Hippotis 39 

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

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

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

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

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



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 34 mm diam., fruiting pedicels 20- 
35 m long, 1-1.5 mm thick. 



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

Amaioua pedicellata is recognized by its long- 



Amphidasya Standley 

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

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



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



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



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



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

lowlands C. latifolia 

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

C. parviflora 



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 24 cm long [petioles 
1-4 mm long] C. semipilosa 

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

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

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



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

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



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



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 (8443'W), and 
the southern Cordillera de Talamanca, Limon 
(8259'W). Collections in addition to the type are 
Bello 17 2 & 872, Bella & Cruz 457, Herrera 3310, 
and Poveda 24. 

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



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

Cinchona Linnaeus 

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

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



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

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



102 



FIELDIANA: BOTANY 



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

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

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



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



Coccocypselum P. Browne, 
nomen conservandum 

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

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



Key to the Species of Coccocypselum 

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

C. lanceolatum 

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

. C. herbaceum 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



103 



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

elevation] C. cordifolium 

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

elliptic 4 

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

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

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

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



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



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



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

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



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



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

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



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



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



104 



FIELDIANA: BOTANY 



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

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

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



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

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



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

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

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



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



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



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



105 



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

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

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



Coffea Linnaeus 

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



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

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



Key to the Common Species of Coffea 

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

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



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



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



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



106 



FIELDIANA: BOTANY 



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

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

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



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

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

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



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



( ondaminea DeCandolle 

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

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



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

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



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



107 



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

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

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



C osmibuena Ruiz & Pav., 
nomen conservandum 

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

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

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



Key to the Species of Cosmibuena 

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

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



108 



FIELDIANA: BOTANY 



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

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



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

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

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

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



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

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

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

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



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



109 



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



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






Coussarea Aublet 



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

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

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



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

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



Key to the Species of Coussarea 

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

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

. 4 



110 



FIELDIANA: BOTANY 



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

flowers, ovary and fruit pilose . c. enneantha 

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

flowers, ovary and fruit glabrous .3 

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

Caribbean lowlands, 0-300 mm elevation c. hondensis 

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

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

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

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

present in C. chiriquiensis 7 

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

lowlands of northern Costa Rica] C impetiolaris 

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

puberulent depressions in the vein axils or absent 6 

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

Rica C. sp. A aff. curvigemmia 

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

C. sp. B aff. curvigemmia 

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

usually with length equaling breadth 10 

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

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

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

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

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

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

at apex C. talamancana 

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

distally C. latifolia 

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

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 mm) tips, but other characteristics 
are very similar and suggest that the material placed 
here is conspecific. 



Coussarea sp. A aff. C. curvigemmia Dwyer, Phy- 
tologia 38: 215. 1978. Figure 47. 

Shrubs or small trees, 2-8 m tall, leafy stems 1.5-3.5 
mm thick, glabrous, grayish or yellowish green when dry; 
stipules 0.5-3 mm long, ovate and rounded distally or 
reduced to an entire ridge, glabrous. Leaves subsessile 
or with petioles 1-3 mm long, 0.5-1.5 mm thick, gla- 
brous; leaf blades 7-16 cm long, 2.5-5 cm broad, nar- 
rowly elliptic-oblong to elliptic-obovate, apex acumi- 
nate, tip ca. 1 6 mm long, gradually narrowed to the acute 
or cuneate base and usually slightly auriculate at the 
petiole, drying thin-chartaceous and grayish green, gla- 
brous above and below, with tufts of hairs in depressions 
(domatia) in the vein axils, 2 veins 4-7/side. Inflores- 
cences 4-7 cm long, 2-6 cm broad, paniculate with a 
single main rachis and short (3-12 mm) opposite or 
subopposite lateral branches, peduncles 1-2.2 cm long, 
glabrous and drying pale yellowish, flowers in distal pairs 
or triads, bracts subtending the flowers absent or less 
than 0.5 mm long. Flowers glabrous externally except 
for short (0.1 mm) erect whitish hairs on the ovary, 
hypanthium ca. 1 mm long, calyx tube 1-2 mm long, 
lobes 0.2-0.5 mm long, narrowly dentate; corolla nar- 
rowly tubular-salverform, white, tube 6-10 mm long, 
0.7-1.5 mm diam., lobes 4, ca. 6-7 mm long; anthers 
4-5 mm long, linear. Fruits not known (probably similar 
to those of C. curvigemmia: 10-13 x 5-8 mm, oblong 
and slightly flattened, whitish). 



116 



FIELDIANA: BOTANY 



Plants of evergreen rain forest of the Caribbean 
lowlands, from near sea level to 300 m elevation. 
Flowering in April and June (Barringer et al. 2642 
&3615cR,F, Gdmez-Laurito 8388 CR). This spe- 
cies is known only from near Suretka in the Tal- 
amanca Valley in southern Limon Province. 

Coussarea sp. A aff. C. curvigemmia is recog- 
nized by the thin subsessile leaves often slightly 
auriculate at the base and with weakly defined 
domatia, delicate whitish inflorescences, and small 
flowers with puberulent ovary and slender gla- 
brous corolla tubes. Coussarea curvigemmia, of 
central Panama, differs from the material placed 
here in having clearly outlined ellipsoid pit-doma- 
tia with few or no hairs, glabrous ovaries, and 
minutely papillate-puberulent corollas that dry 
dark. Both taxa have distinctive thin, slightly 
curved corolla tubes, and it may be that they are 
conspecific. This material also resembles Cous- 
sarea impetiolaris with more robust inflorescences 
and some species of Faramea. 



Coussarea sp. B aff. C. curvigemmia. Figure 48. 

Another species with very similar flowers may 
be represented by Burger & Gentry 8960 F from 
the Osa Peninsula and Bunting & Licht 793 F from 
the lower Rio San Juan, Nicaragua. Both of these 
collections have smaller (4 cm) compact capitate 
inflorescences and larger (20 cm) subsessile ob- 
ovate leaves with long (20 mm) narrow drip tips 
and with slightly auriculate leaf bases. However, 
the Nicaraguan collection has a pedunculate inflo- 
rescence and the Osa collection has a subsessile 
inflorescence, which may be immature. In con- 
trast, Liesner 3225 and Hammel et al. 18604 (all 
at CR) are intermediate with typical C. impetiolaris 
and indicate that the unusual specimens may be 
bridged by intermediates, in which case the de- 
scription given for C. impetiolaris needs to be ex- 
panded to include the collections placed here. 



Coutarea Aublet 

REFERENCE A. Aiello, A reexamination of 
Portlandia (Rubiaceae) and associated taxa. J. Ar- 
nold Arbor. 60: 38-124. 1979. 

Shrubs or small trees, branchlets terete, glabrous or 
puberulent, often with conspicuous elongate lenticels; 
stipules interpetiolar, short and acute, persisting. Leaves 
decussate or somewhat distichous, petiolate; leaf blades 
chartaceous, entire, some species with domatia. Inflo- 



rescences terminal or apparently axillary (terminal on 
short axillary shoots with poorly developed leaves), flow- 
ers usually in open cymose groups of 3 or solitary, pe- 
duncles short, pedicels subtended by narrow bracts. 
Flowers bisexual, monomorphic, large and showy, bi- 
laterally symmetrical due to curvature of the corolla tube 
and asymmetric stamens, hypanthium turbinate, calyx 
lobes 5-6(-8), narrow, often unequal: deciduous; corolla 
funnelform to campanulate and often inflated on the 
lower side, white to rose or purple, corolla tube slightly 
curved, with a glabrous throat, corolla lobes 5-6(-8), 
imbricate in bud; stamens 5 or 6, inserted near the base 
of the corolla tube, filaments long and often twisted in 
bud, anthers basi fixed, linear, exserted or included: ovary 
2-locular, placentas borne on the septum, ovules many 
in each locule. Fruits capsules, ovoid to obovoid or ob- 
long, flattened, coriaceous or woody, 2-locular, dehiscing 
loculicidally from the apex (down the center of the broad 
face of the capsule) to form 2 valves; seeds many, im- 
bricate and ascending, flattened and broadly winged with 
a thin margin around the circumference. 



A genus of 6-10 species ranging from southern 
Mexico to Argentina; only 1 species is found in 
Mexico and Central America. The large curved 
corolla tubes, long free filaments, and the capsules 
splitting down the middle of their flattened sides 
make the genus distinctive. Only a few species of 
Rubiaceae in our flora have similarly large flowers 
(cf. figs. 15 and 31). 



Coutarea hexandra (Jacq.) K. Schum. in Mart., Fl. 
Bras. 6, pt. 6: 196. 1889. Portlandia hexandra 
Jacq., Enum. PI. Carib. 16. 1760; Sel. Stirp. 63, 
pi. 1 82, f. 20. 1 763. C. speciosa Aubl., PI. Guian. 
1: 314, pi. 122. 1775. Figure 31. 

Shrubs or small trees, 2-8(-18) m tall, leaf branchlets 
1.7-4 mm thick, glabrous or sparsely puberulent, be- 
coming dark brown with elongate whitish lenticels; stip- 
ules 1.5-4(-5) mm long, ca. 2 mm broad at the base, 
intrapetiolarand forming a short (0.5- 1.5 mm) tube above 
the petioles, distally triangular and acute, glabrous in 
Central America. Leaves with petioles 2-10(-15) mm 
long, 0.7-1 .5 mm wide, glabrous or minutely puberulent; 
leaf blades 5-1 5 cm long, 2-9 cm broad, ovate to broadly 
elliptic or ovate-oblong, apex acute, short-acuminate or 
caudate-acuminate, base obtuse to rounded and sub- 
truncate (acute), drying thin -chartaceous or membra- 
naceous, glabrous or minutely puberulent on the veins 
above, glabrous or minutely (0.2-0.4 mm) puberulent 
beneath, 2 veins (4-)6-10/side, with tufts of minute 
hairs (domatia) in vein axils beneath. Inflorescences few- 
branched and with (l-)3-9 flowers, peduncles 3-30 mm 
long, bracts 3-5 mm long, pedicels 2-1 5 mm long, merg- 
ing gradually with the base of the flower, glabrous or 
sparsely and minutely puberulent. Flowers to 10 cm long 
and 4 cm broad, mostly glabrous (in Central America), 
hypanthium 4-7 mm long, 2-3 mm diam., calyx tube 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



117 



1.5-2.5 mm long, 4-5 mm broad, calyx lobes 4-6, 4- 
9(-12) mm long, lanceolate to linear, deciduous; corolla 
funnelform-campanulate, white or tinged with pink, gla- 
brous externally, tube 45-80 mm long, 1 0-20 mm diam. 
distally, slightly curved, inflated before anthesis, lobes 
4-6, 1-2 cm long, 7-10 mm broad, apex ovate-triangular 
and obtuse to acute; stamens 6, filaments to 8 cm long, 
anthers 14-19 mm long, 0.5-1 mm thick, yellow, ex- 
serted; style to 85 mm long. Fruits 25-45 mm long, 15- 
28 mm broad, ca. 8 mm thick, oblong-obovate, woody 
when mature, surfaces dark brown and glabrous, with 
or without small white punctate lenticels, the valves 
slightly split in 2 at apex; seeds 7-14(-20) mm long, 6- 
9 mm abroad, oblong to suborbicular, body of the seeds 
3-4 mm diam., wing pale brown. 

Trees and shrubs of both deciduous and ever- 
green forest formations, from near sea level to 900 
m elevation. Flowering in late June-October; 
fruiting in November-April in southern Central 
America. This species appears to be much more 
common in seasonally deciduous forests than in 
evergreen forests in Costa Rica. The species ranges 
from southern Mexico and Belize to Argentina. 

Coutarea hexandra is recognized by its thin 
short-petiolate leaves, few-flowered inflorescences 
with large flowers, the curved corolla tube, and the 
large flattened capsules with winged seeds. The 
fruits are unusual in that they split down the center 
of the broadly flattened halves (fig. 31). The flow- 
ers appear to be filled with gas before anthesis, and 
they are often pendulous at anthesis. The floral 
biology has been discussed by Haber and Frankie 
(1989). In northern Central America, bitter prin- 
ciples in the bark have been used medicinally, 
especially for malaria (Mabberley, 1987). 



Crusea Schlectendal & Chamisso 

REFERENCE W. R. Anderson, A monograph of 



the genus Crusea (Rubiaceae). Mem. New York 
Bot. Gard. 22: 1-128. 1972. 

Annual or perennial herbs, sometimes woody at the 
base, decumbent or erect, stems terete or 4-angled with 
longitudinal ribs, pubescent; stipules interpetiolar and 
united to adjacent petioles to form a thin sheath, with a 
distal cross- vein and 2-12 setae, persisting. Leaves op- 
posite and decussate, sessile or petiolate, petioles adnate 
to the stipular sheath; leaf blades mostly ovate to lan- 
ceolate with strongly ascending pinnate venation, entire, 
domatia absent. Inflorescences terminal or axillary, cap- 
itate, verticillate or of congested dichasial cymes, sessile 
to long-pedunculate, subtended by 2, 4, or 8 (more) leaf- 
like bracts, pedicels short (to 2 mm) or absent. Flowers 
bisexual and radially symmetrical, homostylous, usually 
4-parted, calyx lobes 4 or reduced to 2-3, with minute 
glands in the base of the sinuses between the lobes; co- 
rolla funnelform to campanulate, white to pink, red, or 
purple, corolla lobes 4, valvate in bud; stamens 4, fila- 
ments adnate to the upper half of the tube and free be- 
neath apex of the tube, anthers dorsifixed, exserted; ovary 
2-locular, placenta elongate from the center of the sep- 
tum, with 1 ovule in each locule, stigma 2-lobed or sub- 
capitate. Fruits of 2 dry 1 -seeded mericarps (cocci) borne 
on the sides of and separating from a persisting bifid or 
fenestrated carpophore, mericarps indehiscent and ecos- 
tate, calyx dehiscing circumscissily or persisting; seeds 
with the persisting placenta on the adaxial face. 

A genus of 13 species ranging from Arizona, 
U.S.A., through Mexico and Central America to 
western Panama. The herbaceous habit, congested 
subsessile flowers, often lanceolate leaves with 
strongly ascending secondary veins, narrow co- 
rolla tube (in our species) and unusual fruit help 
distinguish this genus. The mericarps are easily 
mistaken for seeds because of their smooth round- 
ed brownish surfaces and longitudinal adaxial sul- 
ci. The genus reaches its southern limit in Costa 
Rica and western Panama and is represented by 
only a few collections from Costa Rica. This treat- 
ment is based on Anderson's detailed monograph. 



Key to the Species of Crusea 

la. Flowers bright pink to magenta; leaf blades 1-5 cm broad, to 13 cm long 2 

Ib. Flowers white or white-tipped with pink; leaf blades 0.5-3.5 cm broad, to 8 cm long 3 

2a. Corolla tube 20-38 mm long, stigma lobes 1-4 mm long; secondary veins arising from the 
proximal half of the midvein; 1 800-3000 m elevation C. coccinea 

2b. Corolla tube 5-18 mm long, stigma lobes 0-0.5 mm long; secondary veins arising from the 

proximal '/: or 2 /3 of the midvein; 0-200 m elevation C. hispida 

3a. Corolla tube 5-1 1 mm long, stigma lobes 0.2-0.6 mm long; plants usually found around 2000 m 

elevation in Costa Rica C. longiflora 

3b. Corolla tube 1.84 mm long, stigma lobes 0.1-0.3 mm long; plants found in deciduous and partly 

deciduous areas below 1 200 m elevation C. parviflora 



118 



FIELDIANA: BOTANY 



Crusea coccinea DC., Prodr. 4: 567. 1830. C. coc- 
cinea var. chiriquiensis W. R. Anderson, Mem. 
New York Hot. Card. 22: 45. 1972. Figure 31. 



Decumbent or low perennial herbs to 1 m tall, often 
rooting at the nodes, leafy stems 1-3.5 mm thick, gla- 
brous or sparsely puberulent, quadrangular or terete; 
stipule sheath 4-14 mm long (including the setae), 1-6 
mm wide, glabrous or puberulent, setae 3-12 and to 12 
mm long and linear, evenly spaced or in a central group 
with fused bases. Leaves with petioles 4-20(-25) mm 
long, glabrous or puberulent; leaf blades 2.2-10(-l 3) cm 
long, l-4(-5) cm broad, narrowly elliptic to elliptic or 
ovate, apex acute to short- or long-acuminate, base acute 
and decurrent on petiole, drying stiffly chartaceous and 
dark, often slightly scabrous with the major veins im- 
pressed above, glabrous or sparsely puberulent above, 
sparsely hispidulous with short (0.3 mm) hairs along the 
veins beneath, 2 veins 3-6/side, strongly ascending, sub- 
parallel and arising from the proximal half of the mid- 
vein, minutely punctate on both surfaces. Inflorescences 
bracteate/involucrate heads or with verticillate flowers 
in the node below the terminal head, 10-15 mm diam., 
with 15-100 flowers in the head but few-10 flowering at 
the same time, involucral bracts 2 or 4, 2-4 cm long, 
leaf-like and often sessile within the expanded petioles 
and st i pular sheath of the subtending node, often with 
additional smaller bracts. Flowers with hypanthium 1 .4- 
3 mm long, glabrous or puberulent distally, calyx lobes 
to 8 mm long, narrowly triangular, corolla deep red to 
pink, purple, or lavender, funnelform, tube (8-)20-38 
mm long, gradually expanded from a narrow (3 mm) 
base, lobes (3.5-)5-l 1 mm long; stamens with free por- 
tion of the filaments ca. 4-10 mm long, filiform, anthers 
(2-)2.5-3.8 mm long; style as long as the corolla tube. 
Fruits with a broad flat carpophore, 4-7 mm long with 
lobes 0.7-2 mm long, cocci (1.7-)3-6 mm long, 2-2.7 
mm broad, rounded-turbinate to turbinate, brown, calyx 
often coming off as a complete whorl (circumscissile). 



Crusea hispida (Miller) Robinson, Proc. Amer. 
Acad. Sci. 45: 409. 1910. Crucianella hispida 
Miller, Gard. Diet. ed. 8, no. 4. 1768. 

Erect annual herbs to 0.6 m tall, many-branched, leafy 
stems 0.5-4 mm thick, terete with stiff" unicellular trans- 
parent or whitish hairs 1.5-3 mm long, spreading or 
retrorse from a thickened base, smaller (0.2 mm) hairs 
sometimes also present; stipule sheath 2-5 mm long, 4- 
10 mm broad, with 3-7 setae, conspicuously hispid. 
Leaves opposite, often with smaller axillary leaves from 
the same node, petioles 5-25 mm long, hispid; leaf blades 
4-9(-l 1) cm long, l-3.6(-4.8) cm broad, narrowly ovate- 
elliptic to lanceolate or narrowly ovate, apex tapering 
gradually and acuminate, base acute to obtuse, drying 
thin-chartaceous, both surfaces with thin whitish hairs 
ca. 1.3 mm long, 2 vein 5-6/side, strongly ascending. 
Inflorescences solitary terminal bracteate capitulae with 
40-100 closely crowded sessile flowers, subtended by 8 
(4) leaf-like bracts and many linear hispid bracteoles. 
Flowers with glabrous hypanthium, calyx tube 0.3-1 mm 
long, lobes 2-6 mm long, subulate, margins with stiff 
hispid hairs; corolla salverform, light pink to dark red 
or purple, tube 5.5-12(-18) mm long, ca. 0.3 mm diam. 
(dried), lobes 2-5 mm long; anthers 1-1.5 mm long, 
exserted. Fruits with cocci 2-3.5 mm long, 1.1-2.3 mm 
broad, yellowish to dark brown. 

A species of open grassy sites, ranging from 
Mexico to El Salvador and collected only recently 
in Costa Rica at a single locality: Costa de Pajaros, 
Bahia de Nicoya, Puntarenas (C. M. Taylor 249 
and Wilbur 31715, both at DUKE). This population 
was in flower in July; it is variety hispida (with 
shorter corolla tubes). The broader petiolate leaves, 
colorful flowers, and unusual pubescence distin- 
guish this species. 



Herbs of montane evergreen forest formations 
of the Pacific slope and central highlands, from 
1 800 to 3000 m elevation. Probably flowering and 
fruiting mostly in the wet season and beginning of 
the dry season: May-January. The species ranges 
from western Mexico to western Panama. 

Crusea coccinea is recognized by the herbaceous 
habit, short stipular sheath with long narrow setae, 
narrow leaves with subparallel secondary veins, 
and large pink to lavender flowers. Plants of Costa 
Rica and Chiriqui, Panama, belong to variety chi- 
riquensis W. R. Anderson. This variety is similar 
to variety coccinea in having larger corollas, an- 
thers, and mericarps, but variety chiriquensis dif- 
fers in having leaves with major veins deeply im- 
pressed above, a greater number of stipular setae, 
and pink to magenta flowers. While often collected 
in the Chiriqui Highlands, these plants have rarely 
been collected in Costa Rica. 



Crusea longiflora (Willd. ex Roem. & Schult.) W. 
R. Anderson, Mem. New York Bot. Gard. 22: 
89. 1 972. Spermacoce longiflora Willd. ex Roem. 
& Schult., Syst. Veg. 3: 531. 1818. C. brachy- 
phylla Schlechtend. & Cham., Linnaca 5: 165. 
1830. 

Erect annual herbs to 50 cm tall, stems terete or less 
often quadrangular, pubescent to pilose with hairs 1-2 
mm long; stipule sheath 1 .4-6.5 mm long, 2-7 mm broad, 
with 3-7 distal setae, and 0-2 inconspicuous sessile lat- 
eral colleters, longest setae 0.5-3(-5) mm long, equaling 
or shorter than the sheath, with thin hairs 0.5-2 mm 
long. Leaves sessile or subsessile with petioles to 5 mm 
long; leaf blades 8-50 mm long, 3-2 1 mm wide, narrowly 
to broadly elliptic, or ovate, apex acute to obtuse, base 
abruptly narrowed, drying chartaceous and scabrous, 
sparsely hispidulous, 2 veins 2-3/side, strongly ascend- 
ing. Inflorescences 1-2 cm diam., small terminal brac- 
teate heads (or with lateral heads reduced to an axillary 
cluster of 1 or a few flowers), with up to 75(-l 00) flowers 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



119 



in a head, involucral bracts 4 or 8. Flowers sometimes 
cleistogamous and resembling small unopened flower 
buds, hypanthium 0.7-1 . 1 mm long, glabrous, calyx tube 
0.5-1 mm long, lobes 1-3.5 mm long and 0.2-0.8 mm 
wide, broadly to narrowly triangular, often ciliate along 
the edge; corolla white (rarely pink or lavender), tube 5- 
1 1 mm long, narrow at the base and only 0.8 mm broad 
at apex, papillose externally, with long straight hairs within 
distally, lobes 1 .6-3.6 mm long, 0.8-1 .5 mm broad, nar- 
rowly elliptic, becoming strongly reflexed, with few long 
hairs at the base within; stamens with filaments 1.5-3.8 
mm long, anthers 0.6- 1.3 mm long; style 5-16 mm long, 
glabrous, stigmatic lobes 0.2-0.6 mm long. Fruits with 
a bifid carpophore to 1 .5 mm long, the mericarps (cocci) 
1.1-2.3 mm long, ellipsoid-cylindrical, whitish to yel- 
low-brown or dark brown, calyx often coming off as a 
whorl. 



Herbaceous plants of montane evergreen for- 
mations, from ca. 1000 to 2700 m elevation. Flow- 
ering in July-December; fruiting in September- 
December (over the entire range). This species 
ranges from northern Mexico through Guatemala, 
with isolated occurrences in the highlands of Hon- 
duras and Costa Rica. 

Crusea longiflora is distinguished by its erect 
annual habit, short stipular sheath with setae, small 
and narrow (often subsessile) leaves, the narrow 
corolla tube, and the small rounded mericarps. 
The long (1-2 mm) slender unicellular hairs also 
help distinguish this species. It has been collected 
infrequently in the Central Highlands at about 2200 
m elevation. This species is common in Mexico. 



Crusea parviflora Hook. & Arnott, Bot. Beechey 
Voy. 430, pi. 99. 1840. Figure 4. 

Erect, trailing or decumbent herbs to about 50 cm high 
and 1 m long, annual or perennial, leafy stems 0.5-3.5 
mm thick, with 4 longitudinal ridges or wings in early 
stages, with thin whitish hairs 0.3-1 .2 mm long but often 
glabrescent; stipule sheath 1-3 m long, 1.5-4 mm wide, 
often with minute spots, with 3-5 setae 1-7 mm long 
and with hairs to 1 mm long. Leaves with petioles 3- 
9(-15) mm long, with lateral wings continuous with the 
lamina margin, with thin hairs on both surfaces; leaf 
blades 2.4-7(-9) cm long, 1-3.6 cm broad, narrowly to 
broadly elliptic or lanceolate, apex bluntly to sharply 
acute, base gradually (abruptly) narrowed and decurrent 
on petiole, drying thin-chartaceous and often grayish 
green, pilose on both surfaces with thin whitish hairs 
0.3-1 mm long, 2 veins 3-5/side. Inflorescences brac- 
teate heads, with 10-many flowers per head and a ma- 
jority of the flowers in bloom at one time, the heads 
terminal or axillary to distal leaves, subglobose and 1- 
2 cm broad, often on long (to 1 5 cm) peduncles, invo- 
lucral bracts to 3 cm long and leaf-like, pedicels short. 
Flowers with small (0.5-1 mm) glabrous hypanthium, 
calyx tube 0.5-1.3 mm long, lobes 0.6-2.5 mm long, 
0.2-0.5 mm broad and linear-triangular, with thin hairs 



on the outer surface, calyx persisting on the fruit; corolla 
white or the lobes tipped with pink, funnelform, tube 
\.8-4 mm long, 0.7-1.4 mm broad at the throat, usually 
glabrous externally and with longer hairs on the distal 
portion inside, lobes 1.3-2.8 mm long, 0.5-1 mm broad, 
narrowly triangular, erect to spreading (not reflexed), 
usually with small hairs externally at the tips; stamens 
with filaments 1.3-4.5 mm long, glabrous and becoming 
retracted into the corolla after anthesis; style 2.7-8 mm 
long, glabrous, stigmatic lobes 0.1-0.3 mm long. Fruits 
with mericarps 0.8-1.2 mm long, 0.6-0.9 mm broad, 
oblong-cylindrical or subglobose, surface slightly pitted. 

Weedy plants of deciduous and evergreen for- 
mations, from 50 to 1600(-2000) m elevation in 
Central America. Flowering in October-April; 
mature fruits in November-April. The species 
ranges from western Mexico along the Pacific slope 
to isolated localities in Honduras, Nicaragua, and 
Costa Rica. 

Crusea parviflora is recognized by its short weedy 
habit, quadrangular or slightly winged young stems, 
short stipular sheath with few setae, capitate in- 
florescences with usually only 4 broad bracts, and 
unusual carpophore and seed-like mericarps. The 
short-pedicellate flowers contrast with the sessile 
flowers of similar-looking species of Mitracarpus 
and Spermacoce. In our area it has only been found 
in Guanacaste Province below 500 m elevation. 



Declieuxia Humboldt, Bonpland & Kunth 

REFERENCE J. H. Kirkbride, Jr., A revision of 
the genus Declieuxia (Rubiaceae). Mem. New York 
Bot. Gard. 28: 1-87. 1976. 

Herbs or subshrubs, perennial and often woody at the 
base, branches terete or angular, glabrous or puberulent; 
stipules interpetiolar, subulate or reduced to a line bear- 
ing 1-3 linear lobes. Leaves opposite or whorled, sessile 
or short-petiolate, leaf blades entire and usually small, 
linear to elliptic, deltoid or orbicular, usually coriaceous, 
domatia absent. Inflorescences of terminal or axillary 
panicles (compound cymes), often spike-like or race- 
miform, solitary to 3 at the end of the stem, pedunculate, 
branching often dichotomous, flowers in distal cymose 
groups, bracts and bracteoles present or absent. Flowers 
bisexual and radially symmetrical, small, mostly gla- 
brous, hypanlhium turbinate to subglobose or obovoid, 
slightly compressed laterally, calyx lobes 4 (2), equal or 
unequal, small and persisting; corolla funnelform to tu- 
bular, white to blue or purple, tube pilose-villose in the 
throat, corolla lobes 4, short, spreading or reflexed, val- 
vate in bud; stamens 4, inserted in the corolla throat or 
between the lobes, filaments slender, anthers dorsifixed 
and versatile, partly or completely exserted; ovary 
2-locular, ovule solitary in each locule, erect from a near- 
basal placenta, style slender, stigmas 2. Fruits drupa- 
ceous, dry or fleshy, laterally compressed and 2-parted 
or with 2 prominent rounded lobes, black at maturity. 



120 



FIELDIANA: BOTANY 



A genus of about 40 species in tropical America, 
with the largest number of species in Brazil. Our 
representative of this genus is recognized by the 
very short woody stems, stiff subsessile leaves, cy- 
mose branching of the short inflorescences, two- 
lobed fleshy fruits, and restriction to open grassy 
habitats in deciduous or partly deciduous forma- 
tions. 



rocky or savanna-like habitats, the stiff narrow 
sessile leaves often three at a node, the short di- 
chotomously branched panicles, the small flowers, 
and the sessile fleshy deeply two-lobed or rounded 
fruit (laterally compressed when dry). Chacon 
(2258 CR, MO) stated that the flowers are white 
with blue-lavender stamens. Our material belongs 
to variety mexicana, which is distinguished by its 
narrow leaves and lack of pubescence. 



Declieuxia fruticosa (Willd. ex Roem. & Schult.) 
Kuntze, Rev. gen. pi. 1: 279. 1891. Houstonia 
fruticosa Willd. ex Roem. & Schult., Syst. Veg. 
3: 527. 1818. D. mexicana DC, Prodr. 4: 479. 
1830. D. fruticosa var. mexicana (DC.) Standl., 
Field Mus. Nat. Hist., Bot. Ser. 12: 378. 1936. 
Figure 1. 

Erect subshrubs or herbaceous 20-70 cm tall, often 
from a hard woody rootstock, with vertical simple or 
few-branched stems, leafy stems 0.7-3 mm thick, gla- 
brous (in variety mexicana) to pubescent, with 2 or 4 
longitudinal ridges or wings (0.5 mm high), becoming 
terete; stipules 2-5 mm long, linear to linear-subulate, 
glabrous (in variety mexicana) or pubescent, deciduous. 
Leaves 2 or 3/node, sessile or subsessile (petiole to 1 mm 
long); leaf blades 20-^0(-50) mm long, 4-15(-22) mm 
broad, narrowly elliptic-oblong to linear-oblong or ob- 
long, apex acute or obtuse, base cuneate, drying stiffly 
chartaceous to subcoriaceous, glabrous and often lus- 
trous above, glabrous (in variety mexicana) or puberu- 
lent beneath, 2 veins (2-)3-6/side. Inflorescences ter- 
minal or axillary, 1-4 cm long, 1.5-5 cm broad, solitary 
or 3 at the ends of branchlets, open cymose paniculate 
with dichotomous nodes (and a sessile flower at the di- 
chotomy), primary peduncle 5-15(-40) mm long, bracts 
1.5-3 mm long, linear, pedicels 0-1 mm long. Flowers 
heterostylous, hypanthium 0.3-1 mm long, calyx lobes 
0.3-0.8 mm long, linear-oblong, glabrous (in variety 
mexicana) to villous, corolla white to blue or purple, 
4.5-6 mm long, tube 3-4.5 mm long, cylindrical, lobes 
4, ca. 2 mm long; anthers ca. 1 mm long, linear; style 
3-4.5 mm long, stigmas 0.3 mm long. Fruits slightly 
fleshy and prominently 2-lobed (when both ovules de- 
velop), ca. 2 mm long and 3 mm broad, sessile, the lobes 
suborbicular, the fruit subglobose when only 1 ovule 
develops and ca. 2 mm diam., drying black, glabrous. 



Small subshrubs of open grassy savanna-like sites 
in deciduous and semideciduous forest forma- 
tions, from ca. 20 to 1200(-1800?) m elevation. 
In Costa Rica the species is restricted to the Pacific 
slope and is common in Guanacaste and the Bue- 
nos Aires area of the General Valley. Flowering 
throughout the year (primarily in July-Septem- 
ber). The species ranges from southern Mexico to 
Brazil. 

Declieuxia fruticosa is recognized by the short 
stature from a woody base, restriction to open 



Deppea Chamisso & Schlechtendal 

REFERENCE D. H. Lorence and J. D. Dwyer, A 
revision of Deppea (Rubiaceae). Allertonia 4: 389- 
436. 1988. 



Slender shrubs and small trees, stems sparsely to densely 
puberulent; stipules interpetiolar, small, triangular, per- 
sistent. Leaves opposite (rarely in whorls of 3), opposing 
leaves of the same node often unequal, petiolate, leaf 
blades entire and pinnately veined, drying thin-charta- 
ceous, domatia sometimes present. Inflorescences axil- 
lary or terminal, scorpioid, umbellate or corymbiform 
to thyrsoid, the flowers in cymose groups (or rarely of 
solitary flowers), pedunculate, flowers pedicellate, brac- 
teolate. Flowers bisexual and radially symmetrical, hy- 
panthium hemispheric to turbinate or cylindrical, calyx 
lobes 4, minute or large, equal or unequal, usually with 
a small gland in each sinus; corolla short-funnelform to 
salverform, yellow (less often white, orange, or purple), 
corolla tube glabrous within, usually shorter than the 
lobes, corolla lobes 4, spreading or erect at anthesis, 
comoluatc in bud; stamens 4, inserted near the base of 
the tube, filaments short and linear, anthers dorsifixed, 
oblong to narrowly ellipsoid, exserted or partly included, 
a nectariferous disc present; ovary 2-locular, placentas 
elongate and peltate on the septum, ovules many in each 
locule and longitudinally imbricate, style slender, stigma 
entire or bilobed. Fruits a small dry capsule, turbinate 
to obovoid, coriaceous to chartaceous, usually with (6-)8 
longitudinal costae, bisulcate and dehiscing loculicidally 
from apex, valves cleft, calyx persisting; seeds many and 
minute, angulate, testa foveolate and reticulate. 



A genus of about 25 species, centered in Mexico 
and ranging through Central America ( 1 species) 
to southeastern Brazil ( 1 species). This account is 
based on the recent revision by Lorence and Dwyer 
(see reference above). Deppea can be confused with 
Hamelia and Hoffmannia, which have fleshy fruits. 



Deppea grandiflora Schlcchtend., Linnaea 19: 748. 
1847. D. costaricensis Polak., Linnaea 41: 566. 
1877. D. floribunda Hemsl., Diagn. PI. Nov. 
Mexic. 3 1 . 1 879. D. longipes Standl., Contr. U.S. 
Natl. Herb. 18: 138. 1916. Figure 38. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



121 



Shrubs or small trees, 1-4 m tall, leafy branchlets 1- 
4 mm thick, hirtellous with short (0.3 mm) brownish 
hairs, at first with longitudinal ridges but becoming terete 
and glabresent; stipules 0.5-1 mm long, 2-6 mm wide, 
broadly deltoid, glabrate to densely hirtellous, the inner 
surface or margin with 4-8 dark brown digitate colleters. 
Leaves opposite and subequal or unequal at the same 
node (with 1 up to 2 times as long as the other), petioles 
6-35(-50) mm long, 0.5-2 mm thick, glabrate or densely 
hirtellous on the adaxial side; leaf blades (3-)5-17(-21) 
cm long, (1.5-)2-7(-8.5) cm broad, elliptic to narrowly 
elliptic, narrowly ovate or lanceolate, apex tapering grad- 
ually and acuminate, base attenuate to acute (obtuse), 
drying chartaceous, glabrous to sparsely puberulent with 
minute (0.2 mm) thin whitish hairs above, minutely 
strigillose along the veins beneath, occasionally with tuft- 
ed domatia in the vein axils beneath, 2 veins 6-1 I/side 
and weakly loop-connected near the margin. Inflores- 
cence terminal (rarely axillary), 3-12 cm long and 3-15 
cm broad, dichasial and corymbiform, with 15-100 
flowers, peduncles 1.2-7 cm long, densely and minutely 
strigillose-hirtellous, primary branches 1-3 cm long and 
with up to 3 additional orders of branching, distal cy- 
mules of 2-6 flowers, pedicels 1-4 mm long, bracteoles 
present or absent. Flowers with hypanthium 1-2 mm 
long, 0.5-1 mm diam.. obconic or turbinate, with lon- 
gitudinal costae, calyx cup 0. 1-0.2 mm deep, lobes 0.3- 
1 mm long, deltoid; corollas funnelform or rotate, yellow, 
glabrous, tube 1-2 mm long, corolla lobes 4, 47 mm 
long, 3-4 mm wide, obtuse; stamens 4, filaments 2 mm 
long, anthers 3-4 mm long, exserted, basally sagittate; 
style 3.5-4.5 mm long, stigmas 1.5-2 mm long, entire. 
Fruits 2-5 mm long, 2-4 mm diam., obconical to subglo- 
bose, with 6-8 prominent longitudinal costae, opening 
at the top; seeds 0.5-0.7 mm long, discoid and often 
angulate, testa foveolate. 

Shrubs of evergreen montane forest formations, 
from 1 600 to 2700 m elevation. Flowering in Jan- 
uary-July, with a peak in April. Fruiting in Jan- 
uary-September. The species has been collected 
around Monteverde, Volcan Barva, and in the 
western Cordillera de Talamanca (to above San 
Isidro del General) in Costa Rica. The species 
ranges from central and eastern Mexico through 
the highlands of Guatemala and Honduras to the 
Chiriqui Highlands of Panama. 

Deppea grandiflora is recognized by its restric- 
tion to higher montane forest formations, small 
shrubby habit, thin leaves unequal at a node and 
gradually tapering at both ends, bright yellow gla- 
brous corollas with very short tube and broad lobes, 
and small costate capsules opening at the top. Plants 
with pseudoaxillary inflorescences and unopened 
flowers may resemble species of Hoffmannia. Ma- 
terial of Chiococca is also similar. 



Didymaea Hooker f. 

Perennial herbs, scandent or procubent, stems brittle, 
usually much-branched and with long slender inter- 



nodes, glabrous or rarely puberulent; stipules interpetio- 
lar and 2-lobed or apparently free (with 4/node), decid- 
uous or persisting and becoming recurved. Leaves 
petiolate; leaf blades entire and pinnately or subpal- 
mately veined, membranaceous to thin chartaceous, 
domatia absent. Inflorescences of solitary flowers in the 
axils of leaves, the pedicels long but not articulate, be- 
coming recurved in fruit. Flowers bisexual and radially 
symmetrical, hypanthium turbinate-globose, calyx en- 
tire; corolla campanulate to rotate, yellowish to greenish 
brown or purple, glabrous, corolla lobes 4, triangular and 
subacute, valvate in bud; stamens 4, inserted between 
the corolla lobes, filaments short and subulate, anthers 
dorsifixed; ovary 2-locular, ovules solitary in each locule 
and attached on the lower half of the septum. Fruits 
2-parted or 2-lobed, somewhat fleshy, becoming dark 
blue or black and lustrous, the lobes rounded, 1 lobe 
usually smaller and lacking a fully developed seed; py- 
renes rounded. 

A small genus of two to five species, ranging 
from Mexico to Panama. The plants are recog- 
nized by their herbaceous climbing Galium-like 
habit, small thin leaves, minute flowers solitary in 
the leaf axils, and fleshy, often 2-parted rounded 
fruit. The circumscription of species in Central 
America is not resolved (see below). 



Didymaea alsinoides (Cham. & Schlechtend.) 
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 
18: 1291. 1938. Nertera alsinoides Cham. & 
Schlechtend., Linnaea 6: 413. 1831. D. alsi- 
noides var. australis Standl., loc. cit. 1 292. 1 938. 
D. alsinoides var. mollis Standl., loc. cit. 1292. 
1938. D. australis (Standl.) L. O. Williams, 
Fieldiana Bot. 24, pt. 11:61. 1972. Figure 3. 

Herbs or weak-stemmed climbers, leafy stems 0.3-2 
mm thick when dry, quadrangular or with 2 or 4 lon- 
gitudinal ridges or wings ca. 0.5 mm high, sparsely and 
minutely (ca. 0. 3 mm) puberulent or glabrescent; stipules 
0.5-2.5 mm long, narrowly triangular or linear, decid- 
uous. Leaves with petioles ( l-)2-8 mm long, 0.2-0.5 mm 
broad; leaf blades (5-) 7-30 mm long, (3-)4-14 mm broad, 
ovate to ovate-elliptic or narrowly ovate (rarely lanceo- 
late), apex gradually narrowed and acute with sharp tip, 
base abruptly narrowed or rounded and obtuse to trun- 
cate, decurrent on petiole, drying membranaceous 
(translucent), glabrous or minutely puberulent above the 
midvein on the upper surface, sparsely pubescent be- 
neath with thin hairs ca. 0.3 mm long, 2 veins 1-3/side, 
often with subpalmate venation in broadly ovate leaves. 
Inflorescence of solitary flowers in leaf axils, usually only 
1 flower per node, pedicels 1-5 mm long, glabrous. Flow- 
ers ca. 3-4 mm long, hypanthium ca. 0.7 mm long, calyx 
to 0.2 mm long, truncate; corolla ca. 3 mm long, greenish 
purple, tube ca. 1.5 mm long. Fruits 4-6 mm long, sub- 
globose when 1 -seeded, deeply 2-parted and 6-8 mm 
broad when 2-seeded, becoming fleshy and blue at ma- 
turity, lustrous, drying black, usually glabrous; seeds of- 
ten curved and reticulate. 



122 



FIELDIANA: BOTANY 



Climbing plants of the shaded understory in 
montane evergreen wet forest formations, from 
1500 to 2800(-3100) m elevation. Flowering and 
fruiting in September-June. The species (in a wide 
sense) ranges from Mexico to the Chiriqui High- 
lands of Panama. 

Didymaea alsinoides is recognized in Costa Rica 
by its slender-stemmed clambering habit, thin, 
usually ovate leaves, minute flowers usually soli- 
tary at each node, and small fleshy blue-black fruit 
that are subglobose or deeply two-lobed. The pre- 
ceding description is based on Costa Rican and 
Panamanian collections referred to as Didymaea 
australis by Williams. They differ from the more 
northerly collections in having more ovate leaves 
often truncate at the base. Recognizing the various 
morphological and geographic forms of D. alsi- 
noides as separate species seems unwise, insofar 
as nearly all live in the same kind of montane 
habitats and there is considerable morphological 
variation in any one area; compare Williams' 
treatment (in Standley & Williams, 1975, pp. 60- 
63). Compare Nertera granadensis with smaller 
leaves and orange fruit. 



Diodia Linnaeus 

Annual or perennial herbs or small shrubs, erect or 
scandent, usually woody at the base, stems often much- 
branched near the base, terete or 4-angled, glabrous or 
pubescent; stipules interpetiolarand united with the leaf 
bases to form a broad sheath, usually bearing 2-12 slen- 



der awns from the truncated or rounded distal edge of 
the sheath, persisting. Leaves opposite or pseudoverti- 
cillate (with smaller axillary leaves), sessile or short-pet- 
iolate: leaf blades mostly narrow and often scabrous, 
entire or serrulate with minute scabrous hairs along the 
margin. Inflorescences axillary and sessile, capitulate or 
fasciculate (rarely of solitary axillary flowers) and often 
verticillate, long-spicate when the distal subtending leaves 
are reduced, subtended by the leaves and stipular sheaths, 
flowers usually subsessile. Flowers bisexual and radially 
symmetrical, small or minute, hypanthium obovoid to 
turbinate, calyx lobes 2^*; corolla funnelform to cam- 
panulate, white to pink or purplish, corolla tube short 
or long, throat glabrous to villous, corolla lobes 4(3, 5- 
6), valvate in bud; stamens 4 (3, 5-6), inserted in the 
corolla throat, filaments filiform, anthers dorsifixed. lin- 
ear-oblong; ovary 2-locular (rarely 3- or 4-locular), ovule 
solitary and ascending in each locule, affixed to the center 
of the septum, style filiform and exserted, bilobed or 
with 2 short branches. Fruits splitting into 2 mericarps 
(cocci), crustaceous to slightly woody, septicidal from 
apex (rarely splitting at the base), without a central axis, 
each mericarp indehiscent or opening slightly at the base; 
seeds ellipsoid, longitudinally sulcate on the inner face, 
rounded abaxially. 

A genus of about 35 species in the tropical and 
subtropical Americas and with a few species in 
Africa. The weedy growth habit, awned stipular 
sheaths, narrow leaves, small sessile flowers, and 
fruit of two one-seeded indehiscent mericarps help 
to distinguish this genus. Species of this genus may 
resemble species of Crusea and Spermacoce, but 
the leaves do not become pseudoverticillate. This 
treatment has benefited from the annotations and 
advice of C. D. Adams (pers. comm., 1991). 



Key to the Species of Diodia 

la. Plants essentially glabrous, often prostrate and restricted to the Caribbean seashore; leaves often 

closely clustered on short lateral branches (1-4.5 cm long); rarely collected D. serrulata 

Ib. Plants minutely to conspicuously pubescent, rarely prostrate and not restricted to the Caribbean 

shore; leaves not closely clustered on short lateral branches 2 

2a. Largest leaves usually less than 3 cm long, stems erect, stipular sheaths with usually glabrous awns 

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 1 545 mm long, caducous, densely hir- 
sute-sericeous on the outer surfaces. Leaves closely 
crowded at the tips of branchlets, subsessile with petioles 
2-6(-10) mm long and ca. 2 mm thick, densely hirsute; 
leaf blades (7-)9-19 cm long, (3-)4-7.5 cm broad, ob- 
long-obovate to narrowly obovate, widest at or above 
the middle, apex abruptly narrowed and slender acu- 
minate or caudate-acuminate, the narrow (ca. 2 mm) tip 
5-13 mm long, gradually narrowed to the cuneate base, 
drying thin-chartaceous, often brown, with long ( 1 .5-2.5 
mm) thin straight or slightly crooked hairs on upper and 
lower surfaces, 2 veins 5-8/side, 3 veins subparallel 
between the secondaries. Male flowers 8-15 and sub- 
capitate or fasciculate-cymose (tightly clustered at the 
tips of stems), 1 5-20 mm long, outer surfaces of calyx 
and corolla densely sericeous with ascending hairs, calyx 
4-5 mm long, calyx lobes 6-7 and equalling the tube; 
corolla 11-15 mm long, white, corolla lobes 6-8, 5-8 
mm long, equalling or longer than the tube. Female flow- 
ers l(-3) subsessile at apex of branchlets, hypanthium 
ca. 8 mm long, densely sericeous, with hairs 2-3 mm 
long, calyx lobes ca. 4 mm long, linear. Fruits 1-3 at the 
tips of branchlets, subsessile, ca. 22 mm long, 1 2 mm 



Small trees of lowland rain forest formations of 
the Golfo Dulce region, from 10 to 200 m ele- 
vation. Flowering in March and May-June; fruit- 
ing in July-August and October. This species is 
known only from southernmost Costa Rica. 

Duroia costaricensis is recognized by the sub- 
sessile thin hirsute obovate leaves clustered at the 
ends of stems, the terminal clusters of subsessile 
sericeous flowers, and the hirsute subsessile fruits 
at the tips of branches. The type collection (Pittier 
6803 us) came from Sierpe, but most of the other 
collections come from near Rincon de Osa. Duroia 
hirsuta (Poepp. & Endl.) Schumann of South 
America has inflated stems, longer petioles, gen- 
erally longer leaves, pedunculate male flowers, and 
larger fruit. Duroia genipifolia, now Randia geni- 
pifolia (Standl. & Steyerm.) Lorence, of Guatemala 
has less hirsute leaves with better denned petioles 
and more elliptic blades. All three species are sim- 
ilar in appearance. This species also resembles 
Costa Rican material of Hippotis. 



126 



FIELDIANA: BOTANY 



Duroia utleyorum Dwyer, sp. nov. 

Frutices vel arbores 1.5-7 m altae. Foliae lamina ob- 
longa vel oblongo-elliptica, 10-23 cm longa, 6-14 cm 
lata; venis lateralibus 7-1 1, hirsutulis; petiolis 6-23 mm 
longis. Flores lobis calycinis ca. 1 mm longis; corollae 
tubo ca. 12 mm longo, lobis ca. 10 mm longis. Fructus 
immaturi solitarii terminales. 

TYPUS Liesner /774(holotypuscR, isotypus MO), from 
Rincon de Osa, Costa Rica. 



Shrubs or small trees, (1.5-)4-7 m tall, leafy stems 2- 
5 mm thick, densely pubescent with thin straight yel- 
lowish hairs 0.3-0.7 mm long; stipules ca. 9 mm long 
and 4 mm broad, ovate-lanceolate with a sharp acu- 
minate apex, united (intrapetiolar) to form a short (1 
mm) basal tube, with thin brown margins and hirsute. 
Leaves with petioles 6-23 mm long, 1-3 mm thick, hir- 
sutulous with yellowish hairs; leaf blades 10-23 cm long, 
6-14 cm broad, broadly oblong to broadly obovate or 
elliptic-oblong, apex short-acuminate with tip 5-1 5 mm 
long, base obtuse to rounded and subtruncate, drying 
chartaceous, brown, pubescent on the major veins above, 
densely hirsute on the veins beneath with yellowish hairs 
ca. 0.7 mm long, sparsely hirsutulous between the veins, 
2 veins 7- 11 /side. Inflorescences of solitary terminal 
female flowers, on pedicels ca. 8 mm long; male inflo- 
rescences of 3-7 terminal sessile flowers. Flowers whitish 
pilosulous to sericeous; 3 flowers with calyx tube 5-6 
mm long, calyx teeth ca. 1 mm long, remote; corolla 
(preanthesis) white and probably salverform, tube 12 
mm long, lobes 10 mm long, convolute and apparently 
lanceolate. Fruits solitary and terminal, the immature 
fruit ca. 1 5 mm diam. and globose with persisting calyx 
10 mm high and 8 mm diam., densely hirsutulous with 
hairs ca. 0.5 mm long. 



A species of lowland (10-300 m) rain forest for- 
mations on both the Caribbean and Pacific slopes 
of Costa Rica. Young flowers were collected in 
February on the Osa Peninsula (Jimenez et al. 650 
CR, Liesner 1774 & 1853 CR, MO); young fruits 
were collected along the Rio Sarapiqui in May 
(Hartshorn 1486 CR). 

Duroia utleyorum is recognized by its larger 
broad leaves, sessile terminal flowers, and lowland 



evergreen forest habitat. This poorly known spe- 
cies is more likely to be confused with species of 
Randia than with its local cogener. Many impor- 
tant collections have been made in Costa Rica by 
Kathleen and John Utley. 



Elaeagia Weddel 

Trees or large shrubs, branchlets terete and puberulent; 
stipules interpetiolar (also intrapetiolar in some species), 
often covering the shoot apex, caducous or deciduous. 
Leaves opposite, petiolate or subsessile; leaf blades entire 
and pinnately veined, usually puberulent, thin-charta- 
ceous to coriaceous, domatia present or absent. Inflo- 
rescences terminal, paniculate (rarely racemose), many- 
flowered, pedunculate. Flowers bisexual, radially 
symmetrical, usually small, hypanthium hemispheric to 
turbinate, often sulcate on opposite sides, calyx tube cu- 
pular and spreading or inflated, calyx lobes 5 or none, 
short or inconspicuous; corolla campanulate to funnel- 
form, corolla white to yellow-white, tube usually short 
and broad, corolla lobes 5, oblong, rotate to reflexed, 
convolute to slightly imbricate in bud; stamens 5, in- 
serted between the corolla lobes, filament densely pu- 
berulent at the base, anthers dorsifixed, oblong, exserted; 
ovary 2-locular, placentas peltate and attached to the 
septum, ovules many in each locule, crowded and ver- 
tical, style short, stigmas 2 with obtuse tips. Fruits small 
woody capsules, crowned or encircled by the persistent 
calyx, loculicidally and basipetally dehiscent into 2 valves, 
the valves splitting at the top; seeds many, minute, elon- 
gate with membranous or winged testa. 



A genus of about 1 2 species in Mexico, Central 
America, Cuba, and tropical South America. Elae- 
agia utilis Weddel of Colombia is the source of a 
protective lacquer. The genus was misspelled as 
Elaeagnia in the Flora of Panama (1980). Three 
mesoamerican species of Elaeagia were compared 
by Lorence (in Bol. Soc. Bot. Mexico 45: 65-69. 
1 983). The small rounded capsules resemble those 
of Chimarrhis. 



Key to the Species of Elaeagia in Costa Rica 

la. Leaf blades usually 9-25 cm wide, often subsessile and sometimes auriculate at the base, with 1 1- 
19 pairs of secondary veins; inflorescences usually much-branched panicles E. auriculata 

Ib. Leaf blades usually 3-9 cm wide, clearly petiolate and gradually narrowed to the base, with 6-12 
pairs of secondary veins; inflorescences with few lateral branches (paniculate with many branches 
in E. myriantha) 2 

2a. Calyx cup entire distally; flowers separate and borne on long (4 mm) slender pedicels 

E. nitidifolia 

2b. Calyx cup with undulate or minutely lobed margin; flowers crowded and subsessile in small groups 
of2-5 . . 3 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



127 



3a. Corolla tubes 1-1.5 mm long, lobes ca. 1.7 mm long; leaf blades with 7-9 major 2 veins per side, 
without tufted domatia along the 2 veins; Cordillera de Tilaran E. uxpanapensis 

3b. Corolla tubes 2.2-3.4 mm long, lobes 0.4-1 mm long; leaf blades with 9-12 major 2 veins per side, 
often with tufted domatia along the 2 veins; General Valley E. myriantha 



Elaeagia auriculata Hem si.. Diagn. PI. Nov. Mex- 
ic. 32. 1879. Figure 39. 

Shrubs or small trees, 3-10 m tall, leafy branchlets 3- 
8 mm thick, puberulent or glabrescent; stipules 20-50 
mm long, to 20 mm broad, oblong and rounded at apex, 
reddish brown and with subparallcl venation. Leaves 
with petioles 0-10 mm long, 1.5-3 mm thick, often dif- 
ficult to see on auriculate leaves; leaf blades ( 1 2-) 1 5-40 
cm long, (6-)9-25 cm broad, elliptic-obovate to broadly 
elliptic or obovate, apex abruptly narrowed or rounded 
and obtuse to short-acuminate, gradually narrowed to 
the cuneate or auriculate base, drying chartaceous to 
subcoriaceous, dark brown above, upper surface gla- 
brous to short (0.2-0.5 mm) pubescent, lower surfaces 
sparsely to densely pubescent with short stiff hairs, 2 
veins 1 l-16(-19)/side, 3 veins usually subparallel and 
the 4 veins raised beneath. Inflorescences 12-20 cm 
long, to ca. 20 cm broad, many flowered and much- 
branched panicles, solitary or 3 from the terminal leaf- 
bearing node, primary peduncles 3-5 cm long, with 3- 
4 pairs of 1 opposite lateral branches, puberulent, flow- 
ers in racemose or cymose distal branches, pedicels 1-5 
mm long. Flowers 5-7 mm long, hypanthium ca. 1 mm 
long, calyx cup ca. 1 mm long (including the lobes), calyx 
lobes 0.5-1 mm long and 1.5 mm broad, glabrous; co- 
rolla white, tube 1-2 mm long, lobes 1-2 mm long and 
becoming reflexed; stamens with filaments 2 mm long, 
glabrous distally, anthers ca. 1.5 mm long. Fruits 3-5 
mm long and 3-4.5 mm broad, short-ovoid and broadly 
rounded, calyx persistent and visible on the lower part 
of the capsule, opening at the top (but not splitting to 
the base), becoming dark brown or black; seeds 0.6-1 
mm long, body of the seed 0.2-0.4 mm long, with a thin 
membranous wing at opposite ends. 

Trees of evergreen lower montane cloud forest 
and rain forest formations, along the central high- 
lands, from Volcan Tenorio in the Cordillera de 
Guanacaste to San Vito de Goto Brus, and on the 
Caribbean slope, ranging from 350 to 1700 m el- 
evation. Flowering in November-March; fruiting 
in February-June. The species ranges from Gua- 
temala and Honduras to eastern Panama. 

Elaeagia auriculata is recognized by the large 
oblong stipules, short petioles, large broad leaves 
often auriculate at the base, large terminal pani- 
cles, small flowers with truncated calyx lobes and 
very short corolla tubes, and short rounded cap- 
sules opening only at the top. The very large leaves 
with an auriculate base obscuring the petiole are 
very distinctive, but not all specimens have this 
kind of leaf base (see below). The small capsules 
make this species look like species ofChimmarhis. 



Elaeagia karstenii Standley was recorded for 
Costa Rica's flora (Standley, 1938) on the basis of 
a single sterile collection: Valeria 1665 F, from 
Tapanti at 1300 m. The longer (10-14 mm) and 
slender (1.5-2 mm) petioles and the smaller 
sparsely puberulent leaves do resemble some ex- 
amples ofE. karstenii from South America. How- 
ever, longer petioles with cuneate (not auriculate) 
lamina bases do occur in specimens that otherwise 
appear to be typical of E. auriculata: Lent 3734 
and Molina et al. 17346 (note: both of the latter 
collections come from below 1000 m elevation). 
Thus, it is possible that Valeria 1665 is an aberrant 
juvenile shoot of an E. auriculata plant. Alter- 
natively, expanding the concept of E. auriculata 
to include material with longer petioles, cuneate 
leaf bases, and smaller more glabrous leaves may 
require the inclusion of South American speci- 
mens and synonymizing E. karstenii. 



Elaeagia myriantha (Standl.) C. M. Taylor & 
Hammel, Novon (in press). Sickingia myrian- 
tha Standl., Publ. Field Columb. Mus., Bot. Ser. 
7: 27. 1930. Holtonia myriantha (Standl.) 
Standl., Trop. Woods 30: 37. 1932. Simira myr- 
iantha (Standl.) Steyerm., Mem. New York Bot. 
Gard. 23: 306. 1972. Deppea panamensis Dwyer, 
Ann. Missouri Bot. Gard. 67: 145. 1980. 



Trees, 8-25 m tall, leafy stems 2-7 mm thick, gla- 
brous, drying brownish or grayish; stipules 3-5 mm long, 
short-tubular with rounded lobes, often splitting between 
the leaf bases and remaining entire above the petioles 
(ligulate), translucent, persisting. Leaves with petioles 5- 
20(-30) mm long, 0.8-2 mm thick, glabrous, drying red- 
dish brown to dark brown; leaf blades (5-)9-20 cm long, 
(2-)3.5-7(-8) cm broad, elliptic-oblong to elliptic or el- 
liptic-obovate, apex acuminate with tip 10-15 mm long, 
base gradually narrowed and acute or cuneate and de- 
current on petiole, drying chartaceous, greenish to dark 
brown, glabrous above, glabrous or minutely (0.05 mm) 
papillate-puberulent on the veins beneath, 2 veins 9- 
12/side, domatia with short hairs present in the vein 
axils. Inflorescences 10-22 cm long, 12-18 cm broad, 
pyramidal with usually opposite 1 branches, peduncles 
8-40 mm long, 1.5-2.5 mm thick, minutely puberulent, 
larger (2 cm) narrow leaf-like bracts present or absent, 
with smaller (0.5-3 mm) bracts and bracteoles subtend- 
ing branches and flowers, flowers usually subsessile in 
groups of 2-5. Flowers glabrous externally, hypanthium 
1-1.5 mm long, obconic, calyx cup 0.1-0.3 mm long. 



128 



FIELDIANA: BOTANY 



1.3 mm diam., calyx lobes 0.2-0.4 mm long, broadly 
triangular; corolla tubular-campanulate, cream white, 
tube 2.2-3.4 mm long, ca. 2 mm diam., lobes 5, 0.4-1 
mm long, triangular, slightly imbricate in bud; filaments 
exserted 0.5-2 mm, anthers ca. 1.5 mm long; stigmas 1- 
2 mm long. Fruits ca. 3 mm long, 3 mm broad at the 
top when opened, dark brown; seeds 0.3-0.5 mm long, 
broad, reticulate. 

Collected with flowers in January in the General 
Valley at 975 m elevation (Skutch 2387 us, the 
only Costa Rican collection). This species is also 
known from the Andes of Colombia and Vene- 
zuela. 

Elaeagia myriantha is distinguished by the large 
pyramidal terminal panicles of small white flowers 
with broad cylindrical corolla tube and minute 
lobes and the almost glabrous leaves with pubes- 
cent domatia in the vein axils. These are unusual 
in that they are found in the axils of some 3 veins 
as well as along the midvein. The Colombian type 
(H. H. Smith 1810F) has smaller leaves than most 
other collections. 



Elaeagia nitidifolia Dwyer, Ann. Missouri Hot. 
Gard. 67: 157. 1980. Chiococca jefensis Dwyer, 
loc. cit. 67: 88, f. 19. 1980. Figure 39. 

Trees to 1 2 m tall, leafy stems 2-4 m thick, glabrous, 
becoming terete; stipules 3-5 mm long, to 4 mm broad, 
cylindrical at first but splitting, rounded above the pet- 
iole and deeply split between the petioles (becoming lig- 
ulate), glabrous. Leaves closely clustered distally, petioles 
6-17(-25) mm long, 1-2 mm thick, glabrous; leaf blades 
10-23 cm long, 3.5-8 cm broad, oblanceolate to nar- 
rowly oblong-obovate or elliptic-oblong, apex short-acu- 
minate, tip 5-8 mm long, gradually narrowed to the 
cuneate base and decurrent on petiole, drying charta- 
ceous, subglabrous above, minutely (0.05-0. 1 mm) pa- 
pillate-puberulent on the veins beneath, 2 veins 8-12/ 
side. Inflorescences solitary or 3, terminal, 8-1 6 cm long, 
open paniculate with distant cymose flower groups, pe- 
duncles 3-5 cm long, glabrous, pedicels 3-10 mm long, 
slender. Flowers with hy pan t h i urn and calyx continuous, 
ca. 3 mm long and 3-4 mm broad distally, calyx cup 
entire distally; corolla white, tube 1-2 mm long, lobes 
5, 3-4 mm long, 1.3-2 mm broad, oblong; anthers 1.5- 
2 mm long; stigma lobes 2-2.5 mm long. Fruits to 6 mm 
long and 10 mm broad, turbinate, truncated distally. 



Trees of wet evergreen forest formations of the 
Caribbean slope at 400-800 m elevation. The spe- 
cies is only known from the Rara Avis site in 
Heredia and was collected flowering in September 
(O. Vargas 128 CR, MO). The species is also known 
from central and eastern Panama. 

Elaeagia nitidifolia is distinguished by its un- 
usual ligulate stipules, the open few-flowered in- 



florescences, long pedicles, and cupulate calyx with 
entire margin. Specimens may resemble Rustia oc- 
cidentalis (with porate anthers) and Simira myri- 
antha (with smaller corolla lobes and domatia). 



Elaeagia uxpanapensis D. Lorence, Bol. Soc. Bot. 
Mex. 45: 66. 1983 (1984). Figure 39. 

Trees, 15-40 m tall, 55-80 cm dbh, leafy stems 3-9 
mm thick, flattened in early stages, glabrous and becom- 
ing terete; stipules 8-16 mm long. 3-7 mm broad, ovate 
lanceolate with overlapping margins, acute at the apex, 
glabrous and drying dark, deciduous. Leaves with peti- 
oles 3-12 mm long, ca. 1.5 mm thick, glabrous, drying 
dark; leaf blades 6-15(-20) cm long, 3-7(-9) cm broad, 
broadly elliptic to broadly oblong or slightly elliptic- 
obovate, apex abruptly narrowed and bluntly obtuse, 
base cuneate and slightly decurrent on petiole, drying 
stiffly chartaceous to subcoriaceous, dark grayish brown 
above, glabrous above, with a few thin yellowish hairs 
0.3-1 mm long along the side of the midvein beneath 
and in leaf axils (= domatia?), 2 veins 7-9/side. Inflo- 
rescences solitary and terminal or axillary to distal leaves, 
6-14 cm long (perhaps enlarging in fruit), narrowly pa- 
niculate, peduncles 2.5-5 cm long, glabrous, proximal 
lateral branches 1-6 cm long, usually minutely ap- 
pressed-puberulent, flowers sessile in opposite or ter- 
minal glomerules of 2-5 flowers, bracts 0.5-3 mm long. 
Flowers ca. 4 mm long, yellowish green, hypanthium 1- 
1.8 mm long, minutely puberulent at the base, calyx 
lobes 0.5-1 mm long; corolla white, salverform to short - 
funnelform, glabrous externally, tube 1-1.5 mm long, 
scarcely exceeding the calyx lobes, lobes 4-5, ca. 1 .7 mm 
long and 1 mm wide, bluntly rounded distally, with long 
hairs at the mouth of the tube and base of lobes within; 
stamens 4-5, filaments 2-4 mm long, anthers 0.6-0.7 
mm long, oblong; style and stigma 3-4 mm long, stig- 
matic lobes 1 mm long. Fruits not seen (probably ca. 2 
mm long and 2.5 mm broad). 

Only known from the wet Caribbean slopes of 
the Cordillera de Tilaran at ca. 900 m elevation. 
Immature flowers were collected in July (Haber & 
Bella 1928 CR, MO) and mature flowers in May 
(Herrera 600 CR, MO). This species is known only 
from the Rio Penas Blancas valley below Mon- 
teverde and the Reserva Forestal de San Ramon 
(Gomez- Laurito 12065 CR), in Alajuela Province. 
The species is also known from southern Mexico. 

Elaeagia uxpanapensis is recognized by the op- 
posite subsessile flower clusters along the branches 
of the inflorescence, the very small flowers, and 
the two-locular ovule with placentas borne on the 
septum. The large size of the trees, flattened young 
leafy stems, and occasional domatia along the 
midvein are additional distinctive characteristics. 
When first discovered in Costa Rica, specimens 
were thought to be a species related to Warszewic- 
zia schwackei Schum. of South America. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



129 



Exostema L. Richard 

Shrubs or trees, branchlets usually terete, glabrous or 
puberulent; stipules interpetiolar, small, entire or bifid, 
deciduous or persisting. Leaves petiolate or subsessile; 
leaf blades entire, drying membranaceous to subcoria- 
ceous, domatia sometimes present. Inflorescences ter- 
minal or axillary panicles with the distal flowers usually 
in groups of 3, or of solitary axillary flowers, with or 
without bracts, pedicels present. Flowers bisexual, ra- 
dially symmetrical (or somewhat bilaterally symmetrical 
by curvature of the corolla tube), small to large, hypan- 
thium cylindrical or obovoid, calyx with (4-)5(-6) lobes, 
lobes broad or narrow; corolla short or long, salverform, 
tube often long and narrow, throat glabrous or barbate, 
corolla lobes (4-)5, oblong or linear, imbricate in bud 
with 2 exterior; stamens 5 inserted at the base of the 
corolla tube, filaments essentially free, elongate and fi- 



liform, anthers basifixed, linear, usually exserted; ovary 
2-locular, placentas peltate on the septum with many 
ascending ovules, style filiform, usually exserted, stigma 
capitate or bilobed. Fruits capsular, woody, 2-locular, 
ellipsoid to cylindrical, septicidal from apex, bivalved, 
valves entire or bipartite from the apex; seeds many, 
imbricate, flattened and oblong, testa forming a thin 
marginal wing around the body of the seed. 

A genus of ca. 35 species of the American trop- 
ics, especially abundant in drier vegetation of the 
West Indies. The flowers with narrow corolla lobes 
almost equalling the length of the slender corolla 
tube and the long linear exserted anthers make the 
flowers quite distinctive. Our species are confined 
to deciduous or partly deciduous vegetation. 



Key to the Species of Exostema 

la. Flowers solitary in the axils of leaves, 6-8 cm long, corolla lobes 4-5; fruit ca. 1 cm broad, erect; 
leaf blades usually clearly decurrent on the petiole, domatia absent E. caribaeum 

Ib. Flowers more than 1, borne in small panicles axillary to distal leaves or terminal, 2-3 cm long; 
corolla lobes 6; fruit 34 mm broad, pendulous; leaf blades very slightly decurrent on the petiole, 
domatia often present E. mexicanum 



Exostema caribaeum (Jacq.) Roem. & Schult., Syst. 
Veg. 5: 19. 1819. Cinchona caribaeum Jacq., 
Enum. PI. Carib. 16. 1760. E. longicitspeOeTsl., 
Vidensk. Meddel. Dansk. Naturhist. Foren. 
Kjobenhaven 1852: 48. 1853. Figure 31. 

Shrubs or small trees to 8 m tall, leafy branchlets 1 .2- 
3.5 mm thick, glabrous; stipules l-3(-5) mm long, 2-3 
mm broad at the base, triangular to subulate, glabrous 
on the abaxial surface but slightly ciliate along the distal 
edge, persistent. Leaves evenly spaced along the stems, 
petioles 6-1 2(- 16) mm long, 0.4-0.8 mm thick; leaf blades 
4-1 1 cm long, 1.5-5 cm broad, elliptic-ovate to ovate, 
apex gradually narrowed and acuminate, tip 1-2 cm long, 
base obtuse to acute and decurrent on petiole, drying 
thin-chartaceous and dark, glabrous above, glabrous or 
pubescent with thin curved whitish hairs ca. 0.4 mm 
long beneath, often with short (0.5 mm) hairs (domatia) 
in the vein axils beneath, 2 veins 4-5/side. Inflores- 
cences absent, the flowers solitary in distal leaf axils, 
bracts ca. 1 mm long, pedicels 4-10 mm long, ca. 0.5 
mm thick (dry), glabrous. Flowers 6-8 cm long, hypan- 
thium 3-5 mm long, 2-3 mm diam., glabrous, calyx 
lobes 0.5-1 mm long, broadly triangular, corolla white, 
glabrous, tube 23-45 mm long, 2-3 mm diam., often 
curved, lobes 25-40 mm long, ca. 2 broad and linear, 
becoming recurved; stamens 5. long-exserted, anthers 
16-26 mm long, 0.2-0.3 mm broad (dry); stigma 2 mm 
long. Fruits 7-15 mm long, 6-14 mm broad, oblong- 
ellipsoid, smooth and dark brown externally; seeds 3-6 
mm long and 3-4 mm broad, with a thin brownish wing 
on all sides and slightly longer at the 2 ends, body of the 
seed ca. 5 mm long and 3 mm broad. 



Trees of deciduous and partly deciduous forests 
of the Pacific slope, from near sea level to 300 m 
elevation (to 1300 m in Guatemala). Rarely col- 
lected in Costa Rica (Guanacaste and adjacent 
Puntarenas provinces). In Central America flow- 
ering in June-August; fruiting in October-January 
and April. The species ranges from central Mexico 
to northern Costa Rica and the West Indies. 

Exostema caribaeum is recognized by its re- 
striction to seasonally very dry vegetation, the long 
flowers solitary in leaf axils, the long narrow co- 
rolla tube, long corolla lobes, and long slender ex- 
serted anthers. The woody bivalved fruit splitting 
down the broader side and the seeds surrounded 
by a short thin wing are also distinctive. We have 
not seen Oersted's type of E. longicuspe from Pun- 
tarenas (cf. Standley, 1938, p. 1295). This species 
is called "caribee bark tree" and "princewood" in 
the Caribbean, where the wood is used for making 
handles. The bark is used for treating fevers and 
malaria in Mexico. 



Exostema mexicanum A. Gray, Proc. Amer. Acad. 
Sci. 5: 180. 1 86 1 . Figure 40. 

Small to medium-sized trees to 20 m tall with trunks 
to 40 cm dbh, leafy branchlets 1.3-4 mm thick, glabrous 



130 



FIELDIANA: BOTANY 



to sparsely puberulent, lenticels conspicuous, stems be- 
coming grayish; stipules l-3(-4) mm long, triangular and 
acuminate to short-tubular and cuspidate, deciduous. 
Leaves well spaced along the stems, petioles 4-10(-15) 
mm long; leaf blades (5-)6-15(-18) cm long, (2-)3- 
7.5(-10) cm broad, ovate-oblong to elliptic-oblong, apex 
long-acuminate, base obtuse to rounded and subtrun- 
cate, usually glabrous above and below except for the 
minute (0.2 mm) hairs (domatia) in the vein axils be- 
neath, 2 veins 6-9/side. Inflorescences 3-6(-l 2) cm long, 
4-7(-10) cm broad, terminal or axillary to distal leaves, 
primary peduncles 1.5-4 cm long, glabrous or minutely 
puberulent, trichotomous, lenticellate, bracts 1-2 mm 
long, distal flowers in groups of 3, often crowded, ped- 
icles 2-4 mm long, minutely (0. 1 mm) puberulent. Flow- 
ers ca. 2 cm long, hypanthium 1 .5-2.4 mm long, glabrous 
or sparsely and minutely puberulent, calyx lobes 5-6, 
very small (0.5 mm) and deltoid/acute; corolla white to 
yellowish white or yellow, subglabrous or with thin curved 
whitish hairs, tube 7-10 mm long, 0.7-1.3 mm diam., 
lobes 6, ca. 9-12 mm long, 1 .5 mm broad, oblanceolate- 
linear; stamens 6, filaments to 1 6 mm long, anther 3-4 
mm long, 0.2-0.3 mm broad; style to 2 cm long, stigmas 
clavate, 0.6 mm long. Fruits 10-14 mm long, 3-4 mm 
broad, obovoid-clavate to narrowly oblong-obovoid, 
surface glabrous and dark brown with whitish lenticels, 
the 2 valves each splitting in 2 at the top; seeds 6-9 mm 
long, 2-3 mm broad, body of the seed ca. 2.5 x 1 .5 mm, 
oblong, wing often lobed on 1 end. 

Trees of deciduous and partly deciduous forests 
of the Pacific slope, from near sea level to 900 m 
elevation (to 1 300 m in Guatemala). Flowering in 
July-September; fruiting in July and September- 
February. The species ranges from Tamaulipas, 
Mexico, to central Panama. 

Exostema mexicanum is distinguished by its de- 
ciduous habitat and thin leaves, close clusters of 
narrow-tubed flowers in short axillary or terminal 
panicles, the long narrow corolla lobes, exserted 
linear anthers, and woody bivalved capsules with 
winged seeds. The bark has been boiled to make 
a treatment for malaria and fevers. This species is 
rarely collected in Costa Rica. 



trapetiolar, often forming a tubular sheath above the leaf 
base, short-triangular to long-aristate at apex (with 2 
awns per node), persistent or deciduous. Leaves often 
held in a single plane, petiolate (rarely subsessile); leaf 
blades entire, usually oblong to lanceolate, usually gla- 
brous, lacking domatia, pinnately veined. Inflorescences 
terminal (rarely axillary), 1 -many-flowered, corymbose 
to umbellate (rarely capitate), pedunculate, peduncle and 
branches often with a pale blue or purple color like the 
flowers, pedicels usually present. Flowers bisexual and 
radially symmetrical, monomorphic or distylous, hy- 
panthium small, ovoid to turbinate, terete or angular, 
calyx cupular to short-tubular, truncate and entire to 
4-lobed, persistent; corolla usually salverform, white to 
blue or lavender, tube short or long, usually narrow, 
glabrous, corolla lobes 4(-5), linear to lanceolate, valvate 
in bud, spreading or reflexed; stamens 4 (5. 6), inserted 
on the tube or at the throat, anthers linear, dorsifixed. 
linear, included or exserted; ovary 1-locular (rarely 
2-locular in early stages), ovules 2 (less often 1), erect 
from a basal placenta, style filiform, with 2 short branch- 
es. Fruit baccate or drupaceous, transversely oblate (reni- 
form) to subglobose, often broader than long, smooth or 
costate, deep blue to blue-black, 1-locular and 1- (rarely 
2-) seeded; seed (pyrene) horizontal, transversely in- 
dented (excavated) and somewhat reniform, with thin 
testa. 

A large genus of about 1 30 species, ranging from 
Mexico, Central America, and the West Indies 
through tropical South America. The unusual fruit 
(usually broader than long) and single seed help 
to distinguish this genus. A striking sky blue to 
lavender color of both the flowers and inflores- 
cences characterizes a number of our species. The 
usually four-parted flowers, short-tubular stipules 
with only a single apex on each side, and a well- 
developed submarginal vein are additional char- 
acters found in many species. The fruits tend to 
have a leathery exocarp in Faramea, in contrast 
to the succulent exocarps of Coussarea and Psy- 
chotria. Despite these unusual traits, specimens of 
some species may look very similar to some spe- 
cies of Psychotria, Coussarea, and Rudgea. 



Faramea Aublet 

Shrubs or small trees, branchlets terete, 4-angled or 
flattened, usually glabrous; stipules interpetiolar and in- 



Key to the Species of Faramea 

la. Flowers solitary or the inflorescence with 2-3 flowers (l^t flowers per node); rarely collected . 

Ib. Flowers few to many in pedunculate open inflorescences; commonly collected species 4 

2a. Leaf blades 12-20 cm long; peduncles 2-5 cm long, calyx and hypanthium ca. 12 mm long 

[Caribbean slope at 20-1 100 m elevation in Costa Rica] F. pauciflora 

2b. Leaf blades 1-4 cm long; peduncles less than 2 cm long, calyx and hypanthium less than 5 
mm long 3 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



131 



3a. Inflorescences terminal; shrubs 3-4 m tall; Pacific slope, 1200-1700 m elevation 

F. capulifolia 

3b. Inflorescences axillary; herbaceous subshrubs ca. 1 m tall; Caribbean slope, ca. 300 m ele- 
vation F. myrtidfolia 

4a. Leaves sessile or subsessile and the leaf blade usually rounded at the petiole, petioles 0-5 mm long 
5 

4b. Leaves definitely petiolate or if subsessile the leaf blades not rounded at the petiole, petioles 4-20 

mm long 7 

5a. Leaf blades narrowly oblanceolate, 10-16 cm long and 1.5-3.5 cm broad; Chiriqui Highlands 

at ca. 2000 m elevation F. scalaris 

5b. Leaf blades usually oblong, 15-27 cm long and 6-17 cm broad; 0-500 m elevation .... 6 
6a. Leaves strongly tripliveined, secondary veins united by a linear (melastome-like) lateral 

submarginal vein; southern Caribbean lowlands F. trinervia 

6b. Leaves not strongly tripliveined, secondary veins loop-connected near margin; Golfo Dulce 
area F. sessifolia 

7a. Leaves strongly tripliveined (like that of Melastomaceae); flowers and fruit bright blue; 0-800(-1000) 
m elevation F. suerrensis 

7b. Leaves not strongly tripliveined and lacking strong lateral veins near the margin (sometimes present 
in F. eurycarpa), the 2 veins often loop-connected near the margin with the submarginal vein 
arcuate; flowers blue or white 8 

8a. Inflorescences umbellate or fasciculate in the leaf axils, without lateral branches; flowers usually 
white 9 

8b. Inflorescences paniculate, corymbose or cymose, with conspicuous lateral branches from the pri- 
mary peduncle or central rachis; flowers blue or white 10 

9a. Inflorescences fasciculate, some flowers subtended by ovate bracts 5-1 1 mm long; flowers 

not drying black; leaf blades to 1 7 cm long, usually oblong; 0-300 m elevation 

F. parvibracteata 

9b. Inflorescences umbellate, lacking conspicuous bracts; flowers drying black; leaf blades to 1 1 
cm long, elliptic; 1000-2300 m elevation F. ovalis 

lOa. Flowers with the corolla tube 12-22 mm long and corolla lobes 8-18 mm long, white and usually 
drying black; fruit slightly broader than long, often with persisting calyx tube; stipular awns 4-18 
mm long [0-800 m elevation] F. occidentalis 

lOb. Flowers with corolla tubes 4-13 mm long, corolla lobes 3-7 (-10) mm long, blue or white and 
rarely drying black; fruit distinctly broader than long, a persisting calyx tube rarely present; stipular 
awns 1-6 mm long 11 

1 1 a. Leaf blades usually drying yellowish green beneath, with a prominent submarginal vein; inflores- 
cences robust with branches ca. 1 mm thick when dry; stipules early deciduous; corolla tube 6- 
1 mm long; 500-1 700 m elevation F. eurycarpa 

1 1 b. Leaf blades usually drying grayish or pale green beneath; inflorescences delicate with primary 
branches ca. 0.5 mm thick when dry; 0-1600 m elevation 13 

1 2a. Stipules clearly tubular and persisting on the leafy stems; corolla tubes 6-12 mm long, calyx lobes 
0.2-0.4 mm long F. multiflora 

\ 2b. Stipules quickly deciduous, tubes short and inconspicuous; corolla tubes 2-6 mm long, calyx lobes 
0-0.7 mm long 13 

13a. Corolla tube slender, 0.7-1.5 mm diam., blue; 0-400 m elevation F. stenura 

13b. Corolla tube broad, 1.5-2 mm diam., white; 1300-1400 m elevation F. hondurae 

Faramea capulifolia Dwyer, sp. nov. fructum solitarium gerenti. Floras non visi. Fructus glo- 

bosus, 6-9 mm longus, glaber, niger. 

TYPUS Folsom 4262 (holotypus MO, isotypus ?PMA), 

Frutices ad 4 m alti. Foliae lamina lanccolata. 2-6 cm from Cerro Pirre, ridgetop at 1200 m elevation, Darien, 
longa, 0.5-1.6 cm lata, acumine ad 10 mm longo, venis Panama, 
lateralibus 10-16; petiolis 2-4 mm longis; stipulis 2-8 

mm longis, vagina 1-5 mm longa. Inflorescentiae ter- Shrubs, 1-4 m tall, leafy stems 0.5^4 mm thick, gla- 

minales, pedunculo nullo vel 1 5-20 mm longo, glabro, brous with 2 or 4 barely distinct longitudinal ridges; 



132 FIELDIANA: BOTANY 



stipules 2-8 mm long, sheathing tube 1-5 mm long, 
truncated distally with 1 small narrow lobe (2/node), 
glabrous, deciduous. Leaves with petioles 24 mm long, 
ca. 0.5 mm thick, glabrous, sulcate above; leaf blades 2- 
6 cm long, 0.5-1.6 cm broad, lanceolate to narrowly 
ovate-elliptic, apex acuminate, base acute and slightly 
decurrent on petioles, drying chartaceous, olive green 
above, glabrous above and below, 2 veins 5-9/side and 
loop-connected ca. 1 mm from the leaf margin (major 
and intermediate secondaries often difficult to distin- 
guish). Inflorescences terminal and solitary, sessile or 
with peduncles to 2 cm long in fruit (after the loss of the 
distal leaf pair), peduncles ca. 0.4 mm thick and glabrous, 
pedicels 0-2 mm long, fruit solitary. Flowers solitary, 
ca. 10 mm long, glabrous, corolla ca. 8 mm long, rose- 
colored, tube ca. 5 mm long, ca. 0.9 mm diam. Fruits 
6-9 mm diam., glabrous, persisting calyx 0.5-1 mm high, 
surfaces smooth, brilliant blue or becoming black. 



Understory plants in wet cloud forests from 1 000 
to 1700 m elevation. Flowering in May; fruiting 
in June-July and December. This species is known 
only from the Cordillera de Tilaran area in Costa 
Rica and eastern Panama. 

Faramea capulifolia is recognized by its small 
leaves, glabrous parts, and solitary terminal flow- 
ers and fruits. The Costa Rican material has some- 
what smaller leaves (to 4.5 cm long) and blue fruits, 
whereas the type has leaves to 6 cm long and fruits 
becoming black. The fruits are subtended by a 
node where the leaves have fallen, with the distal 
internode appearing to be a peduncle. Flowering 
data was added in proof, based on Gomez- La urito 
12431 F. Also placed here are Barringer et al. 4194 
CR, F, Dryer 1711 F, and Zamora et al. 647 CR, F. 



Faramea eurycarpa J. D. Smith, Bot. Gaz. 44: 1 1 3. 
1907. F. bocaturensis Dwyer, Ann. Missouri Bot. 
Gard. 67: 163. 1980. Figure 45. 

Shrubs or small trees to 4(-8) m tall, leafy branchlets 
1.5-5 mm thick, glabrous and drying yellowish green; 
stipules to 1 6 mm long, united to form a tubular sheath 
3-8 mm long, 2-4 mm broad, with awns 1-3 mm long, 
deciduous. Leaves with petioles 3-7(-10) mm long, 1- 
1.5 mm thick, glabrous; leaf blades (5-)9-16(-19) cm 
long, (2-)3-5.5 cm broad, narrowly oblong to elliptic- 
oblong, apex obtuse or rounded and acuminate to cau- 
date-acuminate, tip 10-14(-20) mm long and ca. 2 mm 
broad, base cuneate (or slightly rounded at the petiole), 
drying chartaceous, usually yellowish green below with 
the midvein yellow, glabrous above and below, 2 veins 
9-14/side and loop-connected near the margin to form 
an arcuate submarginal vein 2-4 mm from the edge, with 
less distinct secondary veins paralleling the major sec- 
ondaries. Inflorescences terminal and solitary, to 14 cm 
long and 1 2 cm broad, pyramidal panicles with opposite 
bracteolate branches, glabrous, peduncles 2-5 cm long, 
stout and 1.3-3 mm thick (dried), the slender pedicels 



4-10 mm long. Flowers with hypanthium 0.7-1.5 mm 
long, calyx cup 0.5-1.5 mm long, calyx teeth 4 or 5, 0.3- 
1 mm long, narrow distally; corolla salverform, blue, 
purple, or white with purple apex, tube 6-10 mm long, 
1-1.5 mm diam., lobes 2-4 mm long, narrowly ovate; 
stamens 4, attached near the middle of the tube and 
subsessile, anthers 3-4 mm long; style 4-8 mm long, 
stigmas ca. 0.7 mm long. Fruits 6-8 mm long and 9-13 
mm broad, transverse-reniform to subglobose, blue, pur- 
ple, or black, drying dark, smooth or slightly rugose when 
dried, persisting calyx less than 1 mm high. 



Plants of the very wet evergreen cloud forests 
of the Caribbean slope, from (300-)500 to 
1200(-1700) m elevation. Flowering in March- 
June, September, and December; probably fruit- 
ing throughout the year. The species ranges from 
Costa Rica and Panama to Colombia and Ecua- 
dor. 

Faramea eurycarpa is recognized by the oblong 
leaves with slender "drip tips," greenish yellow 
color of the veins beneath when dried, deciduous 
stipules, arcuate submarginal vein, and small blue 
flowers with small corolla lobes. The inflorescence 
branches and deciduous tubular stipules are much 
thicker in texture than those of/", multiflora, which 
shares many of the same habitats as F. eurycarpa. 
Collections from the Chiriqui Highlands have a 
greater range of variation than those found in Cos- 
ta Rica, but they do not appear to be specifically 
distinct. The name F. bocaturensis Dwyer applies 
to the distinctive Chiriqui Highland material. 



Faramea hondurae Standl., J. Wash. Acad. Sci. 18: 
169. 1928. Figure 45. 

Shrubs, 3-4.5 m tall, leafy branchlets 1 .3-3 mm thick, 
glabrous; stipules forming a short (3-4 mm) tube, ca- 
ducous and leaving a transverse whitish scar. Leaves with 
petioles 6-16 mm long, 0.8-1.4 mm broad, glabrous; 
leaf blades (6-) 1 0- 1 9 cm long, ( 1 . 5-)3-5 cm broad, nar- 
rowly oblong to narrowly elliptic-oblong, apex long-acu- 
minate to caudate-acuminate, tip 12-20 mm long, base 
acute, drying chartaceous and dark green above, glabrous 
above and below, 2 veins ca. 14/side (but with less 
prominent 2 veins between the major), an arcuate sub- 
marginal vein present near (ca. 2 mm) the margin and 
connecting the secondaries. Inflorescences terminal or 
axillary to distal leaves, solitary or 3, 5-8 cm long, to 6 
cm broad, paniculate with 2 or 3 pairs of opposite 
branches, primary peduncles 20-30 mm long, bracts 2- 
6 mm long and mucronate, distal bracteoles 0.5-3 mm 
long, pedicels 2-5 mm long, slender, glabrous. Flowers 
glabrous, hypanthium ca. 1 mm long, turbinate, calyx 
tube 0.5-1 .5 mm long, campanulatc. with 4 minute lobes; 
corolla salverform, white, tube 2-4 mm long, 1 .5-2 mm 
diam.. lobes 1-2 mm long, obtuse; anthers ca. 1.5 mm 
long, style and stigma ca. 2.3 mm long. Fruits unknown. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



133 



A problematic species known only from two 
collections (Standley 36534 us, 37890 us the type) 
collected between 1 300 and 1 700 m elevation near 
Bajo La Hondura, San Jose, in March 1924. The 
short, relatively broad white corolla tubes make 
this species unique among our species of Faramea 
and make the generic placement doubtful. How- 
ever, a dissection of an ovary (Standley 36534) 
showed two ovules in a single locule, consistent 
with Standley's placement of this species in Fara- 
mea. 



pearance to F. stenura but the tubular stipules of 
F. multiflora are persisting, the calyx lobes are less 
well developed, and distal inflorescence branches 
lack the small bracts characteristic of F. stenura. 
Central American collections differ in minor ways 
(generally smaller thinner leaves and inflores- 
cences) from South American collections, but all 
appear to be part of the same wide-ranging species. 



Faramea myrticifolia Dwyer, sp. nov. 



Faramea multiflora A. Rich., ex DC., Prodr. 4: 
497. 1830. F. talamancarum Standl., Publ. Field 
Columb. Mus., Hot. Ser. 4: 332. 1929. Figure 
45. 

Shrubs or small trees, (l-)2-5(-6) m tall, leafy branch- 
lets 1-3.5 mm thick, terete and glabrous; stipules 4- 
8(-l 3) mm long, 2-3 mm broad but broader beneath the 
inflorescence, the tubular basal sheath 2-5 mm long, 
awns l-5(-8) mm long, persisting. Leaves with petioles 
3-7(-l 1) mm long, 0.7-1.8 mm thick; leaf blades (6-)8- 
13(-17) cm long, 1.5-4(-7) cm wide, oblong to oblong- 
obovate or narrowly elliptic-oblong, apex gradually or 
abruptly narrowed and caudate-acuminate or acuminate, 
tip 7-15 mm long and straight or falcate, base acute or 
obtuse, leaves drying thin-chartaceous to chartaceous 
and dark greenish above, 2 veins 6-12/side (and lesser 
parallel 2 veins between the major), united near the 
margin by a slender arcuate submarginal vein 2-4 mm 
from the leaf edge. Inflorescences terminal, solitary (or 
3), 5-1 4 cm long, to 9 cm broad, paniculate with opposite 
branches 1-2 cm long, bright blue, glabrous, primary 
peduncles 2-3(-6) cm long, basal branches subtended by 
small leaves or broad bracts 8-22 mm long, distal bracts 
absent, pedicels 3-7(-10) mm long, slender. Flowers gla- 
brous externally, hypanthium 0.5-1 mm long, calyx tube 
0.2-0.4 mm long, calyx lobes 4, 0. 1-0.4 mm long; corolla 
bright sky blue, salverform, tube 6-12 mm long, 1.2-2 
mm diam.. lobes 4, 4-9 mm long, 1.3-2.5 mm broad, 
ovate-oblong; anthers 2-3 mm long. Fruits 6-8 mm long, 
(8-) 10- 13 mm broad, oblate or transversely reniform, 
laterally compressed (oblong in cross-section), surface 
smooth and without costae, blue-black at maturity; py- 
renes solitary. 



Understory shrubs of wet evergreen forest in- 
teriors and forest edges, from 20 to 1600 m ele- 
vation. Flowering in every month but October 
(flowering is mainly in May-June at La Selva); 
fruiting throughout the year. The species ranges 
from northern Costa Rica to Brazil and Bolivia. 

Faramea multiflora is recognized by the thin 
smaller leaves (often drying grayish green be- 
neath), narrow acuminate apices, persisting tu- 
bular stipule sheaths, and bright blue flowers and 
inflorescences. This species is very similar in ap- 



Suffrutices 0.3-0.4 m alti; ramulis multis glabris. Foli- 
ae lamina elliptica 2-4 cm longa, 8-15 mm lata, glabra, 
venis lateralibus 6-8; petiolis 3-6 mm longis; stipulis 4- 
5 mm longis. Inflorescentiae axillares, flores 2-5 geren- 
tes, pedunculis 24 mm longis; pedicellis 1-2 mm longis. 
Flores glabri; calycis cupula ca. 0.5 mm longa, lobis ca. 
0.5 mm longis, lilacinis; corolla viridi-alba, ca. 4 mm 
longa. Fructus ca. 8 mm lati, globosus, azureus, glaber. 

TYPUS L. D. Gomez et al. 23401 (holotypus CR, iso- 
typus MO), from Las Brisas de Pacuarito, Limon, Costa 
Rica. 

Herbaceous subshrubs, ca. 40 cm tall, main stem un- 
branched for 20 cm and with many slender distal op- 
posite horizontal branches, leafy stems 0.5-1 .3 mm thick, 
glabrous, prominently 2- or 4-ridged; stipules 4-5 mm 
long, with a short (0.5-1 mm) truncated tube and single 
filiform central awn 3-5 mm long, persisting. Leaves 
with petioles 3-6 mm long, 0.3-0.9 mm thick, glabrous; 
leaf blades 2-4 cm long, 8-15 mm broad, elliptic to 
elliptic-oblong, apex acute or short-acuminate, base acute 
and decurrent on petiole, drying chartaceous, dark green 
above, glabrous above and below, 2 veins 6-8/side and 
loop-connected 1-1.5 mm from the margin. Inflores- 
cences axillary to leaves, 1 or 2/node, cymose umbellate 
with 2-5 flowers on a short (2-4 mm) glabrous peduncle 
drying dark, glabrous, bracts ca. 2 mm long and linear, 
pedicels 1-2 mm long. Flowers glabrous and drying 
blackish, hypanthium 0.5-1 mm long, calyx tube ca. 0.5 
mm long and truncated with linear lobes 0.5 mm long; 
corolla greenish white, ca. 4 mm long and 0.7 mm diam. 
in bud. Fruit globose, ca. 8 mm diam., intense blue but 
drying black, usually I/node. 

Distinctive little plants with many (ca. 1 2) distal 
lateral horizontal branches, small leaves, unusual 
stipules, and minute flowers. Presently known only 
from the type, collected near Siquerres, Limon, 1 8 
April 1985, at about 300 m elevation. It appears 
to be related to F. cobana J. D. Smith of Honduras, 
but that species has larger (8- 1 cm) leaves, longer 
peduncles (to 1 5 mm), and a higher-elevation hab- 
itat. 



Faramea occidentalis (L.) A. Rich., Mem. Fam. 
Rubiac. 96. 1830. Ixora occidentalis L., Syst. 
Nat. ed. 2: 893. 1759. F. zeteki Standl., Contr. 



134 



FIELDIANA: BOTANY 



Arnold Arbor. 5: 147. 1933. F. belizensis Standl., 
Carnegie Inst. Wash. Publ. 461: 90. 1935. F. 
standleyana L. O. Williams, Phytologia 26: 490. 
1973. Figure 44. 



Shrubs or small trees, 2-6(-10) m tall, to 20 cm dbh, 
leafy branchlets 1.5-5 mm thick, glabrous, terete, often 
dichotomously or trichotomously branched; stipule 
sheath 2-8 mm long, 5-8 mm wide at the base, with 
awns 4-12(-18) mm long, deciduous. Leaves with pet- 
ioles 6-15 mm long, 1-2 mm thick, glabrate; leaf blades 
8-18(-21)cm long, (2.5-)3.5-9.5(-l l)cm broad, oblong 
or elliptic to narrowly elliptic-obovate (less often ovate- 
oblong), apex rounded to obtuse and acuminate to cau- 
date-acuminate, the narrow (2 mm) tip 4-17 mm long, 
base acute to obtuse or rounded and subtruncate, drying 
chartaceous to stiffly chartaceous, glabrous above, gla- 
brous or minutely puberulent beneath, 2 veins 6- 107 
side and only loosely loop-connected near the margin (a 
definite submarginal vein absent), with 1-3 more weakly 
defined secondaries between the major secondary veins. 
Inflorescences terminal or less often axillary, solitary or 
several, 5-12 cm long, equally broad, umbelliform or 
trichotomous. few-branched and with 3-9 flowers, often 
drying black, peduncles 1.5-6 cm long, lateral branches 
opposite and few, distal flowers usually in groups of 3, 
bracts 4 mm long, linear, pedicels 3-12(-20) mm long. 
Flowers white but drying black, sweet scented in life, 
probably nocturnal, hypanthium ca. 2 mm long, oblong, 
calyx cup 1.5-3 mm long, teeth absent or minute; corolla 
salverform, tube 10-19(-22) mm long, 2-3 m diam., 
narrowly cylindrical, lobes 4, 8-16(-25) mm long, 1.3- 
3 mm wide near the base, lanceolate to narrowly ovate; 
stamens 4, filaments 0.5 mm long, anthers ca. 8 mm 
long, slightly exserted; style as long as the corolla tube 
or 'Ath as long, stigma 2.5-5.5 mm long. Fruits 6-9 mm 
long, 9-14 mm diam., globose to subglobose-oblate. per- 
sisting calyx 1-2 mm long and 2-3 mm diam., drying 
black. 



Trees and shrubs of evergreen forest formations, 
from near sea level to 1 000 m elevation. Flowering 
in February-July and October in southern Central 
America (primarily in June in Costa Rica); prob- 
ably fruiting throughout the year. The species 
ranges from southern Mexico, Central America, 
and the West Indies through tropical South Amer- 
ica. 

Faramea occidentalis is recognized by the large 
slender flowers drying black, relatively few- 
branched and few-flowered inflorescences also 
drying black, and rounded fruit slightly shorter 
than broad. This species varies greatly in leaf form 
and it does not have a well-developed submarginal 
vein. Though wide-ranging, it has not been col- 
lected very often in Costa Rica. This species is 
closely related to F. luteovirens with larger flowers. 
Fruiting and sterile material may be difficult to 
separate from F. eurycarpa. 



Faramea ovalis Standl., Contr. U.S. Natl. Herb. 
18: 138. 1916. F. quercetorum Standl., J. Wash. 
Acad. Sci. 18: 168. 1928. Figure 45. 

Small trees or shrubs, 2-6(-13) m tall, to 20 cm dbh, 
leafy branchlets 1-4 mm thick, glabrous, becoming gray- 
ish; stipules 3-13 mm long, the basal cupulate tube 1- 
2 mm high, with a single slender awn 3-10 mm long, 
often persisting. leaves not closely congested, petioles 
3-10 mm long. 0.7-1.8 mm thick, glabrous; leaf blades 
3-1 1 cm long, 1.5-4.8 cm broad, elliptic-oblong to ob- 
long or narrowly elliptic, apex abruptly narrowed and 
acuminate or caudate-acuminate, the tip 6-1 3 mm long, 
base acute to obtuse, drying stiffly chartaceous and gray- 
ish green to dark green above (slightly paler beneath), 
glabrous above and below, 2 veins (3-)5-8/side and 
loop-connected distally, with a vein-like edge along the 
leaf margin. Inflorescences terminal (axillary), 3-7 cm 
long, to 4 cm broad, umbellate with 3-5(-9) flowers, 
peduncles 8-24 mm long, 0.5 mm diam., glabrous and 
drying dark, pedicles 4-18 mm long, ca. 0.5 mm thick. 
Flowers glabrous externally, drying dark, hypanthium 1- 

2 mm long, calyx tube 0.5-2 mm high, teeth to 0.5 mm 
long; corolla salverform to somewhat funnelform, white 
or tinted with pink-purple, tube (7-)9-14 mm long, 1 .2- 

3 mm diam., lobes 4-6 mm long; stamens 4. Fruit 6-10 
mm diam., globose, blue drying black and smooth, per- 
sisting calyx tube 1-2.5 mm long and ca. 2 mm diam. 

Plants of moist evergreen cloud forests, 1 000- 
2300 m elevation (400-600 m on Volcan Orosi). 
Flowering in March-June and December; prob- 
ably fruiting throughout the year. The species is 
found in the northwestern cordilleras, eastern parts 
of the Cordillera de Talamanca, and the Chiriqui 
Highlands. 

Faramea ovalis is recognized by its cloud forest 
habitat, lack of pubescence, smaller often caudate- 
acuminate leaves, umbellate inflorescences with 
relatively few large flowers, and globose black fruits. 
The species resembles smaller-leaved specimens 
of F. occidentalis and some species of Coussarea. 



Faramea parvibractea Steyerm., Mem. New York 
Bot. Gard. 17: 376. 1967. Figure 45. 

Shrubs or small trees 2-8 m tall, leafy branchlets 1-4 
mm thick, glabrous and drying greenish, strongly qua- 
drangular; stipules 6-15 mm long, 0.5-1 mm broad, 
encircling the shoot apex, quickly caducous. Leaves with 
petioles 5-14(-20) mm long, ca. 1 mm thick, glabrous; 
leaf blades 7-17 cm long. 1.5-6(-7.5)cm broad, elliptic- 
oblong, ovate-oblong, to oblong-lanceolate, apex acu- 
minate to caudate-acuminate, tip 6-12 mm long, base 
acute to obtuse, drying chartaceous, glabrous above and 
below, 2 veins 6-10/side. weakly loop-connected 3-5 
mm from the margin. Inflorescences fasciculate or pseu- 
doumbellate from distal nodes, to 6(-10) cm long and 
equally broad, with 6-ll(-19) primary peduncles 11- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



135 



20(-30) mm long, flowers usually borne in distal groups 
of (l-)2, 3, or 5 at apex of the primary peduncles, a few 
peduncles with broadly ovate leaf-like bracts 5.5-16 mm 
long and 3-12 mm broad at their apex, bracts subcordate 
and with petiole-like base ca. 1 mm long, white in life, 
pedicles 1-3 mm long. Flowers with hypanthium 0.5- 
1.5 mm long, calyx tube 0.5-1 mm long, calyx lobes 
small and usually unequal (ca. 0.1 mm and 0.5 mm), 
corolla funnelform, white, 7-12(-16) mm long, tube 5- 
8 mm long, 1.5-3 mm diam., lobes 4, 5-10 mm long, 
2-3.5 mm broad, lanceolate to narrowly oblong, obtuse; 
stamens exserted, filaments 3-3.5 mm long, inserted at 
or below the middle of the tube, anthers 3.5-4 mm long, 
linear, bluntly rounded; style 4-6 mm long. Fruits 5-7 
mm long, 8-12 mm broad, oblate and rounded in cross- 
section, smooth, the ribs obscure or prominent when 
dried; pyrene solitary. 

Plants of wet lowland rain forest formations of 
the Caribbean slope, from 4 to 600 m elevation. 
Flowering in January-June and October; fruiting 
in February and May-August. The Costa Rican 
collections are mostly from between Tortugero and 
Limon, with a few from the Osa Peninsula. This 
species is also known from Panama and Vene- 
zuela. 

Faramea parvibractea is recognized by the clus- 
ters of few-flowered peduncles at distal nodes, the 
ovate-subcordate bracts subtending some flower 
groups, the elliptic-oblong leaves with "drip tips," 
the narrow stipules enclosing the shoot apices, the 
glabrous white flowers, and the fruit often in um- 
bellate groups at the ends of leafy stems. The un- 
usual bracts do not appear to be developed be- 
neath all the flower groups, but they are large and 
contradict the specific name. 



Faramea pauciflora Dwyer, Ann. Missouri Bot. 
Gard. 67: 172. 1980. 

Understory shrubs or small trees, 2-5(-10) m tall, 
branches at right angles to main stem, leafy stems 1.5- 
4 mm thick, glabrous; stipules 8-15 mm long and 4 mm 
broad, triangular to tubular with 2 slender awns 3-5 mm 
long, covering the terminal bud, glabrous, drying black, 
caducous. Leaves held in a single plane, petioles 4-7(-l 2) 
mm long, 0.9-1.5 mm thick, glabrous, drying dark; leaf 
blades 11-22 cm long, 4-11 cm broad, elliptic-oblong 
to narrowly oblong or elliptic-obovate, apex acuminate 
or caudate-acuminate, tip 7-17 mm long, base acute, 
drying stiffly chartaceous, dark brown or blackish above, 
glabrous above, glabrous or sparsely and minutely pa- 
pillate-puberulent beneath, 2 veins 8-12/side. Inflores- 
cences ca. 10 cm long with 1-3 flowers, terminal or ax- 
illary, usually I/node, peduncles 2-5 cm long, 0.7-1.5 
mm thick, often with 1-2 pairs of stipule-like bracts near 
the base and terminated by several minute bracts, ped- 
icels 1 5-55 mm long, 0.5 mm thick, glabrous and drying 



black, merging with the hypanthium. Flowers glabrous 
externally and drying black, hypanthium and calyx 10- 
1 5 mm long, 2.5-4 mm diam., tube ca. 4 mm long, lobes 
few or unequal, 1-8 mm long; corolla salverform, white, 
tube 10-22 mm long, lobes 4, 13-30 mm long, rotate 
and with a narrow tip 4-9 mm long often held at 90. 
Fruits 12-14 mm long, 10-14 mm diam., with a per- 
sisting calyx tube 7-9 mm long and 3-4 mm diam., dark 
blue drying black, pendulous. 

Plants of the wet Caribbean slope at elevations 
from 20 to 1100 m. Flowering in March-April; 
fruiting in March-April and August-September in 
Panama. The species ranges from near Nuevo Ar- 
enal southward to Colombia. 

Faramea pauciflora is unusual because of its 
glabrous parts, usually axillary inflorescences with 
one to three long white flowers and long pedicels. 
Two Costa Rican collections have been seen: Her- 
rera 2556 and A. Smith 1644. This species is close- 
ly related to F. luteovirens Dwyer of Panama with 
thicker leaves, fewer 2 veins, and shorter calyx. 
Compare F. occidentalis and the pubescent but 
similarly few-flowered Coussarea enneantha (fig. 
46). 



Faramea scalaris Standl., Contr. U.S. Natl. Herb. 
13: 139. 1916. 

Shrubs or small trees, 1.5-6 m tall, leafy branchlets 
1.5-4 mm thick, terete or quadrangular, drying brown- 
ish, leaves distant (3-6 cm) along the stems; stipules 4- 
8 mm long, with a short-tubular sheath, truncated or 
deltoid at apex, with or without a short (1-2 mm) awn, 
persistent. Leaves subsessile with petioles l-3(-5) mm 
long; leaf blades (5-) 7- 15 cm long, (1-) 1.3-4 cm broad, 
narrowly elliptic-oblong to narrowly oblanceolate, apex 
acuminate, tip 7-15 mm long and straight or falcate, 
base gradually narrowed but often slightly rounded 
and subauriculate at the petiole, drying chartaceous and 
greenish, glabrous above and below, 2 veins 9-13/side, 
these arising at almost 90 angles and united by a linear 
or slightly arcuate lateral vein 2-5 mm from the leaf 
edge, a slender submarginal vein also present 0.3-1 mm 
from the margin in fully developed leaves. Inflorescences 
terminal and solitary, 3-6 cm long, paniculate with short 
opposite branches, primary peduncles 1 5-20 mm long, 
glabrous, bracts caducous, pedicels 2-4 mm long. Flow- 
ers glabrous, hypanthium ca. 1.5 mm long, calyx tube 
ca. 0.5 mm long, lobes 0.1-0.3 mm; corolla salverform, 
blue becoming white, tube 7-10 mm long, 1-1.5 mm 
diam., lobes 3-4 mm long, ca. 3 mm broad, ovate, disc 
1 mm long; style ca. 7 mm long. Fruits unknown. 

Plants of montane wet forest formations, from 
1800 to 2300 m elevation. Flowering in January 
and March. The species is known only from the 
Chiriqui Highlands in western Panama. 



136 



FIELDIANA: BOTANY 



Faramea scalaris is distinguished by its narrow 
subsessile leaves with prominent lateral (mela- 
stome-like) venation, the small inflorescences, and 
the restricted high-elevation habitat. The unusual 
leaf venation suggests that this species is closely 
related to F. trinervia and F. suerrensis of lowland 
formations. 



Faramea sessifolia P. Allen, Rain Forests of Golfo 
Dulce 409. 1956. Figure 44. 



Shrubs or small trees, 3-7 m tall, leafy branchlets 2- 
7 mm thick, the nodes to 10 mm thick, rectangular in 
cross-section (flattened and 4-angular), glabrous, drying 
yellowish; stipules 10-25 mm long, 3-5 mm broad at 
the base, triangular-subulate with a narrow distal tip, 
caducous. Leaves somewhat dimorphic with smaller and 
narrower leaves often subtending the inflorescences, ses- 
sile or subsessile, petioles 0-^4(-6) mm long, ca. 2 mm 
thick; larger leaf blades 17-27 cm long, 6-17 cm broad, 
narrowly oblong to ovate-oblong, smaller leaf blades 8- 
1 7(-l 9) cm long and 2-4(-6) cm broad, narrowly oblong- 
lanceolate to narrowly oblong, apex abruptly narrowed 
(in larger leaves) or gradually narrowed (in smaller leaves) 
and acuminate, tips 8-17 mm long, base rounded and 
subtruncate, drying chartaceous and yellowish green, gla- 
brous above and below, 2 veins 12-16/side and loop- 
connected 2-5 mm from the margin. Inflorescences ter- 
minal, solitary or 3, 10- 15 cm long and 10-1 2 cm broad, 
paniculate with opposite many-flowered branches, pri- 
mary peduncles (2-)6-7 cm long, 0.7-2 mm thick, gla- 
brous, whitish in life and yellowish when dried, bracts 
0.5-1 mm long, pedicels 1-2 mm long above small (0.5 
mm) bracteoles. Flowers glabrous, hypanthium 0.5-1 
mm long, calyx tube 0.2-0.5 mm long, lobes 0.2-0.4 
mm long; corolla salverform, bright blue or bluish pur- 
ple, tube ca. 6-7 mm long and 1 mm diam., lobes 3-4 
mm long, to 2 mm broad at base. Fruits 4-5 mm high, 
7-9 mm broad, oblate, circular or oblong in cross-sec- 
tion, with 8 longitudinal costae, pale grayish brown be- 
coming black. 



Plants of the lowland rain forests of the Osa 
Peninsula region, 20-400 m elevation. Flowering 
in April-May (Allen 5539 F, us type); fruiting in 
March, August-September, and November. The 
species is known only from southwestern Costa 
Rica. 

Faramea sessifolia is recognized by its large sub- 
sessile leaves with many secondary veins and ar- 
cuate submarginal vein, leafy stems rectangular in 
cross-section, lack of pubescence, blue flowers, and 
restricted distribution. The veins on the lower leaf 
surface and inflorescences dry yellowish. This name 
should not be confused with F. sessilifolia (H.B.K.) 
A. DC. of South America. 



Faramea stenura Standl., Publ. Field Columb. 
Mus., Hot. Ser. 4: 331. 1929. Figure 45. 

Small trees to 7 m tall, leafy branchlets 1.7-4 mm 
thick, glabrous, drying greenish; stipules 6-10 mm long, 
3-4 mm broad at the base, triangular to ovate, acute or 
mucronate at apex, caducous. leaves with petioles 5-10 
mm long, 0.6-1.5 mm thick; leaf blades 9-19 cm long, 
2.5-5.5 cm wide, narrowly oblong to narrowly oblong- 
obovate, apex acuminate, tip 5-23 mm long, base acute 
to cuneate, drying chartaceous, dark grayish green above, 
glabrous above and below, 2 veins 9-12/side, arising at 
almost 90 and weakly loop-connected near the margin, 
with thinner secondaries present and parallel with the 
larger. Inflorescences usually terminal and solitary, pa- 
niculate pyramidal, 3-5.5 cm long, 4-7 cm broad, blue, 
primary peduncles 1.5-5(-8) cm long and 1-1.5 mm 
thick, bracts 3-5 mm long, linear or triangular, purplish, 
pedicels 2-4 mm long. Flowers glabrous externally, hy- 
panthium ca. 0.6 mm long, calyx tube 0.2-0.3 mm long, 
lobes 4, 0.3-0.7 mm long; corolla salverform, pale to 
deep blue, glabrous, tube 5-8 mm long, lobes 4, 3-5 mm 
long, 1.2-1.5 mm broad, elliptic to ovate, acute at apex. 
Fruits 4-8 mm long, 12-15 mm broad, transversely reni- 
form (oblong in cross-section), rounded basally and flat- 
tened or depressed above, smooth and usually drying 
green. 

Plants of lowland rain forest formations, 30- 
600 m elevation (but see below). Flowering in Jan- 
uary-September; probably fruiting throughout the 
year. It ranges along the Caribbean coast, from 
Veracruz, Mexico, to western Panama, and in the 
Golfo Dulce area. 

Faramea stenura is distinguished by its small 
blue corollas, caducous stipules, oblate fruit, and 
narrowly oblong leaves with long "drip tips." This 
species is very similar to F. multiflora, but their 
stipules help differentiate most collections. At La 
Selva F. stenura grows in low swales, whereas F. 
multiflora grows on the ridgetops. There are very 
similar plants growing at higher elevations (ca. 
1200 m) in Chiriqui with larger (10 x 14 mm) 
ribbed fruit that dry whitish. It is not clear whether 
they are a high-elevation subspecies or an unde- 
scribed closely related species. 



Faramea suerrensis (J. D. Smith) J. D. Smith, Bot. 
Gaz. 44: 1 1 2. 1 907. Faramea trinervia K. Schum. 
& J. D. Smith var. suerrensis J. D. Smith, Bot. 
Gaz. 31: 115. 1901. Figure 44. 

Shrubs or small treelets, 2-6 m tall, leafy branchlets 
2-6 mm thick (to 12 mm broad at the node), usually 
quadrangular in cross-section, glabrous; stipules 5-10 
mm long, 3-8(-12) mm wide at the base, often united 
to form a short tube, entire or rounded with a small ( 1 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



137 



mm) mucronate tip. Leaves with petioles 4-18 mm long, 
1-2 mm thick; leaf blades 9-20(-28) cm long, 3-8(-12) 
cm broad, narrowly elliptic-oblong to oblong or narrowly 
oblong-oblanceolate, apex short-acuminate to long-cus- 
pidate, tip to 1 8 mm long in some specimens, base acute 
to rounded and subtruncate, drying chartaceous to sub- 
coriaceous, pale grayish green to yellowish green, gla- 
brous above and below, the major veins often becoming 
impressed above, 2 veins 8-16(-25)/side and with less 
prominent parallel 2 veins, 2 veins arising at nearly 90 
from the midvein and joined near the margin by a prom- 
inent linear lateral vein (2-)5-7(-14) mm from the leaf 
edge, with a smaller submarginal vein 0.5-2 mm from 
the leaf edge. Inflorescences terminal and solitary (or 3), 
corymbose panicles with opposite branching, 9-12(-20) 
cm long, to 16 cm broad, bright blue, many-flowered, 
peduncles 2-5(-9) cm long, 2-3 mm thick, glabrous, ped- 
icels 2.5-9 mm long. Flowers distylous, glabrous, hy- 
panthium ca. 1.2 mm long, calyx tube a small (0.3 mm) 
rim, lobes minute or 0.5 mm long and triangular; corolla 
salverform, brilliant pale blue to deep blue, tube 6-10 
mm long, 0.8-1.5 mm diam., lobes 4, 3-5 mm long, ca. 
1.7 mm broad; stamens 4, filaments attached near the 
middle of the tube, anthers 2.2 mm long, included. Fruits 
6-1 1 mm long, 12-16 mm broad, transversely reniform 
and rounded-oblong in cross-section (somewhat flat- 
tened laterally and flat or slightly depressed distally), 
longitudinal ribs slightly developed or obscure, exocarp 
spongy and deep blue in life; pyrenes solitary. 

Plants of lowland rain forest formations on both 
the Pacific and Caribbean slopes, from near sea 
level to 800(-1000) m elevation. Flowering in Jan- 
uary-August and November; fruiting in every 
month except December. The species ranges from 
southern Nicaragua through Costa Rica and Pan- 
ama to northwestern Colombia. 

Faramea suerrensis is distinguished by its larger 
leaves with prominent melastome-like venation, 
brilliant bluish inflorescences and flowers, lack of 
pubescence, and lowland rain forest habitat. The 
unusual quality of the blue coloring of the inflo- 
rescence and the leaf venation make this one of 
our most distinctive species of Rubiaceae. This 
species is very closely related to F. trinervia (q.v.); 
it is possible that the two may be conspecific as 
Schumann and Smith originally thought. The 
breeding system was studied by Bawa and Beach 
(1983). 



Faramea trinervia K. Schum. & J. D. Smith, Bot. 
Gaz. 31: 115. 1901. F. bullata Standl., Publ. 
Field Mus. Nat. Hist., Bot. Ser. 4: 294. 1929. 
Figure 44. 

Shrubs or small trees, leafy stems 2-6 mm thick, gla- 
brous; stipules 6-10 mm long, the basal tube 2-4 mm 
long with rounded or acute apex and narrow awn 3-5 
mm long. Leaves sessile or subsessile with petioles 2-4 



mm long; leaf blades 14-31 cm long, 4-12 cm broad, 
narrowly oblong to narrowly oblong-obovate, apex acu- 
minate or caudate-acuminate, somewhat narrowed be- 
low the middle, base abruptly rounded and subcordate, 
drying stiffly chartaceous to subcoriaceous, grayish green, 
glabrous above and below, 2 veins 10-1 7(-22)/side (dif- 
ficult to separate from the less prominent secondaries), 
united near the margin by a linear lateral vein 5-10 mm 
from the leaf edge, a smaller submarginal vein also pres- 
ent 1-3 mm from the edge. Inflorescence solitary and 
terminal (or 3 with 2 axillary), 4-7 cm long, to 12 cm 
broad, paniculate with opposite or trichotomous branch- 
ing at the apex of the peduncle, basal bracts (small leaves) 
to 2 cm long, peduncles 3-6 cm long, 2.2-3.4 mm thick. 
Flowers glabrous externally, hypanthium ca. 1 mm long, 
obconic, calyx limb ca. 0.3 mm long with minute (0.2 
mm) lobes; corolla color not known, salverform, tube 
6-7 mm long, 0.5-0.9 mm diam., lobes 4, ca. 4 mm long 
and 1 mm broad. Fruits 7-8 mm long and 12-16 mm 
broad, oblate-reniform, flat or depressed centrally above, 
drying smooth and black. 

A species of lowland Caribbean rain forest for- 
mations, 10-600 m elevation. Flowering in March 
(Tonduz 8690 us) and April (Tonduz 9583 us the 
type); fruiting in April (Barringer et al. 2632 CR, 
F) and July (/. Chacon 209 CR). This species is 
known only from the Talamanca valley region, 
southeastern Costa Rica, and Bocas del Toro 
Province in Panama. 

Faramea trinervia is recognized by the stiff nar- 
rowly oblong leaves with melastome-like vena- 
tion, coupled with the short petioles and rounded 
auriculate leaf bases. This species is very similar 
to and may prove to be conspecific with material 
placed under F. suerrensis (a later name). The 
thicker subsessile leaves rounded at the base are 
a unique character combination within the more 
common and more widely ranging F. suerrensis, 
and we treat the two as distinct species, although 
they appear identical in most other respects. The 
type of F. bullata (Cooper 507 F from Panama) 
has short petioles but with a rounded lamina base, 
and we interpret it as an aberrant form of F. tri- 
nervia. 



Ferdinandusa Pohl 

Trees or shrubs, stems glabrous or puberulent; stipules 
interpetiolar, triangular, deciduous or caducous. Leaves 
opposite or verticillate, decussate or distichous, petio- 
late, glabrous or puberulent, entire, pinnately veined, 
without domatia. Inflorescences terminal or axillary to 
the distal leaf pair, paniculate with opposite branching 
and cymose to corymbose in form (rarely fasciculate or 
umbellate), bracteate, flowers pedicillate. Flowers bisex- 
ual and usually radially symmetrical (corolla tube some- 
times curved), monomorphic, calyx cupular, calyx teeth 



138 



FIELDIANA: BOTANY 



short, deciduous or persistent; corolla salverform to fun- 
nelform, white to reddish or greenish, corolla lobes 4(-5), 
convolute or broadly imbricate in bud, rounded distally; 
stamens 4(-5), filaments attached in the upper part of 
the tube, anthers versatile, exserted or included; ovary 
2-locular with few to many ovules vertical on the axile 
placentas, stigmas subcapitate. Fruits a cylindrical to 
oblong (subglobose) woody capsule with septicidal de- 
hiscence from apex; seeds few to many, elliptic and flat- 
tened, with entire to lacerate marginal wings. 

A genus of 20-25 species with 1 species in Costa 
Rica and Panama and the others in South Amer- 
ica. The elongate capsular fruit with small winged 
seeds and broadly overlapping corolla lobes dis- 
tinguish this genus. Compare this genus with ma- 
terial placed in Ladenbergia (with valvate corolla 
lobes) and Macrocnemum. 



Ferdinandusa panamensis Standl. & L. O. Wil- 
liams, Ceiba 3: 34. 1952. Figure 40. 

Trees to 20 m tall, major branches held at 90 angles 
or drooping, leafy stems 2-5 mm thick, terete or slightly 
quadrangular, glabrous or with erect hairs 0.5-1 mm 
long, stems slightly resinous where the hairs are broken 
off; stipules 5-20 mm long, narrowly triangular, acu- 
minate, glabrous, caducous. Leaves opposite or 3/node, 
smaller in size beneath the inflorescence, petioles 4-10 
mm long, 1.5-3 mm thick, glabrous or pubescent; leaf 
blades (6-)10-18(-22) cm long, (4-)6-10 cm broad, ob- 
long to elliptic-oblong or ovate-oblong, apex abruptly 
short-acuminate, tip 3-10 mm long, base obtuse to 
rounded or truncate (subcordate), leaves drying stiffly 
chartaceous, glabrous above and below or with erect 
brownish hairs 0.5-1 mm long on the veins beneath and 
more sparsely on the surfaces beneath, 2 veins 6-9/side, 
arcuate-ascending near the margin but loop-connected 
only near apex, domatia absent. Inflorescences terminal 
(apparently axillary when leaves develop at the first 
branching node of the panicle), solitary or 3, 6-15 cm 
long, 6-22 cm broad, primary peduncles 2-5 cm long, 
bracteolate, bracts ca. 1 mm long, pedicels 5-12 mm 
long and merging with the flower base. Flowers glabrous, 
hypanthium ca. 2 mm long, calyx cup 0.5-1 mm long, 
2-4 mm diam., calyx teeth 4 or 5, ca. 0.5 mm long; 
corolla funnelform, yellowish green or white, fleshy, tube 
(4-)6-18(-25) mm long, 2-3 mm diam., slightly ex- 
panded at the base, and much expanded distally, lobes 
5, 4-6 mm long, 3-6 mm broad at the base; stamens 5, 
filaments linear, attached near the middle of the tube, 
anthers sometimes dimorphic (long and short); style to 
11 mm long, stigmas ca. 1.5 mm long. Fruits (2-)3-6 
cm long, 6-1 2 mm thick, narrowly oblong or cylindrical, 
abruptly rounded at the base and apex, with a short (0.5- 
1 mm) persistent calyx, surface smooth and with obscure 
longitudinal ribs, brown; seeds 1-2 cm long, elliptic. 

Trees of lowland Caribbean rain forest forma- 
tions, from near sea level to 200 m elevation. 
Rowers have been collected in January-May and 



November-December; fruits were collected in 
February-July. The species is known only from 
Costa Rica and near Chiriqui Lagoon in Panama 
(von Wedel 2232 F the type). 

Ferdinandusa panamensis is recognized by the 
elongate woody capsules with small winged seeds, 
fleshy flowers with broadly overlapping corolla 
lobes, broad leaves often rounded at the base, and 
the occasional presence of stiffbrownish hairs. The 
names cafe macho and cafecillo have been used 
for this species in Costa Rica. The wood is very 
hard and young trees are used for boat poles. 



Galium Linnaeus 

REFERENCES L. Dempster, The genus Galium 
(Rubiaceae) in Mexico and Central America. Univ. 
Calif. Publ. Bot. 73: 1-33. 1978. The genus Galium 
(Rubiaceae) in South America, IV. Allertonia 5: 
283-345. 1990. F. Ehrendorfer, Revision of the 
genus Relbunium (Endl.) Benth. & Hook. (Rubi- 
aceae-Galieae). Bot. Jahrb. Syst. 76: 516-553. 
1955. 

Herbs or slender vines, annual or perennial, often 
woody at the base (rarely shrubs), erect, decumbent or 
climbing, monoecious or dioecious, stems usually slen- 
der and with 4 prominent longitudinal ridges (square in 
cross section), glabrous or puberulent with thin hairs; 
stipules apparently absent but represented by leaf-like 
parts, indistinguishable from the true leaves (except that 
they lack axillary buds). Leaves in whorls of (3-)4-8(-10 
to many) at each node, comprising the true leaves and 
the transformed stipules, sessile or subsessile, often nar- 
rowly oblong, entire or with sharp retrorse trichomes 
along the edge (serrulate), with 1 or 3 major veins, with- 
out domatia. Inflorescences terminal or axillary, often 
paniculate with dichasial or trichotomous branching, or 
with 3 terminal flowers from the distal node, inflores- 
cences often with bract-like reduced leaves and thin- 
divaricate branching (with a 4-parted involucre in spe- 
cies formerly placed in Relbunium), pedicels articulate 
beneath the flower. Flowers bisexual or unisexual, ra- 
dially symmetrical, usually very small, calyx tube minute 
or lacking; corolla rotate to campanulate or urceolate, 
white to yellow, green, pink, or red, corolla lobes (3-)4(-5), 
valvate in bud; stamens (2-)3-4(-5), filaments attached 
to the short tube of the corolla, anthers versatile and 
exserted; ovary 2-locular with 1 ovule borne on the sep- 
tum in each locule, styles 2 (sometimes united at the 
base), stigmas capitate. Fruits usually 2-lobed or 2-parted 
(singular when 1 ovule fails to develop), dry or fleshy, 
smooth and glabrous to tuberculate or densely hispidu- 
lous, small, finally separating into 2 1 -seeded mericarps, 
the mericarps rounded and indehiscent; seeds convex 
dorsally, attached to the pericarp, testa membranous. 

A large genus of some 300-400 species, es- 
pecially well represented in the North Temperate 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



139 



zone and at higher elevations in the tropics. 
Dempster listed 4 1 species in Mexico and Central 
America (3 of which are probably early introduc- 
tions); the genus is especially well represented in 
Baja California and the central highlands of Mex- 
ico. Galium, as a genus, is easy to recognize with 
its small whorled leaves on slender herbaceous 



clambering 4-angled stems, minute flowers on 
slender pedicles, and 2-lobed little fruit. Material 
of Galium may be mistaken for species ofDidym- 
aea and Nertera. This treatment is based in large 
part on the annotations and publications of Laura- 
may Dempster. 



Key to the Species of Galium 

la. Leaves in whorls of 6 or 8(-10); fruit covered with ascending curved hairs G. mexicanum 

1 b. Leaves in whorl of 4 at each node; fruit glabrous or with uncinate (hooked at the tip) hairs ... 2 
2a. Flowers solitary in the leaf axils and subtended by a whorl of 4 leaf-like or calyx-like bracts 

G. hypocarpium 

2b. Flowers rarely solitary in leaf axils, rarely subtended by 4 bracts 3 

3a. Fruits glabrous, leaves with 1 primary vein and no lateral veins [usually glabrous; stems with minute 

(0. 1-0.2 mm) hairs] G. aschenbornii 

3b. Fruits covered with ascending uncinate hairs; leaves with 1 primary vein and 2 lateral veins usually 

visible 4 

4a. Leaf blades usually ovate to elliptic; stems and leaves with few to many longer (0.4 mm) hairs; 

inflorescences usually with few (ca. 5) flowers; corollas often hispidulous G. uncinulatum 

4b. Leaf blades usually oblong-lanceolate to elliptic, stems and leaves sparsely short-puberulent (ca. 0.2 

mm); inflorescences usually with more than 5 flowers; corollas glabrous G. orizabense 



Galium aschenbornii Schauer, Linnaea 20: 701. 
1 847. Relbunium aschenbornii (Schauer) Hemsl., 
Biol. centr. amer. Bot. 2: 62. 1881. Figure 3. 



Creeping or climbing herbs to 1 .2 m long, with slender 
woody or herbaceous stems arising from a small root- 
stock, sometimes rooting from distal nodes, with both 
long (5 cm) and short (4 mm) internodes, leafy stems 
0.3-1 mm thick, glabrous or with short retrorse hairs 
0. 1-0.2 mm long; stipules leaf-like. Leaves usually 4/node, 
petioles 0. 1-0.5 mm long; leaf blades 4-8(-14) mm long, 
1 .2-3(-4) mm broad, narrowly oblong or elliptic-oblong 
(to lanceolate or ovate-oblong), apex obtuse to acute with 
a short (0.3 mm) tip, base acute to obtuse (rarely round- 
ed), margin entire or with a few retrorse aculeolate tri- 
chomes, midvein prominent, secondary and lateral veins 
obscure or weakly developed. Inflorescences often of 3 
terminal flowers subtended by a whorl of 4 reduced leaves, 
or of several flowers in compound dichasia, pedicels 1- 
5 mm long, filiform. Flowers ca. 1.5 mm long, greenish 
yellow, said to be <J, 2, and bisexual on the same plant 
or on different plants, hypanthium 0.5-0.7 mm long, 
calyx ca. 0.3 mm long; corolla rotate, lobes 0.5-1 mm 
long, white to yellowish or red. Fruits 3-4 mm long, 4- 
7 mm broad when dry, 2-lobed distally, orange becoming 
black, glabrous and wrinkled when dried, borne on slen- 
der pedicels 2-5 mm long. 



Plants of evergreen montane forest formations, 
from 1 200 to 2900(-3400) m in Central America. 
Probably flowering and fruiting throughout the 



year. The species ranges from the states of Jalisco 
and San Luis Potosi in Mexico to western Panama. 
Galium aschenbornii is recognized by the usu- 
ally narrow leaves in whorls of four, short petioles, 
and glabrous fruit. It is infrequently collected in 
Costa Rica. 



Galium hypocarpium (L.) Clos in Gay, Fl. Chil. 3: 
186. 1847. Vaillantia hypocarpia L., Syst. Nat. 
ed. 10: 1307. 17 59. Relbunium hypocarpium (L.) 
Hemsl., Biol. cent. amer. Bot. 2: 63. 1881. G. 
hypocarpium (L.) Fosberg, Sida 2: 386. 1966. 
Figure 3. 



Herbs to 60 cm high, prostrate to procumbent or 
climbing over low objects, leafy stems 0.4-1 .3 mm thick, 
with 4 prominent longitudinal ridges, nodes usually well 
spaced, puberulent with thin whitish hairs 0.3-1 mm 
long; stipules leaf-like. Leaves 4/node, sessile; leaf blades 
3-1 3(-22) mm long, 2-4(-8) mm broad, oblong to ovate- 
oblong or broadly oblong-obovate, apex obtuse to 
rounded, with a minutely apiculate tip, base obtuse, dry- 
ing chartaceous to subcoriaceous, both surfaces with stiff 
ascending hairs to 0.7 mm long or the surfaces glabrous 
and the margins hirsutulous. 2 veins 2-3/side (or the 
venation obscure). Inflorescences of solitary flowers in 
the leaf axils, usually with 2 flowers per node, peduncles 
5-15 mm long, with an involucre of 4 bracteoles sub- 
tending each flower, bracteoles subequal, 2-4 mm long, 



140 



FIELDIANA: BOTANY 



ovate to oblong or lanceolate, hirsutulous. Flowers small, 
hypanthium rounded, calyx tube and teeth undeveloped; 
corolla campanulate, white, 1.5-2.5 mm long, tube cy- 
lindrical, lobes 4, shorter than the bracteoles and alter- 
nating with them, surfaces glabrous and with marginal 
hairs; stamens 4, filaments short, anthers small. Fruits 
2-3 mm long, to 3.5 mm broad, broadly 2- or 3-lobed, 
or with only 1 seed and globose, orange or reddish or- 
ange, glabrous to puberulent. 

Plants of moist evergreen montane forest for- 
mations and high elevation paramos, from 1800 
to 3400 m elevation. Probably flowering and fruit- 
ing throughout the year. The species ranges from 
Veracruz, Mexico, southward through the higher 
elevations of Central America into the Andes 
mountains of South America as far south as north- 
ern Chile and Argentina. 

Galium hypocarpium is recognized by its slender 
stems with four leaves at each node, creeping or 
clambering habit, the solitary axillary flowers sub- 
tended by an involucre of four leaf-like bracteoles. 
These plants closely resemble our other species of 
Galium, but the other species rarely have single 
flowers subtended by a four-parted involucre. 



Galium mexicanum H.B.K., Nov. gen. sp. 3: 337 
(quarto). 1818. Figure 3. 

Trailing or climbing herbs to l(-2.4?) m long, leafy 
internodes 0.3-2.8 mm thick, with minute (0. 1-0.3 mm) 
retrorse aculeolate hairs, often with dense longer (0.5 
mm) whitish hairs at the node; stipules leaf-like. Leaves 
usually 6 or 8(-12)/node, essentially sessile; leaf blades 
(4-)8-20(-25) mm long, 1-3 mm broad, narrowly ob- 
lanceolate to narrowly oblong, apex obtuse (rounded), 
usually with an apiculate tip ca. 0.5 mm long, base grad- 
ually narrowed, with curved aculeolate retrorse hairs (ca. 
0.2 mm long) along the margin and midvein beneath, 
mostly glabrous above and on the flat surfaces beneath, 
primary vein prominent, 2 weakly denned lateral veins 
often present. Inflorescences usually terminal, paniculate 
arrangements of small cymose groups of 3-7 flowers sub- 
tended by reduced leaf-like bracts, with divaricate 
branching and slender pedicels 1-3 mm long. Flowers 
1.5-2 mm long, bisexual, hypanthium 0.5-0.7 mm long, 
turbinate, densely hirsutulous with minute (0. 1 mm) as- 
cending hairs, calyx reduced; corolla 1-1 .5 m long, white 
to pink or red, campanulate or rotate, glabrous exter- 
nally, puberulent within. Fruits dry, ca. 3 mm broad, 
covered with ascending slightly curved hairs or with a 
few uncinate hairs at the tip (in Costa Rica). 



Plants of evergreen montane forest formations, 
from ( 1 200-) 1 600 to 3 1 00 m elevation. Flowering 
in April-July and December-January in southern 
Central America. Rarely collected in Costa Rica 
but apparently common in the Chiriqui High- 



lands. The species ranges from the southwestern 
United States to Panama. 

Galium mexicanum is recognized by the slender 
clambering stems with whorls of six or eight nar- 
rowly oblong or oblanceolate leaves and the small 
fruit with dense pubescence of curved hairs. The 
aculeolate trichomes on stems and leaves help the 
plants to climb and makes them adhesive and very 
difficult to disentangle from shrubbery or clothing. 



Galium orizabense Hemsley, Diagn. PI. Nov. 
Mexic. 3: 54. 1878. Figure 3. 

Erect or spreading herbs, 20-75 cm long, with several 
to many stems from a small root stock, leafy stems 0.3- 
1 mm thick, with thin white curves hairs 0.1-0.3 mm 
long; stipules leaf-like. Leaves 4/node, subsessile or short- 
petiolate, petioles to 2 mm long; leaf blades 6-18(-25) 
mm long, 2-4(-5) mm broad, narrowly oblong to nar- 
rowly elliptic-oblong or lanceolate, apex obtuse or short- 
apiculate, drying thin-chartaceous or membranaceous, 
with thin hairs ca. 0.3 mm long, on the upper surface, 
margin and major veins beneath, with a prominent 1 
vein and 2 straight lateral veins (usually readily appar- 
ent). Inflorescences paniculate, with divaricate branches 
subtended by slightly reduced leaves, often with 5-15 
flowers on open lateral branches, pedicels 1-10 mm long. 
Flowers 1-1.5 mm long, hypanthium ca. 0.5 mm long, 
covered with minute hairs; corolla rotate, lobes ca. 0.4 
mm long and equally broad, white or greenish yellow. 
Fruits dry, 1-2 mm broad, covered with minute uncinate 
hairs, borne on thin (to 0.5 mm) pedicels. 

Plants of evergreen montane forest formations, 
from 1 500 to 2500 m in southern Central Amer- 
ica. Flowering in March, July-August, and De- 
cember-January in southern Central America. The 
species ranges from eastern and central Mexico to 
Panama. 

Galium orizabense is recognized by the uncinate 
hairs on the fruit and the narrow verticillate leaves. 
This species is very similar to G. uncinulatum and 
the two may be conspecific. However, most spec- 
imens can be differentiated by the key, and we 
follow Dempster's treatment. 



Galium uncinulatum DC., Prodr. 4: 600. 1830. 
Figure 3. 

Prostrate or procumbent herbs 1 5-90 cm long, leafy 
stems 0.3-1 .5 mm thick, with thin whitish hairs 0.3-0.6 
mm long, slightly scabrous; stipules leaf-like. Leaves 4/ 
node, sessile or subsessile with petioles to 1 (-2) mm long; 
leaf blades (4-)6-12(-22) mm long, (2-)2.5-6(-13) mm 
broad, ovate to narrowly ovate or narrowly oblong, apex 
obtuse (rounded) with a short (0.5 mm) tip, base obtuse, 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



141 



drying membranaceous or chartaceous, margin and sur- 
faces with thin ascending or spreading hairs 0.2-0.4 mm 
long, with 1 vein and 2 well-defined lateral veins, the 
lateral veins 0.5-1 mm from the margin in larger leaves. 
Inflorescences terminal or axillary, with opposite or cy- 
mose branching, to 3 cm long, flowers usually few (3-5) 
on the slender peduncles, often subtended by whorls of 
reduced leaves, pedicels 1-10 mm long, 0.1-0.2 mm 
thick when dried. Flowers 1-1.5 mm long, hypanthium/ 
ovary ca. 0.5 mm long, covered by minute hairs that will 
expand in fruit, calyx reduced; corolla campanulate to 
rotate, white to greenish or yellow, usually puberulent 
externally. Fruits ca. 1.5 mm long and (l-)2-3 mm broad, 
usually 2-lobed and rounded, dry and covered by pale 
yellowish or whitish hooked (uncinate) hairs ca. 0.3-0.4 
mm long. 

Plants of evergreen montane forest formations 
from 1000 to 2800(-3300) m elevation. Flowering 
collections have been made in all months of the 
year except May and October-November in Cen- 
tral America. The species has been little collected 
in Costa Rica. The species ranges from southern 
Arizona and Texas (U.S.A.) through Mexico and 
highland Central America to Panama. 

Galium uncinulatum is recognized by having 
four, often broad, little leaves at each node, small 
few-flowered inflorescences with thin peduncles 
and pedicels, and fruits densely covered with thin 
uncinate hairs. This species is very similar to G. 
orizabense(q.v.), which appears to live in the same 
habitats but tends to have narrower leaves and 
shorter puberulence. Specimens referred to as Ga- 
lium obovatum H.B.K. by Standley, both in the 
herbarium and in his flora (1938), are G. uncin- 
ulatum. 



Gardenia Linnaeus 

Trees or shrubs, branches terete, glabrous or puber- 
ulent; stipules interpetiolar and intrapetiolar, triangular, 
apex acute to acuminate, often forming a short sheath 
at the base. Leaves opposite or in whorls of 3, subsessile 
to short-petiolate, coriaceous to chartaceous, entire, pin- 
nately veined, domatia often present. Inflorescences ter- 
minal or axillary, of 1 or 3 flowers (rarely more and 
corymbose), sessile or short-pedicellate. Flowers radially 
symmetrical, bisexual, usually large, hypanthium ovoid 
to ellipsoid or obconic, calyx tube short (sometimes 
spathe-like), calyx lobes 5-8 when present; corolla sal- 
verform to campanulate or funnelform, white or yellow, 
corolla tube glabrous or puberulent, corolla lobes 5-11, 
convolute in bud, spreading or recurved; stamens 5-9, 
inserted on the upper half of the tube, filaments short or 
absent, anthers dorsifixed, linear to linear-oblong, in- 
cluded or partly exserted, disc annular to crenate; ovary 
1 -locular (rarely 2- or 6-locular at apex), ovules many 
and horizontal on parietal placentas, style linear and 



terete, stigma linear to clavate, 1 - or 2-lobed. Fruits ob- 
long to ovoid, pyriform or globose, terete or costate, the 
outer wall fleshy to leathery or woody, rupturing irregu- 
larly or the endocarp breaking into 2-5 valves; seeds 
very many, imbedded in a fleshy pulp, horizontal, an- 
gulate, embryo small. 

A genus of about 200 species in the tropics and 
subtropics of the Old World. A few species are 
important as ornamental trees in warm climates; 
they are also grown under glass in cold climates 
for their large aromatic flowers, which are often 
used for corsages. The genus is similar to Genipa. 
The genus has not become naturalized in Central 
America, where one species is commonly seen in 
gardens. 



Gardenia augusta (L.) Merr., Interpr. Herb. Am- 
boin. 485. 1917. Varneria augusta L., Amoen, 
Acad. 4: 136. 1759. G. jasminoides Ellis, Phil. 
Trans. 51, pt. 2: 935. 1761. G.florida L., Sp. PI. 
ed. 2: 305. 1762. 



Shrubs or many-branched small trees to 5 m tall, leafy 
branchlets 1.5-5 mm thick, sparsely and minutely (0.2 
mm) puberulent, glabrescent; stipules 5-10 mm long, at 
first enclosing the apex and splitting down one side to 
become spathe-like, with a short tube 2-4 mm long at 
the base, persisting. Leaves 2(-3)/node, petioles 14 mm 
long and little differentiated from the leaf base; leaf blades 
3-12 cm long, 1.5-5 cm broad, elliptic-obovate to ellip- 
tic-oblong or broadly elliptic (in smaller leaves), apex 
acuminate, base acute and decurrent on petiole, glabrous 
above and below (but sometimes with pit domatia and 
a few hairs in the leaf axils), 2 veins 6-9 /side. Inflores- 
cences often of 3 terminal flowers, or solitary flowers in 
the axils of near-terminal leaves, pedicels ca. 1 mm 
long. Flowers large (6-10 cm long), apparently differing 
in size in different cultivars, sweetly aromatic, hypan- 
thium ca. 10 mm long, calyx lobes 8-30 mm long, nar- 
rowly oblong, spur-like; corolla white, tube 2-5 cm long, 
corolla lobes usually 6 (some cultivars with 2 series), ca. 
25 mm long and 18 mm broad, obovate; anthers ca. 18 
mm long. 



Plants of parks and gardens cultivated for their 
large white sweetly aromatic flowers. These plants, 
native to Asia, are called jazmin, jazmin delcabo, 
and "gardenia." 



Genipa Linnaeus 

Trees, branchlets usually thick, terete, glabrous or pu- 
berulent; stipules interpetiolar and intrapetiolar, connate 
to form a short tube, caducous or deciduous with the 
leaves. Leaves opposite and decussate, subsessile or pet- 



142 



FIELDIANA: BOTANY 



iolate, often large, pinnately veined, chartaceous to co- 
riaceous, without domatia. Inflorescences terminal or 
axillary, flowers solitary or in few-flowered cymes or 
subcapitate, pedicels present and continuous with the 
hypanthium base. Flowers radially symmetrical, bisex- 
ual (rarely unisexual and dioecious), 5- or 6-parted, hy- 
panthium turbinate to campanulate, calyx tube truncat- 
ed and entire or with 5-6 short lobes; corolla salverform 
to funnelform, carnose, white to yellowish white, tube 
short to long, glabrous or puberulent externally, barbate 
in the throat within and at the base of the lobes, corolla 
lobes 5-6, convolute in bud, spreading; stamens 5-6, 
inserted in the upper part of the tube, anthers subsessile, 
dorsifixed, linear, partly exserted; ovary 1-locular or be- 
coming 2-locular, style thick, stigmas fusiform, placen- 
tation parietal, ovules many and horizontal in vertical 
files. Fruits baccate, large, ovoid to subglobose or ob- 
ovoid, calyx tube persistent at apex of the fruit, pericarp 
thick, fleshy to coriaceous; seeds many, large, com- 



pressed (with two parallel flattened sides), the testa slight- 
ly fibrous. 



A genus of 5-10 species, ranging from southern 
Florida (U.S.A.) and Mexico through Central 
America into tropical South America. The genus 
is distinguished by its few-flowered terminal and 
subterminal inflorescences, large flowers with thick 
corolla lobes, parietal placentation, and many large 
horizontal seeds in vertical files within the large, 
often solitary fruit. Cenipa vulcanicola Stand!, of 
Mexico and Guatemala has been transferred to 
Glossostipula concinna (Standl.) Lorence; it has 
axile placentation. 



Key to the Species of Genipa 

la. Corolla densely sericeous distally, peduncles to 25 mm long; stipules acute at apex, persisting with 
the leaves; leaves glabrous or pubescent, with 7-18 major secondary veins on each side; widespread 
G. americana 

Ib. Corolla glabrous on the outer surfaces, peduncles to 10 mm long; stipules broadly ellipsoid and 
rounded at apex, usually caducous; leaves glabrous above and with appressed hairs on the veins 
beneath, with 69 major secondary veins on each side; not recorded north of southern Costa Rica 

. G. williamsii 



Genipa americana L., Syst. Nat. ed. 10. 2: 931. 
1759. G. oblongifolia Ruiz & Pav., Fl. Peruv. 
Chil. 2: 67, pi. 220. 1798. G. caruto H.B.K., 
Nov. gen. sp. 3: 407 (quarto). 1820. G, ameri- 
cana var. caruto (H.B.K.) Schum. in Mart., Fl. 
Bras. 6(6): 352. 1889. G. codonocalyx Standl., 
Contr. U.S. Natl. Herb. 1 7: 446. 1 9 1 4. G. venosa 
Standl., J. Wash. Acad. Sci. 18: 168. 1928. Fig- 
ure 26. 



Small to large trees 4-27 m tall, often with a spreading 
hemispheric crown, trunk to 50 cm thick, bark smooth 
and lenticellate, leafy branchlets 4-9 mm thick, densely 
pubescent in early stages or glabrous; stipules 10-25 mm 
long, triangular, the basal sheathing tube 1-3 mm long, 
acute, deciduous with the leaves. Leaves with petioles 
2-13 mm long, 2-3 mm thick, glabrous or pubescent; 
leaf blades 1 2-42 cm long, (4-)6-19 cm broad, obovate 
to elliptic-obovate or broadly oblanceolate, apex acu- 
minate to obtuse or rounded, gradually narrowed to a 
cuneate or slightly decurrent base, drying chartaceous 
and often very dark above, glabrous and lustrous above, 
glabrous to densely pubescent beneath with thin soft 
hairs ca. 0.5 mm long, 2 veins 9-1 8/side. Inflorescences 
terminal or subterminal, 4-10 cm long and with 1-9 
flowers, cymose, peduncles to 25 mm long, glabrous, 
pedicels 4-12 mm long. Flowers 2.5-4 cm long, appar- 



ently bisexual but perhaps functionally unisexual, hy- 
panthium difficult to distinguish from the calyx tube and 
together 7-17 mm long, calyx tube to 10 mm diam. 
distally, truncate or with broad short lobes, glabrous on 
the exterior and puberulent within; corolla ca. 4 cm broad, 
salverform, carnose, densely descending-sericeous ex- 
ternally (except at the base of the tube), white or yellow- 
ish white, darkening with age, tube 5-15 mm long, 4-7 
mm diam. (to 10 mm at the lobes), lobes 5-6, 11-28 
mm long, 5-12 mm broad, obovate and rounded at apex; 
anthers 6-14 mm long, becoming recurved between the 
lobes; stigmas ca. 5 mm long. Fruits 4-1 1 cm long, 3- 
1 1 cm diam., obovoid to subglobose, smooth and grayish 
brown or yellowish brown, the persisting calyx 3-6 mm 
long, crateriform on apex of the fruit and 8-10 mm 
diam., pedicels up to 5 cm long in fruit; seeds 6-12 mm 
long, 4-7 mm broad, ca. 2.3 mm thick. 



Trees of both wet evergreen rain forests and 
seasonally very dry deciduous forest formations 
in the Caribbean and Pacific lowlands, from near 
sea level to 900 m elevation. Flowering occurs 
primarily in March-August; fruiting throughout 
the year. This species ranges from southern Flor- 
ida and the West Indies, through Mexico and Cen- 
tral America through tropical South America to 
Paraguay. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



143 



Genipa atnericana is recognized by the larger 
leaves with short petioles clustered at the ends of 
stems, usually solitary large fruit with many hor- 
izontal seeds in vertical files, and large sericeous 
flowers with short corolla tubes and large lobes. 
This is a common and distinctive tree, especially 
conspicuous in deciduous forest formations in the 
dry season because of its large terminal fruit (but 
compare Alibertia edulis). This species may be 
confused with species of Borojoa, but those tend 
to have sessile terminal flowers and stipules with 
parallel venation. Guaitil, caruto, jagua, andjagua 
negro are common names for this species. The 
juice of the young pulpy fruit turns black or dark 
blue and is used by Native Americans as a dye or 
body paint. The species is sometimes cultivated, 
and the fruit is eaten. The wood is easy to work 
but strong and resistant; it is used for making fur- 
niture and carts and in building construction. 

Genipa americana is here interpreted to be a 
very variable species, following Dwyer (1980) and 
Steyermark (1974). The types of Standley's G. co- 
donocalyx(Pittier 12085 us) and G. venosa (Stand- 
ley & Valeria 45269 us) appear to represent no 
more than unusual forms of G. americana and 
were described when the full pattern of variation 
in G. ameriana was not apparent. The type of G. 
venosa has prominent petioles (3.54 cm long) and 
unusually long fruit, but it seems better to treat it 
as a variant of G. americana rather than as a dis- 
tinct species. Collections with the leaves densely 
pilose beneath have been referred to variety caruto 
(H.B.K.) K. Schum. 



Genipa williamsii Standl., J. Wash. Acad. Sci. 8: 
642. 1918. Figure 26. 

Small to medium-sized trees, 4-20 m tall, with boles 
ca. 25 cm dbh, leafy stems 3-7 mm thick, glabrescent, 
becoming pale brown; stipules 10-32 mm long, 6-18 
mm broad, ovate from a narrowed base, flattened, ap- 
pressed-sericeous. Leaves with petioles 7-22(-60) mm 
long, 1 .5-2.7(-4) mm thick, glabrous, often drying black- 
ish and lustrous; leaf blades 8-17(-33)cm long, 5-10(-16) 
cm broad, obovate-oblong, to broadly elliptic or elliptic- 
oblong, apex rounded and lacking a narrowed tip, base 
obtuse to acute and slightly decurrent on petiole, drying 
stiffly chartaceous to subcoriaceous and dark reddish 
brown above, glabrous above, appressed-sericeous on 
the major veins beneath with hairs ca. 0.3 mm long, 2 
veins 7-9/side and weakly loop-connected distally, 3 
venation obscure. Inflorescences of ca. 3 (4-7) terminal 
flowers subtended by 2 ovate-lanceolate bracts (stipules) 
ca. 1 2 mm long with glabrous surfaces but ciliolate along 
the edge, peduncles 5-10 mm long, pedicels 8-10 mm 
long and continuous with the hypanthium, drying black, 



bracteoles 3 mm long or reduced to ridges. Flowers gla- 
brous externally, drying black, hypanthium 3-4 mm long, 
4-5 mm broad at apex, obconic, calyx tube 0.5-1 mm 
high, entire or slightly undulate; corolla salverform, white 
and carnose, corolla tube 1.8-3 cm long, 4-5 mm diam., 
lobes 5, 18 mm long, 8-12 mm broad distally, obovate- 
oblong and rounded distally; anthers sessile, stigmas 4, 
to 4 mm long, unequal. Fruits subglobose, ca. 7 cm 
diam., drying black; seeds ca. 10 mm long, 5-6 mm 
broad, imbedded in white pulp. 

This species has been collected only in south- 
ernmost Limon Province at 450-650 m elevation 
(Hammel et al. 1 7597 CR, MO, Herrera 3208 CR.MO) 
in Costa Rica. Flowering in July; fruiting in April- 
June and October in Panama. The species ranges 
to Colombia. 

Genipa williamsii is recognized by the leaves 
rounded distally, clavate flower buds, and both the 
fleshy flowers and the large fruits that dry black. 
It is similar to species of Ladenbergia, but those 
have domatia and valvate corolla lobes. 



Geophila D. Don 

REFERENCE L. O. Williams, Geophila (Rubi- 
aceae) in North America. Phytologia 26: 263-264. 
1973. 

Creeping perennial herbs, stems slender and puberu- 
lent or glabrous, rooting at the nodes; stipules interpetio- 
lar, small, rounded-ovate to triangular, entire to shal- 
lowly bilobed, persisting. Leaves usually with long 
petioles; leaf blades rounded and often cordate at the 
base, membranaceous to thin-chartaceous, venation pin- 
nate, domatia absent. Inflorescences terminal or pseu- 
doaxillary, few-flowered heads or cymes, peduncles short 
or long, flowers subtended by an involucre of small bracts, 
pedicels short or absent. Flowers radially symmetrical 
and bisexual, usually 5-parted (less often 4-7-parted), 
calyx tube with 4-7 narrow lobes, persistent; corolla fun- 
nelform to salverform, white, corolla tube narrow, pilose 
in the throat, corolla lobes 4-7, valvate in bud, spreading 
or recurved; stamens 4-7, filaments filiform and inserted 
in the floral tube, anthers dorsifixed, linear, half exserted; 
ovary 2-locular, ovules solitary in each locule and basal, 
style slender with 2 stigmas. Fruits a juicy berry, usually 
containing 2 1 -seeded pyrenes (nutlets); pyrenes plano- 
convex and smooth or costate on the dorsal surface, with 
a ventral sulcus. 



A genus of 20-30 species native to the American 
tropics, Africa, and Asia. The slender creeping 
stems, long-petiolate leaves with rounded blades, 
few-flowered inflorescences, and fleshy, two-seed- 
ed fruits characterize this genus. These plants may 
be mistaken for species ofCoccocypselum, but that 
genus has many-seeded fruit. 



144 



FIELDIANA: BOTANY 



Key to the Species of Geophila 

la. Fruit black at maturity, pyrenes weakly costate; peduncles 2-10 cm long; basal lobes of the leaf 

blades separated by a sinus (cordate with non-overlapping lobes) G. macropoda 

Ib. Fruit red at maturity (if fruit are blue go to the genus Coccocypselum), pyrenes strongly costate; 

peduncles 0.2-10 cm long; basal lobes of the leaf blades separate to overlapping 2 

2a. Ovary, fruit and leaves conspicuously pilose with thin hairs 0.5-2 mm long [leaf blades subcordate 

with a small basal sinus; peduncles to 7 cm long in fruit] G. cordifolia 

2b. Ovary and fruit glabrous, leaves glabrous or puberulent with short (0.1-0.3 mm) hairs; peduncles 

to ca. 2 cm long 3a 

3a. Leaf blades ovate and slightly longer than broad, often narrowed at apex, cordate but usually without 

a visible basal sinus (because the lobes overlap slightly) G. repens 

3b. Leaf blades ovate-triangular and distinctly longer than broad, usually acute at apex, cordate to 

subcordate at the base and with a small sinus G. gracilis 



Geophila cordifolia Miq., Stirp. Surin. Sel. 176. 
1 850. Mapouria trichogyne Muell.-Arg. in Mart., 
Fl. Bras. 6(5): 426. 1881. Geophila trichogyne 
(Muell.-Arg.) Standl., Publ. Field Columb. Mus., 
Bot. Ser. 7: 423. 1931. Figure 2. 

Creeping herbs, leafy stems 0.7-1 .5 mm thick, densely 
pubescent with slender pale straight or crooked hairs 0. 5- 
2 mm long; stipules 2-4(-6) mm long, 1.5-4 mm broad 
(broadest beneath the inflorescences), usually glabrous, 
persisting. Leaves with petioles 3-7(-l 3) cm long (short- 
er on leaves subtending the inflorescences), conspicu- 
ously pubescent with slender crooked or straight mul- 
licellular hairs 0.7-1.5 mm long; leaf blades 3-7(-ll) 
cm long, 2-6(-8.5) cm broad, broadly ovate to oblong- 
ovate or narrowly ovate (triangular-ovate), apex obtuse 
to short-acuminate, base cordate with rounded lobes 8- 
35 mm broad, basal sinus 3-15 mm deep, drying thin- 
chartaceous and brownish, both surfaces covered with 
thin usually crooked hairs 0.7-2 mm long, 2 veins 3- 
5/side, not usually loop-connected near the margin. In- 
florescences terminal, capitate, 1-3 cm long (to 6 cm in 
fruit), with 5-1 7 flowers, peduncles 5-1 5 mm long, elon- 
gating in fruit, densely pubescent, bracts 3-10 mm long, 
lanceolate and pubescent, pedicels 0.5-3 mm long. Flow- 
ers with an urceolate hypanthium, calyx tube 0.5 mm 
long and 1 .5 mm diam.. lobes ca. 3 mm long, with slen- 
der hairs ca. 1 mm long; corolla 5-6 mm long, white 
often tinged with pink distally, tube 2-4.5 mm long, 
glabrous externally, with a short collar of hairs at the 
point of filament attachment within, lobes 5, 1 .5-2.5 mm 
long; stamens with anthers 0.8-1 mm long, included. 
Fruits ca. 8 mm long, ovoid-globose, orange to red, with 
scattered slender hairs, pyrenes ca. 4 mm long and 3 mm 
broad, with 3-5 prominent longitudinal dorsal ribs. 

Plants of evergreen forest formations, from near 
sea level to 500(-1100) m elevation. Probably 
flowering and fruiting primarily in the wet season 
(May-December). The species ranges from Belize 
along the Atlantic slope of Central America to 
Colombia, Venezuela, and the Amazon basin in 
Brazil and Peru. 



Geophila cordifolia is recognized by the long, 
often crooked, thin multicellular hairs that cover 
almost all parts of the plants. The long-petiolate 
leaves with cordate bases, short inflorescences 
elongating in fruit, and glabrous stipules are ad- 
dition distinctions. In Costa Rica the species, is 
known only from the La Selva area and from Vol- 
can Rincon de la Vieja. 



Geophila gracilis (Ruiz & Pav.) DC, Prodr. 4: 537. 
1 830. Psychotria gracilis Ruiz & Pav., Fl. Peruv. 
2: 63, pi. 211, f. C. 1799. G. croatii Steyerm., 
Phytologia35:401. 1977. 

Creeping herbs to 20 cm tall or slender-stemmed vines 
to 1.5 m long, leafy stems 0.3-1.3 mm thick, glabrous, 
horizontal internodes 3-7 cm long, with adventious roots 
near the nodes; stipules 2-4 mm long. 1-2 mm broad, 
usually rounded at apex, curving outward in age and 
persisting. Leaves with petioles 2-8 cm long (but shorter 
below the inflorescences), 0.3-1 mm thick, glabrous 
abaxially and with 2 rows of short (0.2-0.5 mm) stiff 
retrorse or erect hairs along either side of the adaxial 
sulcus; leaf blades 1.8-5 cm long, 1-3.5 cm broad, tri- 
angular-ovate to ovate, apex gradually narrowed and 
acute (or obtuse), base cordate to subcordate, sinus 3-8 
mm deep, the basal lobes usually separate but occasion- 
ally overlapping and the sinus obscured, drying thin- 
chartaceous, glabrous above or with few thin hairs 0.3- 
0.8 mm long, usually glabrous beneath, 2 veins 3-47 
side, usually loop-connected near the margin. Inflores- 
cences terminal, 12-15 mm long, capitate with ca. 3-7 
flowers, peduncles 2-8 mm long (apparently longer when 
the leaves of the subtending node are reduced and bract- 
like, bracts 4-7 mm long, 1-1.5 mm broad, united at 
the base and persisting, pedicels 0-1 mm long. Flowers 
with hypanthium ca. 2 mm long, calyx lobes ca. 3 mm 
long and 0.5 mm broad, narrowly oblong, persistent and 
enlarging in fruit; corolla white. Fruits red at maturity, 
3-5 mm long, subglobose. pyrenes ca. 3.5 mm long, with 
3 prominent longitudinal dorsal ridges. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



145 



Plants of evergreen forest formations, from near 
sea level to ca. 500 m elevation. Flowering in May- 
December (primarily in June and July in central 
Panama; Croat, 1978). The species is known from 
southeastern Nicaragua, the Canal area of Pana- 
ma, and the upper Amazon basin of Brazil, Peru, 
and Bolivia. 

Geophila gracilis is recognized by its slender 
stems rooting at most nodes, triangular-ovate leaf 
blades with little or no puberulence, and short 
terminal capitate inflorescences subtended by 
bracts fused at the base. Variation in specimens 
from South America clearly encompass the dis- 
tinctions used to separate G. croatii. This species 
is apparently common on Barro Colorado Island, 
Panama, and has been collected in Nicaragua, but 
it has yet to be collected in Costa Rica. 



Geophila macropoda (Ruiz & Pav.) DC., Prodr. 4: 
537. 1830. Psychotria macropoda Ruiz & Pav., 
Fl. Peruv. 2: 63, pi. 21 1, f.6. 1799. Figure 2. 

Creeping herbs, leafy stems 0.8-2 mm thick, glabrous 
or very minutely (0.1 mm) puberulent, often with 2 
prominent longitudinal ridges; stipules 2-6 mm long, 
ovate-oblong, glabrous, deciduous or obscured by the 
adventitous roots. Leaves with petioles (2-) 3-1 4 cm long 
(shorter below the inflorescences), 0.7-1.2 mm thick, 
glabrous abaxially but with 2 adaxial ridges with short 
(0.1-0.4 mm) dense hairs; leaf blades 3-9 cm long, 2.5- 
8 cm broad, broadly ovate to ovate-orbicular, apex 
rounded to broadly obtuse, base cordate with lobes 1-4 
cm broad, basal sinus 2-15 mm deep, drying membra- 
naceous or thin-chartaceous and often grayish green, gla- 
brous above, glabrous beneath except for the minute 
puberulence on the major veins near the base, 2 veins 
3-5/side and weakly loop-connected near the margin. 
Inflorescences usually axillary, 2-4 cm long and elon- 
gating in fruit, capitate with 3-7 flowers, peduncle 1.5- 
5(-7) cm long, minutely puberulent with whitish hairs 
ca. 0. 1 mm long, bracts 3-6 mm long, united at the base, 
pedicels to 2 mm long. Flowers with hypanthium ca. 1.5 
mm long, essentially glabrous, calyx lobes 2-3 mm long; 
corolla 3-7 mm long, tube 3-4 mm long, corolla lobes 
5, 2-3 mm long. Fruits black or blue, sessile, 5-10 mm 
long, 3-7 mm diam.. ellipsoid to ovoid; pyrenes 4-7 mm 
long, 2-3 mm broad, without raised longitudinal ribs 
(costae) on the convex surface. 

Plants of the lowland Caribbean rain forest for- 
mations, from near sea level to 600 m. Flowering 
in April-November. The species ranges from 
southern Mexico through Central America to Bo- 
livia and Paraguay. 

Geophila macropoda is recognized by the axil- 
lary and long-pedunculate inflorescences, closely 
clustered flowers, black fruit, and pyrenes without 
prominent longitudinal costae. 



Geophila repens (L.) I. M. Johnston, Sargentia 8: 
281. 1949. Rondeletia repens L., Syst. ed. 10: 
928. 1759. Psychotria herbaceajacq., Enum. PI. 
Carib. 16. 1760. Geophila herbacea (Jacq.) 
Schumann in Engl. & Prantl., Nat. Pflanzenfam. 
4,4: 119. 1891. Figure 2. 



Creeping herbs to ca. 10 cm high, leafy stems 0.5-1 
mm thick, glabrous or very sparsely and minutely pu- 
berulent; stipules 0.5-2 mm long, 1-2 mm broad, broad- 
ly ovate, glabrous, persisting or deciduous. Leaves with 
petioles l-6(-8.5) cm long (sometimes shorter in leaves 
subtending the inflorescences), 0.4-1.3 mm thick, gla- 
brous on the abaxial surface but with short (0.2-0.5 mm) 
retrorse or crooked hairs along the adaxial (upper) side; 
leaf blades 1.2-5.5 cm long, 1-5 cm broad, broadly ovate 
to ovate-suborbicular, apex bluntly obtuse or rounded- 
obtuse, base cordate with lobes 5-25 mm broad, basal 
sinus 1-7 mm deep and usually obscured by the over- 
lapping basal lobes, drying membranaceous, upper sur- 
face glabrous or with a few short hairs near the margins, 
glabrous beneath or with a few short (0. 1-0.3 mm) hairs 
on the veins near the petiole, 2 veins 3-5/side, cystoliths 
visible (as short whitish lines) or obscure on the lower 
surface. Inflorescences solitary and terminal on short 
leafy shoots (apparently 2-3 and axillary when directly 
subtended by small leaves), with (l-)2-5 flowers, pe- 
duncles 5-35 mm long, puberulent with short retrorse 
hairs, subtending bracts 3-6 mm long, united at the base 
and lanceolate distally, pedicels 0-2 mm long. Flowers 
white or becoming pink in age, hypanthium ca. 1 mm 
long, calyx tube 1-1.5 mm long, glabrous, calyx lobes 
1-3 mm long, 0.3-0.7 mm broad, glabrous; corolla 8- 
14 mm long, glabrous or puberulent, funnelform, tube 
6-9 mm long and 1-1.5 mm diam., lobes 5, 3-5 mm 
long, 1.5-2.5 mm broad, ovate and obtuse to acute; sta- 
mens with filaments ca. 0.5 mm long, attached near the 
middle of the tube, anthers ca. 2 mm long; style 5-7 mm 
long. Fruits 8-10 mm long, ovoid or globose, bright red 
at maturity, subsessile or short (1-2.5 mm) pedicellate; 
pyrenes 3.5-5 mm long, 2.5-3 mm broad, with 3 slightly 
raised longitudinal ribs (costae) on the convex face. 



Plants of the shaded forest floor in evergreen or 
partly deciduous forest formations of both the Ca- 
ribbean and Pacific lowlands of Costa Rica, from 
near sea level to 800 m elevation. Flowering in 
June-October; fruiting in July-November. The 
species ranges from Mexico and the West Indies 
to Peru and Bolivia in the New World; it is also 
found in West Africa, the Philippines, and the 
western Pacific. 

Geophila repens is recognized by its creeping 
habit and short stature, small cordate leaves with 
the sinus obscured, petioles with puberulence along 
one side, bright red fruit, and pyrenes with weakly 
developed costae. This is our most commonly en- 
countered species of Geophila; it has been col- 
lected at La Selva and from near Canas in Guana- 
caste Province to the Golfo Dulce area along the 



146 



FIELDIANA: BOTANY 



Pacific. Lechuga is a name used for this species in 
the Golfo Dulce region. Breeding biology was stud- 
ied by Bawa and Beach (1983). 



Gonzalagunia Ruiz Lopez & Pavon 

Shrubs or small trees, distal branches often curved, 
pendant or scandent, leafy stems usually slender and 
pubescent, terete, nodes usually thickened; stipules in- 
terpetiolar, usually broad at the base and triangular with 
a narrow distal awn (rarely intrapetiolar and tubular). 
Leaves distichous, petiolate or subsessile; leaf blades of- 
ten thin-chartaceous, usually narrow and with ascending 
secondary veins, domatia absent or obscure. Inflores- 
cences solitary, usually terminal and narrowly long-ra- 
cemiform, spiciform or thyrsiform, the flowers solitary, 
cymose or fasciculate on short lateral branches of the 
central axis, bracteoles present, flowers sessile or pedi- 
cellate. Flowers radially symmetrical, bisexual, mono- 
morphic or distylous, small, 4- (less often 5-) parted, 
hypanthium urceolate to rounded, calyx tube very short, 
calyx lobes 4 or 5, small, equal or unequal, persisting in 
fruit; corolla salverform to funnelform, white or pink, 
corolla tube narrow, villose in the throat, corolla lobes 



4(-5), imbricate or valvate; stamens 4(-5), filaments short 
or absent, anthers dorsifixed, 2-lobed at the base, ex- 
serted or partly exserted; ovary 2-or 4-locular, ovules 
numerous, placentation peltate on the septum, style with 
2 or 4 stigmatic lobes. Fruits baccate, fleshy or spongy, 
subglobose, 2- or 4-locular and usually with 2 or 4 lobes 
or sulci, with 2 or 4 hard cocci; seeds 4-many within 
the cocci, minute. 

A genus of 25-35 species, ranging from Mexico, 
Central America, and the West Indies through 
tropical South America. The long narrow spike- 
like distal inflorescences, slender drooping distal 
stems, small narrow-tubed flowers and baccate 
fruits with 2-4 pyrenes make this a very distinctive 
genus. Some species of Rondeletia with long-nar- 
row inflorescences may be confused with species 
of Gonzalagunia in the absence of fruit; Rondeletia 
has capsular fruit. Several of our species are weedy 
shrubs of open secondary growth and closely re- 
lated; they can be difficult to distinguish in the 
absence of mature flowers or fruit. A few species 
resemble species of Buddleia (Loganiaceae). 



Key to the Species of Gonzalagunia 

la. Stipules united or overlapping above the petioles to form a short tubular sheath or broad tube-like 

base 4-12 mm long 2 

Ib. Stipules not forming a tube above the node, or the broad margins not overlapping at the base, stipule 

only 1-3 mm long before being narrowed into the awn-like apex 3 

2a. Stipular sheath to 1 cm long; flower clusters and lateral branches of the inflorescences subtended 

by caducous bracts 4-12 mm long and 1-2 mm broad, corolla 34 mm long; leaf blades with 

5-8 pairs of major secondary veins; small treelets of wet forest understory .... G. bracteosa 
2b. Stipular sheath 0-5 mm long or the stipule margins slightly overlapping; bracts less than 2 mm 

long, corolla 4-6 mm long; with 10-14 pairs of secondary veins; trees to 18 m tall 

Rondeletia brenesii 

3a. Petioles usually less than 4 mm long; corolla tube usually less than 7 mm long 4 

3b. Petioles usually more than 4 mm long: corolla tube usually more than 7 mm long 7 

4a. Leaf blades with 4-7 pairs of major secondary veins, laminae thin-textured; flowers thin-textured 

and often solitary, corolla glabrous externally G. rudis 

4b. Leaf blades with 7-1 3 pairs of major secondary veins; thinly to stiff-chartaceous; flowers thick- 
textured, corolla densely sericeous externally 5 

5a. Leaf blades subsessile, narrowly lanceolate, 9-26 cm long, with 9-15 pairs of secondary veins 

arising at angles of about 3040 [only known from the Golfo Dulce area] G. brenesii 

5b. Leaf blades subsessile or short petiolate, ovate to elliptic and rarely lanceolate, to 1 6(-30?) cm 

long, with 4-12 pairs of secondary veins arising at angles of 40-60 5 

6a. Corolla lobes ca. 1.5 mm long; leaf blades with 4-1 1 pairs of major secondary veins, ovate to 

ovate-elliptic, thinly chartaceous; Caribbean and Pacific slope, 0-1200 m elevation 

G. ovatifolia 

6b. Corolla lobes 3-4 mm long; leaf blades with 7-12 pairs of major secondary veins, ovate-elliptic 

to lanceolate, stiffly chartaceous; wet Caribbean slope 300-1200 m elevation 

G. stenostachya 

7a. Mature fruit becoming blue-black; corollas white or white tinged with pink, corolla tubes 8-13 mm 
long; inflorescences with the flower clusters sessile; 0-1200(-1500) m elevation . . G. panamensis 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



147 



7b. Mature fruit white; corollas reddish to pink, corolla tubes 6-10 mm long; inflorescences with the 
flower clusters on short (1-3 mm) peduncles (but note that some inflorescences may only have 
solitary flowers and no apparent secondary peduncles); 900-2200 m elevation G. rosea 



Gonzalagunia bracteosa (J. D. Smith) B. L. Rob- 
inson, Proc. Amer. Acad. Arts 45: 405. 1910. 
Gonzalea bracteosa J. D. Smith, Hot. Gaz. 33: 
252. 1902. Figure 21. 



rescences appear to be restricted to northeastern 
Costa Rica. This species differs greatly from our 
other members of the genus; it resembles Psy- 
chotria pilosa. 



Shrubs or small treelets, 1.5-4 m tall, leafy branchlets 
2-4(-6) mm thick, terete, with many ascending strigose 
hairs 1-1.5 mm long; stipules 8-18(-24) mm long, 3-6 
mm broad, with a tubular sheath to 12 mm long, dark 
brown and pubescent along the midvein, acute to acu- 
minate and with a slender tip 1-7 mm long. Leaves with 
petioles 2-10 mm long, 1-2 mm thick, broad, pubescent; 
leaf blades 6-19(-22) cm long, 2.5-7.5(-8.5) cm broad, 
narrowly elliptic-obovate to obovate-oblong, oblong or 
elliptic, usually broadest above the middle, apex acu- 
minate (acute), tip to 1 5 mm long, gradually narrowed 
to the acute or obtuse base, leaves usually drying char- 
taceous and dark brown above (rarely subcoriaceous), 
sparsely pubescent above with thin appressed hairs 0.5- 
1 mm long (densely strigulose on the midvein), more 
densely appressed-pubescent beneath with brownish hairs 
0.5-1.3 mm long, 2 veins 5-7 /side. Inflorescences sol- 
itary, axillary or terminal, 6-25 cm long, spike-like or 
thyrsiform panicles with small (5-15 mm) alternate or 
opposite flower clusters 2-14 mm distant along the rachis 
(rarely with lateral branches to 4 cm long), primary pe- 
duncles 2-8 cm long, with ascending hairs to 2 mm long, 
several bracts 5-12 mm long and 1-2 mm broad sub- 
tending the flower clusters, flowers sessile. Flowers with 
hypanthium ca. 1 mm long, densely hirtellous, calyx tube 
minute, calyx lobes ca. 0.5 mm long, difficult to see 
among the hairs, glabrous on the inner surface; corolla 
funnelform, greenish white to white, with few straight 
hairs 0.5 mm long externally, tube 2-3 mm long, corolla 
lobes 5, 1-2 mm long; stamens 5, anthers ca. 1 mm long; 
style ca. 2.5 mm long, stigma 0.4 mm long. Fruits 3-5 
mm long, 3-5 mm broad, depressed globose, becoming 
blue or blue-black, pubescent, sessile; pyrenes 2-4. 

Plants of lowland rain forest formations on the 
Caribbean and Pacific slopes in Costa Rica, from 
10 to 850 m elevation. Flowering in January-Sep- 
tember; fruiting throughout the year. The species 
ranges from northeastern Costa Rica to Colombia. 

Gonzalagunia bracteosa is characterized by the 
hirsutulous pubescence on many parts, obovate- 
oblanceolate leaves drying dark above, narrow in- 
florescences with short flower clusters subtended 
by conspicuous bracts, small sessile flowers, and 
tubular stipules. Two collections are noteworthy 
because the inflorescences have lateral branches 
2-4 cm long and with many bracts along their 
length: Folsom 9778 (DUKE, F) and Zamora & San- 
chez 469 (CR, F). Specimens exhibiting such un- 
usual variation in bract development and inflo- 



Gonzalagunia brenesii Standl., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 18: 1302. 1938. Figure 20. 



Shrubs, 1.5-3 m tall, leaf branchlets 1.5-6(-8) mm 
thick, rounded-quadrangular in cross-section, sparsely 
pubescent with appressed hairs ca. 0.5 mm long or gla- 
brous, drying reddish brown; stipules 4-8 mm long, 2.5- 
7 mm broad at the base, with a very short (1-2 mm) 
broadly triangular base and a slender awn 4-6 mm long. 
Leaves subsessile, petioles 0-3(-5) mm long; leaf blades 
9-26 cm long, 2.7-6 cm broad, narrowly lanceolate to 
narrowly elliptic-oblong or elliptic-oblanceolate, apex ta- 
pering gradually and long-acuminate, base obtuse, drying 
chartaceous and dark brown above, lustrous above in 
life, glabrous or sparsely pubescent above, with thin 
whitish ascending hairs beneath, the hairs longer (ca. 0.4 
mm) on the major veins beneath, 2 veins 9-1 3/side and 
weakly loop-connected distally, 3 veins subparallel. In- 
florescences solitary and terminal, 20-50 cm long, spi- 
ciform with short (1-5 mm) lateral branches bearing 2- 
6 flowers, peduncles 0-6 cm long, 1.5-3 mm thick, with 
ascending appressed hairs, bracts 1-3 mm long, linear, 
pedicels 1-2 mm long. Flowers 6-12 mm long, hypan- 
thium 0.5-1 mm long, sericeous at the base, calyx tube 
ca. 0.5 mm long, calyx lobes 0.2-0.3 mm long, glabrous; 
corolla white or pink, tube 7-10 mm long and 0.7-1 mm 
diam., pubescent, lobes 3 mm long and 1.8 mm broad, 
obtuse; ovary 4-locular. Fruits 3-5 mm long, 3-6 mm 
broad, usually 4-lobed, becoming white, minutely and 
sparsely puberulent. 

Plants of rain forest formations in Costa Rica's 
southern Pacific lowlands, from near sea level to 
500 m elevation. Flowering in April-December; 
fruiting in January and August-September. This 
species is endemic to Costa Rica, ranging from the 
forests of the Pacific slope above Quepos eastward 
to the Osa Peninsula. 

Gonzalagunia brenesii is distinguished by its re- 
stricted range, long narrow subsessile leaves, long 
spicate inflorescences, and short narrowly tubular 
white or pink flowers. This species appears to be 
related to the G. panamensis-G. rosea complex. 



Gonzalagunia ovatifolia (J. D. Smith) B. L. Rob- 
inson, Proc. Amer. Acad. Arts 45: 405. 1910. 



148 



FIELDIANA: BOTANY 



Gonzalea ovatifolium J. D. Smith, Bot. Gaz. 27: 
336. 1899. Figure 20. 

Shrubs to 3(-4) m tall, leafy branchlets 1.3-4 mm 
thick, with thin ascending brownish hairs ca. 0.5 mm 
long, terete, glabrescent; stipules 6-10 mm long, subulate 
with a short (1-2 mm) base and long (4-7 mm) awn-like 
apex. Leaves subsessile with petioles l-4(-5) mm long, 
ca. 1.3 mm thick, pubescent; leaf blades 5-12(-16) cm 
long, 2-6(-7) cm broad, ovate to ovate-elliptic, apex 
short-or long-acuminate, tip 3-20 mm long, abruptly 
narrowed to rounded at the obtuse base, unequal at the 
base, the leaves drying thin-chartaceous and dark above, 
pale grayish to pale greenish beneath, essentially glabrous 
above, appressed-pubescent on the veins beneath with 
thin short (ca. 0.5 mm) hairs, 2 veins 6-1 I/side. Inflo- 
rescence solitary and terminal, 12-45 cm long, narrowly 
spike-like with flowers in distant (3-10 mm) clusters, 
rachis slender 0.5-1 mm thick, with thin whitish as- 
cending hairs ca. 0.5 mm long, flowers in sessile or sub- 
sessile groups of 1-3, subtended by linear bracts 2-4 mm 
long, flowers usually solitary in the distal half of the 
inflorescence, pedicels to 1 mm long. Flowers with hy- 
panthium ca. 1 mm long, densely sericeous, calyx lobes 
1-2 mm long, glabrous; corolla white, tube 4-5 mm long, 
lobes 1-2 mm long; anthers ca. 1 mm long. Fruits 2-3 
mm long and 3-4 mm broad (dried), white, usually 
4-lobed, pubescent with thin hairs ca. 0.3 mm long. 

Shrubs of wet evergreen forest formations of 
both the Caribbean and Pacific slopes in Costa 
Rica, from near sea level to 1200 m elevation. 
Flowering in January-April and July-August; 
fruiting in the same months and in October and 
December. The species ranges from Nicaragua to 
Colombia. 

Gonzalagunia ovatifolia is recognized by its sub- 
sessile ovate acuminate leaves, the long inflores- 
cences with few-flowered cymules or solitary flow- 
ers along its length, and the small corollas. This 
species may be difficult to separate from some 
specimens of G. rosea (q.v.). 



Gonzalagunia panamensis (Cav.) K. Schum. In 
Mart., Fl. Bras. 6(6): 292. 1889. Buena pana- 
mensis Cav., Anales Hist. Nat. 2: 279. 1800. 
Gonzalea panamensis (Cav .) Spreng., Syst. Veg. 
1:417. 1825. Figure 20. 

Shrubs to 3(-5) m tall, erect or scandent, leafy branch- 
lets 0.8-4 mm thick, terete or slightly quadrangular, 
sparsely to densely sericeous with thin whitish ascending 
hairs ca. 0.5 mm long; stipules 4-7 mm long, the broad 
base 0.5-2 mm long, subulate with a narrow awn-like 
tip, pubescent on the midrib and edges. Leaves with 
petioles 6-27 mm long (shorter on young axillary shoots), 
ca. 1 mm thick, pubescent, with lateral margins contin- 
uous with the decurrent leaf margin; leaf blades 5-15 
cm long, 1-6 cm broad, narrowly lanceolate to lanceo- 



late-elliptic or ovate-lanceolate, apex tapering gradually 
and acute or acuminate, base acute and often decurrent 
on petiole, drying thin-chartaceous, densely pubescent 
on the veins and more sparsely between the veins with 
appressed hairs ca. 0.3 mm long above and below, 2 
veins 5-7/side, ascending. Inflorescences solitary and 
terminal (or axillary by later lateral growth of side shoots), 
6-22(-40) cm long, peduncles 1-5 cm long, 1-2 mm 
thick, pubescent, the flower clusters ca. 5 mm diam., 
essentially sessile (but solitary flowers pedicellate), bracts 
to 4 mm long and linear, pedicels 0-2 mm long. Flowers 
4-parted, hypanthium 0.7-1 mm long, sparsely pubes- 
cent, calyx ca. 1 mm long, calyx lobes 0.5-1 mm long 
and 0.5 mm broad at the base; corolla white or white 
tinged with pink, tube (8-)10-13 mm long, 0.7-1 mm 
diam., sparsely pubescent, lobes 2-3 mm long, ca. 2 mm 
broad at the base, puberulent within; stamens 4, anthers 
2-2.2 mm long; style 7-10 mm long, stigma ca. 0.7 mm 
long. Fruits 2.5-4 mm long, 3-8 mm broad, depressed 
globose to 4-lobed, glabrous or sparsely puberulent, red 
becoming purple black or black. 

Shrubs of evergreen or partly deciduous forest 
formations of the Pacific slope in Costa Rica, from 
near sea level to 1 200 m elevation. Flowering in 
all months except March-April and November; 
fruiting in January, February, September-Octo- 
ber, and December. The species ranges from Mex- 
ico to Colombia and in the West Indies. 

Gonzalagunia panamensis is recognized by the 
usually lanceolate leaves on slender well-devel- 
oped petioles, the flower clusters sessile on the 
inflorescence rachis (or the flowers solitary and 
pedicellate), the long narrow white corolla tube, 
and the fruit turning red or black. Collections from 
higher elevations have broader leaves and may 
represent introgression from another species; com- 
pare G. ovatifolia and G. rosea. 



Gonzalagunia rosea Standl., Ann. Missouri Bot. 
Gard. 25: 836. 1938. G. longithyrsa Fosberg, 
Sida 2: 387. 1966. Figure 20. 

Shrubs or small trees, 2-4(-6) m tall, branches erect 
or scandent, leafy branchlets 1-4 mm thick, terete, with 
stiff appressed-ascending pale yellowish or grayish hairs 
0.2-0.5 mm long; stipules 3-7 mm long, ca. 3-4 mm 
wide at base, broadly triangular with a short (1-4 mm) 
narrow tip, pubescent on the edge and midrib. Leaves 
with petioles 4-20 mm long, ca. 1 mm thick, densely 
pubescent; leaf blades 7-18(-22) cm long, 2-6(-8) cm 
broad, narrowly ovate-elliptic to lanceolate, lanceolate- 
oblong or narrowly elliptic-oblong, apex gradually or 
abruptly acuminate (acute) with tip 5-1 5 mm long, base 
acute to obtuse, drying thin-chartaceous to chartaceous, 
dark grayish brown above, puberulent on the upper sur- 
face (denser on the major veins) with short (0.2-0.3 mm) 
straight hairs, more densely puberulent beneath with hairs 
to 0.5 mm beneath, 2 veins 5-9(-l l)/side, strongly as- 
cending (and not loop-connected near the margin). In- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



149 



florescences solitary, terminal (axillary by further growth 
of lateral branches), 12-35 cm long, flowering part ca. 
2.5 cm broad, peduncles 5-35 mm long, 1-2.5 mm thick, 
densely short hirsute, flowers in groups of l-3(-5) and 
borne on secondary peduncles 1-5 mm long, alternate 
or opposite on the rachis and 2-10 mm distant, bracts 
1-3 mm long and linear, pedicels l-3(-4) mm long. 
Flowers with hypanthium 0.5-1.5 mm long, ca. 1 mm 
diam., densely pubescent, calyx tube ca. 0.5 mm long, 
calyx lobes 4 or 5, 0.5 mm high, triangular; corolla red 
in early bud, becoming rose red or pink, tube (6-)8-10 
mm long, 0.7-1.6 mm diam., slender and widening be- 
low the lobes, sparsely to densely puberulent, lobes 2-3 
mm long, ca. 2 mm wide at the base, glabrous distally 
on the inner surface and villous near the mouth; stamens 
4, anthers 2-2.5 mm long, anthers partly exserted; style 
9-1 1 mm long, stigma 0.3-0.7 mm long. Fruits 3-5 mm 
long, 3-6 mm broad, white, usually 4-lobed, glabrous or 
sparsely puberulent. 

Common shrubby plants of lower montane ev- 
ergreen forest formations, from (900-)1 100-2200 
m elevation. Probably flowering and fruiting 
throughout the year, but flowering primarily in 
February-September and fruiting primarily in 
June-September. The species ranges from the Cor- 
dillera de Tilaran in Costa Rica to eastern Panama. 

Gonzalagunia rosea is recognized by the thin- 
petiolate narrowly ovate to lanceolate leaves, the 
long inflorescences with pedunculate flower clus- 
ters, the pink corollas, and the spongy white fruit. 
This is a common weedy shrub of open habitats. 
Some specimens of this species may be difficult to 
distinguish from G. panamensis and G. ovatifolia, 
and it is possible that hybridization occurs. 



Gonzalagunia rudis (Standl.) Stand 1.. J. Wash. 
Acad. Sci. 17: 170. 1927. Duggenia rudis Standl., 
Contr. U.S. Natl. Herb. 18: 125. 1916. 

Shrubs, 1.5-4 m tall, leafy stems 0.7-3 mm thick, with 
minute (0.2 mm) thin appressed-ascending whitish hairs, 
terete and glabrescent; stipules 4-7 mm long, with a short 
(1-2 mm) broad base and long narrow awn. Leaves with 
petioles 1-3 mm long, ca. 1 mm thick, hertellous; leaf 
blades 2.5-10 cm long, 1-4 cm broad, narrowly ovate 
to ovate-oblong or lanceolate, apex tapering gradually 
and acute to long-acuminate, base acute to obtuse, drying 
thin-chartaceous, with scattered thin appressed hairs ca. 
0.5 mm long on both surfaces, with denser pubescence 
on the veins beneath, 2 veins 4-7/side. Inflorescences 
terminal or axillary, solitary, 4-14 cm long, spicate, pe- 
duncles 1-3 cm long, 0.3-0.7 mm thick, densely pubes- 
cent with ascending hairs, flowers usually solitary or in 
groups of 2-3 ca. 5 mm diam., subtended by linear bracts 
2-3(-5) mm long, pedicels 1-2 mm long. Flowers 
5-parted, hypanthium ca. 1 mm long, urceolate, densely 
puberulent, calyx lobes 1-2 mm long, linear, corolla white, 
tube ca. 6 mm long and 0.7 mm diam., lobes ca. 5 mm 
long and 1 mm broad, glabrous externally, puberulent 



within; stamens included. Fruits 2-4 mm long, 2.5-5 
mm broad, white, depressed globose, 4-lobed, with thin 
erect hairs 0.5 mm long. 

Plants of evergreen forest formation on the Pa- 
cific slope of southern Costa Rica, from near sea 
level to 500 m elevation (to 1 000 m in Panama). 
Flowering primarily in the wet season (June-Sep- 
tember). This species ranges from about 84W in 
Costa Rica to eastern Panama. 

Gonzalagunia rudis is recognized by its smaller 
thin lanceolate leaves, the slender spike-like inflo- 
rescences with mostly solitary subsessile flowers, 
and the unusual calyx and corolla with long narrow 
lobes. Costa Rican collections differ somewhat in 
their narrower more lanceolate leaves, but collec- 
tions with such leaves are also found in Panama. 



Gonzalagunia stenostachya (Standl.) W. Burger, 
comb. nov. Rondeletia stenostachya Standl., 
Publ. Field Mus. Nat. Hist., Bot. Ser. 18: 1372. 
1938. Arachnothryx stenostachya (Standl.) Bor- 
hidi, Acta Bot. Hung. 33: 303. 1987. 

Shrubs to 3 m tall, leafy stems 2.3-6 mm thick, with 
short dense yellowish or reddish brown hairs ca. 0.5 mm 
long; stipules 4-15 mm long, triangular, acute, with yel- 
lowish hairs along the midrib. Leaves with petioles 2-4 
mm long, ca. 1.5 mm thick; leaf blades 7-16 cm long, 
2-6 cm broad, elliptic, elliptic-oblong to lanceolate or 
oblanceolate, apex acute or short-acuminate, base grad- 
ually narrowed and cuneate base (rounded in Dryer 1274), 
drying stiffly chartaceous and dark reddish brown above 
(grayish green beneath), with thin hairs 0.4 mm long or 
glabrescent above, densely puberulent on the midvein 
beneath, 2 veins 7-12/side, ascending. Inflorescences 
(7-) 15-25 cm long, 1.3-2.5 cm broad, spiciform thyrsoid 
panicles, peduncles 1.3-6 cm long, 1.2-2 mm thick, stri- 
gose, lateral cymules sessile and separate along the rachis, 
of (l-)2-5 flowers, bracts ca. 3 mm long, linear, pedicels 
0-4 mm long. Flowers 4-parted, hypanthium ca. 1.5 mm 
long and 1.5 mm diam., densely sericeous, calyx lobes 
1-2 mm long, narrowly triangular; corolla white, densely 
sericeous with hairs 0.5-0.9 mm long, tube 4-9 mm long, 
slender, lobes 3-4.5 mm long, oblong. Fruits ca. 3 mm 
long and 4 mm broad (?immature) with sericeous hairs 
ca. 0.5 mm long. 

Plants of wet evergreen forest formations of the 
Caribbean slope, 300-1200 m elevation. Flower- 
ing in February-March, July, and October; fruit- 
ing in October. This endemic species is known 
from near Monteverde and the La Selva-Braulio 
Carillo area. 

Gonzalagunia stenostachya is recognized by its 
subsessile leaves with many ascending secondary 
veins, the long slender spikes with sessile and well- 



150 



FIELDIANA: BOTANY 



separated cymules, and the sericeous flowers. The 
Panamanian G. kallunkii Dwyer (?= G. veraguen- 
sis Dwyer) has rather similar inflorescences but 
the petioles of that species are well developed and 
the leaves have fewer secondary veins. 



Guettarda Linnaeus 

Trees or shrubs, branchlets terete, puberulent or gla- 
brous, occasionally with spines; stipules interpetiolar, 
simple or slightly connate above the petioles (intrapetio- 
lar), often slightly overlapping above the node, triangular 
and acuminate to rounded distally, persisting or decid- 
uous. Leaves opposite (rarely 3 or 4/node), petioles short 
to long; leaf blades entire, with pinnate venation, the 
distal secondaries often strongly ascending and the 3 
veins often subparallel, domatia sometimes present. In- 
florescences solitary and axillary (1 or 2/node), usually 
pedunculate and with cymose branching, branches of the 
inflorescences often dichotomous (bifurcate) and with 
sessile flowers along 1 side, bracts and bracteoles present 
or reduced. Flowers bisexual (rarely unisexual), radially 
symmetrical, 4-9-parted (usually 5- or 6-parted), hy- 
panthium ovoid to globose or tubular, calyx tube cu- 
pulate to campanulate or short-tubular, calyx entire or 
with 2-9 poorly developed lobes/teeth; corolla funnel- 
form or salverform, white, yellowish, purple or bluish, 



corolla lobes 4-9, imbricate or subvalvate, the margins 
often undulate; stamens 4-9, anthers narrow, sessile or 
subsessile, dorsifixed, included; ovary 2-9-locular, with 
1 pendulous ovule from apex of each elongate-tubular 
locule, stigma capitate or lobed. Fruits drupaceous, glo- 
bose to elongate, rounded or angulate in cross-section, 
the exocarp fleshy but thin, endocarp woody to stony, 
2-9-locular. 

A genus of 60-80 species in the New World 
tropics, with a few species in the southwest Pacific 
and a species widespread on tropical coasts (G. 
speciosa L.). Some species of Guettarda have in- 
florescences with two equal cincinnoid branches; 
these are scorpioid cymes (cincinni) in which the 
sessile flowers are all in a close line along a single 
side of the rachis. The leaves of Guettarda are 
generally thin, often clustered at the ends of 
branchlets, and with the distal secondaries strong- 
ly ascending. Domatia are often present, and the 
3 veins are usually subparallel. The inflorescences 
are always axillary, and the flowers are all salver- 
form in Costa Rican species. This treatment ben- 
efited from the annotations made by Alfredo Gri- 
jalva in 1982. 



Key to the Species of Guettarda 

la. Inflorescences subsessile; plants of Cocos Island and the evergreen Pacific lowlands . . .G. conferta 

Ib. Inflorescences short-to long-pedunculate; plants of mainland Central America 2 

2a. Bracts subtending the flowers 3-6 mm long, thin-brownish, narrowly ovate-oblong; Pacific slope 

and lowlands 3 

2b. Bracts subtending the flower absent or less than 3 mm long and caducous 4 

3a. Inflorescences with peduncles 2.5-6 cm long; leaf blades usually rounded at the base; Gulf 
of Nicoya G. brenesii 

3b. Inflorescences with peduncles 1-3 cm long; leaf blades acute at the base; western Costa Rica 

G. foliacea 

4a. Inflorescences with short (to 1 cm) branches, the branches not cincinnoid (scorpioid-cymose) in 

appearance 5 

4b. Inflorescences with conspicuous lateral cincinnoid (scorpioid) branches more than 1 cm long (with 

the flowers all along 1 side 6 

5a. Corolla tubes 8-12 mm long, peduncles less than 3 cm long; petioles 4-30(-40) mm long; 
fruit ca. 12 mm long, subglobose; Pacific and Caribbean lowlands G. macrosperma 

5b. Corolla tubes 13-18 mm long, peduncles more than 3 cm long; petioles 20-70 mm long; 

fruit ca. 20 mm long, oblong, truncated distally; Golfo Dulce and Panama 

G. sanblasenesis 

6a. Trees to 30 m tall; leaves and twigs glabrous; flowers becoming 2-4 mm distant on the rachis 

[corolla tubes 20-30 mm long] G. turrialbana 

6b. Trees to 10(-25) m tall; leaves and stems densely to sparsely puberulent; flowers 0-2 mm distant 

on the rachis 7 

7a. Stipules glabrous, often broadly overlapping; midvein sparsely appressed strigose along the sides 

beneath [corolla tube 16-20 mm long; 1500-2500 m elevation] G. poasana 

7b. Stipules pubescent (at least along the midrib); midvein densely pubescent over the entire surface 

on the lower side of the leaf blade . 8 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



151 



8a. Leaf blades narrowly oblong and coriaceous, densely tomentulose beneath; corolla tube 4-7 mm 

long [1200-1900 m elevation] G. tornefortiopsis 

8b. Leaf blades elliptic to broadly ovate, drying chartaceous, lacking a densely matted tomentum 

beneath; corolla tubes 6-18 mm long 9 

9a. Leaf blades rounded and often truncate at the base, to 30 cm long; peduncles to 10 cm long [corolla 

tubes 15-18 mm long]; northern Caribbean lowlands G. combsii 

9b. Leaf blades usually acute to obtuse at the base, to 22 cm long; peduncles to 4 cm long; evergreen 

formations 300-1700 m elevation 10 

lOa. Leaf blades with 6-10 pairs of major secondary veins, 3 veins clearly differentiated from the 

smaller 4 veins; corolla tubes 13-18 mm long; Central Highlands and evergreen Pacific lowlands 

G. crispiflora 

lOb. Leaf blades with 3-5 pairs of major secondary veins, 3 and 4 veins little differentiated and parallel 

with each other; corolla tubes 6-9 mm long; Caribbean slope Chomelia venulosa 



Guettarda brenesii Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 18: 1303. 1938. Figure 33. 

Trees to 10 m tall, leafy branchlets 2-4 mm thick, at 
first with straight ascending hairs 0.5-1 mm long, terete, 
soon glabrescent and becoming very dark with numerous 
narrow lenticels ca. 0.5 mm long; stipules 7-12 mm long, 
narrowly ovate-triangular, acute, sericeous along the 
midrib and base, deciduous. Leaves clustered at the end 
of branchlets, petioles 5-12 mm long, ca. 1 mm thick, 
with thin ascending hairs 0.5-1.3 mm long; leaf blades 
(4-)6.5-2 1 cm long, (3-)4.5-l 2 cm broad, broadly ovate 
to broadly elliptic, apex obtuse or subrotund, base obtuse 
(and rounded at the petiole) to broadly rounded and 
subcordate, drying chartaceous and dark above, sparsely 
hispidulous above with hairs ca. 0.5 mm long, more 
densely pubescent beneath with thin whitish hairs to 1 
mm long, 2 veins 6-8/side, with denser tufts of hairs 
(domatia) in the vein axils, 3 veins subparallel basally 
and joining with the opposing tertiary veins at an angle. 
Inflorescences 3-10 cm long, equally wide, axillary, pe- 
duncle 2-6.5 cm long, ca. 0.8 mm thick, pubescent, usu- 
ally bifurcate with 2 main branches and further cymose 
branching, bracts and bracteoles 4-8 mm long, lanceo- 
late, pubescent along the midrib, distal axes to 4 mm 
long often bearing single flowers and resembling pedicels 
(but with bracteoles at their apex beneath the flowers). 
Flowers with hypanthium ca. 1 mm long and 1 mm 
broad, short-tubular and sericeous, calyx tube ca. 1 mm 
long, entire and more sparsely pubescent distally; corolla 
white, tube (9-) 14-1 8 mm long, 0.5-0.8 mm diam., with 
minute appressed-ascending whitish hairs externally, 
lobes 5, ca. 3 mm long and 1.7 mm broad, margin broad- 
ly rounded and subentire. Fruits unknown. 

Trees of lowland deciduous forest formations 
near the Bay of Nicoya. Immature inflorescences 
were collected in June (Brenes 15694 the type), 
and inflorescences with falling corollas were col- 
lected in July. Fruiting in July-August and Oc- 
tober-November. The species is known only from 
along the Pacific coast of central Costa Rica. 

Guettarda brenesii is recognized by the bracteate 
inflorescences, the long narrow corollas, the broad- 
ly ovate leaves, and the rocky, seasonally very dry 



seaside habitat. This species is probably related to 
G. foliacea, which shares characters of the inflo- 
rescence. The type appears to have immature leaves 
and inflorescences; hence, Standley's description 
represents minimal measurements. A highly re- 
stricted range and short flowering season may ex- 
plain the paucity of collections. 

Guettarda combsii Urban, Symb. Ant. 6: 48. 1 909. 

Small to large trees (shrubs), 5-30 m tall, trunks to 60 
cm dbh, leafy branchlets 2-6 mm thick, with soft whitish 
erect or ascending hairs 0.5-1 mm long but soon gla- 
brescent, terete; stipules 6-14 mm long, ovate to lan- 
ceolate, acuminate, pubescent along the midrib. Leaves 
often clustered at the ends of branchlets, petioles 2-9 cm 
long, 1.3-2 mm thick, minutely pubescent; leaf blades 
7-20(-28) cm long, 5-13 (-20) cm broad, very variable 
in shape, from broadly ovate to ovate-oblong or subor- 
bicular, apex abruptly narrowed and obtuse or short acu- 
minate, base broadly obtuse to rounded and truncate to 
subcordate, drying thin-chartaceous, subglabrous or 
sparsely pubescent above with thin hairs ca. 1 mm long, 
more densely pubescent beneath with thin whitish hairs 
0.5-1 mm long, 2 veins 8-1 I/side, 3 veins parallel and 
prominent. Inflorescences 12-18 cm long, with long (5- 
1 5 cm) peduncles and 2 or 4 distal cincinnoid branches 
2-5 cm long, minutely (0.1-0.4 mm) pubescent, bracts 
2-4 mm long and ca. 1 mm broad, flowers sessile and 
closely spaced. Flowers with hypanthium ca. 1 mm long 
and 1.3 mm diam., with a dense greenish white or pale 
grayish white tomentum, calyx tube 1.5-2.5 mm long, 
entire or 2-lobed; corolla greenish white to cream white, 
tube 15-18 mm long, 1-1.5 mm diam., with dense re- 
trorse sericeous hairs, corolla lobes 5-7, ca. 3-4 mm long; 
ovary 4- or 5-locular. Fruits 6-8 mm long, subglobose, 
covered with a dense minute (0.05 mm) tomentum, gray- 
ish green, the surface becoming wrinkled. 

Trees of evergreen Caribbean rain forest for- 
mations, from near sea level to 900 m elevation. 
Flowering in May; fruiting in September in Belize. 
The species ranges from Belize to southeastern 
Nicaragua. 



152 



FIELDIANA: BOTANY 



Guettarda combsii is recognized by the broad 
pubescent leaf blades usually rounded and trun- 
cate to subcordate at the base, the long-pendun- 
culate inflorescences with four short cincinnoid 
branches, and the subglobose fruit. The fact that 
these plants become very tall trees may account 
for the paucity of collections in southern Central 
America. A sterile collection from a tall tree at 
Minefields. Nicaragua (Proctor et al. 27130 F), sug- 
gests that this species is also likely to occur in 
northern Costa Rica. 



Guettarda conferta Benth., Hot. voy. Sulph. 106. 
1845. 

Trees to 10 m tall, with dense, ferruginous hairs on 
the branchlets, petioles, peduncles, and nerves of the 
leaves; stipules broadly obovate, 12 mm long, hirsute 
externally at base, otherwise glabrous, about equaling 
the petioles, deciduous. Leaves with blades 10-20 cm 
long, 6-9 cm broad, ovate, apex acuminate, base acute, 
hirsute on both sides. Inflorescences 2.5-3 cm long, cy- 
mose, subsessile, the branches recurved and 2.5 cm long 
or less. Flowers ca. 8 mm long, calyx tube 1-2 mm long, 
shallowly 3- or 4-dentate; corolla white, sericeous-hir- 
tellous, tube ca. 10 mm long, corolla lobes 4, ca. 4 mm 
long, obtuse crispate. Fruits ovoid-tetragonous, 4 mm 
long, 3 mm diam.. hirsute, 4-locular. 

Plants of Cocos Island and the Pacific slope of 
southern Costa Rica and the Osa Peninsula, 0- 
300 m elevation. Flowering in August and Decem- 
ber; fruiting in January. 

Guettarda conferta is distinguished by its short 
sessile inflorescences. In general aspect this species 
resembles G. crispiflora. 



Guettarda crispiflora Vahl, Ecolog. Amer. 36: pi. 
6. 1797. G. chiriquiensisSlandl., Ann. Missouri 
Hot. Gard. 25: 838. 1938. Figure 32. 

Small to medium-sized trees 4-20 m tall, leafy branch- 
lets 1.5-6 mm thick, at first quadrangular but soon be- 
coming terete, with short (0.3 mm) thin appressed-as- 
cending hairs but glabrescent; stipules 8-18(-22) mm 
long, to 1 cm broad, ovate-elliptic to slightly obovate, 
apex acute to acuminate, with thin ascending sericeous 
hairs along the midrib and at the base. Leaves clustered 
distally, petioles 2-7 cm long, 1.3-2 mm thick, minutely 
puberulent and with longer hairs along the adaxial side; 
leaf blades (6-)8-22 cm long, (3-)5-l 1 cm broad, ovate- 
elliptic or ovate-oblong to ovate-lanceolate or broadly 
ovate, apex tapering gradually and acuminate or acute, 
base obtuse to slightly rounded or attenuate and acute, 
drying stiffly chartaceous, glabrous or minutely (0.2 mm) 
puberulent above, pubescent beneath with larger (0.4-1 
mm) hairs on the major veins and smaller (0.3 mm) thin 



whitish hairs on the 3 veins, tufts of hairs (domatia) 
sometimes present in the vein axils beneath, 2 veins 6- 
10/side, distal 2 veins strongly ascending, 3 veins often 
parallel. Inflorescences 2-6(-8) cm long, equally wide, 
with a short (4-25 mm) peduncle and 2 diverging cin- 
cinnoid branches 1 5-30(-60) mm long and enlarging in 
fruit, the rachis minutely grayish white tomentulose, the 
flowers sessile and closely (0-4 mm) spaced. Flowers 
sweet-scented, hypanthium 1.5-2.5 mm long, 1.5-2 mm 
diam., densely grayish white tomentulous, calyx tube ca. 
0.5 mm long, calyx lobes 4, ca. 0.5 mm high; corolla 
white or pinkish, tube 13-18 mm long, 1-1 .5 mm diam.. 
narrowly tubular, densely short-sericeous with retrorse 
or spreading hairs, lobes 5-6 mm long, the lobes with 
smaller undulate marginal lobes. Fruits ca. 8 mm long 
and 6 mm diam., oblong and with 4 prominent longi- 
tudinal ribs, becoming purple and with a white pulp. 

Trees of the Caribbean slope cloud forest for- 
mations and southern Pacific wet forest forma- 
tions, from 300 to 1700 m elevation. Flowering 
in January, April, and June-September, with Jan- 
uary and August collections being most frequent; 
fruiting in March and JuneJanuary. In our area 
the species is known from the Caribbean slope of 
the Central Highlands (from near Monteverde 
eastward to Tapanti, Cartago, and San Joaquin de 
Dota, San Jose), on the Osa Peninsula, near San 
Vito, and in the Chiriqui Highlands. The species 
also occurs in the Lesser Antilles and Trinidad. 

Guettarda crispiflora is recognized by the short- 
pedunculate inflorescences with bifurcate cincin- 
noid branching, the white corollas with crisped 
and undulate corolla lobes, the four-angled fruit, 
longer petioles, and the subparallel (almost lineo- 
late) minor venation. Guettarda poasana is closely 
related to G. crispiflora, and material of the two 
species should be compared when making iden- 
tifications. Guettarda chiriquiensis was distin- 
guished by its more densely pubescent vegetative 
parts, but there are a few intermediate collections 
in Costa Rica. Nevertheless, the distinctive pop- 
ulations of the Chiriqui Highlands and adjacent 
Costa Rica may be worthy of subspecific rank. 



Guettarda foliacea Standley, Contr. U.S. Natl. 
Herb. 18: 139. 1916. Figure 33. 

Shrubs or small trees, 3-6(-10) m tall, often branching 
from the base and with clambering branches, leafy stems 
0.9-4 mm thick, at first strigose with thin ascending hairs 
0.3-1 mm long, glabrescent, becoming brown and terete, 
spines often present; stipules 3-5(-12) mm long, trian- 
gular to lanceolate, strigulose. caducous. Leaves oppo- 
site, petioles 3-25 mm long, 0.4-1.2 mm thick, ap- 
pressed strigose to sericeous; leaf blades 3-16 cm long, 
2-7 cm broad, elliptic to ovate or obovate, apex acute 
to short-acuminate, base cuneate to slightly rounded or 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



153 



subtruncate, drying thin-chartaceous and greenish, with 
thin whitish hairs 0.2-0.9 mm long on both surfaces 
(dense only on the major veins), 2 veins 4-8/side, 4 
veins often parallel, domatia of dense hairs present in 
the vein axils beneath. Inflorescences 2-7 cm long, to 5 
cm broad, peduncles 5-30 mm long, 0.5-1.1 mm thick, 
with thin ascending hairs, usually with a single pair of 
dichotomous distal branches (each with 3-7 flowers), 
subtended by lanceolate ciliate bracts 3-8 mm long, flow- 
ers sessile and crowded in cymes. Flowers densely mi- 
nutely sericeous externally, hypanthium ca. 1 mm long, 
calyx tube 1-1.5 mm long, ca. 1.3 mm diam., entire; 
corolla white, tube 12-20 mm long, lobes 4(-5), 2-5 mm 
long, rounded distally. Fruits 1-3 cm diam., globose, 
with a dense minutely velutinous surface, becoming red. 

Plants of evergreen or partly deciduous forests, 
0-300(-1000) m elevation. In central Panama 
flowering occurs primarily in late June-early July 
(Croat, 1978) and fruiting in September-Novem- 
ber. This species ranges from the Cordillera de 
Tilaran to Colombia. 

Guettarda foliacea is recognized by its small in- 
florescences, lack of calyx lobes, thin variable leaves 
on slender petioles, and minor venation, often with 
a small group of parallel veins (sublineolate). The 
spines are rarely seen on herbarium collections. 
The westernmost collection (8453'W, Haber et al. 
4775 CR, MO) came from the edge of cloud forest 
at 1 000 m elevation with immature flowers in May. 



tomentulose, calyx tube 0.5-1 mm long, entire; corolla 
white or yellowish, tube 8-13 mm long, 0.5-1.3 mm 
diam., narrowly tubular, lobes 3-4 mm long, rounded 
distally. Fruits 10-18 mm long, globose to oblong, yel- 
lowish brown or grayish with a dense covering of minute 
(0.1-0.2 mm) velvet-like or matted hairs. 

Trees and shrubs of both evergreen and decid- 
uous forest formations, from near sea level to 1 000 
m elevation on the Pacific slope and from near sea 
level to ca. 500 m on the Caribbean slope. Flow- 
ering in March-November, with the majority col- 
lected in May; fruiting in July-January. The spe- 
cies ranges from southern Mexico to Panama. 

Guettarda macrosperma is recognized by its 
small cymose inflorescences, rounded fleshy fruit 
usually over 1 cm diam., and often smaller leaves 
with the subparallel 3 veins usually meeting at 
angles between the 2 veins (> -shaped). Trees in 
the Caribbean lowlands may reach 30 m in height; 
the same trunk and bark characteristics are found 
in trees of both seasonally dry forest and evergreen 
rain forests (N. Zamora, pers. comm.). Some spec- 
imens may resemble Chomelia panamensis. It ap- 
pears that Guettarda divaricata (Roem. & Schult.) 
Standl. of Mexico is closely related, and the two 
may be part of a more broadly defined taxon. Ma- 
lacahuite is a common name. 



Guettarda macrosperma J. D. Smith, Bot. Gaz. 1 8: 
204. 1893. Figure 33. 

Shrubs or more often trees, 4-12(-30) m tall, trunks 
often fluted or with deep depressions, with dark bark 
exfoliating in patches, leafy branchlets 1.2-4 mm thick, 
sparsely puberulent with thin ascending hairs 0.5-1.5 
mm long, soon glabrescent and becoming dark brown 
or blackish with short (0.3-1.2 mm) grayish lenticels; 
stipules 3-8(-12) mm long, triangular-lanceolate, dense- 
ly sericeous on the back with longer hairs, caducous. 
Leaves often crowded at the ends of branchlets, petioles 
(4-)10-45 mm long, 0.8-1.7 mm thick, with straight 
ascending hairs; leaf blades (4-)6-18 cm long, (2-)3-l 1 
cm broad, broadly elliptic, broadly ovate-elliptic, obo- 
vate-elliptic, or elliptic-oblong, apex abruptly narrowed 
and short-acuminate to acute or obtuse, base obtuse to 
rounded and subtruncate, drying chartaceous to stiffly 
chartaceous, sparsely pubescent with thin short (0.3-0.7 
mm) hairs and glabrescent, with thin short (ca. 0.3 mm) 
ascending hairs along the 3 veins beneath and with lon- 
ger and denser hairs along the major veins, 2 veins 4- 
8/side, 3 veins rarely subparallel, often with tufts of hairs 
(domatia) in the vein axils. Inflorescences 2-6 cm long, 
primary peduncles 4-38 mm long, usually with 2 short 
distal branches, densely ascending sericeous, bracts 2-3 
mm long, flowers sessile and closely crowded (not clearly 
cincinnoid in arrangement). Flowers with hypanthium 
1-1.5 mm long and ca. 1 mm diam., densely whitish 



Guettarda poasana Standl., J. Wash. Acad. Sci. 1 8: 
182. 1928. Figure 32. 

Small trees, 3-10(-15) m tall, trunk to 40 cm thick, 
bark shredding off in oblong patches, leafy branchlets 2- 
5 mm diam., somewhat flattened in early stages and 
glabrous, drying dark but becoming grayish in age, len- 
ticels difficult to see; stipules 12-20 mm long, 8-10 mm 
broad, ovate and long-acuminate at apex, the stipules 
overlapping on the sides, glabrous. Leaves clustered near 
the ends of branches, petioles (2-)3-7(-10) cm long, 0.8- 
1.6 mm thick, glabrous and drying dark; leaf blades 7- 
14(-16) cm long, 2.5-6(-9) cm broad, broadly elliptic- 
ovate to broadly elliptic or elliptic, apex gradually ta- 
pering and short-acuminate, base obtuse to acute and 
slightly decurrent on petiole, drying stiffly chartaceous, 
glabrous and lustrous above, minor venation with thin 
whitish ascending hairs ca. 0.3 mm long beneath, with 
longer (0.5-1 mm) straight hairs on the mid vein and 
secondaries beneath, 2 veins 5-8/side, the distal strongly 
ascending, 3 veins often subparallel but not prominent 
beneath, small tufts of hairs (domatia) often present in 
the vein axils beneath. Inflorescences axillary, 3-6 cm 
long and equally wide, peduncles 1-3 cm long, glabrous 
or sparsely puberulent, with 2 cincinnoid branches 2- 
3.5 cm long, the flowers 5-11 on each branch and sessile, 
ca. 1-4 mm distant. Flowers sweet-scented, hypanthium 
1.5-2 mm long, 1.3 mm diam., glabrous or sparsely 
puberulent, calyx tube 0.5-1 mm long, lobes ca. 0.3 mm 
long; corolla white, reddish, or lavender, tube 1 6-20 mm 



154 



FIELDIANA: BOTANY 



long 1-1.4 mm diam., densely tomentulose externally, 
lobes 4-5, 4-6 mm long, usually white with fringed- 
undulate smaller lobes; tips of the anthers exserted 1-2 
mm from the mouth of the tube. Fruits becoming 8 mm 
long and 6 mm diam., oblong and with 4 prominent 
longitudinal ridges, reddish purple. 

Trees of evergreen cloud forest formations, 
1300-2200(-2700?) m elevation (down to 1 100 m 
on the northern vocanoes). Flowering in March 
and May-November (mostly in June); fruiting 
probably throughout the year. This species is en- 
demic to Costa Rica and ranges from the Cordille- 
ra de Guanacaste in the west to the eastern slopes 
of Volcan Barva. 

Guettarda poasana is recognized by its glabrous 
stipules and stems, long-petiolate leaves, narrowly 
tubular flowers with whitish fringed lobes, four- 
angled fruit, and restricted cloud forest range. The 
petioles dry dark and are sometimes pink in life. 
This species is closely related to G. crispiflora and 
might be considered a subspecific element of that 
species. However, though their ranges overlap 
slightly, G. crispiflora and G. poasana do not grow 
in the same locality; G. poasana is generally found 
at higher altitudes. 



Guettarda sanblasensis Dwyer, Ann. Missouri Bot. 
Gard. 67: 204. 1980. Figure 33. 

Trees, 12-20 m tall, leafy branchlets 1.5-4 mm thick, 
with short (0.3 mm) appressed-ascending hairs, quickly 
glabrescent, terete; stipules 3-6 mm long, densely seri- 
ceous with pale yellowish ascending hairs 0.5-1 mm long, 
early caducous. Leaves 1-5 cm distant at the ends of 
branchlets, petioles 2-7 cm long, 0.7-1 .3 mm thick, with 
slender appressed-ascending hairs; leaf blades 1 1-22 cm 
long, (4-)6-l 2 cm broad, broadly elliptic to broadly ovate- 
elliptic, apex abruptly narrowed and short-acuminate (or 
acute), base obtuse to slightly rounded, drying charta- 
ceous and dark brown above, lustrous above and with 
scattered thin appressed hairs 0.2-0.3 mm long, the hairs 
more numerous and longer (0.3-0.5 mm) beneath, 2 
veins 5-9/side, the distal arcuate-ascending, 3 veins 
prominent above and below and paler in color beneath, 
subparallel or > -shaped between the secondaries, with 
minute tufts of hair (domatia) in the vein axils beneath. 
Inflorescences 6-10 cm long, primary peduncles to 6 cm 
long, ca. 1.2 mm thick and minutely appressed-puber- 
ulent, bifid but the 2 branches with additional dichot- 
omous branches (not scorpioid/cincinnoid), pedicels 0- 
2 mm long, bracts minute (0.5 mm) and caducous. Flow- 
ers with hypanthium ca. 1.5 mm long and 1.2 mm diam., 
with longitudinal ribs, calyx tube 2-3 mm long and ca. 
2 mm diam., minutely velutinous, subentire distally; 
corolla white or pink, tube 13-18 mm long, 1-1.3 mm 
diam., minutely (0.2 mm) ascending-sericeous, lobes 5- 
6, 4-5 mm long and 1-1.5 mm broad, oblong and entire, 
stigma 0.5 mm long, subglobose. Fruits 2-2.2 cm long 



and 8-12 mm thick, oblong-obovoid, abruptly rounded 
(truncated) at apex, persisting calyx 1-3 mm long, 1.5- 
2 mm diam., surface minutely velutinous and yellowish 
or grayish brown. 

Trees of evergreen forest formations of the Pa- 
cific lowlands, from near sea level to 600 m ele- 
vation and usually on limestone. Flowering in July- 
September; fruiting in September-November. This 
species is known only from a few collections in 
the Golfo Dulce region of Costa Rica; it ranges to 
eastern Panama. 

Guettarda sanblasensis is recognized by the 
broadly elliptic leaves on long slender petioles, the 
long-pedunculate inflorescences with dichoto- 
mous or cymose distal branching, long-tubular co- 
rollas with five or six entire oblong perianth lobes, 
and oblong fruit with truncated apex and usually 
persisting calyx tube. 



Guettarda tournefortiopsis Standl., Publ. Field 
Columb. Mus., Bot. Ser. 7; 293. 1931. Tour- 
nefortiopsis reticulata Rusby, Bull. New York 
Bot. Gard. 4: 369. 1907, non G. reticulata Gri- 
seb., 1863. Figure 32. 

Small trees, 4-10 m tall, leafy branchlets 1.5-6 mm 
thick, at first densely tomentose with soft wooly hairs to 
2 mm long, 4-angular, soon .glabrescent and dark with 
lenticels 0.3-0.7 long, becoming terete and pale grayish; 
stipules 1 2-20 mm long, narrowly ovate-triangular and 
acuminate, tomentose, usually early deciduous. Leaves 
with petioles 13-35 mm long, 1.5-2.2 mm thick, gla- 
brescent and drying dark; leaf blades 9-16 cm long, 2- 
5 cm broad, lanceolate to narrowly oblong-lanceolate or 
narrowly elliptic-oblong, apex acute to short acuminate, 
base obtuse to acute and usually with the margin revo- 
lute, drying subcoriaceous, dark brown above, glabrous 
and usually lustrous above, with the major and minor 
veins slightly impressed above and the surface slightly 
rugose, densely tomentulous between the veins beneath 
and whitish to pale brown in color, 2 veins 9-1 I/side, 
domatia present. Inflorescences 2-5 cm long and equally 
wide, becoming 8 cm long in fruit, peduncles 1 2-20 mm 
long (to 30 mm in fruit), 1 .5 mm thick and tomentulous, 
with 2 cincinnoid branches 2-6 cm long and each branch 
bearing up to 20 flowers in 2 rows along 1 side, flowers 
sessile and closely crowded, bracts absent. Flowers with 
hypanthium 1 .5-2 mm long, covered by a dense tomen- 
tum, calyx lobes ca. 1 mm broad, broadly obtuse and 
difficult to see; corolla reddish to coral pink, white with- 
in, tube 4-7 mm long, ca. 1.5 mm diam., densely pu- 
berulent with retrorse hairs externally, lobes 5, 1-2 mm 
long. Fruits 5-8 mm long, 4-6 mm diam., globose to 
rounded-oblong, 5-angled during development, sessile, 
becoming purple or black. 

Trees of wet montane cloud forest formations, 
from 800 to 1900 m elevation. Collections with 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



155 



flowers (or flower buds) and fruits have been made 
in January-March and August-October. Known 
only from the Cordillera de Tilaran and the west- 
ern parts of the Cordillera de Talamanca, prov- 
inces of Cartage and San Jose, in Costa Rica. The 
species is also known from the Chiriqui Highlands 
and was originally described from Bolivia. 

Guettarda tournefortiopsis is easily recognized 
because of its stiff narrow leaves densely tomen- 
tulose beneath (when young), the two-branched 
scorpioid inflorescences (resembling those found 
in the Boraginaceae), small tomentulose red flow- 
ers, and small sessile fruits. 



and long corolla tubes. The tall size of this dis- 
tinctive species may explain why it had not been 
collected before 1986. The collections from the 
Osa Peninsula differ in a number of ways from the 
type; more material is needed to assess the pattern 
of variation. 



Hamelia Jacquin 

REFERENCE T. S. Elias, A monograph of the 
genus Hamelia (Rubiaceae). Mem. New York Bot. 
Gard. 26: 81-144. 1976. 



Guettarda turrialbana Zamora & Poveda, Ann. 
Missouri Bot. Card. 75: 1 157. 1988. Figure 30. 

Trees to 30 m tall and with trunks 40 cm dbh, bark 
exfoliating, leafy branchlets 1.5-6 mm thick, glabrous 
and dark in early stages, terete, becoming pale grayish 
and with few broadly ellipsoid lenticels 0.8-1 .5 mm long; 
stipules ca. 1 2-25 mm long, triangular and acuminate, 
overlapping, glabrous, deciduous. Leaves clustered at the 
ends of branchlets, petioles 2-3.5 cm long, 1-2 mm thick, 
glabrous and drying dark; leaf blades 9-26 cm long, 4- 
1 2 cm broad, oblong to broadly elliptic-oblong or ovate- 
oblong, apex abruptly narrowed and short-acuminate, 
base abruptly narrowed or rounded and obtuse, drying 
stiffly chartaceous, dark brown above, glabrous and lus- 
trous above, subglabrous beneath with tufts of hairs 
(domatia) in the vein axils, 2 veins 8-10/side, 3 veins 
slightly raised above but not clearly subparallel. Inflo- 
rescences 5-11 cm long, primary peduncles 3-5 cm long, 
1-1.3 mm thick, glabrous and drying dark, cymose with 
2 primary branches, the branches 2-3 cm long and with 
3-5 flowers 2-5 mm distant on the rachis (or the branch- 
es very short and all the flowers close together in Hammel 
et al. 16848 CR, MO), pedicels short (1 mm) or absent, 
flower subtended by short (0.5 mm) broad bracts. Flow- 
ers glabrous externally, with the sweet odor of Coffea 
flowers, hypanthium ca. 2 mm long and 1.2 mm diam.. 
calyx cup ca. 2 mm long, ca. 3 mm diam. distally and 
entire; corolla white, tube 2040 mm long, 2-2.8 mm 
diam., puberulent within, lobes 5, ca. 5 mm long and 2 
mm broad, oblong and entire, distally rounded. Fruits 
24-28 mm long, 6-12 mm diam., oblong, glabrous. 

Trees of lowland rain forest formations; col- 
lected at 600 m elevation, below the CATIE site 
near Turrialba on the Caribbean slope, and on the 
Osa Peninsula at 200-300 m elevation. Flowering 
in May (Hammel et al. 16848 CR, F, MO) and June 
(H err era 4246 MO, Zamora et al. 1263 CR holo- 
type, F); fruiting in August and September. En- 
demic. 

Guettarda turrialbana is recognized by its tall 
stature, mostly glabrous parts, oblong leaves, few- 
flowered bifurcate inflorescences, entire calyx cup, 



Shrubs or small trees, branchlets usually slender, terete 
or 4-angled in cross-section, glabrous to densely puber- 
ulent; stipules interpetiolar, usually small, entire or with 
an awn (or trilobate), caducous or less often persistent. 
Leaves opposite and decussate or in whorls of 3-5/node, 
usually regularly spaced by well-developed internodes, 
often long-petiolate, usually thin in texture, glabrous or 
puberulent, pinnately veined, domatia present in some 
species, conspicuous raphides visible on the dried leaf 
surfaces in a few species. Inflorescences terminal or less 
often axillary, 3-many-flowered, often thyrse-like with 
cymose branches or with helicoid branches, pedunculate, 
the flowers often all along 1 side of the distal branches, 
sessile or short-pedicellate. Flowers bisexual and radially 
symmetrical, monomorphic, hypanthium urceolate to 
tubular, calyx tube minute or absent, calyx lobes 5, small, 
rounded to elongate, usually persisting; corolla narrowly 
tubular to funnelform or campanulate-urceolate (with a 
short narrow base), bright yellow to orange, orange-red, 
or deep red, with 5 longitudinal ribs alternating with the 
lobes, corolla lobes 5, erect to recurved, slightly imbri- 
cate in bud; stamens 5, filaments inserted at the base of 
the corolla tube, flattened, anthers long-linear, sagittate 
at the base and usually with the connective prolonged 
distally, included or partly exserted; ovary (4-)5-locular, 
with axile placentation and many ovules in each locule, 
style narrowly cylindrical, stigmas 1-5. Fruit a fleshy 
berry, oblong to ovoid or subglobose, with 5 longitudinal 
ribs, terminated by the circular calyx scar and a conical 
disc; seeds numerous, plano-convex or angled, foveolate. 

Hamelia ranges from southern Florida, U.S.A., 
through Mexico, Central America, and the West 
Indies into tropical South America. Elias recog- 
nized 16 species in his fine monograph. The nar- 
rowly tubular or funnelform yellow to orange or 
red flowers are usually all aligned on the upper- 
most sides of the inflorescence branches. In some, 
the inflorescences have longer cincinnus-like 
branches. Many of our species have three to four 
leaves at distal nodes and a number have long- 
petiolate leaves; most have small tufts of hairs 
(domatia) in the vein axils beneath. Individual 
plants may vary greatly within many species, and 
this makes identification difficult. In addition, there 



156 



FIELDIANA: BOTANY 



may be intermediates or hybrids between some of 
our species. 

The genus is divided into two subgenera. Sub- 
genus Hamelia has narrowly tubular flowers that 
do not enlarge distally at anthesis and are red, 
orange, or yellow in color. Subgenus Amphituba 
has the yellow corolla tube slightly to conspicu- 
ously expanded distally at anthesis. However, the 



flowers of subgenus Amphituba remain narrowly 
tubular until just before anthesis and may be dif- 
ficult to distinguish from those of subgenus Ha- 
melia on this account. Species of Hoffmannia may 
be similar, but they have consistently axillary in- 
florescences; compare Deppea, with papery cap- 
sules. 



Key to the Species of Hamelia 

la. Young stems and undersides of leaves conspicuously pubescent with hairs 0.2-0.5 mm long .. 2 
Ib. Young stems and undersides of leaves glabrous or with minute (0.1-0.2 mm) inconspicuous hairs 

4 

2a. Stipules 7-17 mm long; leaf blades with 8-18 major 2 veins on each side, with hairs to 1 mm 
long; corolla expanded distally at anthesis, corolla lobes 2-6 mm long [calyx lobes 0.5-2 mm 

long] H. xerocarpa 

2b. Stipules 2-5(-8) mm long; leaf blades with 4-9 major 2 veins on each side, with hairs to 0.5 

mm long; corolla narrowly tubular, corolla lobes 1-2 mm long 3 

3a. Calyx lobes 0-0.7 mm long; seeds 0.6-0.9 mm long; petioles 10-80 mm long; common wide- 
spread plants, 0-1 500 m elevation H. patens 

3b. Calyx lobes 24 mm long; seeds 1-1.2 mm long; petioles 3-20 mm long; uncommon plants of 

the Caribbean lowlands, 0-300 m elevation H. rovirosae 

4a. Corolla tube 8-1 3 mm long at anthesis; fruits 5-10 mm long 5 

4b. Corolla tube 1 3-35 mm long at anthesis; fruits 7-16 mm long 6 

5a. Inflorescences 3-9 cm long, corolla tube becoming expanded distally at anthesis; leaf blades 
with 5-9 major 2 veins on each side, 5-1 7(-23) cm long, usually drying greenish to pale grayish; 

fruits 4-7 mm long; Caribbean and evergreen Pacific lowlands H. axillaris 

5b. Inflorescences 8-18 cm long, corolla tube narrowly tubular at anthesis; leaf blades with 7-12 
major 2 veins on each side, 10-27(-32) cm long, usually drying reddish brown to pinkish gray; 

fruits 6-10 mm long; evergreen Pacific slope H. magnifolia 

6a. Leaf blades with 8-13 major 2 veins on each side; corolla tube distally widened (4-7 mm) at 

anthesis; corolla lobes 2-4 mm long 7 

6b. Leaf blades with 3-9 major 2 veins on each side; corolla tube narrowly (2-4 mm) tubular at anthesis 

(in H. patens, often broader in H. calycosa) 

7a. Leaf blades 3-6 cm broad; corolla tube 1 3-22 mm long; 0-800 m elevation ...//. xerocarpa 

7b. Leaf blades 4-12 cm broad; corolla tube 25-35 mm long; 700-1600 m elevation 

H. macrantha 

8a. Leaf blades with 6-9 pairs of 2 veins, petioles 10-80 mm long; sepal lobes 0-0.8 mm long, corolla 

lobes 1-2 mm long; common, 0-1 500 m elevation H. patens 

8b. Leaf blades with 3-7 pairs of 2 veins, petioles 8-20 mm long; sepal lobes 1-2.5 mm long, corolla 
lobes 3-6 mm long; not reported from Costa Rica H. calycosa 



Hamelia axillaris Sw., Prodr. 46. 1788. H. lutea 
Rohr ex Smith in Rees, Cyclop. 17.1811. Figure 

42. 

Shrubs, 1-5 m tall or small treelets to 5 m tall, leafy 
branchlets 1 .2-4 mm thick, glabrous, with 4 longitudinal 
ribs and 4-angular in cross-section but becoming terete; 
stipules 2-6(-8) mm long, ca. 1 mm broad, triangular to 
narrow with folded margins. Leaves opposite (rarely 
4/node), petioles l-4(-7) cm long, ca. 1 mm wide, gla- 



brous to sparsely and minutely papillate-puberulent; leaf 
blades 5-17(-23) cm long, 2-8 cm broad, narrowly el- 
liptic, elliptic-oblong or obovate to narrowly obovate- 
oblong, apex acuminate with tip ca. 1 cm long, base 
attenuate and decurrent on petiole, drying chartaceous 
or membranaceous, glabrous above, glabrous or mi- 
nutely (0. 1 mm) papillate-puberulent beneath, with tufts 
of hairs (domatia) in the vein axils, 2 veins 5-9/side 
and loop-connected near the margin, short (0. 1-0.3 mm) 
linear cystoliths visible on the lower surface when dry. 
Inflorescences axillary or terminal, 3-8 cm long, 3-8 cm 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



157 



broad, compound dichasia with lateral branches 1-4.5 
cm long, often scorpioid with 3-15 secund flowers on 
the uppermost side, peduncles 5-15 mm long, glabrous 
or minutely and sparsely puberulent, bracts 0.5-1 mm 
long, narrow, flowers sessile or with pedicels. Flowers 
with hypanthium 1.5-3 mm long, ca. 1.2 mm diam., 
glabrous, calyx lobes 0.5-1.5 mm long, 1 mm wide at 
the base, glabrous to puberulent; corollas yellow, nar- 
rowly urceolate to funnelform, tube 8-13 mm long, ca. 
1 mm diam. near the base and 3-5 mm wide distally, 
glabrous, lobes 1-2 mm long, broadly triangular; sta- 
mens with filaments 4-5 mm long, anthers 5-8 mm long 
with apical connective 0.5 mm long, style 8-10 mm long, 
stigmas 3-4 mm long. Fruits 4.5-7 mm long, 3-4 mm 
diam., ovoid-oblong to subglobose, disc 0.5 mm high 
and 0.7 mm broad; seeds ca. 1 mm long. 



Plants of the evergreen Caribbean slope and the 
Osa Peninsula, from near sea level to 600(-1000) 
m elevation. Flowering primarily in late June-Oc- 
tober; fruiting in February and June-December. 
The species is known from southern Mexico, Be- 
lize, Guatemala, the larger West Indian islands, 
and from Nicaragua southward to Venezuela and 
Bolivia. 

Hamelia axillaris is recognized by its short yel- 
low corollas funnelform at anthesis, short rounded 
fruit, general lack of pubescence, and usually com- 
pact inflorescences with flowers along one side of 
the distal branches. There may be intermediates 
between this species and H. magnifolia on the Osa 
Peninsula; compare the extreme upper-right figure 
in Figure 42 (based on Utley & Utley 1208 F). 



Hamelia calycosa J. D. Smith, Bot. Gaz. 12: 132. 
1887. 

Shrubs or small trees to 12 m tall, leafy branchlets 1- 
3 mm thick, glabrous or glabrescent; stipules l-2.5(-4) 
mm long, ca. 1 mm broad at the base, subulate or with 
a narrow awn, minutely puberulent. Leaves opposite or 
3-4/node on distal branches, petioles 6-20 mm long, 
0.4-0.8 mm broad, glabrate; leaf blades 3-1 1(-15) cm 
long, l-3.5(-5) cm broad, lanceolate to narrowly elliptic- 
oblong or elliptic-oblanceolate, apex tapering gradually 
and acuminate, tip ca. 7 mm long, base acute to cuneate 
and decurrent on petiole, drying membranaceous to 
chartaceous, glabrous above, glabrous or minutely (0. 1- 
0.2 mm) pubescent beneath or rarely with longer (0.7 
mm) thin hairs, with tufts of hairs (domatia) in the vein 
axils beneath, 2 veins 3-7/side. Inflorescences terminal 
or rarely axillary, 3-10 cm long, with 4-24 flowers, pe- 
duncles to 2 cm long and often with 2 dichotomous 
branches distally, usually minutely puberulent, bracts 1- 
2 mm long, subulate or triangular, flowers secund and 
with pedicels l-5(-8) mm long. Flowers with hypanthi- 
um 2-4 mm long, oblong, glabrous or sparsely pubescent, 
calyx lobes 1-3 mm long, ca. 0.5 mm wide, narrowly 
oblong, caducous; corolla funnelform, yellow or pale or- 
ange (striped with maroon), tube 1 5-24(-32) mm long, 



constricted (1.5 mm) near the base and 8-10 mm diam. 
distally, pubescent externally, lobes 3-6 mm long, ovate; 
stamens 5, filaments 46 mm long, anthers 15-18 mm 
long, 1 mm wide, distal rounded connective ca. 1 mm 
long, style to 14 mm long, stigmas 5 and connate, ca. 
1.2 mm long. Fruits 7-16 mm long, 4-8 mm thick, cy- 
lindrical, disc conical and 1-3 mm long; seeds 1-1.2 mm 
long. 

Trees and shrubs of Caribbean rain forest for- 
mations, from near sea level to 1500 m elevation. 
Flowering is in April-September in northern Cen- 
tral America. The species ranges from southern 
Mexico, Belize, Guatemala, Honduras, Panama, 
and Colombia to Peru. 

Hamelia calycosa is recognized by the larger 
sepal lobes, longer corollas expanded distally, ped- 
icellate fruit, and smaller leaves with short peti- 
oles. Although not yet collected in Costa Rica, this 
species is likely to be present. 



Hamelia macrantha Little, Carib. Forester 9: 274. 
1948. 

Shrubs or small trees to 8(-l 2) m tall and 1 3 cm dbh, 
leafy branchlets 2-6 mm thick, 4-angular in cross-sec- 
tion, glabrous or minutely puberulent; stipules 3-7 mm 
long, 1-2.5 mm broad at the base, triangular to awl- 
shaped, glabrous or rarely puberulent. Leaves opposite 
but 3-4 at distal nodes, petioles (1 .5-)6-l 1 cm long, 1 .3- 
2.7mm broad, glabrous or puberulent; leaf blades (7-) 1 2- 
23(-27) cm long, (3-)4-12(-15) cm broad, broadly ob- 
long or elliptic-oblong to ovate or obovate, apex obtuse 
to abruptly short-acuminate, base rounded and subtrun- 
cate to obtuse (cuneate), drying membranaceous to char- 
taceous, glabrous above, glabrous to minutely (0. 1 mm) 
papillate puberulent beneath in Costa Rica, domatia 
sometimes present, 2 veins 8-12/side and loop-con- 
nected near the margin. Inflorescences terminal or ax- 
illary, 6-17 cm long and wide, to 15 cm broad, panic- 
ulate, 20-many-flowered, peduncles 2-8 cm long, 1-3 
mm thick, lateral branches to 6 cm long, bracts 0.4-0.7 
mm long, ovate and acute, glabrate or puberulent, flow- 
ers sessile to short (2 mm) pedicellate. Flowers with hy- 
panthium 3-5 mm long, 1.5-2 mm diam., glabrous, calyx 
lobes 0.7-1 .5(-2) mm long, 1-1.5 mm broad at the base, 
triangular-subulate; corolla yellow, tubular-funnelform, 
tube 23-35 mm long, constricted near the base and 5- 

7 mm diam. distally, glabrous, lobes 3-5 mm long, ovate, 
glabrous; stamens with filaments 6-8 mm long, anthers 
16-20 mm long, included, connective prolonged ca. 1 
mm beyond the thecae; style to 20 mm long, stigmas 5- 

8 mm long. Fruits 11-15 mm long, 3.5-5 mm diam., 
ellipsoid, red becoming black and lustrous, ovarian disc 
conical; seeds 0.8-1 mm long. 

Trees and shrubs of evergreen cloud forest and 
lower montane forest formations on both the Ca- 
ribbean and Pacific slopes, from 700 to 1500 m 
elevation in Costa Rica. Flowering in June-Sep- 



158 



FIELDIANA: BOTANY 



tember in Costa Rica and Panama; fruiting in Au- 
gust-September. The species ranges from Costa 
Rica and Panama to Colombia and Ecuador. 

Hamelia macrantha is recognized by the large 
yellow corolla tube slightly widened distally at an- 
thesis, larger leaves with many secondary veins 
and often with long narrow petioles, and the mon- 
tane habitat (in Costa Rica). Note that the widened 
corolla tube is only seen at anthesis. The foliage 
of this species resembles that of some specimens 
of H. xerocarpa variety costaricensis, and the two 
species may be difficult to separate. 



Hamelia magnifolia Wernham, J. Bot. 49: 210. 
1911. Figure 42. 

Shrubs or small trees to 5(-6) m tall, trunks to 1 2 cm 
dbh, leafy branchlets 2-7 mm thick, with 4 longitudinal 
ridges and quadrangular in cross-section, glabrous or 
minutely (0. 1 mm) papillate-puberulent; stipules 4-9 mm 
long, to 2 mm wide at the base, narrowly triangular 
(rarely bifid), glabrate. Leaves opposite (rarely 4/node), 
petioles l-5(-7) cm long, 1.7-2.3 mm broad; leaf blades 
10-27(-32) cm long, 4-ll(-15) cm broad, broadly ob- 
long to ovate-oblong, elliptic-oblong or ovate-elliptic, 
apex short-to long-acuminate with tip to 2 cm long, base 
rounded and truncate to obtuse, drying stiffly charta- 
ceous, glabrous above, glabrous beneath, 2 veins 7- 
1 2(-l 5)/side, some 3 veins subparallel, domatia absent. 
Inflorescences terminal, 8-12(-18) cm long, to 15 cm 
wide, paniculate with opposite branching, with more than 
60 flowers, peduncles 1-3 cm long, distal branches to 7 
cm long and with secund flowers, minutely papillate- 
puberulent or glabrous, bracts 0.5-1 mm long, narrow 
pedicels 0-1 mm long. Flowers with hypanthium 2-3 
mm long, 0.8-1 .5 mm diam., minutely puberulent, calyx 
lobes 0.3-0.6 mm long, 1 mm broad at the base, deltoid 
or broadly rounded, puberulent along the margin; corolla 
narrowly tubular at anthesis (also somewhat narrowed 
above the base and below the middle), yellow, tube 10- 
13 mm long, 2-3 mm diam., lobes 1.2-3 mm long; sta- 
mens with filaments 2-3 mm long, anthers 6-8 mm long, 
apical connective ovate and ca. 1 mm long. Fruits 6-10 
mm long, 3-4 mm diam., oblong to broadly ellipsoid, 
glabrous, ovarian disc ca. 1 mm long, red; seeds 0.4-0.5 
mm diam. 



Plants of evergreen forest formations on the Pa- 
cific slope, from 1 00 to 900 m elevation in Costa 
Rica. Flowering in January-August; fruiting in 
June-September and December-January. The 
species is restricted to the evergreen formations of 
the Pacific slopes of southern Costa Rica and ad- 
jacent areas in Chiriqui Province, Panama. 

Hamelia magnifolia is recognized by the small 
narrowly tubular flowers in large inflorescences, 
the large leaves often rounded at the base and with 
many secondary veins, the smaller fruits, and the 



restricted geographic range. Standley (1938) listed 
zorillo Colorado as a common name. 



Hamelia patens Jacq., Enum. PI. Carib. 16. 1760. 
H. patens var. glabra Oersted, Vidensk. Meddel. 
Naturhist. Foren. Kjobenhavn 1852: 42. 1853. 
H. viridifolia Wernham, J. Bot. 49: 213. 1911. 
Figure 42. 



Shrubs or small trees, 2-7 m tall, leafy branchlets 1- 
7 mm thick, glabrous or pubescent with crooked trans- 
lucent or yellowish hairs to 0.6 mm long (in variety 
patens), 4-angled in early stages but becoming terete; 
stipules 2-6 mm long, narrowly oblong to linear, pu- 
bescent, as many as the leaves at each node. Leaves 
usually 3(-4) at distal nodes (less often opposite or rarely 
5/node), often unequal at the same node, separated by 
well-developed internodes, petioles (8-)20-55(-80) mm 
long, 0.6-1.6 mm wide, glabrous to densely pubescent; 
leaf blades 5-17(-23) cm long, l-7(-10) cm broad, el- 
liptic-oblong, ovate-elliptic, ovate-oblong, or elliptic ob- 
ovate-elliptic, apex usually short-acuminate, base acute 
to attenuate (obtuse in larger leaves) and decurrent on 
petiole, drying membranaceous to thin-chartaceous, gla- 
brous (or sparsely pubescent in variety patens) above, 
glabrous to densely villosulous beneath with thin straight 
or curved whitish hairs 0.2-0.5 mm long (rarely with 
scurfy yellowish hairs), tufts of hairs (domatia) often 
present in vein axils beneath, 2 veins 6-9/side, small 
(0.2 mm) linear raphides (cystoliths) resembling ap- 
pressed hairs often visible on ,the dark upper surface of 
dried leaves. Inflorescences terminal, solitary or 2-4, 4- 
9(-15) cm long, to 12(-20) cm broad, an open panicle 
with 2-3 nodes on the rachis and 2-4 branches at each 
node, peduncles 1-3 cm long (to 5 cm in fruit), minutely 
papillate-puberulent (less often with conspicuous hairs), 
often orange to coral red, primary branches often ending 
in a flower and 2 long distal secondary branches bearing 
a row of sessile or subsessile flowers along the upper side, 
bracts 0.3-1 mm long, triangular, distal flowers with 
pedicels 1-5 mm long. Flowers with hypanthium 1.5-3 
mm long, 1-2 mm diam., longitudinally ribbed, deep 
red, glabrate to densely short-villous. calyx lobes 0.5-1 
mm high, rounded or broadly triangular, persisting; co- 
rolla narrowly tubular at anthesis, orange to reddish or- 
ange, tube (12-)14-18(-23) mm long, 1.5-3 mm diam., 
lobes 1-2.5 mm long, 1-2 mm broad at the base, tri- 
angular; stamens with filaments 5-7 mm long, anthers 
8-12 mm long, included or slightly (3 mm) exserted. the 
apiculatc connective 0.5 mm long; stigmas 3-5 mm long. 
Fruits 6-1 3 mm long, 4-10 mm diam., oblong to ovoid- 
oblong (subglobose), red becoming black or bluish, gla- 
brous or with hairs ca. 0.2 mm long, raphides often 
visible on the surface, with a ring around the top formed 
by the calyx scar ca. 3 mm diam., disc to 1 mm high 
and inconspicuous; seeds 0.5-1 mm long. 

Common shrubs and treelets of open early sec- 
ondary growth in evergreen and partly deciduous 
forest formations, from near sea level to 1 600 m 
elevation. Flowering and fruiting in all months of 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



159 



the year in Costa Rica. The species ranges from 
southern Florida, Mexico, the West Indies, and 
Central America southward to northern Argentina 
and Paraguay (the species is not found in the 
Guianas, northeastern Brazil, the central Amazon 
basin, or central Brazil). 

Hamelia patens is recognized by its bright or- 
ange or red-orange flowers with narrowly tubular 
corollas with small lobes and included anthers and 
the usually three- or four-leaved distal nodes. The 
open inflorescences with dichotomous or dichasial 
branching and with secund flowers often in a row 
along the upper side of distal inflorescence branch- 
es are also distinctive. These shrubs and little tree- 
lets of open evergreen lowland secondary sites are 
among the most common and conspicuous of Cos- 
ta Rica's woody flora. They appear to germinate 
only in open sunny sites. The breeding biology of 
this species was studied by Bawa and Beach ( 1 983). 
Standley ( 1 938) listed the common names anileto, 
azulillo, coralillo, palo camaron, pissi, zorillo, and 
zorillo real, and he cited Pittier for the Indian 
names pili-tso (Guatuso) and tsus-krd (Brunka). 

Hamelia patens variety patens has the leaves 
sparsely to densely villous (especially on the lower 
surface), and the flowers are sparsely to densely 
villous externally. Variety glabra Oersted has the 
leaves glabrous above and sparsely villous or pu- 
berulent on the veins beneath, and the flowers are 
usually glabrous externally. There seem to be no 
ecological or geographic distinctions between the 
varieties, and they may be no more than glabrous 
and puberulent forms found within the same pop- 
ulations. 



Hamelia rovirosae Wernham, J. Bot. 49: 211. 1911. 
Figure 34. 

Shrubs or slender treelets to 5(-10) m tall, leafy 
branchlets 0.8-3.5 mm diam., with curved or crooked 
multiccllular hairs 0.3-1 mm long and often in longi- 
tudinal rows, with 4 longitudinal ribs and quadrangular 
in early stages but becoming terete and glabrescent; stip- 
ules 2-6(-8) mm long, with a very short (1 mm) broad 
base and long narrow linear awn, puberulent, caducous 
or persisting with the leaves. Leaves usually 3/node (rare- 
ly opposite), petioles 3-14(-20) mm long, 0.5-1 mm 
thick, pubescent with short crooked hairs; leaf blades 
(2.5-)5-15 cm long, (1.5-)2-6 cm wide, elliptic-oblong 
to elliptic-ovate, elliptic or elliptic-obovate, apex acute 
to short-acuminate, base acute to attenuate and slightly 
decurrent on petiole, drying thin-chartaceous, glabrous 
to sparsely pubescent above, sparsely to densely villous 
beneath with thin straight or crooked hairs 0.2-0.5 mm 
long, with denser tufts of hairs (domatia) in the vein 
axils, 2 veins 3-7 /side and weakly loop-connected near 



margin. Inflorescences 4-12 cm long, equally broad, pe- 
duncles to 3 cm long, villous, floral rachis with dichot- 
omous distal branches bearing 2-8 flowers, bracts to 0.5 
mm long, caducous, flowers sessile or short (1-2 mm) 
pedicellate. Flowers with hypanthium 2-4.5 mm long, 
villous with curly hairs ca. 0.5 mm long, calyx lobes 2- 
4 mm long and enlarging in fruit, 1 mm wide and oblong, 
villous; corolla narrowly tubular at anthesis, reddish or- 
ange to bright red or dark red, tube 16-22 mm long, 2- 
3 mm diam., minutely villous with hairs ca. 0.5 mm 
long (often in longitudinal rows), lobes 1-2 mm long, 1- 
1.5 mm broad at the base, ovate; stamens with filaments 
7-9 mm long, anthers 10-12 mm long, slightly exserted, 
connective appendage ca. 0.5 m long; stigmas connate, 
2-3 mm long. Fruit 8-14 mm long, 4-8 mm diam., 
cylindrical to oblong-ellipsoid, red becoming black, cov- 
ered with small (ca 0.5 mm) crooked hairs, persisting 
sepals to 6 mm long and 2 mm broad; seeds ca. 1 mm 
long. 

Shrubs of Caribbean lowland evergreen forest 
formations, from near sea level to 200 m elevation. 
Flowering in February, April, and July-October 
in Central America. The species ranges along the 
Caribbean lowlands, from Tabasco, Mexico, to 
Bocas del Toro, Panama. 

Hamelia rovirosae is recognized by the unusual 
crooked multicellular hairs, corollas narrowly tu- 
bular at anthesis, puberulent fruit with prominent 
calyx lobes, and restriction to the Caribbean low- 
lands. This species is frequently confused with H. 
patens. 



Hamelia xerocarpa Kuntze, Rev. gen. pi. 1: 284. 
1 89 1 . H. costaricensis Standl., Contr. U.S. Natl. 
Mus. 20: 207. 1919. H. panamensis Standl., loc. 
cit. 208. 1919. H. rowlei Standl., J. Wash. Acad. 
Sci. 15: 7. 1925. H. storkii Standl., loc. cit. 7. 
1925. Figure 42. 

Shrubs or small trees to 5 m tall, leafy branchlets 1.5- 
5 mm thick, with 4 longitudinal ridges and quadrangular 
in cross-section, glabrous to sparsely pilose with stiff 
erect hairs to 0.9 mm long; stipules 6-1 3(-l 7) mm long, 
cuspidate and often with 2 small lateral teeth, glabrate 
or pubescent along the edge, drying black. Leaves usually 
3 or 4/node, petioles (10-15)-85 mm long, about 1 mm 
broad, glabrous to densely pubescent; leaf blades 8.5- 
1 7(-37?) cm long, 3.5-9(-l 4?) cm broad, ovate to broad- 
ly elliptic-oblong or ovate-rotund, apex acute to acu- 
minate, base obtuse to cuneate and decurrent on petiole, 
leaves drying thin-chartaceous, glabrate above, usually 
minutely papillate-puberulent or with straight or curved 
hairs 0.2-0.9 mm long beneath, 2 veins (7-)9-13(-18)/ 
side and loop-connected near the margin, 3 veins often 
subparallel, domatia present or absent. Inflorescences 
terminal on short lateral branches (and apparently ax- 
illary), 5-15 cm long and equally wide, peduncles 1- 
2.5(-5) cm long, the dichasia to 6(-l 5) cm long and with 
3-9(-26) flowers along 1 side, glabrous or with yellowish 



160 



FIELDIANA: BOTANY 



hairs to 1 mm long, bracts 0.6-0.8 mm long, the secund 
flowers sessile or subsessile. Flowers with hypanthium 
2-3 mm long, glabrous to pilose, calyx lobes 0.5-2 mm 
long, ovate, glabrous to pilose; corolla funnelform (but 
narrowly tubular until anthesis), yellow, tube 1 8-36 mm 
long, 4-7 mm diam., glabrous to densely puberulent with 
hairs 0.2-0.4 mm long, lobes 2-5(-8) mm long, 2-5 mm 
broad at the base, ovate and acute; stamens with fila- 
ments 8-16 mm long, anthers 10-16 mm long, connec- 
tive little ( 1 mm) extended distally; style ca. 20 mm long, 
stigmas 5, to 5 mm long. Fruits 11-14 mm long and 3- 
5 mm diam., oblong to ovoid-oblong; seeds 0.5-1 mm 
long. 



Plants of evergreen lowland formations and gal- 
lery forests in deciduous areas, from near sea level 
to 800 m elevation. Flowering in May-December 
(throughout the year in Panama). The species 
ranges from Nicaragua to northern Colombia. 

Hamelia xerocarpa is recognized by the broader 
corolla tube (at anthesis) with longer corolla lobes, 
the unusual yellowish puberulence (when present), 
the lowland evergreen habitat, and the leaves usu- 
ally with many secondary veins and often with 
long petioles. Unfortunately, few herbarium col- 
lections exhibit the broader funnelform corollas 
in anthesis, and it is easy to misidentify this spe- 
cies. This species may be difficult to separate from 
H. macrantha. The breeding biology was studied 
by Bawa and Beach (1983). 

Hamelia xerocarpa variety xerocarpa is distin- 
guished by the conspicuous hairs on many parts 
of the plant and the leaves with 11-18 pairs of 
secondary veins. Variety costaricensis (Standl.) 
Elias is recognized by the lack of pubescence and 
leaves with 9-12 pairs of secondary veins. 



1 1 illia Jacquin 

REFERENCE C. M. Taylor, Revision of Hillia 
subg. Ravnia (Rubiaceae: Cinchonoideae). Sel- 
byana 11: 26-34. 1989. 



Shrubs, small trees, or lianas, epiphytic or less often 
terrestrial, branchlets thick and terete, glabrous; stipules 
interpetiolar and intrapetiolar but splitting apart along 
the edges. Ungulate and blunt at apex, caducous. Leaves 
opposite, equal or unequal at each node, subsessile to 
short-petiolate; leaf blades elliptic to obovate, entire, 
decurrent on the petiole, semisucculent and drying co- 
riaceous, without domatia. Inflorescences of solitary ter- 
minal flowers (or 3-flowered dichasia in H. thflora), bracts 
reduced or absent (the flowers at first enclosed within 
the large untied stipules), pedicels short or absent. Flow- 
ers bisexual, monomorphic, often large, glabrous exter- 
nally, hypanthium continuous with the pedicel, calyx 
tube often absent, calyx lobes 2-5 and distant (or none), 
sometimes with a secondary smaller set of calyx lobes 
alternate with the larger lobes; corolla salverform to fun- 
nelform or tubular, white to yellowish, pinkish, orange, 
or red, semisucculent, corolla lobes (3-)5-7(-9), con- 
volute in bud, becoming reflexed; stamens (4-)5-7, fil- 
aments very short and inserted below the throat, anthers 
basifixed, elongate-linear, obtuse at each end, included 
(except in H. longifilamentosa); ovary 2-locular, ovules 
many and ascending in each locule on scptal placentas, 
style as long as the corolla tubes, stigmas subcapitate or 
2. Fruits woody capsules, narrowly cylindrical to very 
narrowly oblong, truncated distally, dark brown and 
smooth, dehiscing septicidally and basipetally into 2 flat- 
tened valves; seeds many and imbricated, rhombic and 
flattened, with a minute circumferential wing, appen- 
daged at the base and with a tuft of hairs at the distal 
apex. 

Hillia is a genus of about 20 species, ranging 
from southern Mexico to Brazil and Peru. The 
genus is recognized by the larger flowers with long 
tubes, many ascending imbricated ovules, long tu- 
bular ("cigar-shaped") capsules, and flattened 
winged seeds with a tuft of hairs at one end. The 
more colorful flowers of subgenus Ravnia (see fol- 
lowing key) appear to be an adaptation to bird 
pollination. Specimens lacking flowers or fruit can 
be very difficult to identify to species. In addition, 
one group of our species may be part of a poly- 
morphic complex; see the discussion under H. 
maxonii. Some of our species of Hillia are very 
similar to Cosmibuena, but that genus has seeds 
lacking the tufted hairs at one end. 



Key to the Species of Hillia 

la. Corolla pinkish to red or orange, funnelform or tubular with the central part of the tube inflated 

and narrowed at both ends, flowers not scented; subgenus Ravnia . 

Ib. Corolla greenish or white, long-tubular with rotate lobes or salverform, corolla tube expanded only 
at the corolla lobes, flowers often sweet-scented; subgenus Hillia . 4 

2a. Corolla tube rose red and inflated in the middle, 3-5 cm long, flowers often in groups of 3; a 

commonly collected species " triflora 

2b. Corolla tube yellow-orange to peach or rose, funnelform and widest at the mouth, 4-6.5 cm 

long, flowers solitary; rarely collected species .... 
3a. Corolla lobes 6-12 mm long; free portion of the filaments ca. 1 mm long H. allenii 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



161 



3b. Corolla lobes 13-17 mm long; free portion of the filaments 9-10 mm long 

H. longifilamentosa 

4a. Plants of lower elevation forest, not known from above 300 m in Costa Rica; larger leaf blades 

more than 10 cm long and 5 cm broad 5 

4b. Plants of higher elevations, rarely collected below 600 m; larger leaf blades rarely more than 1 cm 

long or 5 cm broad 6 

5a. Leaf blades with thin texture, the 5-6 pairs of 2 veins arising at ca. 60 angles from the midvein; 
corolla tubes 8-10 cm long H. macrophylla 

5b. Leaf blades thick-textured, the 4-5 pairs of 2 veins arising at ca. 30 angles from the midvein; 

corolla tubes 4-5 cm long H. grayumii 

6a. Leaf blades 4-10 cm long, usually tapering gradually to a bluntly acute apex, broadest at the middle 

or below; corolla tubes 4-5 cm long; seed hairs ca. 1 7 mm long H. loranthoides 

6b. Leaf blades 0.7-3.7 cm long, usually bluntly obtuse to rounded at the apex; corolla tubes rarely 

exceeding 4 cm in length; seed hairs ca. 10 mm long 7 

7a. Leaf blades 12-37 mm long, stipules 6-16 mm long; corolla lobes suborbicular, corolla tubes 24- 

42 mm long; seeds ca. 3 mm long; 1400-2400 m elevation H. maxonii 

7b. Leaf blades 7-14 mm long, stipules 3-7 mm long; corolla lobes broadly to narrowly ovate, corolla 

tubes 15-35 mm long; seeds ca. 2 mm long; 600-1600 m elevation H. panamensis 



Hillia allenii C. M. Taylor, Selbyana 1 1: 32. 1989. 
Ravnia panamensis Steyerm., Ceiba 3: 22. 1 952, 
not Hillia panamensis Standl. 

Epiphytic shrubs to 1.5 m tall, leafy stems 2.5-7 mm 
thick, glabrous, terete; stipules 10-12 mm long, Ungulate, 
quickly caducous. Leaves with short (14 mm), thick (2- 
3 mm) petioles, glabrous and drying dark; leaf blades 4- 

1 1 cm long, 1.5-5 cm broad, elliptic to narrowly elliptic- 
oblong, apex slender acuminate, base cuneate or slightly 
rounded, drying subcoriaceous, dark brown above, gla- 
brous above and below, 2 veins 5-9/side, usually ob- 
scure. Inflorescences of solitary terminal flowers borne 
on short (34 mm) thick (2 mm) glabrous pedicels drying 
black. Flowers glabrous externally, hypanthium 4-7 mm 
long and 3.5 mm thick, calyx tube not developed, calyx 
lobes 6, 6-14 mm long, 2-3 mm broad, narrowly spat- 
ulate-oblong; corolla funnelform, pale red and pale yel- 
low to salmon-pink, tube 25-40 mm long, lobes 6, 6- 

1 2 mm long and 1 mm broad at the base, bluntly acute; 
anthers 5-7 mm long on filaments ca. 1 mm long. Fruits 
not seen. 

Plants of montane cloud forest formations at ca. 
1000 m elevation. Flowering in June and Septem- 
ber. This species known only from the Cordillera 
de Tilaran and in western Panama. 

Hillia allenii is recognized by the brightly col- 
ored funnelform corolla and the short filaments. 
Fruiting material may be very difficult to distin- 
guish from that of H. longifilamentosa. 



Hillia grayumii C. M. Taylor, Selbyana 12: 137. 
1991. Figure 28. 

Epiphytic shrubs, ca. 1 m tall, leafy stems 4-7 mm 
thick, slightly quadrangular, glabrous, brownish, mi- 



nutely grooved; stipules ca. 40 mm long, 6-8 mm broad, 
lanceolate, caducous. Leaves isophyllous, usually decus- 
sate, petioles 6-20 mm long, 2-3 mm thick, articulated 
at the stem; leaf blades 9-19 cm long, 3-6.5 cm broad, 
elliptic to elliptic-oblong, apex acuminate with tip 1-2 
cm long, base obtuse or acute, drying stiffly chartaceous 
or subcoriaceous and dark brown above, glabrous above 
and below, 2 veins 4-6/side and strongly ascending (ca. 
30), obscure beneath. Inflorescences of solitary terminal 
flowers, subtending stipules caducous, peduncles 2-3 mm 
long, bracts 1-3 mm long, triangular, acute. Flowers gla- 
brous, hypanthium 7-8 mm long, cylindrical, calyx limb 
to 0.5 mm long, truncate or slightly lobed; corolla tu- 
bular- funnelform, bright pale green to yellow, tube 43- 
50 mm long, lobes 6, 8-9 mm long, triangular, obtuse 
to rounded; filaments ca. 10 mm long, anthers 6, ca. 9 
mm long. Fruits 10-12 cm long and 8-14 mm broad, 
drying dark brown, stipe ca. 3 mm long; seeds 3 mm 
long, 0.5 mm broad. 



Plants of lowland rain forest and swamp forest 
formations, collected from near sea level to 600 
m elevation. Flowering in May-June; fruiting in 
March and May-June. The species is known only 
from the Caribbean lowlands of northern and cen- 
tral Costa Rica. 

Hillia grayumii is distinguished from its con- 
geners by the lowland habitat, yellowish funnel- 
form flowers, and large fruit. Among Costa Rican 
species, it is similar to H. macrophylla, but that 
species grows in cloud forests and has thin-tex- 
tured leaves and tubular flowers. 



Hillia longifilamentosa (Steyerm.) C. M. Taylor, 
Selbyana 1 1: 32. 1989. Ravnia longifilamentosa 
Steyerm., Ceiba 3: 21. 1952. 



162 



FIELDIANA: BOTANY 



Epiphytic or clambering shrubs to 6 m tall, leafy stems 
2-7 mm thick, glabrous, smooth, brown to gray; stipules 
22-37 mm long, elliptic to oblanceolate, caducous. Leaves 
with petioles 2-8 mm long, thick, glabrous; leaf blades 
6-16 cm long, 2-7 cm broad, elliptic or elliptic-oblong, 
apex acute or slightly acuminate, base obtuse to cuneate, 
drying subcoriaceous, grayish green, glabrous above and 
below, 2 veins 4-6/side, strongly ascending. Inflores- 
cences of solitary terminal flowers, pedicles ca. 1 mm 
long, bracts 2-3 mm long, triangular, acute. Flowers with 
hypanthium 4-10 mm long, obconic to ellipsoid, calyx 
lobes 6, 6-14 mm long, narrowly triangular to Ungulate 
or oblanceolate; corolla tubular-rotate, orange-red to 
salmon-pink, or white marked with pink, tube 32-43 
mm long, ca. 3 mm diam., lobes 6, 13-17 mm long, 
triangular to Ungulate; stamens 4-6, anthers ca. 5 mm 
long, well exserted, dark green. Fruits 10 cm long. 

Plants of montane cloud forest formations, 1 1 00- 
1700 m elevation. Flowering in April, July-Au- 
gust, and November. This species is known only 
from a few collections, ranging from Zarcero, Ala- 
juela, to Chiriqui, Panama. 

Hillia longifilamentosa is distinguished by its 
solitary terminal flowers with anthers extended 8- 
10 mm beyond the throat of the tube. Compare 
H. allenii. 



Hillia loranthoides Standl., J. Wash. Acad. Sci. 
18: 165. 1928. Figure 27. 

Epiphytic shrubs, 0.4-1.5 m tall but with vining 
branches, leafy stems 3-8 mm thick, glabrous, grayish, 
terete or quadrangular; stipules 18-22 mm long (to 35 
mm beneath the flowers), 6-10 mm broad, oblong-ob- 
ovate, obtuse or rounded distally. Leaves closely clus- 
tered or distant, decussate, petioles 4-8(-15) mm long, 
1.5-2.8 mm thick; leaf blades (3-)4-10 cm long, (1 .3-)2- 
4.5 cm broad, elliptic to elliptic-oblong, ovate or ovate- 
lanceolate, apex bluntly acute or obtuse, base obtuse to 
cuneate, drying coriaceous, dark grayish, 2 veins 4-57 
side, strongly ascending or obscure. Inflorescences of sol- 
itary terminal sessile flowers, subtended by a pair of 
enlarged (8-25 mm) oblong-obovate bract-like reddish 
stipules, 1-2 smaller (1-4 mm) pairs of triangular or 
Ungulate bracteoles often present at the base of the short 
pedicel. Flowers 6-7 cm long, hypanthium ca. 4 mm 
long, calyx tube minute, calyx lobes 4, 8-10 mm long, 
1-2 mm broad, linear-Ungulate; corolla tubular with ro- 
tate lobes, white or cream, carnose, tube 40-60 mm long, 
2-5 mm diam., lobes 4, 10-25 mm long, 8-16 mm broad, 
rounded distally. Fruits 3-8 cm long, 7-10 mm diam.; 
seeds ca. 2.5 mm long and 0.7 mm thick, distal hairs 6- 
17 mm long. 

Plants of evergreen montane forests, from 
(300-)700 to 1400 m elevation. Flowering in Feb- 
ruary-May; fruiting in January and May. In Costa 
Rica, this species is known only from the Cordil- 
lera de Tilaran, near San Ramon, and El Retiro 



(Cartago) and from a single lower elevation col- 
lection on the Osa Peninsula. The species is also 
known from southern Mexico. 

Hillia loranthoides is distinguished by its thick 
leaves, usually tapering gradually to the apex, larg- 
er flowers with slender tubes, and seeds with longer 
bristles. Its distribution is unusual and may be an 
artifact of the difficulty of collecting epiphytes. 



Hillia macrophylla Standl., Publ. Field Columb. 
Mus., Hot. Ser. 7: 201. 1931. Figure 28. 

Epiphytic shrubs or vines, 3-7 m tall, leafy branchlets 
3-7 mm thick, glabrous, pale brown and lenticelate; stip- 
ules 10-35 mm long, 12-25 mm broad, narrowly lan- 
ceolate (in Costa Rica) to oval-oblong, the basal sheath 
1-2 mm long, caducous. Leaves decussate, petioles 6- 
20 mm long, 2-3.5 mm thick, terete, clearly differenti- 
ated from the lamina base; leaf blades 9-2 1 cm long, 6- 
10 cm broad, ovate-elliptic to elliptic-oblong or elliptic, 
apex acuminate (acute), base obtuse, drying membra- 
naceous to chartaceous, dark brown above, glabrous 
above and below, major veins with a rugose texture. 2 
veins 6-9/side arising at ca. 60 angles and loop-con- 
nected near the margin. Inflorescences of large solitary 
sessile terminal flowers, subtended by elongated (20-50 
x 5-24 mm) caducous stipules, bracteoles usually ab- 
sent. Flowers with hypanthium 6-7 mm long, 3-3.5 mm 
diam., cylindrical, calyx tube 0-0.5 mm long, subentire; 
corolla tubular-salverform, white, tube 5.5-1 1 cm long, 
2-4.5 mm diam., lobes 5 or 6, 2.5-5 cm long. ca. 8 mm 
broad at the base and narrow (4 mm) distally, linear- 
lanceolate to narrowly triangular. Fruits 7-12 cm long, 
the opened valves becoming 14-20 mm broad, rounded 
at the base, acute at apex, sessile; seeds 1.5-4 mm long 
with distal hairs 14-18 mm long. 

Plants of moist cloud forests from 800 to 1 800 
m elevation. Flowering in March-June; a single 
fruiting collection was made in December. This 
species is known only from near Monteverde, Ca- 
taratas de San Ramon, and above the Rio Gato, 
Cartago, in Costa Rica. It is also found in Colom- 
bia, Ecuador, and Peru. 

Hillia macrophylla is unique among our species 
of Hillia because of the larger thin-textured leaves, 
more numerous ascending secondary veins, and 
long-tubular flowers with corolla lobes that have 
long (3-4 cm), narrow tips. It is a rarely collected 
species. South American collections appear to have 
larger (3-4 mm) seeds and larger stipules sub- 
tending the flowers. 



Hillia maxonii Standl., J. Wash. Acad. Sci. 18: 
163. 1 928. H.palmana Standl. Joe. cit. 18: 164. 
1928. H. hathewayi Fosberg, Sida 2: 387. 1966. 
Figure 27. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



163 



Epiphytic shrubs, 0.7-2. 5(-5) m tall, often pendant to 
5 mm long, leafy stems 1-5 mm thick, dark or pale 
grayish, glabrous, older nodes articulate with transverse 
ridges; stipules 6-16(-32) mm long, 2-6 mm broad, ob- 
long-obovate to obovate, rounded, larger and bract-like 
beneath the flowers. Leaves decussate and often crowded 
on short (3-15 mm) internodes, petioles 2-6(-15) mm 
long; leaf blades (12-)18-37(-60) mm long, 6-20(-30) 
mm broad, obovate to oblong-obovate or ovate-elliptic, 
apex bluntly obtuse to rounded, base obtuse to cuneate, 
decurrent on the petiole, drying subcoriaceous and dark 
grayish or brownish, glabrous above and below, 2 veins 
3-4/side, strongly ascending or obscure. Inflorescences 
of solitary, terminal, sessile flowers subtended by a pair 
of enlarged bract-like stipules rounded at apex. Flowers 
glabrous externally, fragrant, hypanthium ca. 2.5 mm 
long and 1 .8 mm thick, calyx tube very short, calyx lobes 
absent or 4-6 mm long, 0.5-1.8 mm broad; corolla tu- 
bular with rotate lobes, slightly fleshy, tube 24-42 mm 
long, 1.4-4 mm diam., greenish or white, lobes 4, 10- 
14(-22) mm long and usually equally broad or broader 
than long, suborbicular and rounded distally, bright white 
or yellowish white. Fruits (22-)30-60 mm long, 5-9 mm 
thick, the valves to 8 mm broad when opened and ex- 
panded; seeds 3-4 mm long, 0.5-1 mm diam., distal 
hairs 6-13 mm long. 

Plants of evergreen montane forest formations 
from 500 to 2400 m elevation. Flowering in April- 
September and December; fruiting in September 
and November-March. This species has been col- 
lected near Managua, Nicaragua, in the Cordillera 
de Tilaran, Cordillera Central, in the western part 
of the Cordillera de Talamanca, and in Chiriqui 
and Veraguas, Panama. 

Hillia maxonii is recognized by its usually epi- 
phytic habit, smaller stiff leaves, bract-like stipules 
subtending the flowers, and tubular corollas with 
broadly rounded lobes. Smaller-leaved specimens 
of this species may represent intermediates with 
H. panamensis. Larger-leaved specimens resem- 
ble H. tetrandra Sw. of the West Indies and north- 
ern Central America. However, H. tetrandra has 
larger (5-1 1 x 2.5-6 cm) leaves that are more 
often obovate, quite unlike those of southern Cen- 
tral America. All three taxa exhibit a wide range 
of variation, and it is possible that they are ele- 
ments of a single polymorphic species. (A similar 
problem is found in the epiphytic species of Psy- 
chotria; see the discussion under P. guadalupen- 
sis.) 



Hillia panamensis Standl., N. Amer. Fl. 32: 117. 
1921. H. chiapensis Standl., J. Wash. Acad. Sci. 
16: 16. 1926. Figure 27. 

Epiphytic shrubs or small treelets to 4 m tall, leafy 
stems 1-2.7 mm thick, glabrous and grayish, often de- 



veloping opposite longitudinal sulci and expanded nodes; 
stipules 3-7 mm long, 1-2 mm broad, oblong to narrowly 
obovate and rounded distally, translucent. Leaves de- 
cussate and usually closely crowded on short (0.5-5 mm) 
internodes, petioles 1-4 m long; leaf blades 7-12(-14) 
mm long, 4-8(-10) mm broad, elliptic to elliptic-oblong 
or obovate, apex acute to bluntly obtuse or rounded, 
base cuneate to obtuse, drying stiffly chartaceous to sub- 
coriaceous, grayish to very dark, margins often becoming 
slightly revolute, glabrous above and below, 2 veins 2- 
3/side, strongly ascending or obscure. Inflorescences of 
solitary terminal flowers, the distal stipule pair slightly 
expanded (6 x 2.5 mm) and bract-like smaller (1-2 mm) 
rounded bracts sometimes present at the base of the 
sessile flowers. Flowers glabrous externally, hypanthium 
ca. 1.5 mm long, 1 mm thick, tubular, calyx lobes to 1 1 
mm long, 0.3-0.6 mm broad, linear, caducous; corolla 
tubular with rotate lobes, white, slightly succulent, tube 
15-30(-38) mm long, 0.9-1.5 mm diam., lobes 4, 5-7 
mm long, 3.5-6 mm broad, ovate to lanceolate, bluntly 
obtuse. Fruits 20-42 mm long, opened expanded valves 
3-4 mm wide; seeds ca. 2 mm long and 0.6 mm thick, 
distal hairs 10 mm long. 

Plants of evergreen montane forest formations 
from 600 to 1 600 m elevation. Flowering in May- 
August; fruiting in December. This species is found 
in southern Mexico-Guatemala, in the Cordilleras 
de Guanacaste and Tilaran in Costa Rica, and in 
western and central Panama. 

Hillia panamensis is distinguished by its epi- 
phytic habit, very small stiffclosely crowded leaves, 
fragrant flowers with long slender corolla tubes, 
and rotate lobes usually narrower than long. It has 
been called jasmin del volcdn. This species may 
not be specifically distinct from the very similar, 
and partly sympatric, H. maxonii. But the differ- 
ences used in the keys do seem to separate a great 
majority of specimens. See the discussion under 
H. maxonii. 



Hillia triflora (Oersted) C. M. Taylor, Selbyana 
1 1: 30. 1989. Ravnia triflora Oersted, Vidensk. 
Meddel. Dansk Naturhist. Foren. Kjobenhavn 
1852: 49. 1853. Ravnia pittien 'Standl., N. Amer. 
Fl. 32: 1 14. \92l.LagenanthusparviflorusE\van, 
Mutisia 4: 5. 1952. H. triflora var. pittieri 
(Standl.) C M. Taylor, Selbyana 11:31. 1989. 
Figure 27. 

Epiphytic (rarely terrestrial) shrubs, 0.5-1 .5(-3) m tall 
(branches to 2 m long and pendulous), leafy stems 1.8- 
5 m thick, glabrous; stipules 15^43 mm long, 4-10 mm 
broad, largest beneath the inflorescences, elliptic and 
acute, glabrous and caducous. Leaves usually decussate, 
isophyllous or anisophyllous at a node, petioles 3-7(-l 2) 
mm long, 1.5-2 mm thick, poorly differentiated from 
the base; leaf blades 5-1 3(-l 9) cm long, 1 .5-4.5 cm broad, 
narrowly elliptic-oblong to oblanceolate, elliptic-oblong 



164 



FIELDIANA: BOTANY 



or oblong (linear-lanceolate in variety pittieri), apex ta- 
pering gradually and narrowly acuminate, tapering grad- 
ually to the acute or cuneate base, drying coriaceous, 
dark green or grayish green above, 2 veins 3-5/side. 
Inflorescences of 1 or 3 terminal flowers borne on a very 
short (0.5-2 mm) thick peduncle, bracts ca. 3 mm long 
or absent, pedicels 1-6 mm long but difficult to distin- 
guish from the hypanthium. Flowers glabrous externally, 
hypanthium ca. 5 mm long, 1.2-1.8 mm diam. calyx 
lobes 6 (5, 7), 4-14 mm long, 0.5-1 mm broad, linear- 
oblong; corolla red, tubular with relatively small lobes, 
tube 40-65 mm long, 1.5-10 mm broad, lobes 6 (5, 7), 
3-4 mm long, to 5 mm broad at the base; stamens 5-6, 
anthers 3-5 mm long, partially exserted. Fruits 5-10 cm 
long, ca. 8 mm diam., valves becoming up to 12 mm 
broad; seeds 0.8-2.5 mm long, distal hairs 15-30 mm 
long. 

Plants of montane and premontane evergreen 
forest formations from (100-)800 to 2400 m ele- 



vation. The lowest elevation collections are from 
La Selva; this species has not been collected from 
below 1000 m on the Pacific slope. Flowering in 
all months, but most commonly February-Au- 
gust; fruiting in October-February. The species 
ranges from southern Mexico to northwestern Co- 
lombia. 

Hillia triflora is distinguish by its epiphytic hab- 
it, the tubular reddish flowers usually in terminal 
triads, long narrow capsules, and seeds with a dis- 
tal tuft of hairs. There are few fruiting collections. 
There may be some local differentiation between 
the two varieties; at Monteverde, variety pittieri 
is found at slightly lower elevations than is variety 
triflora. The following key distinguishes the two 
varieties. 



la. Leaves at each node approximately equal in size at the same node on flowering shoots; mature 
corolla tube not or only slightly inflated, up to 1.6 times broader in the broadest portion as at the 
apex H. triflora var. triflora 

Ib. Leaves at each node strongly differing in size on flowering shoots, larger leaves ca. 1.5 times longer 
than the other leaf of the same node; mature corolla tube strongly inflated, ca. 1 .8 times broader 
in center than at the apex H. triflora var. pittieri 



Hippotis Ruiz Lopez & Pavon 

Trees or shrubs, puberulent (in ours) or glabrous; stip- 
ules interpetiolar, triangular to obovate, large and ca- 
ducous. Leaves opposite, decussate, petiolate; leaf blades 
usually thin, pinnately veined, without domatia, with 
the minor venation parallel (lineolate) within the areolae 
formed by the 3 and 4 veins. Inflorescences axillary to 
leaves or undeveloped leaves and axillary in fruit, short 
cymose and 2-3-flowered, capitate or of solitary flowers. 
Flowers bisexual, monomorphic, radially symmetrical 
or bilaterally symmetrical by curvature of the corolla, 
usually large, sericeous externally, calyx bilobate or spa- 
thaceous; corolla tubular or funnelform, white, rose, red, 
or pale orange, corolla tube straight or curved, corolla 
lobes 5, short and truncated or emarginate, plicate-val- 
vate in bud; stamens 5, filaments usually unequal, borne 
at the middle or lower half of the tube, anthers included; 
ovary 2-locular, placentas peltate from the center of the 
septum, with many horizontal ovules in each locule, 
stigmas 2, subcapitate. Fruits baccate, ovoid to ellipsoid 
or globose; seeds many, small, angular. 

A genus of around 12 species in northern South 
America, Ecuador, and Peru; 1 species reaches 
northernmost Costa Rica. The larger thin hairy 
leaves, unusual lineolate minor venation, few larg- 
er pseudoterminal flowers, spathaceous calyx, and 
hairy fleshy fruit with many small seeds help to 
distinguish our species. 



Hippotis albiflora Karst., Fl. Colomb. 1: 33, pi. 
1 7. 1 858. Duroia panamensis Dwyer, Ann. Mis- 
souri Bot. Gard. 55: 38. 1968. Figure 25. 

Small trees, 3-17 m tall, leafy stems 2.5-8 mm thick, 
densely pilose with yellowish hairs 1-3 mm long, terete 
or slightly quadrangular, older stems glabrescent and pale 
grayish; stipules 15-30 mm long, 5-12 mm broad, nar- 
rowly ovate or lanceolate, with long yellowish hairs ex- 
ternally, leaves with petioles 10-28(-35) mm long, 1.5- 
2.5 mm thick, densely pilose with erect or ascending 
hairs 1-2 mm long; leaf blades 14-30(-35) cm long, 6- 
14(-18) cm broad, obovate to oblong-obovate, apex ta- 
pering gradually or abruptly and acuminate or caudate- 
acuminate, tip 1-2 cm long, base obtuse to rounded and 
subtruncate, drying thin-chartaceous to chartaceous, 
densely pilose on both surfaces with thin straight or curved 
hairs 0.7-2 mm long, 2 veins 7-10/side, 3 veins often 
subparallel, the smallest veins distinctly parallel (lineo- 
late) in small groups. Inflorescences of 1-3 flowers in the 
axils of the distal leaves but often appearing terminal, 
peduncles absent or very short, bracts 3-5 mm long, 
ovate, pedicels 3-8 mm long, 1-2 mm thick and densely 
yellowish velutinous. Flowers 5-6 cm long, hypanthium 
and calyx 3.3-3.8 cm long, ca. 8 mm broad, densely 
pilose, spathe-like and split ca. 10 mm down 1 side; 
corolla ca. 5 cm long and 2 cm broad distally, funnel- 
form, white to cream, pilose externally, tube 35-45 mm 
long, greenish white, straight or slightly curved, lobes 5- 
8 mm long and 10 mm broad, obtuse. Fruits 3-4 cm 
long (not including the persisting calyx), 1-2 cm diam., 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



165 



ellipsoid and hirsute, becoming purple; seeds 2 mm long, 
foveolate. 



Plants of rain forest formations in the Caribbean 
lowlands, from 10 to 400 m elevation (to 1600 m 
in Venezuela). Flowering in February, April, and 
July-October; fruiting in May-September. This 
species has been collected near Rio Colorado in 
northern Costa Rica, from La Selva and the Hitoy 
Cerere reserve. It is also known from the province 
of Bocas del Toro, in Panama, and from Colombia 
and Venezuela. 

Hippotis albiflora is recognized by the conspic- 
uous pubescence, larger thin leaves with lineolate 
minor venation, few large pseudoterminal flowers, 
spathaceous calyx, and larger, hirsute, many-seed- 
ed fruit. Our material may differ in some details 
from that described from Venezuela; it could be 
that Costa Rican populations are worthy of sub- 
specific recognition. Compare this species with 
Duroia costaricensis. 



Hoffmannia Swart/ 

Herbs, herbaceous subshrubs, or slender shrubs, main 
vertical stems usually with few or no lateral branches, 
glabrous or pubescent, terete or quadrangular, internodes 
often hollow when dried; stipules interpetiolar, usually 
small and triangular, often succulent and divergent, ca- 
ducous. Leaves opposite and decussate or rarely verti- 
cillate, equal at the node or sometimes slightly unequal, 
petiolate or occasionally subsessile, petioles with inflated 
chambers (ant vesicles) in a few species; leaf blades usu- 
ally drying membranaceous to chartaceous (rarely sub- 
coriaceous), entire, often decurrent on the petiole, pin- 
nately veined, often with conspicuous raphides on the 
dried leaf surfaces, domatia rarely present. Inflores- 
cences axillary, sessile or pedunculate, with few to many 
flowers, usually cymose or dichasial and often with heli- 
coid branches, fasciculate to capitate in a few species, 
flowers usually pedicellate, bracts and bracteoles rarely 



present. Flowers bisexual and monomorphic, glabrous 
or pubescent externally, hypanthium often with longi- 
tudinal ribs, calyx tube usually very short, calyx lobes 
4(-5), usually short and often triangular, persisting; co- 
rolla salverform to funnelform or rotate, white, yellow, 
orange, or rose to deep red or purple, corolla tube short 
and usually narrow, glabrous on the interior, corolla lobes 
4(-5), imbricate along the edges or apparently valvate, 
apex usually acute; stamens 4 (3-5), borne within the 
tube of the corolla, filaments very short, anthers linear 
to narrowly oblong, usually white; ovary 2- (3, 4) locular, 
each locule with many ovules borne longitudinally on 
bilamellate axile placentas, style slender, stigmas bilo- 
bate or clavate. Fruits baccate, usually small, 2- (3, 4) 
locular, often becoming enlarged and spongy at maturity, 
with many minute multiseriate horizontal seeds; seeds 
angulate, their surfaces often reticulate to foveolate. 



A genus of about 1 00 species, ranging from Mex- 
ico and the West Indies to South America. A ma- 
jority of the species are found in Mexico and north- 
ern Central America. The usually single-stemmed 
habit, presence of raphides in many parts, small 
triangular stipules (often caducous), axillary inflo- 
rescences, four-parted flowers, and many-seeded 
fleshy fruits help to distinguish the species of Hoff- 
mannia from other Rubiaceae in Costa Rica. Most 
of the Costa Rican species are restricted to areas 
with very high rainfall (none are found in decid- 
uous vegetation), and relatively few species grow 
below 800 m elevation. 

Hoffmannia is probably Costa Rica's taxonom- 
ically most difficult genus of Rubiaceae. These 
semisucculent subshrubs do not dry well, and the 
flowers, often in small dense clusters, are also not 
well preserved. In addition, there appears to be 
great variation from plant to plant within many 
species. The genus is related to Hamelia and Dep- 
pea; it is currently being studied by Dr. John Dwyer 
(MO), and the review presented here should be con- 
sidered no more than tentative. 



Key to the Species of Hoffmannia 

la. Petioles with lateral longitudinal inflated vesicles; plants to 1.2 m tall, surfaces with longer (1.5 
mm) multicellular hairs H. vesiculifera 

Ib. Petioles without lateral inflated structures; plants 0.5-3 m tall, glabrous to villous with multicellular 
hairs 2 

2a. Inflorescences sessile and capitate in the leaf axils in early stages (rarely becoming pedunculate or 
branched in later stages), often verticillate at the nodes 3 

2b. Inflorescences sessile and fasciculate to pedunculate and paniculate or cymose, neither densely 

capitate nor verticillate 4 

3a. Leaf blades neither bullate nor areolate, to 30( 40) cm long, glabrous or pubescent; corolla 
ca. 7 mm long; commonly collected plants H. congesta 



166 



FIELDIANA: BOTANY 



3b. Leaf blades bullate or areolate, to 20(-25) cm long, pubescent; corolla ca. 4.5 mm long; rare 

H. areolata 

4a. Leaves usually 3/node, leaf blades cuneate at the base with long-decurrent lateral margins on the 
petiole and these slightly expanded near the stem, lateral margins of the petiole 2-4 mm broad 
[leaf blades with 1 1-18 pairs of 2 veins]; Vara Blanca to the upper Rio Grande de Orosi, 1400- 

1600 m elevation H. amplexifolia 

4b. Leaves opposite (rarely with 3 leaves per node), leaf blades obtuse to cuneate at the base, if long- 
decurrent on the petiole never forming a lateral margin 2-4 mm wide along the petiole and not 

expanded near the base; species widely distributed, 10-2300 m elevation 5 

5a. Plants of Cocos Island 6 

5b. Plants of mainland Central America 7 

6a. Petioles 2-5 cm long; corolla glabrous on the exterior H. piratarum 

6b. Petioles 6-8 cm long; corolla puberulent on the exterior H. nesiota 

7a. Young stems glabrous or subglabrous, or with very small (0.2 mm) hairs in longitudinal lines along 

the young stern 8 

7b. Young stems densely pubescent with crooked multicellular hairs, the hairs varying 0.3-2 mm long, 

older stems pubescent or glabrescent 19 

8a. Inflorescences short, the 1 peduncle and rachis usually less than 3 cm long (if plants grow 
below 600 m elevation and have petioles 2-6 cm long and leaves 4-15 cm broad, go to 

dichotomy 12) 9 

8b. Inflorescences small to large, peduncle and rachis of some inflorescences usually exceeding 

4 cm in length 12 

9a. Corolla tubes 3-4 mm diam. and 3-4 mm long; leaf blades 15-35 cm long with 9-15 

pairs of 2 veins, cuneate basally and long-decurrent on the petiole H. dolae 

9b. Corolla tubes 1-3 mm diam. (dried) and 1-6 mm long; leaf blades 5-15 cm long with 
5-9 pairs of 2 veins, abruptly narrowed at the base (but leaves to 23 cm long, with 7- 

12 pairs of 2 veins, and cuneate long-decurrent bases in H. hamelioides) 10 

lOa. Corolla 9-14 mm long, corolla lobes 3-6 mm long; calyx lobes ca. 1.5 mm long, ovary 
2-3 mm long; flowers fasciculate (inflorescences rarely pedunculate), usually more than 

1 5/node; fruit elongate when dried H. psychotriifolia 

lOb. Corolla less than 9 mm long, corolla lobes 1.5-4 mm long; calyx lobes 0.5-1 mm long, 
ovary 1-2 mm long; flowers fasciculate or borne on slender pedunculate; fruit rounded 

when dried 11 

1 la. Inflorescences compact, fasciculate or with short stiff peduncles, often less than 2 cm 
long; leaf blades to 1 5 cm long, rarely tapering gradually at the base and decurrent on 

the petiole; a commonly collected species H. longipetiolata 

lib. Inflorescences open with thin peduncles and pedicels, usually more than 2 cm long; 
leaf blades to 23 cm long, usually tapering gradually at the base and decurrent on the 

petiole; uncommon plants H. hamelioides 

12a. Sepal lobes 2-4 mm long; corolla tube 1-3 mm long, corolla lobes 3-7 mm long; plants not 
collected from about 700 m elevation [leaf blades usually with an arcuate submarginal vein 

and abruptly narrowed at the base] H. liesneriana 

12b. Sepal lobes rarely more than 1 .8 mm long; corolla tube usually more than 2 mm long (except 
in H. laxa), corolla lobes 2-6 mm long; plants rarely found below 800 m elevation (except 

H. pallidiflord) 13 

1 3a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein; in- 
florescences with a prominent erect peduncle 4-10 cm long and usually terminated by 3 

branches; 1200-1500 m elevation H. davidsoniae 

13b. Secondary veins loop-connected only in the distal part of the lamina, not forming an arcuate 
submarginal vein; inflorescences often pendant, with shorter slender peduncles (400-)800- 

2300 m elevation -I 4 

14a. Corolla tubes 4-8 mm long; inflorescences short to long and with many lateral branches; leaf 
blades drying stiffly chartaceous and often large, 12-37 cm long with 9-15 pairs of 2 veins; 
1000-2200 m elevation ! 5 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 167 



14b. Corolla tubes 1 .5-4 mm long; inflorescences short (to 6 cm) with few lateral branches [corolla 
tube 1.54 mm long]; leaf blades drying membranaceous to thin-chartaceous, 7-20(-25) cm 

long with 7-10 pairs of 2 veins; (400-)800-1 500 m elevation 17 

1 5a. Leaves usually drying grayish and stiffly chartaceous, petioles 2-10 cm long; leaf blades 
cuneate and slightly decurrent on the petiole; (corolla tubes 4-8 mm long); fruit rose- 
red to white; central Costa Rica H. leucocarpa 

1 5b. Leaves usually drying dark brown and chartaceous, petioles 2-5 cm long; leaf blades 
usually cuneate and long-decurrent at the base; fruit red; central Costa Rica to the 

Chiriqui Highlands (note: the following 2 species may intergrade) 16 

16a. Corolla tubes 6-10 mm long; calyx lobes 1.2-3 mm long; leaf blades often narrowly 
obovate and long attenuate at the base; central Costa Rica to Panama, 1000-2500 m 

elevation H. arborescens 

16b. Corolla tubes 3-7 mm long; calyx lobes 0.5-1.5 mm long; leaf blades not usually 
narrowly obovate and long attenuate at the base; westernmost Costa Rica and the 

Chiriqui Highlands, 1 100-1 700 m elevation H. pittieri 

1 7a. Inflorescences usually unbranched (sometimes bifurcate) [drying yellowish and not very slen- 
der, probably erect; pedicels 0-3 mm long and ca. 0.5 mm thick (dried); corolla tube 2.54 
mm long; 400-1300 m elevation on the Caribbean slope] H. pallidiflora 

1 7b. Inflorescences usually branched 18 

18a. Inflorescences with very thin (0.2-0.3 mm) branches that usually dry black, pendulous, 
flowering portion 24 cm long, pedicels 4-9 mm long and 0.2-0.3 mm thick (dried); 1000- 

1 500 m elevation H. laxa 

18b. Inflorescences with thicker branches that do not dry black, flowering portion 1-2 cm long, 

pedicels 2-6 mm long, not thin and black; 0-200 m elevation, Osa Peninsula 

H. hammelii 

1 9a. (from 7b) Leaf blades rounded to bluntly obtuse at the apex, often bullate, many 4 veins parallel 
(sublineolate but difficult to see); inflorescences with few (3-7) subsessile flowers on long (3-7 cm) 

peduncles; plants to 40 cm tall [200-1200 m elevation] H. bullata 

1 9b. Leaf blades acute to acuminate (rarely obtuse) at the apex, never bullate, 4 veins partly parallel 
only in H. aeruginosa; inflorescences with short (0-2 cm) peduncles or with more than 7 flowers 

if peduncles are long, pedicels 0-2 cm long; plants usually more than 40 cm tall 20 

20a. Secondary veins loop-connected near the margin to form an arcuate submarginal vein [inflores- 
cences subsessile and usually less than 2 cm long] 21 

20b. Secondary veins loop-connected only in the distal part of the blade, a submarginal vein absent 

(except in the distal third) 22 

2 la. Leaf blades cuneate and slightly decurrent at the base, with 9-1 5 pairs of 2 veins; 400-1000 

m elevation H. inamoena 

21b. Leaf blades acute to obtuse at the base, not usually decurrent on the petiole, with 7-12 pairs 

of 2 veins; 1000-1500 m elevation in north-central Costa Rica H. aeruginosa 

22a. Plants not found above 900 m elevation; leaf blades usually less than 15 cm long, rarely cuneate 

and long-decurrent at the base, with 6-10 pairs of 2 veins H. valerii 

22b. Plants not found below 1 000 m elevation; leaf blades often exceeding 1 5 cm in length, usually 

gradually cuneate and long-decurrent on the petioles, with 7-12 pairs of 2 veins 23 

23a. Inflorescences with thick densely villous peduncles and pedicels; calyx lobes 1.5-3 mm long; corolla 

tube 6-8 mm long (lobes 3-4.5 mm long) H. asclepiadea 

23b. Inflorescences with slender glabrescent or sparsely puberulent peduncles and pedicels; calyx lobes 
0.3-1.5 mm long; corolla tubes 1.5-4 mm long (note: the following 2 species may be synonymous) 

24 

24a. Inflorescences short but open and few-flowered; corolla lobes mostly 24 mm long 

H. hamelioides 

24b. Inflorescences short but crowded and the flowers clustered around the node; corolla lobes 3-7 mm 
long H. decurrens 

168 FIELDIANA: BOTANY 



Hoffmannia aeruginosa Stand!. , Publ. Field Mus. 
Nat. Hist., Hot. Ser. 18: 1313. 1938. Figure 9. 

Herbaceous subshrubs or slender shrubs, 0.5-2.5(-4) 
m tall, with few branches, leafy stems 2-6 mm thick, 
quadrangular or rounded, densely pilose with reddish 
brown crooked hairs 0.2-0.7 mm long; stipules 1-2 mm 
long, 2-3 mm broad at the base, broadly triangular, vil- 
lous, persisting. Leaves with petioles (10-)20-70 mm 
long, 0.8-2 mm thick, densely villous, often with petioles 
of the same node differing in length; leaf blades (6-)9- 
20 cm long, (3-)4-10 cm broad, elliptic-oblong to ovate- 
elliptic or broadly elliptic, apex acuminate or short-acu- 
minate to acute, base obtuse to acute (not or only slightly 
decurrent on the petiole); drying thin-chartaceous, gray- 
ish or grayish brown above, glabrous above, puberulent 
beneath with short (0.1-0.2 mm) hairs on 2 veins and 
longer (0.5-2 mm) hairs on the midvein, 2 veins 7-12/ 
side and evenly loop-connected 2-4 mm from the mar- 
gin, many 4 veins parallel (forming rectangular areolae). 
Inflorescences solitary and axillary (2/node), 1-2 cm long 
with 2-10 flowers, glomerulate, sessile or on short (3 
mm) peduncles, unbranched, flowers sessile or subses- 
sile. Flowers densely covered with curly-crooked hairs 
0.3-0.5 mm long, dark reddish brown when dried, hy- 
panthium 2.5-3 mm long, 2 mm diam. distally, obconic, 
calyx lobes 1 .3-2.5 mm long, ca. 0.6 mm wide, narrowly 
oblong; corolla rotate, reddish, tube 1-2 mm long, lobes 
4-6 mm long, 2-3 mm broad; anthers ca. 2.5 mm long; 
style and stigma white. Fruits not seen. 

Plants of wet cloud forest formations on the 
Caribbean slope from (?100-)800 to 1600 m ele- 
vation. Flowering in April-August. The species is 
known from north-central Costa Rica, near Ciu- 
dad Quesada and Zarcero, and from western Pan- 
ama. 

Hoffmannia aeruginosa is recognized by the 
dense reddish (in life) multicellular hairs on all 
parts, long-petiolate leaves (with blades scarcely 
decurrent), small dense inflorescences, and re- 
stricted range. The petioles often unequal at a node, 
the subparallel minor venation and arcuate sub- 
marginal vein are also distinctive features. Com- 
pare this species with material placed under H. 
inamoena with less dense pubescence, especially 
on the petioles. 



Hoffmannia attinis Hemsley, Diagn. PI. Nov. 31. 
1879. 

TYPE Endres 150, without locality. 

Branches terete, puberulent when young. Leaves 
with petioles ca. 6 mm long; leaf blades 10-12.5 
cm long, ovate-oblong, obtusely acuminate, atten- 
uate at the base, minutely puberulent beneath. In- 



florescences umbellate-cymose, fasciculate, with 
ca. 6 flowers, peduncles 8-16 mm long, slender, 
pedicels 2-4 mm long. Flowers 6-8 mm long, pu- 
berulent. Based on Standley (1938), who had only 
seen the description. 



Hoffmannia amplexifolia Standl., J. Wash. Acad. 
Sci. 15: 8. 1925. Figure 8. 

Herbs or subshrubs, l-2(-3) m tall, leafy stems 3-10 
mm thick, glabrous or minutely puberulent on the ridges 
in early stages, with 3 longitudinal ridges; stipules 1-2 
mm long and 2 mm broad at the base, triangular, pu- 
berulent along the edge. Leaves 3 (4)/node, petioles ab- 
sent or short (3 mm) when the decurrent leaf base is 
reduced to a lateral ridge, minutely puberulent beneath; 
leaf blades (11-) 16-37 cm long, 3-13 cm broad, nar- 
rowly elliptic to oblanceolate or very narrowly elliptic, 
gradually narrowed to the long-acuminate apex, gradu- 
ally narrowed to the cuneate and long-decurrent base 
with a margin 2-4 mm wide along the petiole, often 
expanded (to 6 mm) and auriculate at the base, leaves 
drying chartaceous and dark brown to grayish above, 
glabrescent above, minutely (0.1-0.2 mm) puberulent 
below, 2 veins 11-1 8(-22)/side, usually loop-connected 
1-4 mm from the leaf edge. Inflorescences axillary to 
leaves, 2-12/node, to 10 cm long, with short (3-15 mm) 
or long (to 60 mm), thick (1.5-2 mm) peduncles and 
long (to 7 cm) slender lateral branches from the apex of 
the peduncle, minutely puberulent, pedicels 0-5 mm long 
with hairs 0. 1-0.4 mm long. Flowers sparsely to densely 
puberulent with minute crooked hairs that dry reddish, 
hypanthium 2-3 mm long, 1.5-2 m diam. distally, tur- 
binate to obconic, calyx reddish, calyx cup ca. 0.7 mm 
deep, calyx lobes 1-2.5 mm long and 0.5-1 mm broad, 
triangular to oblong; corolla rotate, greenish white to pale 
yellow, tube 1-4 mm long, 1-2 mm diam., lobes 4-9 
mm long, 1.5-3 mm broad; anthers 3-5 mm long and 
0.7-1.5 mm broad, connivent, yellow. Fruits becoming 
pink or red, to 8 mm diam. in life (4-5 mm long dried), 
ellipsoid to subglobose. 



Plants of the very wet forests of the Caribbean 
slope, 800-2000 m elevation. Flowering in March- 
July; fruiting in October-December and March. 
The species is known only from central Costa Rica 
(Vara Blanca de Sarapiqui to Tapanti). 

Hoffmannia amplexifolia is recognized by the 
nodes with three leaves, long-decurrent leaf base 
forming a leafy lateral margin along the petiole 
and often slightly expanded (auriculate) at the base 
(appearing to be amplexicaul), distinctive puber- 
ulence (but not on the stems), much-branched in- 
florescences, and broad connivent anthers. Al- 
meda el al. 6665 (CAS, F) from near Las Alturas, 
Puntarenas, is tentatively placed here; it has very 
pubescent inflorescences with corolla tubes 5-6 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



169 



mm long. Pressed specimens can be similar to H. 
subauriculata with two leaves per node. 



Hoffmannia arborescens J. D. Smith, Bot. Gaz. 
37: 417.1 904. H.josefina Standl., J. Wash. Acad. 
Sci. 15: 8. 1925. (?//. steinworthii Fosberg, Sida 
2: 388. 1966.) Figure 10. 

Herbaceous subshrubs, 1.5-2 m tall, leafy stems 2-6 
mm thick when dried, glabrescent; stipules ca. 2 mm 
long. Leaves with petioles 3-8(-25) mm long; leaf blades 
7-18 cm long, 2.5-7 cm broad, oblanceolate to narrowly 
obovate, base acute to attenuate and long-decurrent on 
petiole, glabrous above and below, drying stiffly char- 
taceous, often dark reddish brown above, 2 veins 6- 127 
side. Inflorescences 1-3 in each leaf axil (2-6/node), 3- 
5 cm long, to 4 cm broad, open cymose with many 
branches, peduncles 3-20 mm long, puberulent with stiff 
hairs, flowers crowded distally, pedicels 2-6 mm long. 
Flowers with hypanthium 2-3 mm long, ca. 2 mm diam., 
calyx lobes 1-3 mm long, 1 mm broad at the base, blunt 
at apex; corolla funnelform, white to yellow or flushed 
with red, glabrous or puberulent, tube 6-8(-l 0) mm long, 
1-2 mm diam., lobes 3-6 mm long. Fruits ca. 8 x 4.5 
mm, red. 



In evergreen montane forests from 1 600 to 2500 
m elevation. Flowering in February, April-Au- 
gust, and November-December; fruits were col- 
lected in February (immature fruits in August). 
This species ranges from Costa Rica to western 
Panama. 

Hoffmannia arborescens is recognized by the 
usually narrowly obovate leaves gradually long- 
attenuate at the base, flowers on short inflores- 
cences in leaf axils (or below), larger flowers with 
prominent calyx lobes, long corollas, and the high- 
er altitude range. Occasional individual plants may 
be puberulent (Q. Jimenez 231 CR). This species 
may resemble H. hamelioides, while large-leaved 
specimens may be confused with the more com- 
mon and closely related H. leucocarpa. The dif- 
ferences between H. arborescens and H. pittieri are 
minor, and the two species may intergrade in the 
highlands of western Panama, where H. pittier is 
common and distinctive. 

This species was described from material orig- 
inating from Sta. Rosa del Copey at ca. 1 800 m 
elevation (Tonduz 12230 = J. D. Smith n. 8121 
us holotype). An isotype labeled as H. arborescens 
at CR lacks any indication of having the leaf-like 
floral bracts 1 cm long mentioned in the original 
description. This may have been an error on the 
part of J. D. Smith, as no such bracts have been 
seen in any Costa Rican material of Hoffmannia. 
The type bears the same Herb. Nat. C.R. number 



(12230) as does the type of H. decurrens Standl. 
(q.v.), and they probably represent a mixed col- 
lection of two different species. 



Hoffmannia areolata Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 22: 52. 1940. Figure 9. 

Herbaceous subshrubs or slender stemmed shrublets, 
0.6-1 .5(-2) m tall, leafy stems 2-9 mm thick, glabrescent 
or sparsely pilose with crooked hairs 0.3-0.7 mm long; 
stipules 3-6 mm long and 2-4 mm broad at the base, 
glabrous, ligulate, caducous. Leaves with petioles 3- 
40(-90) mm long, difficult to distinguish from the de- 
current lamina base; leaf blades (6-)8-20(-30) cm long, 
3-8(-12) cm broad, elliptic to ovate-elliptic or oblan- 
ceolate, apex tapering gradually and acute or short-acu- 
minate, more abruptly narrowed to an obtuse or cuneate 
base and long-decurrent on petiole, drying membrana- 
ceous to stiffly chartaceous, dark greenish above, the 
surface distinctly bullate and with short (0.2-0.5 mm) 
scabrous hairs above, with thinner hairs ca. 0.3 mm long 
beneath, 2 veins 1 1-16/side. Inflorescences verticillate 
in leaf axils, glomerulate, to 2 cm long and 8-15 mm 
broad, sessile, often obscuring the node, with pink hairs, 
the flowers sessile or subsessile. Flowers closely con- 
gested within the inflorescence, reddish tomentulous, ca- 
lyx lobes 3-5 mm long and 0.5 mm broad, greenish to 
magenta; corolla funnelform, white, rose or translucent, 
ca. 4.5 mm long; tube ca. 2.5 mm long with lobes ca. 2 
mm long; anthers ca. 1.5 mm long. Fruits 4-8 mm long, 
3-6 mm diam. (to 10 mm in life), ellipsoid to subglobose, 
rose to deep red, spongy. 

A little-collected species of montane cloud forest 
formation, 900-2100 m elevation. Flowering in 
March-December; fruiting in April, July-Septem- 
ber, and January. The species ranges from the Cor- 
dillera de Tilaran eastward to the Chiriqui High- 
lands of Panama. 

Hoffmannia areolata is recognized by its rugose- 
bullate leaves (areolate beneath), densely flowered 
axillary sessile glomerulate (usually verticillate) in- 
florescences, very small flowers, and hairs with 
thickened bases. The leaves are very dark green 
above in life. In general aspect, these plants re- 
semble H. congesta and some Psychotria spp. with 
axillary inflorescences. 



Hoffmannia asclepiadea Standl., J. Wash. Acad. 
Sci. 15: 7. 1925. Figure 10. 

Slender herbaceous shrubs, 1.5-2(-3) m tall, usually 
unbranched, leafy stems 2.5-7 mm thick, sparsely to 
densely pilose with crooked hairs 0.3-0.5 mm long (?rarely 
glabrous as in Williams el al. 28096 CR, F); stipules 2-4 
mm long and 3 mm broad at the base, triangular, pu- 
berulent, caducous. Leaves with petioles 1.5-5 cm long, 



170 



FIELDIANA: BOTANY 



1-3 mm thick, densely to sparsely puberulent; leaf blades 
6-21 cm long, 4-9(-ll) cm broad, elliptic to elliptic- 
oblong or ovate-elliptic, apex acuminate with tip 1-2 cm 
long, base rounded to obtuse to cuneate-decurrent, dry- 
ing chartaceous, dark brown above (much paler be- 
neath), glabrous or sparsely pubescent above with hairs 
to 0.7 mm long, more densely pubescent beneath with 
crooked reddish hairs, 2 veins 9-12/side, often loop- 
connected near the margin. Inflorescences 2/node, open 
cymose, 2-4 cm long, with many closely spaced flowers 
on pubescent peduncles 3-10 mm long and ca. 0.7 mm 
thick, usually with 3 major branches and small (0.6 mm) 
bracteoles, pedicels 1-3 mm long, short villous. Flowers 
pubescent, with 5 distal longitudinal lines of hairs on the 
unopened buds, hypanthium 1-1.5 mm long, calyx lobes 
0.5-1 mm long, narrowly triangular; corolla funnelform, 
pale yellow, tube 6-8 mm long, ca. 1.5 mm diam., lobes 
2-3(-4-5?) mm long, to 2 mm broad. Fruits said to be 
subglobose and red. 

Plants of wet cloud forest formations from 1 900 
to 2300 m elevation. Flowering in April-July. The 
species is known only from along the Cordillera 
Central and western portion of the Cordillera de 
Talamanaca in central Costa Rica. 

Hoffmannia asclepiadea is recognized by the 
dense puberulence on leaves and inflorescences, 
compact many-flowered inflorescences on very 
short peduncles, longer corolla tubes, and restric- 
tion to higher-elevation cloud forests. This species 
may be no more than a very puberulent form of 
another species, such as H. arborescens. 



6 mm long and 2 mm broad; anthers ca. 3.2 mm long. 
Fruits becoming red or orange-red, 6-10 mm long and 
4-9 mm diam., subglobose but with a truncated apex 
and persisting calyx. 

Plants of wet evergreen forest formations from 
50 to 1200 m elevation in Costa Rica. Sometimes 
forming a ground cover under the deep shade of 
the forest. Flowering in April-August; fruiting Jan- 
uary-July in Costa Rica. This species ranges from 
Veracruz, Mexico, to central Panama. 

Hoffmannia bullata is recognized by its smail 
stature with roots at lower nodes, prominently bul- 
late leaves that are obtuse or rounded apically, 
few-flowered inflorescences with long peduncles, 
and puberulence on many parts. The sublineolate 
4 venation is distinctive but difficult to see in 
some collections. The midvein is often white on 
the dark green upper surface in living material. 
This species displays much variation within its 
range and within Costa Rica. Collections from the 
Pacific slope and near the Panama border tend to 
have smaller stature and narrowly elliptic-oblong 
leaves with more deeply impressed venation and 
acute apices (fig. 9, uppermost left). These differ- 
ences are worthy of more detailed study. In ad- 
dition, H. discolor (Lemaire) Hemsl. of Veracruz, 
Mexico, is closely related, but the leaves dry darker, 
the flowers have longer (2-4 mm) linear calyx lobes, 
and the fruits are larger. 



Hoffmannia bullata L. O. Williams, Fieldiana Bot. 
36: 52. 1973. Figure 9. 

Herbs or herbaceous subshrubs, 10-60 cm tall, erect 
or decumbent, stems unbranched, lower nodes with roots, 
leafy stems l-4(-7) mm thick, densely pilosulous with 
curved hairs 0.1-0.3 mm long drying reddish brown, 
terete; stipules 2-3 mm long, triangular, deciduous. Leaves 
opposite, petioles 4-45(-55) mm long, 1-2.5 mm thick, 
pilosulous with curved or crooked multicellular hairs; 
leaf blades 4-22 cm long, 2-9(-12) cm broad, obovate 
to oblong-obovate, apex rounded to bluntly obtuse, base 
obtuse to cuneate (not or only slightly decurrent on pet- 
iole), drying chartaceous, grayish or grayish brown above, 
glabrous above, minutely puberulent on the veins be- 
neath with curved reddish hairs ca. 0.2 mm long, 2 veins 
5-12/side, many 4 veins subparallel (sublineolate) and 
demarking narrow rectangular areas on the upper dried 
surface (but often difficult to see). Inflorescences usually 
solitary in leaf axils ( 1 -2/node), 3-8 cm long, glomerulate 
or subumbellate, peduncles 2-7 cm long, 0.5-1 mm thick, 
glabrous or sparsely pilose, with 3-9 flowers along the 
same side of bifurcate branches, pedicels 1-5 mm long. 
Flowers sparsely puberulent or subglabrous, hypanthium 
2-3 mm long, 1.5-2 mm broad distally, urceolate or 
turbinate, red, calyx lobes 1-2.5 mm long, 0.4 mm broad 
and acute; corolla rotate, pale red or rose, 7-10 mm long 
in bud, tube 3-4 mm long and 0.8 mm diam., lobes 3- 



Hoffmannia congesta (Oerst.) Dwyer, comb. nov. 
Xerococcus congestus Oerst., Vidensk. Meddel. 
Dansk Naturhist. Foren. Kjobenhavn 1852: 52. 
1853. Figure 9. 

Herbs or subshrubs, 0.5-2(-3) m tall, stem erect and 
usually unbranched, terete, leafy stems 3-15 mm thick, 
glabrous or with a few large (1 mm) crooked hairs; stip- 
ules 6-14(-20) mm long, 4-6 mm broad at the base, 
triangular-ovate, with or without an awn, glabrous, co- 
riaceous, deciduous. Leaves with petioles 2-8(-15) cm 
long, 1.2-3 mm broad, glabrous or with crooked mul- 
ticellular hairs to 2 mm long; leaf blades 15-30MO) cm 
long, 8-15(-25) cm broad, broadly elliptic to broadly 
ovate-elliptic or elliptic-obovate, apex short-acuminate, 
somewhat rounded to abruptly narrowed at the obtuse 
to acute base and slightly decurrent on petiole (rarely 
long-decurrent or sometimes rounded to a truncated base 
in large leaves), drying stiffly chartaceous, dark brown 
or dark grayish above, glabrous or with scattered crooked 
hairs, 2 veins 7-14(-17)/side. Inflorescences solitary in 
leaf axils (2/node), sessile and usually glomerulate, many- 
flowered, 1-2 cm long and to 6 cm broad (across the 
node), subglobose to verticillate, reddish, pedicels 0-2.5 
mm long. Flowers tightly congested, sparsely and mi- 
nutely puberulent, hypanthium 3 mm long and 1-2 mm 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



171 



diam.. calyx red, calyx lobes 3-6 mm long, 1-1.5 mm 
broad, obtuse at the tip, with parallel venation; corolla 
salverform, white, tube 2-3.5 mm long, lobes 2-3.5 mm 
long and 1 mm broad. Fruit becoming white and spongy, 
ca. 5 x 3 mm when dried, oblong. 

Understory plants of the wet Caribbean slopes 
from 700 to 1800(-2400?) m elevation. Flowering 
in January-October; fruiting in October-March. 
The species ranges from the Cordillera de Tilaran 
eastward to Veraguas, Panama. 

Hoffmannia congesta is characterized by its ses- 
sile (often verticillate) glomerulate reddish inflo- 
rescences, long calyx lobes, white fruit, un- 
branched stems, large leaves, and prominent 
stipules. This is one of our most commonly col- 
lected and most distinctive species of Hoffmannia. 
In the past it was assigned to its own genus (Xero- 
coccus); specimens are often found in the uniden- 
tified section of Rubiaceae collections. 



Hoffmannia davidsoniae Standl., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 22: 53. 1946. (?= H. lan- 
cistigma Dwyer, Ann. Missouri Bot. Gard. 56: 
279. 1969). Figure 7. 

Herbs or slender subshrubs, 0.5-1 .7 m tall, stems usu- 
ally unbranched, leafy stems 1.5-5 mm thick, glabrescent 
(rarely densely puberulent, cf. Davidse et al. 29175 CR); 
stipules 1.5-3 mm long, ca. 1 mm broad at the base, 
narrowly triangular, caducous. Leaves with petioles 1-7 
cm long, 0.8-2 mm thick, glabrous; leaf blades 8-17 cm 
long, 3-9(-12) cm broad, broadly oblong to elliptic-ob- 
long or ovate-oblong (rarely lanceolate), apex narrowed 
abruptly and short-acuminate, base obtuse to acute and 
slightly decurrent on petiole, drying grayish to dark brown 
or dark grayish green above, glabrous above and below 
or minutely puberulent, 2 veins 10-14/side, sometimes 
arising from the midvein at nearly 90, loop-connected 
distally and forming an arcuate submarginal vein 1-4 
mm from the edge. Inflorescences borne on lower leafless 
nodes, 1-2/node, 5-15 cm long, paniculate and with a 
prominent peduncle 4-6(-12) cm long and 1-2.3 mm 
thick, glabrous, with 3(-4) distal branches and each branch 
with 7-17 flowers, pedicels 2-4(-6) mm long. Flowers 
glabrous, hypanthium 2-3 mm long, ca. 1 mm diam. 
distally, turbinate, calyx lobes 0.3-1 mm long, triangular; 
corolla rotate to fimnelform, yellow or flushed with pur- 
ple, tube 1-4 mm long, lobes 2-4(-6) mm long; anthers 
to 3.8 mm long. Fruit ca. 8 mm long and 4 mm diam., 
ellipsoid to subglobose, becoming red. 

Plants of wet cloud forest formations at 700- 
2300 m elevation. Flowering in January, April, 
July, and September; fruiting in September and 
December-January. The species ranges from cen- 
tral Costa Rica eastward to the Chiriqui High- 
lands. 



Hoffmannia davidsoniae is a little-collected spe- 
cies characterized by the long stiff peduncles usu- 
ally terminated by three branches and the long- 
petiolate leaves with an arcuate submarginal vein. 
This species is poorly known in Costa Rica, and 
there appears to be great variation from collection 
to collection. The secondary veins often arise at 
nearly 90 from the midvein, but there are collec- 
tions in which this is not the case. This species is 
closely related to H. liesneriana. 



Hoffmannia decurrens Standl., Contr. U.S. Natl. 
Herb. 20:205. 1919. 

Slender shrubs or subshrubs, 1.5-3 m tall, leafy stems 
1.5-4 mm thick, densely puberulent with crooked yel- 
lowish brown or reddish brown hairs 0.1-0.3 mm long; 
stipules 1.5-3 mm long, ca. 2 mm broad at the base, 
triangular, hirstulous, deciduous. Leaves with petioles 
5-40 mm long, ca. 1 mm thick, difficult to distinguish 
from the lamina base, densely hirsutulous with hairs 0.3- 
0.5 mm long; leaf blades 7-21 cm long, 2-7 cm broad, 
elliptic-obovate to oblanceolate or elliptic, apex acu- 
minate, gradually narrowed to a narrowly cuneate base 
and long-decurrent on petiole, drying membranaceous 
or thin-chartaceous and dark greenish brown or grayish 
green above, glabrous on the upper surface, pubescent 
on the veins beneath with hairs 0.2-0.7 mm long, 2 
veins 7-10/side. Inflorescences 1-3/axil, l-2(-3)cm long, 
cymose or fasciculate with few (1-5) flowers or rarely 
2-branched and scorpioid with 5-10 flowers, peduncles 
5-10(-24) mm long, 0.2-0.3 mm thick, puberulent, ped- 
icels 2-6 mm long, slender. Flowers sparsely to con- 
spicuously pubescent, hypanthium 0.7-1 .3(-2) mm long, 
0.6-1(-1.7) mm diam. distally, turbinate, calyx lobes 
0.7-1.5 mm long, triangular; corolla yellow or whitish, 
tube 2-4 mm long, lobes 3-7 mm long, ca. 2 mm wide 
at the base. Fruits becoming 8-9 mm long, 5-6 mm 
diam. (dried), ovoid to globose, red, borne on slender 
peduncles and pedicels. 

Plants of montane evergreen forest formations, 
1 100-2000 m elevation. Flowering in April-June 
and December; fruiting in December-February. 
This poorly characterized species is endemic to 
Costa Rica and presently known to range from 
Monteverde to Sta. Maria de Dota. 

Hoffmannia decurrens is recognized by the pu- 
berulent young stems, narrowly obovate leaves with 
cuneate long-decurrent base, and very short inflo- 
rescences with very slender peduncle and pedicels. 
This species is poorly characterized at present, with 
a diverse array of specimens placed here. This 
species may include material presently assigned to 
H. hamelioides. The type, Tonduz 12230 us, bears 
the same collection number as the type of H. ar- 
borescens; these types (not seen) probably repre- 
sent a mixed collection. 



172 



FIELDIANA: BOTANY 



Hoffmannia dotae Standl., J. Wash. Acad. Sci. 18: 
181. 1928. H. ramonensis Standl., loc. cit. 180. 
1928. Figure 10. 

Single-stemmed shrubs or herbaceous subshrubs, 
(0.5-)l-3(-6?) m tall, leafy stems 2-10 mm thick, gla- 
brous to pilosulous, older stems terete and hollow, woody; 
stipules 3-4 mm long and 4 mm broad at the base, 
glabrous, deciduous. Leaves 2 (rarely 3) at a node, pet- 
ioles 4-20 mm long (to 5 cm if including the decurrent 
lamina base), ca. 1.5 mm broad, glabrous; leaf blades 
1 5-30(-35) cm long, (4-) 7-1 2 cm broad, elliptic-obovate 
to obovate-oblong or oblanceolate, apex short-acumi- 
nate, tapering gradually or abruptly to the cuneate and 
long-decurrent base, the narrowed basal part of the lam- 
ina to 6 cm long, drying membranaceous to thin-char- 
taceous, usually dark brown or gray above, glabrous 
above, pubescent in very early stages beneath and be- 
coming glabrous or with reddish brown hairs 0.1-0.4 
mm long along the major veins, 2 veins 9-15/side and 
often loop-connected near the edge. Inflorescences ax- 
illary or at older leafless nodes, l-3/axil(2-6/node), 1.5- 
3(-5) cm long, ca. 1 5 mm broad, usually paniculate with 

3 primary branches and 4-9 flowers, peduncles 0-5 mm 
long (to 35 mm when the primary peduncle is reduced 
and the 1 branches function as peduncles), minutely 
puberulent, pedicels 3-6 mm long. Flowers conspicu- 
ously (0. 1-0.3 mm) puberulent or glabrous, hypanthium 
3-4 mm long, 2-3 mm diam. distally, obconic and red- 
dish, calyx tube ca. 0.5 mm long and rotate, lobes (l-)2- 

4 mm long, 1-2 mm broad at the base, bluntly triangular 
to oblong, often held horizontally; corolla rotate, usually 
red or orange at the base and orange or yellowish distally, 
fleshy and glabrous, buds ca. 10 mm long, tube 2-5 mm 
long, 2.5-4 mm diam., lobes 4-9 mm long, 2-3 mm 
broad, yellowish within. Fruits 6-8 mm long and 4-6 
mm diam. (dried), ca. 8 mm diam. in life, orange or 
reddish. 

Plants of wet montane cloud forest formations, 
from 850 to 2100 m elevation. Flowering in Feb- 
ruary-September; fruiting in September-January. 
It is uncommon in Costa Rica, except for the San 
Vito area. This species ranges from the Cordillera 
de Tilaran to the Chiriqui Highlands of Panama. 

Hoffmannia dotae is recognized by its larger leaf 
blades with long-decurrent base forming narrowly 
winged margins along the sides of the petiole (but 
not auriculate at the base), the often cauliflorus 
few-flowered inflorescences, and the short thick 
corolla tube with larger stiff yellow petals. The 
flower buds with their short conical corolla tubes 
and broad (3-4 mm) base distinguish this species 
from all other Costa Rican species, none of which 
have such thick corolla tubes. Compare H. leu- 
cocarpa with more numerous slender flowers and 
leaves acute to cuneate at the base. 



HofTmannia gesnerioides (Oerst.) Kuntze, Rev. gen. 
pi. 285. 1891. Ophryococcus gesnerioides Oerst., 



Vidensk. Meddel. Dansk. Naturhist. Foren. 
Kjobenhavn 1852: 53. 1852. 

This species ranges from Guatemala to central 
Nicaragua at elevations of 1000-1 500 m. The small 
stature (to 60 cm), long reddish hairs, and few- 
flowered cymes on peduncles that elongate to 3 
cm in fruit help to distinguish this species, which 
is not known from Costa Rica. 



HofTmannia hamelioides Standl., J. Wash. Acad. 
Sci. 15: 8. 1925. Figure 11. 



Herbs or shrubs, (0.5-)1.5-3 m tall, stems often 
branched, leafy stems 1.2-3 mm thick, usually densely 
covered with crooked reddish brown multicellular hairs 
0.3-1 mm long (sometimes glabrescent); stipules 2-3 
mm long, triangular, drying reddish brown. Leaves with 
petioles 6-25 mm long, 0.7-1 .5 mm thick, usually dense- 
ly puberulent; leaf blades 8-23 cm long, 3-8 cm broad, 
oblanceolate-elliptic to narrowly obovate or elliptic-ob- 
long, apex acuminate, base gradually narrowed and acute 
or long-cuneate, decurrent on petiole, drying membra- 
naceous to thin-chartaceous, dark brown to blackish 
brown above, glabrous or pubescent beneath with crook- 
ed hairs 0.2-0.7(-1) mm long, 2 veins 7-12/side. Inflo- 
rescences axillary or at older nodes, solitary (1-2/node), 
1-3 cm long, usually on short (1-10 mm) slender pe- 
duncles, with 3-7 flowers, pedicels 0.7-1.5 mm long, 
0.2-0.4 mm thick, minutely (0.1-0.2 mm) puberulent. 
Flowers with short (0.2-0.7 mm) crooked hairs, buds ca. 
8 mm long, hypanthium ca. 2 mm long and 1.5-2 mm 
diam., calyx lobes 0.7-1.5 mm long, 0.3-0.6 mm broad, 
narrowly oblong to linear; corolla yellowish white or 
yellow, tube 1.5-2 mm long, 1.5-2 mm diam., lobes 2- 
4 mm long, 1.5-2 mm broad, apex acute, becoming re- 
flexed. Fruits 4-8 mm long, 3-6 mm diam. (dried) red- 
dish at maturity, with persisting calyx lobes. 



Plants of evergreen forests in the central high- 
lands, from 450 to 2200 m elevation. Flowering 
in February and May-July; fruiting in July and 
December. This species is known only from the 
Central Volcanic highlands and the Cordillera de 
Talamanca in Costa Rica. 

Hoffmannia hamelioides is recognized by the 
usually puberulent young stems, narrowly oblan- 
ceolate leaves gradually tapering to the long-at- 
tenuate base on slender petioles, small inflores- 
cences with filamentous peduncles and pedicels, 
and short corolla tubes. Compare this species with 
H. decurrens (perhaps synonymous) and H. valerii 
with cuneate to truncated leaf bases and lower 
elevation habitat. Glabrous individuals resemble 
a number of species, especially H. longepetiolata. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



173 



Hoffmannia hammelii C. M. Taylor, sp. nov. 

Species Hoffmanniae laxae similis, sed ab ea stipulis 
brevioribus (0.7-1.5 mm longis) ac cymis secundis dif- 
fert; etiam H. pallidiflorae similis, sed foliis basi acutis 
ac lobulis corollinis brevioribus (3-4 mm longis) differt. 

TYPUS Liesner2907 (holotypus CR, isotypus MO), from 
near Sirena, Corocovado National Park, 0-200 m alt., 
5 July 1977, Puntarenas, Costa Rica. 

Herbaceous subshrubs, 0.3-0.5(-1) m tall, stems 
branched only near the ground, leafy stems 1.3-3 mm 
thick, glabrous and terete, often drying dark; stipules 0.7- 
1.5 mm long, ca. 1 mm broad at the base, glabrous, 
deciduous. Leaves with petioles 6-27 mm long, 0.7-1.4 
mm broad, glabrous; leaf blades (4-) 7- 18 cm long, 
(1.5->4-7 cm broad, elliptic to elliptic-obovate or ellip- 
tic-oblanceolate, apex acuminate, base acute (not de- 
current), drying stiffly chartaceous, grayish above, usu- 
ally paler or yellowish beneath, glabrous above and below, 
2 veins 7-13/side. Inflorescences solitary and axillary 
to leaves (1-2/node), 2-7 cm long but the flowering por- 
tion only 1-2 cm long, with a single rachis or cymose 
with a terminal flower and 2 lateral branches with 5-12 
flowers each, peduncles 1 5-50 mm long, rachis with 2- 
7(-l 1) flowers along 1 side, glabrous, pedicels 2-6 mm 
long. Flowers with ovary/hypanthium 2 mm long and 
1 .5 mm diam., calyx tube 0.5 mm long, calyx lobes 0.5- 
0.8 mm long, ca. 1 mm broad at the base, triangular; 
corolla rotate, white to yellow-green, tube 2-3 mm long, 
lobes 3-4 mm long. Fruits ca. 5 mm diam., ellipsoid to 
subglobose, red, glabrous. 

Plants of the evergreen Pacific lowlands, from 
near sea level to 200 m elevation. Flowering in 
July-September and November; fruiting in Jan- 
uary. This species is endemic to the Golfo Dulce 
area and the Osa Peninsula. 

Hoffmannia hammelii is recognized by its small 
stature, leaves tapering equally to apex and base, 
small flower groups on long slender peduncles, 
leaves often drying grayish or reddish brown, and 
restricted geographic range. This species appears 
to be part of a species group including H. laxa, H. 
pallidijlora, H. bullata, and H. discolor (Lemaire) 
Hemsl. of Mexico. The species is named in honor 
of Barry Hammel, who has made many important 
contributions to our knowledge of Costa Rica's 
Rubiaceae. Other collections seen were M. Cha- 
varria et al. 254 CR, Kernan 813 & 1249 CR, MO, 
and Knapp 2184 CR, MO. 



Hoffmannia inamoena Standl., J. Wash. Acad. Sci. 
18: 179. 1928. H. fimbrianthera Dwyer, Ann. 
Missouri Bot. Gard. 56: 277. 1969. Figure 11. 

Herbaceous subshrubs. 1-2 m tall, with few branches, 
leafy stems 2.5-5 mm thick, sparsely to densely minutely 
(0.1-0.2 mm) puberulent in early stages but soon gla- 



brous and drying grayish, quadrangular; stipules 2-3 mm 
long. Leaves with petioles 10-60 mm long, 0.8-2 mm 
thick, minutely and sparsely puberulent or glabrous; leaf 
blades 8-22 cm long, 3.5-10 cm broad, elliptic-oblong 
to oblong or obovate, apex acuminate with tip 3-1 5 mm 
long, base obtuse to acute and usually slightly decurrent 
on petiole, drying membranaceous or thin-chartaceous 
and grayish green above, glabrous above, very minutely 
(0.05 mm) puberulent on the veins beneath or glabrous, 
2 veins 9-14/side and loop-connected near the margin. 
Inflorescences axillary or on older leafless nodes (1-2/ 
node), ca. 1 cm long and 1-5 flowers, glomerulate, sessile 
or with peduncles 1-3 mm long, pedicels 1-3 mm long, 
pubescent with hairs 0. 1-0.4 mm long or glabrous. Flow- 
ers with crooked reddish hairs to 0.3 mm long (rarely 
glabrous), hypanthium ca. 2 mm long and 1.7 mm diam. 
distally, urceolate, calyx lobes 1.3-2.3 mm long, nar- 
rowly oblong to narrowly triangular, apex obtuse; corolla 
rotate, deep red, puberulent, tube ca. 1.5 mm long, lobes 
ca. 4 mm long. Fruits 5-7 mm long, subglobose, becom- 
ing white, puberulent. 



Plants of evergreen forest formations from 400 
to 1000 m elevation. Flowering in January and 
May-July; fruiting in June and November-De- 
cember. The species ranges from Bijagua, Alajue- 
la, eastward along the Caribbean slope to Rio Re- 
ventazon, and in the General Valley of the Pacific 
slope. 

Hoffmannia inamoena is recognized by the 
minute puberulence on many parts, secondary 
veins clearly loop-connected near the margin of 
the leaf, very small sessile inflorescences, and rel- 
atively large narrow calyx lobes. The material 
placed here is rather poor and may not represent 
a single species. Holm and Iltis (58 F) noted that 
both flowers and fruit fall easily from the stems. 



Hoffmannia laxa Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 22: 1 16. 1940. Figure 11. 

Herbaceous subshrubs or slender shrublets 1-2 m tall, 
branched only near the base, leafy stems 1-6 mm thick, 
glabrous; stipules 1-3 mm long, triangular, glabrous, de- 
ciduous. Leaves with petioles 1 .5-6(-9) cm long, 0.8-2.2 
mm broad, glabrous; leaf blades 7-18(-21) cm long, 4- 
9.5(-ll) cm broad, elliptic to elliptic-ovate or ovate, 
apex acuminate, tapering gradually or abruptly to the 
cuneate or acute base and slightly decurrent on petiole, 
drying thin-chartaceous, grayish green to dark brownish 
green above, glabrous above and below, 2 veins 7-1 1/ 
side. Inflorescences 1-3 in each leaf axil (2-6/node), 3- 
8 cm long, ca. 2 cm broad, pendant, open paniculate or 
racemose with few (8-15) distant flowers, reddish in life, 
peduncles 10-45 mm long, 0.2-0.8 mm thick, glabrous, 
rachis often elongate with alternate flowers and minute 
(0.5 mm) bracteoles, pedicels 4-9 mm long (to 16 mm 
in fruit), 0.2-0.3 mm thick (dried). Flowers glabrous, 
hypanthium ca. 2 mm long and 1.8 mm diam., subglo- 
bose, calyx lobes 0.2-0.7 mm long, triangular; corolla 



174 



FIELDIANA: BOTANY 



funnelform, white to pale green, tube l-2(-3) mm long, 
lobes 3-5 mm long and 1 mm broad at the base; anthers 
ca. 2 mm long. Fruits to 8 mm long, 4-5 mm diam., 
globose to ellipsoid, greenish white when immature, 
whitish purple or red to bluish black or purple-black at 
maturity, with a small persisting calyx tube; seeds black. 



Plants of evergreen lower montane forest for- 
mation on both the Caribbean and Pacific slopes, 
200-1600 m elevation. Flowering in February- 
June and August-October; fruiting in January and 
March-October. This species is known only from 
Monteverde eastward to the slopes of Volcan Poas 
and in western Panama. 

Hoffmannia laxa is recognized by the often long- 
petiolate somewhat larger leaves, the lack of pu- 
bescence, the open few-flowered inflorescences with 
long thin filamentous (when dried) peduncles and 
pedicels, and the small flowers with very short 
corolla tubes. The name H. capillacea Dwyer has 
been applied incorrectly to material of this species. 



Hoffmannia leucocarpa Stand!.. J. Wash. Acad. 
Sci. 1 5: 9. 1925. //. carpinteraeStendl, N. Amer. 
Fl. 32: 199. 1934, nom. nov. for//, macrophylla 
Standl., J. Wash. Acad. Sci. 15: 9. 1925, non //. 
macrophylla Hemsl. (?= H. trichocalyx Standl., 
J. Wash. Acad. Sci. 18: 181. 1928.) Figure 10. 



Slender shrubs or small treelets, l-2.5(-4) m tall, with 
few or no lateral branches, leafy stems 2.5-8 mm thick, 
glabrous (rarely ferruginous-villous), usually hollow; 
stipules 1.5-4 mm long, 2-4 mm broad at the base, 
triangular, glabrous, deciduous. Leaves with petioles 1.5- 
7(-10)cm long, 1.3-3.8 mm broad, glabrous: leaf blades 
14-24(-35) cm long, 5-12(-15) cm broad, elliptic to 
broadly elliptic, apex tapering abruptly and acuminate, 
tapering more gradually to the cuneate base and slightly 
decurrent on petiole, drying chartaceous to stiffly char- 
taceous, pale greenish brown to dark grayish green above, 
glabrous above and below, 2 veins 11-14/side. Inflo- 
rescences 1-3 /axil below the leaves (2-6/node), 4-8(-l 5) 
cm long, with 1 or 2 orders of branching (rarely with a 
single rachis), often pendant, peduncles 2-5(-8) cm long, 
0.5-2 mm thick, usually terminated by 3 subequal 1 
branches, glabrous or less often minutely puberulent, 
pedicels 2-8(-10) mm long, sparsely to densely puber- 
ulent or villose. Flowers with hypanthium ca. 2 mm long 
and 1.5 mm diam., obconic, puberulent with thin hairs 
ca. 0.2 mm long, calyx green to pink, orange or red, calyx 
tube 1-2 mm long, lobes 0.5-2 mm long, bluntly tri- 
angular; corolla rotate to funnelform, pale pink to yellow 
marked with orange or red, glabrous to villosulous, tube 
4-8 mm long, 1.5-2 mm diam., lobes (2-)3-7 mm long, 
2 mm broad at the base; anthers ca. 4 mm long. Fruits 
8-10 mm long and 7-8 mm diam., globose to obovoid 
(ca. 1 cm diam. in life), often bisulcate in development, 
white to pink or red. 



Plants of the wet Caribbean cloud forests and 
nearby areas, from 1000 to 2300 m elevation. 
Flowering in February-July; fruiting in February, 
April, June-July, and October-December. The 
species ranges from the eastern part of the Cor- 
dillera de Tilaran (Monteverde) to the western 
slopes of Volcan Irazu in central Costa Rica (but 
see below). 

Hoffmannia leucocarpa is recognized by the 
larger firm-textured leaves on long petioles, pe- 
dunculate and branched (usually pendant) inflo- 
rescences, and the larger flowers with corolla tubes 
and lobes usually equal in length. The fruit can be 
white or pink to red. This species differs from 
larger-leaved specimens of H. longepetiolata by 
the more rigid leaf blades with short-acuminate 
apices, larger inflorescences, and larger corollas. 
This species may intergrade with H. arborescens, 
forming a complex that includes H. pittieri. Dried 
specimens can be similar to H. dotae, but that 
species has smaller inflorescences and much 
broader and more succulent corolla tubes and the 
leaves have winged petioles. Most material placed 
here has branched inflorescences, but a few have 
monopodial circinnate inflorescences; this is an- 
other example of the great variability within Hoff- 
mannia species and why determining species in 
the genus is so difficult. 

The preceding circumscription excludes a num- 
ber of specimens from the San Vito area formerly 
placed here. These are treated as unusual elements 
of//, pittieri or H. arborescens. Hoffmannia tricho- 
calyx probably represents an unusually pubescent 
representative of this species; it is based on Stan- 
dley & Torres 47690 (holotype), 47480, 47819. & 
47924, all us. 



Hoffmannia liesneriana L. O. Williams, Fieldiana 
Bot. 36: 54. 1973. Figure 8. 

Herbs or slender subshrubs, 0.3-1(-1.5) m tall, stems 
usually unbranched, quadrangular with opposing flat and 
sulcate surfaces, leafy stems 4-12 mm thick, glabrous 
(strigillose): stipules 1-3 mm long, 3 mm broad at the 
base, triangular, glabrous. Leaves with petioles (2-)6-l 3 
cm long, 1.5-3 mm thick, glabrous; leaf blades (1 1-)13- 
24 cm long, (6-) 7- 13 cm broad, broadly ovate-elliptic 
to broadly elliptic-oblong, apex abruptly narrowed and 
short acuminate, abruptly narrowed to an obtuse or 
rounded base, drying chartaceous, dark grayish or gray- 
ish brown above, glabrous above, sparsely to densely 
minutely (0. 1-0.2 mm) puberulent on the veins beneath, 
2 veins (7-)10-14(-l 7)/side, usually loop-connected near 
the margin. Inflorescences borne at older leafless nodes 
or near the ground, 1-4/node, 2-11 cm long, with many 
flowers in 1 -several dense clusters or on helicoid branch- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



175 



es but rarely expanded, reddish to reddish purple, pe- 
duncles 4-50 mm long, glabrous or sparsely puberulent, 
pedicels 0-4 mm long, slender. Flowers subglabrous or 
puberulent with hairs ca. 0.2 mm long, reddish, hypan- 
thium 3-4 mm long, 2-2.6 mm diam., calyx lobes 2-5 
mm long, ca. 0.4 mm broad; corolla salverform, rose or 
pinkish red to red-brown, tube 1-3 mm long, lobes 3-7 
mm long, ca. 1.3 mm broad; anthers ca. 4 mm long, 
white. Fruits 4-6 mm long, 2-3 mm diam. (dried), red; 
seeds ca. 0.2 mm long. 

Plants of lowland rain forest formations on the 
Caribbean slope, 50-600(-1 100) m elevation. 
Flowering in March and May-November; fruiting 
in the same months and in January. This species 
apparently ranges from Rio San Lorenzo, Alajuela, 
eastward to Guapiles, Limon, in Costa Rica. 

Hoffmannia liesneriana is recognized by the large 
leaves with long petioles, the short unbranched 
stems, flowers in close clusters on cauliflorous in- 
florescences, long calyx lobes, and lower-elevation 
habitat. Unlike so many of our other Hoffmannia 
species, this species has leaves that are often 
abruptly narrowed at the base (but some collec- 
tions do have long-decurrent leaf blades). This spe- 
cies resembles Psychotria capacifolia with white 
flowers. Compare H. davidsoniae and H. eliasii 
Dwyer of Panama. 



Hoffmannia longipetiolata Polak., Linnaea 4 1 : 567. 
1877. H. tonduzii Standl., Contr. U.S. Natl. 
Herb. 20: 205. 1919. H. woodsonii Standl., Ann. 
Missouri Dot. Card. 28: 471. 1941. Figure 11. 

Small shrubs or herbaceous subshrubs, 1-2 m tall, 
usually few-branched and slender-stemmed, leafy stems 
0.8-5 mm thick, glabrous or minutely puberulent along 
narrow longitudinal ridges or with crooked hairs 0.5 mm 
long in early stages; stipules 0.7-3 mm long, 2-3 mm 
broad at the base, triangular, glabrous, deciduous. Leaves 
with petioles 3-15(-40) mm long, 0.5-1 mm broad, gla- 
brous or puberulent; leaf blades 4- 1 2(- 1 5) cm long, 1 . 5- 
4(-6) cm broad, elliptic-obovate to elliptic or ovate-el- 
liptic, apex short- to long-acuminate, tip to 1 5 mm long, 
base acute to cuneate and somewhat decurrent, drying 
membranaceous to thin-chartaceous, brownish green to 
grayish green above, glabrous above, glabrous or mi- 
nutely puberulent on the veins beneath, 2 veins 4-7 (-9)7 
side. Inflorescences 1-3/axil (2-6/node), 1-2 cm long, 
cymes or fascicles with (l-)3-6(-12) flowers, peduncles 
2-5 mm long, pedicels 2-8(-10) mm long, ca. 0.3 mm 
thick, glabrous or rarely sparsely and minutely (0. 1 mm) 
puberulent. Flowers greenish yellow, glabrous, hypan- 
thium 1.5-2 mm long, 1.3 mm diam. distally, obconic, 
calyx lobes 0.5-1 (-2) mm long, ca. 1 mm broad at the 
base, triangular and acute; corolla rostrate, yellow, pink 
or cream white, tube 1.5-3 mm long, 1.3-1.5 mm diam., 
lobes 2-4 mm long, 1 .5-2 mm broad at the base, reflexed; 
anthers 3-3.5 mm long. Fruits 5-6 mm diam., subglo- 
bose, red to pink. 



Plants of wet evergreen forest formations on both 
the Caribbean and Pacific slopes, 600-1 800(-2000) 
m elevation. Flowering in January-October, with 
the majority in May-July; fruiting in April and 
September-February. The species ranges from the 
Cordillera de Guanacaste eastward to central Pan- 
ama. 

Hoffmannia longipetiolata is recognized by the 
smaller or medium-sized leaves that taper grad- 
ually to apex and base, little or no pubescence, 
terete stems, very small few-flowered inflores- 
cences, and small glabrous yellow flowers. The thin 
peduncles and pedicels, short corolla tubes, and 
larger corolla lobes are also helpful in recognizing 
this species. This is one of the most commonly 
collected species of Hoffmannia in Costa Rica. 
Originally it was thought that the smaller-leaved 
specimens determined as H. tonduzii were dis- 
tinct, but there are too many intermediate collec- 
tions to be able to segregate the larger-leaved spec- 
imens (some of which have long petioles). 
Collections from lower (600-900 m) elevations 
and from western Panama often have larger leaves 
than collections from central highland Costa Rica. 
Two species that are very similar to H. longipe- 
tiolata are H. decurrens (with densely puberulent 
young stems) and H. psychotriifolia (with larger 
fasciculate flowers and narrowly ellipsoid fruits). 



Hoffmannia nesiota J. D. Smith, Bot. Gaz. 61: 374. 
1916. 

Shrublets, branchlets subterete, glabrous; stipules not 
seen. Leaves with petioles 7-8 cm long, 2-3 mm thick, 
pubescent; mature leaf blades 18-26 cm long, 10-13 cm 
broad, elliptic to broadly elliptic-oblong, acuminate, base 
obtuse and slightly decurrent, stiffly chartaceous and pu- 
bescent or glabrescent, 2 veins 12-16/side. Inflores- 
cences cymose with peduncles 1-5 cm long, pubescent 
or glabrescent, lacking bracts and bracteoles, pedicles 6- 
10 mm, slender. Flowers 15-16 mm long, with calyx 
lobes 1.5-2 mm long, triangular; corolla 11 mm long, 
rotate, puberulent, tube 5-6 mm long, lobes 5-6 mm 
long, linear; ovary trilocular, style 12 mm long. Fruits 
ca. 1 cm diam.; seeds ellipsoid ca. 0.5 mm long. 

A poorly known species endemic to Cocos Is- 
land. The species is only known from the type 
collection: Pittier 12387 us, collected at Wafer Bay, 
June 1898, and Barclay 2178 us, April 1838. The 
Barclay collection is much more pubescent than 
the type but appears to represent the same species. 
Fruits of the Barclay collection are two-locular, 
not three-locular as described for the type. 

Hoffmannia nicotianifolia (Mart. & Gal.) L. O. 



176 



FIELDIANA: BOTANY 



Williams is a species of Mexico and northern Cen- 
tral America; the name has been incorrectly ap- 
plied to Costa Rican collections. 



Hoffmannia pallidiflora Standl., J. Wash. Acad. 
Sci. 15:9. 1 925. H. rexmontis Dwyer, Ann. Mis- 
souri Hot. Gard. 67: 241. 1980. Figure 11. 

Herbs or subshrubs, 0.5-1.5 m tall, erect or leaning 
on others, usually unbranched, leafy stems 2-6 mm thick, 
glabrous, apparently succulent in life and distorted on 
drying; stipules 2-3 mm long, 2 mm broad at the base, 
glabrous, caducous. Leaves with petioles (1.2-)2-6 cm 
long, 0.8-1 .8 mm broad, petioles of the same node often 
unequal in length, glabrous; leaf blades 10-25 cm long, 
4-10 cm broad, broadly elliptic to elliptic-oblong or el- 
liptic-obovate, apex acuminate with a narrowed tip 5- 
15 mm long, tapering gradually to a cuneate base (or 
abruptly obtuse) and decurrent on petiole, drying mem- 
branaceous, dark yellowish green or grayish green, gla- 
brous above and below, 2 veins 8- 11 /side. Inflores- 
cences solitary and axillary to basal leaves or cauliflorus 
(2/node), 3-8 cm long, with a single unbranched rachis 
or with 2 lateral helicoid branches 1-3 cm long, pedun- 
cles 15-35 mm long, 0.3-1 mm thick, flowers 3-30, 
pedicels 0-3 (-6) mm long, ca. 0.5 mm thick. Flowers 
glabrous, hypanthium 2-3 mm long and 1 mm diam. at 
apex, narrowly obconic, calyx lobes 1-2 m long, trian- 
gular to ligulate, acute; corolla funnelform, white or pale 
yellowish green, tube 2-4 mm long, ca. 1.3 mm diam., 
lobes 4.5-8 mm long, 0.8-1.7 broad. Fruits white be- 
coming red or dark reddish purple, 4-6 mm diam. (to 1 
cm in life), globose or ellipsoid-oblong during develop- 
ment. 

Plants of the wet cloud forests along the Carib- 
bean escarpment, (400-)700-1400 m elevation. 
Flowering in January-June; fruit in February-June, 
August, and November-December. Collections 
have been made from Bijagua, Alajuela, south- 
eastward to the upper Rio Grande de Orosi, Car- 
tago, and in western Panama. 

Hoffmannia pallidiflora is recognized by the gla- 
brous, slender, usually simple unbranched or two- 
branched pedunculate inflorescences, very thin 
leaves usually decurrent at the base and often long- 
petiolate, narrowly obconic hypanthium, and re- 
striction to the middle elevations of the Caribbean 
slope. The thin foliage dries yellowish to very dark 
green. The slender unbranched inflorescences have 
flowers along only one side and usually dry yel- 
lowish. 



Hoffmannia piratarum Standl., J. Wash. Acad. Sci. 
18: 180. 1928. 

Shrublets, branchlets obtusely tetragonal, glabrous; 



stipules caducous. Leaves with petioles 25-45 mm long; 
leaf blades 10-20 cm long, 5-6 cm broad, elliptic-lan- 
ceolate, long-acuminate, the tip often falcate, drying 
membranaceous, dark green and glabrous above, sparse- 
ly short villous and glabrescent beneath, with 8-9 2 
veins on each side. Inflorescences in leaf axils, 1-3/nodc, 
to 3 cm long, peduncles 1-2.5 cm long, short villous, 
pedicels 2-5 mm long. Flowers puberulcnt proximally, 
hypanthium 3 mm long, calyx lobes 2-3 mm long, tri- 
angular or rounded; corolla glabrous, tube ca. 4 mm long. 
2 mm diam. at base and 5 mm at apex, corolla lobes ca. 
6 mm long. Fruits ca. 6 mm long. 

Plants of Wafer Bay, Cocos Island. Flowering 
in January. Standley knew it only from the type 
collection Pittier 16259 us, and the preceding de- 
scription is based on his description. 



Hoffmannia pirtieri Standl., Contr. U.S. Natl. Herb. 
18: 140. 1916. H. panamensis Standl., Contr. 
U.S. Natl. Herb. 20: 204. 1919. 

Shrubs or subshrubs, 1.5-3(-4) m tall, usually with 
unbranched main stems, leafy stems 2-9 mm thick, be- 
coming woody and 12 mm thick, glabrous or densely 
pubescent on the young stems; stipules ca. 1 .5 mm long, 
ca. 2.5 mm broad at the base, broadly triangular, gla- 
brous. Leaves with petioles 1.5-7 cm long, 1.5-5 mm 
broad, glabrous or rarely puberulent in early stages, with 
narrow lateral margins continuous with the lamina mar- 
gins; leaf blades 1 2-26(-35) cm long, 4-1 2(-l 6) cm broad, 
elliptic-obovate to elliptic-oblong or oblanceolate, apex 
acuminate with tip 8-18 mm long, gradually narrowed 
to the cuneate base and long-decurrent on the petiole, 
drying chartaccous and dark brown or dark grayish green 
above, glabrous above and below (rarely puberulent in 
early stages), 2 veins 9-14/side and weakly loop-con- 
nected near the margin. Inflorescences 1-5/axil from 
thickened reduced peduncles, 2-5(-9) cm long, with 
(5-)10-15 flowers, peduncles 4-40 mm long, with 1 
branches 3-30 mm long (often 2-branched with alternate 
flowers along 1 side), glabrous to densely reddish pu- 
berulent, pedicels to 9 mm long. Flowers glabrous to 
reddish puberulent, hypanthium 2-3 mm long, 1.5-2 
mm diam., oblong-obconic, calyx orange-red, calyx lobes 
0.5-1 mm long; corolla yellowish tipped with red or 
orange, tube 3-6 mm long, 0.7-1.6 mm thick, lobes 3- 
5 mm long, 1.7 mm broad at the base. Fruits ca. 5 mm 
diam., deep green becoming deep red. 

Plants of evergreen montane forest formations, 
1 100-1 700(-2200) m elevation. Flowering in Feb- 
ruary-August; fruiting in January-February and 
August. The species (as here interpreted, see be- 
low) ranges from the San Vito de Coto Brus area 
into the adjacent Chiriqui Highlands of Panama. 

Hoffmannia pittieri is recognized by the taller, 
more woody stems, larger leaves with cuneate long- 
decurrent leaf bases, inflorescences with slender- 
pedicellate flowers, short corolla lobes, and re- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



177 



stricted latitudinal range. The inflorescences vary 
from short to moderately long with few to many 
branches. The interpretation presented here as- 
sumes that the very similar H. leucocarpa does not 
range eastward of Volcan Irazu. There is probably 
intergradation with H. arborescens (which usually 
has larger more branched inflorescences), and it 
may be necessary to place all this material under 
a more broadly denned H. arborescens. 



Hoffmannia psychotriifolia (Benth. in Oerst.) Gri- 
seb., Fl. Brit. W. Ind. 321. 1861. H igginsia psy- 
chothaefolia Benth. in Oerst., Vidensk. Meddel. 
Dansk. Naturhist. Foren. Kjobenhavn 1852: 50. 
1853. Figure 11. 

Slender shrubs or herbaceous subshrubs, l-2.5(-4) m 
tall, with few lateral branches, leafy stems 0.8-4.5 mm 
thick, glabrous, terete; stipules 1-1.5 mm long or reduced 
to a ridge, 1-2 mm broad at the base, triangular, glabrous. 
Leaves with petioles 8-35(-50) mm long, 0.7-1.5 mm 
thick, glabrous; leaf blades 5-14(-20) cm long, 1 .5^.5(-6) 
cm broad, narrowly elliptic to oblanceolate or narrowly 
elliptic-obovate, apex acuminate with tip 1-2 cm long, 
gradually narrowed to the acute or cuneate and decurrent 
base, drying membranaceous to thin-chartaceous, dark 
green or dark grayish brown above, distinctly paler be- 
neath, glabrous above and below, 2 veins 5-9/side. In- 
florescences usually fasciculate from a thickened solitary 
short-shoot (brachyblasts) in the leaf axils, with 5-30 
flowers per node, peduncles rarely present or 0-3(-8) mm 
long, pedicels 2-8 mm long, ca. 0.3 mm thick, glabrous 
or minutely puberulent. Flowers glabrous, to 18 mm 
long, hypanthium 2-3 mm long, 0.7-1.7 mm diam. dis- 
tally, oblong-obconic, with longitudinal ridges, calyx lobes 
0.6-2(-3) mm long, triangular to ligulate; corolla fun- 
nelform, pale yellow to white or pink, tube 4-9 mm long, 
1-3 mm diam., lobes 3-6 mm long, 1.4-2 mm broad at 
the base. Fruits 6-12 mm long, 5-8 mm diam., oblong- 
ellipsoid, becoming red and translucent. 

Plants of evergreen montane forest formations, 
(900-)1 100-2100(-2400) m elevation. Flowering 
in January-July and September-November; fruit- 
ing in February-September and November. The 
species ranges from southern Mexico and Guate- 
mala to Panama and occurs in the West Indies. 

Hoffmannia psychotriifolia is recognized by the 
thin-glabrous leaves tapering gradually to both apex 
and base, stems rounded and slender, small, usu- 
ally fasciculate inflorescences, slender pedicels, 
larger flowers with well-developed calyx lobes, 
longer ridged ovary, and oblong fruit. This species 
may be difficult to separate from H. longipetiolata 
(including H. tonduzii), but that species has small- 
er flowers, few-flowered, often pedunculate inflo- 
rescences, and young stems that may have puber- 
ulent longitudinal ridges. 



Hoffmannia refulgens (Hooker) Hemsley was 
based on cultivated material thought to have been 
collected in South America but most probably from 
northern Central America or southern Mexico (D. 
Lorence, pers. comm.). This name has been used 
for Costa Rican collections now placed under H. 
bullata. 



Hoffmannia subauriculata Stand! .. J. Wash. Acad. 
Sci. 18: 179. 1928. H. haydenii Dwyer, Ann. 
Missouri Bot. Gard. 56: 277. 1969. Figure 8. 

Herbaceous subshrubs, 1-2 m tall few-branched, leafy 
stems 1 .5-5(-7) mm thick, subglabrous; stipules 2-3 mm 
long. Leaves subsessile with poorly denned petioles be- 
cause of the long-attenuate lamina base and winged lat- 
eral margins 2-4 mm broad (short petioles 1-3 mm long 
sometimes present below the auricles); leaf blades 8- 
2 5 (-40?) cm long, 3-10(-13) cm broad, obovate to ob- 
lanceolate or broadly elliptic, apex short-acuminate, ta- 
pering very gradually to the cuneate or long-attenuate 
base (5-7 x l cm) and usually slightly auriculate near 
the stem, drying dark, glabrous above, subglabrous or 
minutely puberulent on the veins below, 2 veins 9- 137 
side. Inflorescences 2-4 on lower leafless nodes, 3-9 cm 
long, usually with few or no lateral branches (cymose to 
cincinnoid), with ca. 6-12 flowers, peduncles 2-4 cm 
long, glabrous or with reddish curved hairs to 0.2 mm 
long, pedicels 2-6(-12) mm long. Flowers with minute 
curved hairs or glabrous, with calyx lobes ca. 1.5 mm 
long, triangular; corolla not seen at anthesis. Fruits ca. 
8x6 mm, ellipsoid, bright red. 

Plants of wet evergreen forests in the central 
mountains and on the Caribbean slope, from 100 
to 1 800 m elevation. Flowering in June-July and 
November; fruiting in December-January. The 
species ranges from near Monteverde to central 
Panama. 

Hoffmannia subauriculata is distinguished by 
the long-decurrent leaves with broad (ca. 5 mm) 
lateral margins (along what would otherwise be a 
petiole) and their slight auriculate expansion near 
the base. The inflorescences appear to develop only 
at leafless nodes. This species is similar to H. am- 
plexifolia, which has three leaves per node. 



Hoffmannia valerii Standl., J. Wash. Acad. Sci. 
18: 178. 1928. Figure 9. 

Weak-stemmed shrubs or subshrubs, 0.5-1. 5(-3?) m 
tall, usually without lateral branches, leafy stems 1-4 
mm thick, with thin crooked multicellular hairs 0.3-1.5 
mm long; stipules 1-4 mm long, rounded distally or with 
an awn, caducous. Leaves with petioles 8-60 mm long, 
0.8-1.6 mm thick, villous with hairs 0.3-1 mm long; 
leaf blades 4-14(-19) cm long, 1.8-7(-9) cm broad, 



178 



FIELDIANA: BOTANY 



broadly elliptic to narrowly elliptic-oblong or ovate-el- 
liptic, apex acuminate, base obtuse to acute and some- 
what decurrent on petiole, drying membranaceous to 
thin-chartaceous, dark brown or grayish green above, 
glabrous or with few scattered hairs above, with more 
numerous crooked hairs 0.3-1.5 mm long beneath, 2 
veins 6-10/side. Inflorescences axillary or at leafless 
nodes, 2-4/node, l-2(-3) cm long, fasciculate or con- 
tracted cymes with 3-9 flowers, peduncles 1-4 mm long, 
pedicels 2-8(-12) mm long, filiform, villous or subgla- 
brous. Flowers puberulent with crooked hairs that dry 
reddish, hypanthium 2-3 mm long, 1.5-2 mm diam. 
distally, calyx lobes l-3(-4) mm long, narrowly trian- 
gular; corolla rotate, pale yellow, tube 1-3 mm long, 
lobes 3-5(-7) mm long; anthers 2-3 mm long. Fruits 4- 
6 mm long, 4-5 mm diam., ellipsoid, becoming orange 
or bright red. 

Plants of the wet evergreen Caribbean slope and 
lowlands, often along streams and in wet areas, 
50-950 m elevation. Flowering in February-Au- 
gust and November; fruiting in January, July, and 
November. The species ranges from near Arenal, 
Alajuela, to western Panama. 

Hoffmannia valerii is recognized by the long hairs 
on many younger parts, slender unbranched stems, 
small short-pedunculate or fasciculate inflores- 
cences, short corolla tubes, and lower-elevation 
habitats. The lype (Valeria 57 us) was collected at 
600 m elevation near Arenal, Guanacaste Prov- 
ince. This species resembles H. gesnerioides (Oer- 
sted) Kuntze of Nicaragua at 800-1500 m eleva- 
tion, but that species has longer (2-4 cm) peduncles, 
attenuate leaf bases, and dense spreading villose 
pubescence on all parts. Costa Rican material 
identified as H. boraginoides Dwyer ined. appear 
to be plants of//, valerii that have unusually long 
(2.5-3 mm) calyx lobes. The excellent DUKE col- 
lections from La Selva display considerable vari- 
ation in sepal lobe length, and it seems best to 
consider these plants all part of H. valerii. 



Hoffmannia vesiculifera Standl., Publ. Field Co- 
lumb. Mus., Bot. Ser. 4: 285. 1929. H. kirkbridei 
Dwyer, Ann. Missouri Bot. Gard. 56: 278. 1969. 
//. morn Dwyer, loc. cit. 67: 236. 1980. Figure 8. 

Herbs or subshrubs, 0.3-1.5 m tall, main stems usually 
unbranched, leafy stems 2.5-1 mm thick, usually densely 
villous with multicellular crooked reddish brown hairs 
to 1.5(-3) mm long; stipules to 2-5 mm long, little de- 
veloped and difficult to see among the pubescence, ca- 
ducous. Leaves with petioles 14-45 mm long, with a 
lateral longitudinal inflated chamber (vessicle or pouch) 
12-20 mm long and 4-9 mm broad on each side (2/ 
petiole), with conspicuous reddish brown hairs; leaf blades 
12-27(-40) cm long, 7-15(-20) cm broad, elliptic to el- 
liptic-obovate (less often obovate-oblong or oblanceo- 



late), apex acuminate to short-acuminate, base obtuse to 
cuneate and decurrent (to the inflated vessicles), drying 
thin-chartaceous, dark brown or grayish above, paler 
grayish or reddish brown beneath, with few scattered 
crooked multicellular hairs above and below, more 
densely hirsutulous on the midveins above and below, 
the crooked reddish hairs 0.4-1 .3(-3) mm long, 2 veins 
8-17/side. Inflorescences axillary or terminal on short 
lateral axillary shoots and subtended by leaf-like bracts 
ca. 2 cm long, solitary (2/node), sessile or with peduncles 
(= lateral branches?) to 3 cm long, flowers 10-many. in 
dense subsessile capitulae or on condensed helicoid cymes 
(with the inflorescences occasionally expanding to 7 cm 
and becoming branched in fruit, as in the type: Cooper 
231 F), pedicels to 4 mm long. Flowers yellowish but 
drying reddish brown, with conspicuous hairs 1-2 mm 
long, hypanthium ca. 2.5 mm long, and 1.5 mm diam., 
calyx lobes 2.5-7 mm long, linear, with few to many 
reddish hairs; corolla rotate, glabrous, cream yellow to 
reddish, tube 2-4(-8) mm long, corolla lobes 4-17 mm 
long, to 3 mm broad at the base, lanceolate; anthers 3.5- 
4 mm long. Fruits 4-6 mm long, 3-5 mm diam., be- 
coming bright red or white, thin-walled with longitudinal 
ribs, often with long ( 1 mm) crooked hairs. 

Plants of wet evergreen forests of the Caribbean 
slope and lowlands, (20-)300-1500 m elevation. 
(Note: All Costa Rican collections came from above 
300 m elevation.) Flowering in February-June and 
September; fruiting in April and October-Febru- 
ary. The species ranges from central Costa Rica 
to central Panama. 

Hoffmannia vesiculifera is our most distinctive 
species of Hoffmannia. The unusual inflated struc- 
tures of the petiole are found in no other Costa 
Rican species of Rubiaceae and resemble those 
found on some Melastomaceae. The long crooked 
multicellular hairs, condensed inflorescences, and 
long-linear calyx lobes are additional distinctions. 
The variation in leaf size and form and inflores- 
cences (dense heads to helicoid cymes or, rarely, 
branching infructescences) is considerable but does 
not warrant segregation of species or subspecies. 
The inflorescences may all be borne on lateral ax- 
illary branches that are variously reduced, as ev- 
idenced by the paired leaf-like bracts subtending 
the inflorescences. Only one of our collections, Lent 
91 1 F, cited small ants in the petiole chambers. 



Holtonia Standley 

Holtonia Standley is now considered part of 
Elaeagia. 



Isertia Schreber 

REFERENCE B. Boom, A revision of the genus 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



179 



Isertia (Isertieae: Rubiaceae). Brittonia 36: 425- 
454. 1984. 

Shrubs or trees, branchlets slender and subterete or 
thick and quadrangular, glabrous or puberulent; stipules 
interpetiolar or intrapetiolar, sometimes deeply divided 
and apparently 2/node, persisting. Leaves opposite, pet- 
iolate, often large; leaf blades drying thin-chartaceous to 
coriaceous, venation pinnate, domatia absent. Inflores- 
cences terminal and solitary, paniculate-thrysiform or 
racemose-thyrsiform, often large, the secondary branch- 
es terminating in dichasia or scorpioid cymes, flowers 
sessile or pedicellate. Flowers bisexual, radially sym- 
metrical, hypanthium ovoid to subglobose, calyx lobes 
4-6, small, equal or unequal, persisting; corolla tubular- 
funnelform to salverform, corolla tube short or elongate, 
villous in the throat, corolla lobes 4-6(7), short and 
spreading, valvate or imbricate in bud; stamens 4-7, 
filaments inserted near the mouth of the tube, anthers 
dorsifixed, loculate with the interior of the thecae divided 
into small chambers, included or exserted; ovary 2-6(-7)- 



locular, ovules numerous on axile placentas, style linear, 
stigma with 2-6(7) oblong lobes. Fruits berry-like with 
fleshy exocarp and 2-6 pyrenes, each pyrene (nutlet) with 
a bony endocarp and 2-many seeds, globose; seeds min- 
ute, angular, brownish, the testa deeply foveolate. 

A genus of 14 species, ranging from Central 
America through northern South America and oc- 
curring naturally only in western Cuba and Gua- 
deloupe in the Caribbean. Boom has divided the 
genus into two sections: section Cassupa with 2- 
3( 4) locules and stigmas and fleshy fruit and sec- 
tion Isertia with (4-)5-7 locules and stigmas and 
hard fruit. Our two species represent both sections 
of the genus. The large leaves, large many-branched 
solitary terminal inflorescences, flowers with long 
narrow corolla tubes, and loculate anthers distin- 
guish this genus. 



Key to the Species of Isertia 

la. Leaf blades usually dull greenish or grayish beneath, usually acute at the base and decurrent on the 
petiole; corolla yellow to orange or red, corolla tubes to 28 mm long; fruit oblate, ca. 7 mm diam. 
I. haenkeana 

1 b. Leaf blades usually whitish gray beneath, subtruncate to obtuse at the base and not conspicuously 
decurrent on the petiole; corolla white, corolla tubes to 55 mm long; fruit ellipsoid, ca. 10 mm 
diam. . /. laevis 



Isertia haenkeana DC., Prodr. 4: 437. 1830. 7. 
deamii Bartlett, Proc. Amer. Acad. Sci. 33: 59. 
1907. 7. deamii var. stenophylla J. D. Smith, 
Bot. Gaz. 61: 374. 1916. Figure 50. 

Shrubs or small trees, 2-6(-20) m tall, branches thick 
and often slightly narrower at the node, leafy branchlets 
3-10 mm thick, quadrangular with rounded edges, be- 
coming terete, densely puberulent with short (0.2-0.6 
mm) grayish hairs; stipules 4/node (or interpreted as 2 
with 2 almost separate lobes), 7-14(-45) mm long, 2-8 
mm broad at base, narrowly triangular with long-acute 
apex, glabrous abaxially or sericeous along the midvein 
(strigulose), drying dark. Leaves often smaller beneath 
the inflorescence, petioles 5-50 mm long, 1.8-4 mm thick, 
with adaxial margins continuous with the lamina mar- 
gins, densely puberulent; leaf blades (7-)14-45(-64) cm 
long, (4-)7-16(-28) cm broad, elliptic to elliptic-oblong 
or elliptic-obovate, apex short- or long-acuminate, base 
gradually narrowed and acute to attenuate, decurrent on 
petiole, drying chartaceous, dark brown above, glabrous 
and often lustrous above, with thin erect or appressed 
whitish hairs 0.2-0.5 mm long on the veins beneath, 2 
veins 14-22/side and loop-connected near the margin. 
Inflorescences 8-22 cm long, 6-12 cm broad, peduncle 
a thick extension of the stem, to 5 cm long, with a thick 
central rachis and many lateral branches, strigulose, the 
lateral branches (dichasia) with a short (1-2 cm) reddish 
orange peduncle and a terminal flower at the dichotomy 



of 2 longer scorpioid branches with 4-9 flowers, the sec- 
ondary branches subtended by narrow bracts 3-1 1 mm 
long, pedicles 0-2 mm long, bracteoles 1-2 mm long. 
Flowers with hypanthium and calyx tube 2-3 mm long 
and equally broad, cupulate, subentire with 4 minute 
lobes, sparsely strigulose externally; corolla bright yellow 
turning orange or reddish, tube 17-25(-28) mm long, ca. 
1.5 mm diam. at base and 2-3 mm distally, lobes 5-6, 
5-7 mm long, with prominent dense yellowish hairs ca. 
2 mm long on the inner face basally; stamens 5 or 6, 
filaments and anthers 3.5-6 mm long; ovary 4-6(-7) loc- 
ular. Fruits berry-like, 4-5 mm long and 6-8 mm diam., 
oblate, smooth and strigulose, usually with 5 or 6 car- 
tilaginous lobes and 5-6 multiseeded pyrenes; seeds 0.6- 
0.9 mm long. 



Plants of evergreen lowland wet forest forma- 
tions, from near sea level to 600 m elevation in 
southern Central America. Probably flowering 
throughout the year; in central Panama flowering 
is primarily in the early rainy season (May-July), 
with fruit maturing in the late wet season and early 
dry season (Croat, 1978). This species ranges from 
Mexico to Panama, Columbia, and Venezuela and 
is found in westernmost Cuba. 

Isertia haenkeana is recognized by its large op- 
posite leaves with many secondary veins and de- 



180 



FIELDIANA: BOTANY 



current lamina base, four persisting stipules at each 
node, cupular calyx with minute lobes, narrow yel- 
low to red corolla tubes, short corolla lobes with 
bearded hairs within, and rounded berry-like fruit 
with hardened pyrenes within. This species resem- 
bles Palicourea guianensis. 



Isertia laevis (Triana) B. M. Boom, Brittonia 36: 
433. 1984. Cassupa laevis Triana, Ann. Sci. Nat. 
Paris, Ser. IV 9: 44. 1858. C. panamensis Standl., 
Contr. U.S. Natl. Herb. 18: 135. 1916. /. pana- 
mensis (Standl.) Standl., Publ. Field Columb. 
Mus., Bot. Ser. 8: 346. 1931. Figure 30. 

Small trees to 10(-1 5) m tall, often with several trunks 
ca. 10 cm diam., leafy branchlets 4-12 mm thick, qua- 
drangular with rounded edges, densely appressed-pu- 
berulent or strigulose with short (ca. 0.3 mm) yellowish 
or brownish hairs; stipules apparently 4/node but united 
at the base (for 1-2 mm) and leaving a scar around the 
stem above the leaf bases, 6-12 (20-40) mm long, 3-6 
mm broad, triangular-subulate, coriaceous, glabrous or 
sparsely puberulent on the abaxial surface. Leaves op- 
posite, petioles 1.6-7.5(-12) cm long, 2-5 mm thick, 
minutely (0. 1-0.2 mm) puberulent; leaf blades smaller 
beneath the inflorescences, 15-60 cm long, 7-22 cm 
broad, oblong to ovate-oblong or elliptic-oblong, apex 
acuminate or acute apex, tip to 2 cm long, base obtuse 
to rounded and truncate (rarely slightly decurrent on 
petiole), drying chartaceous to subcoriaceous, usually dark 
brown above, glabrous or subglabrous above and with 
the minor venation often impressed, minutely puberu- 
lent on the veins beneath, whitish canescent between the 
veins and the major and minor venation clearly outlined, 
2 veins 1 5-22/side and loop-connected near the margin, 
proximal 2 veins arising at 90 to the midvein, 3 veins 
subparallel. Inflorescences (7-)14-35 cm long, (4-)6-15 
cm broad, ovoid to ellipsoid in outline and with many 
short (4 cm) lateral branches, peduncle 2-6 cm long, to 
6 mm thick, lateral branches usually a 5-flowered di- 
chasium (the distal branches with 2 flowers on the upper 
side), with short (1-3.5 cm) secondary peduncles, sparse- 
ly and minutely puberulent, flowers sessile or short (2- 
4 mm) pedicellate, bracteoles 1-3 mm long, broadly ovate. 
Flowers sweet smelling and apparently opening in the 
evening, hypanthium and calyx tube 5-7 mm long, 3-5 
mm diam., an elongate cup glabrous externally, the calyx 
lobes minute and 3-6 or obscure; corolla salverform, 
white, tube 32-55 mm long, 2-5 mm diam., sparsely 
and minutely puberulent externally, corolla lobes 6-7, 
1 0- 1 4 mm long, with long (1mm) yellowish hairs inside 
near the mouth; stamens 6-7, filaments ca. 3 mm long 
and laminar, anthers 7-9 mm long, included; ovary 2- 
3-locular, style 32-55 mm long, stigma with 2 oblong 
lobes 3-5 mm long. Fruits berries to 12 mm long, 8-1 1 
mm diam., ellipsoid to subglobose, smooth and usually 
glabrous, becoming black; seeds 0.7-1 mm long. 

Trees of open secondary vegetation in evergreen 
rain forest formations in both the Caribbean and 
southern Pacific lowlands, from near sea level to 



about 800 m elevation in Costa Rica. Probably 
flowering throughout the year but with flowers and 
fruit collected mostly in May-September. This 
species ranges from Costa Rica and Panama along 
the Andes to Bolivia and in the adjacent Amazon 
Basin. 

Isertia laevis is recognized by its large opposite 
leaves with whitish surfaces beneath, four persist- 
ing stipules at each node, solitary terminal inflo- 
rescences with long-tubed white flowers, subentire 
calyx tube, and subglobose berries. Panamanian 
material of this species was called Isertia hypoleu- 
ca by Croat (1978) and Dwyer (1980), but I. hy- 
poleuca Bentham is a different species occurring 
in Venezuela, the Guianas, and the Amazon basin. 



Ixora Linnaeus 

Shrubs or small trees, branchlets terete or angular, 
mostly glabrous; stipules interpetiolar (also slightly unit- 
ed above the petioles to form a very short intrapetiolar 
ridge), simple, usually broad at the base and acuminate, 
persisting. Leaves opposite and decussate or in whorls 
of 3, sessile or petiolate, laminae often coriaceous, ve- 
nation pinnate, domatia absent. Inflorescences terminal 
or less often axillary, cymose, thyrsoid or paniculate, 
branching often trichotomous, often with a rounded or 
flattened top (corymbose), flowers pedicellate and sub- 
tended by 2 bracteoles. Flowers bisexual and mono- 
morphic, hypanthium ovoid, calyx tube short, calyx lobes 
4 (5), short or extended, corolla salverform, white to 
pinkish, red, scarlet, or yellow, corolla tube narrow, gla- 
brous or puberulent at the throat, corolla lobes 4 (5-8), 
linear-lanceolate to ovate, imbricate or convolute in bud 
and rotate at anthesis; stamens 4 (5-8), inserted in the 
throat or mouth of the tube, filaments short or none, 
anthers oblong to linear, dorsifixed, exserted or partly 
exserted; ovary 2-locular with 1 ovule in each locule, 
peltately attached to the middle of the septum, style 
filiform, stigmas 2. Fruits baccate, red becoming black, 
the pericarp fleshy or leathery, with 1-2 pyrenes; seeds 
concave-convex or plano-convex to subglobose. 

A pantropical genus of ca. 400 species with the 
largest number of species in Africa and the Indo- 
Pacific area. The colorful inflorescences, narrow 
corolla tubes, stamens borne near the apex of the 
corolla tube, and two-seeded fruit help to char- 
acterize the genus. The flowers are protandrous in 
a distinctive manner: pollen is transferred from 
the clustered anthers into a concave area of the 
stigmatic head; elongation of the stigma disperses 
the pollen. Later, the stigma lobes open to expose 
their receptive surfaces for pollination. Four spe- 
cies are known to occur in Central America; two 
are native and two are widely cultivated as or- 
namentals. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



181 



Key to the Species of Ixora 

la. Flowers closely crowded together in capitate inflorescences; corolla tubes 25-45 mm long; plants 
cultivated for ornament 2 

Ib. Flowers crowded in distal cymose groups in an open paniculate inflorescence; corolla tubes 3-8 mm 

long; native wild species 3 

2a. Flowers red or orange; leaves sessile or subsessile /. coccinea 

2b. Flowers white; leaves petiolate /. finlaysoniana 

3a. Fruits ca. 10 mm diam.; flowers sessile in small glomerules, peduncles 0.7-3 mm thick; leaf blades 
1 2-26 cm long /. floribunda 

3b. Fruits ca. 4 mm diam.; flowers in panicles and pedicellate, peduncles 0.3-1 mm thick; leaf blades 
7-16 cm long /. nicaraguensis 



Ixora coccinea L., Sp. PI. 110. 1753. Figure 31. 

Shrubs 0.5-3(-5) m tall, much branched, leafy branch- 
lets 1-4 mm thick, glabrous, terete; stipules 0.5-1.5 mm 
long at the broad base and with a narrow tip 2-14 mm 
long, glabrous. Leaves opposite, sessile or subsessile with 
petioles to 2 mm long; leaf blades (2-)3-l 0(-1 6) cm long, 
( 1-) 1 .5-4.5(-6) cm broad, oblong to elliptic-oblong, ovate- 
oblong or oblong-obovate, apex rounded to obtuse or 
acute and often with minute (0.5-1 mm) slender tip, base 
rounded and subtruncate or subcordate to obtuse, drying 
stiffly chartaceous or subcoriaceous, glabrous above and 
below. Inflorescences terminal, to 10 cm long, flowers 
closely clustered (subcapitate), peduncles 1-3 cm long, 
flowers usually sessile at the apex of short-stipitate triads 
(cymes). Flowers with hypanthium 1-1.5 mm long, tur- 
binate, calyx lobes 4, ca. 1 mm long; corolla red or or- 
ange-red, tube 25-45 mm long, only 0.5-1 mm diam., 
glabrous or subglabrous, corolla lobes 4, 10-15 mm long, 
4-8 mm broad; stamens 4, anthers exserted and soon 
deciduous; style usually exserted. Fruits 8-1 5 mm diam. 
(rarely developed in cultivars). 

Ixora coccinea is a widely cultivated species 
throughout Central America, from near sea level 
to about 1500 m elevation. The rounded clusters 
of brilliant reddish flowers with long slender tubes 
and the evergreen leaves make it an attractive or- 
namental. The species originated in India and is 
now grown throughout the tropics and subtropics. 
Several varieties and forms have been recognized; 
see F. R. Fosberg and H.-H. Sachet, Three culti- 
vated Ixoras. Baileya 23: 74-85. 1989. This spe- 
cies is called cruz de Malta, flor defuego, jazmin, 
jazmin rojo, and jazmin de coral. 



Ixora finlaysoniana Wallich ex G. Don, Gen. hist. 
3: 572. 1834. Figure 31. 

Shrubs or small trees to 5 m tall, leafy branchlets 2- 
6 mm thick, glabrous, becoming terete with age; stipules 
2-7 mm long, triangular and acuminate or broad with 
1 or 2 awns. Leaves opposite, petioles 4-20 mm long, 



1-2 mm thick, glabrous; leaf blades 6-18 cm long, 2- 
6.5 cm broad, narrowly oblong to narrowly oblong-ob- 
ovate, apex bluntly obtuse or rounded (acute), base grad- 
ually narrowed and obtuse or acute, drying subcoria- 
ceous, glabrous above and below. Inflorescences terminal, 
5-10 cm long, to 10 cm broad and rounded, of densely 
congested cymose branches forming a capitate cluster, 
peduncles 0-3 cm long. Flowers glabrous, hypanthium 
1-2 mm long, calyx lobes 3-4 mm long 1 .5-2 mm broad, 
petaloid in texture; corolla white, tube 20-30 mm long, 
0.3-1 mm diam., lobes 4-6(-8) mm long, l-2(-3) mm 
wide; stamens 5, exserted, anthers narrow; stigma ca. 2.2 
mm long, slender and narrowly 2-lobed. 

Ixora finlaysoniana is widely cultivated in the 
tropics and planted at lower (0-1 000 m) elevations 
in Central America. The rounded inflorescences 
of many white flowers with long slender tubes, the 
lack of pubescence, the foliaceous sepal lobes, and 
the short-petiolate coriaceous oblong leaves char- 
acterize this species. Native of southeast Asia; this 
species is referred to as corona de la reina and 
mono de reina. 



Ixora floribunda (A. Rich.) Griseb., Cat. PI. Cub. 
134. 1866. Siderodendronfloribundum A. Rich, 
in Sagra, Hist. Cuba 1 1: 24. 1850. Figure 43. 

Small trees to 1 5 m tall, short-shoots frequently pres- 
ent, leafy branchlets 2.2-6 mm thick, smooth and gla- 
brous, terete; stipules 4-10 mm long, united above the 
leaf bases to form a short (1-4 mm) sheath, triangular- 
subulate and often shifted to the side above the petiole 
(not strictly interpetiolar), glabrous abaxially. Leaves op- 
posite, petioles 10-25 mm long, 1.5-2.5 mm thick, gla- 
brous; leaf blades 12-26 cm long, 4-10 cm broad, oblong 
to elliptic-oblong, or obovate, apex obtuse to short-acu- 
minate, base obtuse to attenuate and decurrent on pet- 
iole, drying stiff-chartaceous to subcoriaceous, concol- 
orous, glabrous above and below, 2 veins 6-1 0/side, not 
loop-connected distally. Inflorescences terminal, 1-3, 5- 
1 2 cm long, panicles with 2-3 pairs of opposite branches, 
primary peduncles 1-3 cm long, secondary peduncles 
equally long and densely puberulent with short (0.2-0.3 



182 



FIELDIANA: BOTANY 



mm) grayish hairs, flowers sessile and 3-12 in distal 
cymose clusters. Flowers with hypanthium ca. 1.5 mm 
long and 1.2 mm diam., turbinate, covered with short 
(0.2 mm) thin erect whitish hairs (and with some smaller 
glandular hairs); calyx tube 0.5-1 mm long, calyx lobes 
minute; corolla white, tube ca. 4 mm long, 0.3-0.6 mm 
diam. in proximal half, glabrous externally, white-villous 
within, lobes 4, 3-4 mm long, ca. 1 .5 mm wide, glabrous; 
filaments 0.6-1.5 mm long, anthers ca. 3.3 mm long, ca. 
1.5 mm broad; stigma lobes ca. 1.5 mm long and un- 
equal. Fruits 6-10 mm long, subglobose, sessile, sparsely 
puberulent, drying red and smooth, persistent calyx tube 
0.4-0.7 mm high and ca. 1.2 mm broad. 

Trees of both lowland rain forest formations and 
seasonally deciduous formations, from 1 to 800 
m elevation in Central America. Flowering in Jan- 
uary-February; fruiting in March-May in Central 
America. This species, apparently uncommon in 
Central America, ranges from Honduras and El 
Salvador to Colombia and some of the West In- 
dies. 

Ixora floribunda is recognized by its stiff-gla- 
brous elliptic-oblong leaves, terminal inflores- 
cences of white salverform flowers, puberulent ca- 
lyx, and sessile red berries. Our collections come 
from the area between Canas and Bagaces and 
Monte Aguacate on the seasonally deciduous Pa- 
cific slope of central Costa Rica. The species has 
also been collected in the Caribbean rain forest 
formations of Honduras and Nicaragua. It has been 
called pah de Maria. 



Ixora nicaraguensis Wernham, J. Hot. 50: 243. 
1912. 7. rauwolfioides Standl., Trop. Woods 1 1: 
25. 1927. Figure 43. 

Shrubs or small trees to 10 m tall, often with many 
branches, leafy branchlets 1.5-3 mm thick, glabrous, te- 
rete, grayish; stipules 3-6(-8) mm long, 2-3 mm broad 
at the base, broadly triangular and acuminate to cuspi- 
date with a short awn to 3 mm long. Leaves opposite, 
petioles 3-9 mm long, 0.7-2 mm broad, glabrous, with 
lateral (or adaxial) ridges; leaf blades 6-1 3(-l 6) cm long, 
2-5(-7) cm broad, elliptic-oblong, oblong or ovate-ob- 
long, apex abruptly narrowed and acute or short-acu- 
minate, base obtuse to somewhat attenuate and slightly 
decurrent on petiole, drying chartaceous, glabrous on 
both surfaces, 2 veins 6-12/side, often obscure and 
weakly loop-connected distally. Inflorescences 1-3 and 
terminal, 3-9 cm long and equally broad, open pyra- 
midal panicles with 2-3 pairs of opposite branches, pe- 
duncles 1-25 mm long, 0.3-1 mm thick, minutely (0.1- 
0.2 mm) puberulent, flowers on slender pedicels 0-10 
mm long in distal triads, distal bracts ca. 0.5 mm long. 
Flowers with hypanthium and calyx tube 1-1.7 mm long, 
minutely puberulent or glabrous, calyx lobes obsolete or 
minute (0. 1-0.5 mm); corolla white, glabrous externally, 
tube 3-6 mm long, ca. 1 mm diam., lobes 4, 3-4 mm 



long, 1-2 mm broad, oblong and rounded distally; fila- 
ments 0.5-1 mm long, anthers 2-3 mm long, 0.3 mm 
thick; stigma ca. 1.5 mm long, exserted. Fruits 5-6 mm 
long, 4-5 mm diam., subglobose to ovoid, glabrous and 
red; pyrenes 5 mm long. 



Plants of evergreen rain forest formations on 
both the Caribbean and southern Pacific slope of 
Costa Rica, from near sea level to 300 m elevation. 
Flowering in February-March and June-October; 
fruiting in March and July-November. This spe- 
cies ranges from Belize to eastern Panama. 

Ixora nicaraguensis is recognized by its small 
white flowers on open thin-branched inflores- 
cences, slender corolla tubes, and small globose 
two-seeded fruit, borne on thin-branched infruc- 
tescences. This species may be mistaken for a Psy- 
chotria, but the corolla lobes are valvate in bud in 
Psychotria (and related genera). 



Ladenbergia Klotzsch 

Small to large trees, the bark with bitter substances; 
stipules usually large, interpetiolar or also united distally 
(intrapetiolar) and forming a ring around the stem above 
the leaf bases after falling, triangular to obovate, with 
colleters at the adaxial base, caducous. Leaves opposite 
(occasionally whorled), petiolate, often subcoriaceous, 
pinnately veined, some species with domatia. Inflores- 
cences terminal, paniculate with opposite branching or 
cymose, ebracteolate. Flowers bisexual, monomorphic, 
medium to large, fragrant, hypanthium turbinate to cy- 
lindrical, calyx tube cupular, calyx lobes short or long; 
corolla funnelform or salverform, sericeous externally, 
corolla lobes 5-6, valvate in bud, minutely papillose 
within and on the margins; stamens 5-6, filaments very 
short, inserted near the center of the tube, anthers linear, 
dorsi fixed, included; ovary 2-locular, placentas elongate, 
spongy and borne on the septum, ovules numerous in 
each locule and vertically imbricate, style slender, stig- 
matic lobes 2. Fruits elongated capsules, cylindrical or 
flattened, with septicidal dehiscence (but the septum very 
thin and dehiscence often appearing to be loculicidal). 
splitting from the top into 2 woody or coriaceous valves; 
seeds numerous, longitudinally imbricate and peltate, 
flattened and elongate, body oblong and surrounded with 
a thin flattened dentate or laciniate wing. 



A genus of about 30 species ranging from Costa 
Rica to Bolivia; most of the species are Andean. 
The genus is similar to Cinchona and Condami- 
nea. The large salverform and sericeous flowers 
with long valvate corolla lobes, large stiff usually 
broad leaves, large broad stipules, and 2-valved 
capsules with winged seeds help to characterize 
the genus. 

Paul Standley's original separation of Costa Ri- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



183 



can material into three species appears to represent 
three valid morphological entities that are not 
sympatric, and intermediate collections are not 



apparent. Nevertheless, the species are very sim- 
ilar, and they might prove to be three subspecies 
of a single species. 



Key to the Species of Ladenbergia 

la. Tufts of stiff hairs (0.5-1 mm long) usually present at the base of the petiole in young leaves; leaf 

blades often acute at the base [corolla tubes 10-23 mm long, corolla lobes 11-17 mm long]; 

(1200-)! 500-2 100 m elevation L. valerii 

Ib. Base of the petiole glabrous or with minute (0.2 mm) appressed hairs; leaf blades rarely acute at 

the base; 0-1 500 m elevation 2 

2a. Corolla tubes 15-20 mm long, corolla lobes 8-12 mm long; fruit 2-5.5 cm long; central highlands 

1000-1400 m elevation L. brenesii 

2b. Corolla tubes 25-50 mm long, corolla lobes 16-20 mm long; fruit 6-9 cm long; evergreen lowlands, 

0-1000 m elevation L. sericophylla 



Ladenbergia brenesii Standl., Pub. Field Mus. Nat. 
Hist., Bot. Ser. 18: 1323. 1938. Figure 29. 

Trees, 5-25 m tall, leafy branchlets 4-12 mm thick, 
glabrous or sparsely puberulent at the nodes with short 
(0.1-0.3 mm) hairs, quadrangular, becoming terete and 
pale grayish; stipules 16-35 mm long, 4-16 mm broad, 
ovate to obovate, obtuse to acute at the apex, glabrous 
or with few slender ascending hairs ca. 0.5 mm long. 
Leaves with petioles 10-35 mm long, 1.8-3.5 mm thick, 
glabrous and drying dark; leaf blades 12-25 cm long, 5- 
1 3 cm broad, broadly elliptic to elliptic-oblong or ellip- 
tic-obovate, apex abruptly narrowed and obtuse or bluntly 
acute, base obtuse or slightly rounded, drying subcor- 
iaceous, glabrous above, glabrous or with a few scattered 
hairs below, 2 veins (6-)7-10/side. Inflorescences 6-18 
cm long, pyramidal, peduncles 2-7 cm long, glabrous, 
with opposite branches and bracts to 5 mm long, pedicels 
0-3 mm long. Flowers with hypanthium 3-6 mm long, 
2.5-5 mm diam., clavate-tubular, densely sericeous with 
yellowish brown ascending hairs, calyx tube ca. 1 mm 
long, calyx lobes l-2(-4) mm long, ca. 2 mm broad, 
broadly rounded, glabrous or sparsely puberulent distally 
and with a minutely ciliate edge; corolla white, tube 1 5- 
20 mm long, 5-6 mm diam., densely sericeous, lobes 8- 
12 mm long, 2-3.5 mm broad. Fruits (2-)3-6 cm long, 
5-10 mm broad (to 16 mm when fully flattened), sub- 
terete and narrowly oblong before dehiscing, valves pu- 
berulent externally and lustrous within; seeds 12-15 mm 
long, 34 mm broad, surrounded by a thin translucent 
erose wing, body of the seed 3-4 mm long and ca. 2 mm 
broad. 

Trees of wet evergreen cloud forest formations 
of the central highlands, 1000-1400 m elevation. 
Flowering in March-July; fruiting in July-Novem- 
ber. The species is endemic and ranges from the 
Cordillera de Tilaran to the western parts of the 
Cordillera de Talamanca. 

Ladenbergia brenesii is recognized by its cloud 
forest habitat, large puberulent flowers, long nar- 



row capsules splitting into valves that become flat, 
and broad glabrous leaves. Compare this species 
to L. valerii with which it can easily be confused. 
Also known as aquijilla and quina. 



Ladenbergia sericophylla Standl., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 18: 1324. 1938. Figure 29. 

Trees, 8-35 m tall, trunks to 60 cm. dbh, leafy inter- 
nodes 5-12 mm thick, young stems quadrangular and 
drying dark, quickly becoming pale gray and terete; stip- 
ules 15-45 mm long, 10-22 mm broad, oblong or ob- 
ovate-oblong, rounded at the apex, minutely puberulent 
with thin appressed-ascending hairs 0.1-0.3 mm long. 
Leaves opposite, petioles 15-4 5 (-60) mm long, 2-3.5 
mm thick, glabrous and drying very dark, reddish in life; 
leaf blades 14-24(-30) cm long, 8-16(-18) cm broad, 
very broadly elliptic to broadly elliptic-oblong or slightly 
obovate, apex abruptly narrowed and rounded or bluntly 
obtuse, base abruptly narrowed and obtuse or slightly 
attenuate, decurrent, drying stiffly chartaceous to sub- 
coriaceous, usually dark brown above, glabrous above, 
minutely (0. 1-0.3 mm) appressed-puberulent on the veins 
beneath and with larger (1 mm) hairs in the vein axils 
(domatia), 2 veins 5-8/side. Inflorescences 10-20 cm 
long, to 1 8 cm broad, pyramidal, peduncles 1-5 cm long, 
4-8 mm thick, glabrous or minutely appressed-puber- 
ulent, lateral branches opposite and subtended by bracts 
3-6 mm long, bracteoles subtending the flowers 1-2 mm 
long, rounded apically, pedicels 1-6 mm long. Flowers 
with hypanthium 5-7 mm long, 2-3 mm diam., tubular, 
densely sericeous with ascending yellowish hairs, calyx 
tube 2-4 mm long, cupulate, sparsely puberulent or gla- 
brous, calyx lobes 2-3 mm long, 2.5-3 mm broad, round- 
ed at the apex, glabrous and drying dark; corolla white 
or yellowish white, tube 25-50 mm long, 3-5 mm diam., 
densely sericeous, lobes 16-30 mm long, ca. 4 mm wide 
and narrowly oblong to lanceolate. Fruits 6-1 1 cm long, 
10-14 mm broad, oblong-cylindrical and rounded-rect- 
angular in cross-section, straight or curved (falcate), mi- 



184 



FIELDIANA: BOTANY 



nutely appressed-puberulent, persisting calyx 3-6 mm 
long and 6-8 mm broad, narrowed at the base to form 
a pedicel 5-1 5 mm long; seeds 1 1-20 mm long, 4-5 mm 
broad with thin wing, body of the seed ca. 3 x 1.5 mm. 

Tall trees of evergreen rain forest formations of 
the Caribbean lowlands and on the southern Pa- 
cific slope, 20- 1 000 m elevation. Flowering in Jan- 
uary-February, July, and September; fruiting in 
January-May. This species is only known from La 
Selva, the western parts of the General Valley, the 
mountains bordering the Pacific near Canas Gor- 
das, and the Osa Peninsula. While endemic to Cos- 
ta Rica, it may also occur in westernmost Panama. 

Ladenbergia sericophylla is characterized by its 
large broad leaves with appressed hairs on the veins 
beneath, large broad stipules, long salverform se- 
riceous corollas, and large capsular fruit. The taller 
height of these trees may explain why our collec- 
tions of this species are so few. 



Ladenbergia valerii Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 18: 1324. 1938. Figure 29. 

Small trees, 3-20 m tall, leafy branchlets 2-10 mm 
thick, quadrangular or terete in early stages, essentially 
glabrous but with stiff retrose or erect hairs ca. 0.5 mm 
long at the base of the petiole attachment; stipules 8-16 
mm long, 5-8 mm broad, broadly elliptic or ovate, apex 
rounded, sparsely puberulent with appressed hairs or 
glabrous and with a few hairs at the base and along the 
midrib abaxially. Leaves opposite, petioles 6-25 mm 
long, 1.5-2.5 mm thick, glabrous or very sparsely ap- 
pressed-puberulent (except for tufts of longer hairs below 
the base), drying dark; leaf blades 6-17(-28) cm long, 
3-8(-13) cm broad, broadly elliptic to elliptic-obovate 
or narrowly elliptic, apex abruptly narrowed and short- 
acuminate or bluntly acute, base obtuse to acute, drying 
stiffly chartaceous to subcoriaceous. glabrous or with a 
few thin hairs above, with thin straight hairs 0.3-0.7 mm 
long on the major veins beneath, 2 veins 5-9/side. In- 
florescences 7-20 cm long, pyramidal with opposite 
branching, peduncles 2-7 cm long, sparsely puberulent 
or glabrous, bracts ca. 3 mm long, distal flowers in triads 
and subtended by bracteoles 0.5-1 mm long, pedicels 0- 
3 mm long and merging with the hypanthium. Flowers 
with hypanthium and calyx tube 5-8 mm long, 2-3 mm 
diam., densely appressed yellowish sericeous, calyx lobes 
3-6 mm long, 2.5-3.5 mm broad at the base, broadly 
rounded to obtuse distally and becoming reflexed, gla- 
brous; corolla white or white with longitudinal pink 
stripes, salverform, tube 10-23 mm long, 1.5-4.5 mm 
diam., densely sericeous with ascending hairs, lobes 5, 
(11-)12-17 mm long, 2.5-4.5 mm broad, lanceolate to 
narrowly oblong, papillate-puberulent within, becoming 
recurved. Fruits 3-8(-l 1) cm long, 8-14 mm broad; seeds 
10-14 m long, 2-4 mm wide, body of the seed 3 m long 
and 1.5 mm wide. 

Trees of montane cloud forest formations from 



(1200-)! 600 to 2100 m elevation. Flowering in 
January-September and November; fruiting in 
May-August. The species ranges from the Cor- 
dillera de Tilaran to the Cordillera de Talamanca 
(as far east as 8304' W) and will probably be found 
in nearby Panama. 

Ladenbergia valerii is recognized by its higher- 
elevation habitat, unusual tufts of stiff short hairs 
beneath the petioles on otherwise glabrous or gla- 
brescent stems, sericeous corolla with relatively 
long corolla lobes, and capsular fruit. This species 
is very closely related to L. brenesii of similar for- 
ests at somewhat lower elevations. 



Lasianthus Jack 

Herbaceous subshrubs (in Central America), shrubs or 
rarely small trees, glabrous or pubescent; stipules inter- 
petiolar, broadly triangular to lanceolate, persisting or 
deciduous. Leaves opposite and decussate, petiolate, 
usually acuminate at the apex, pinnately veined and of- 
ten with many arching secondary veins, without doma- 
tia. Inflorescences axillary, mostly sessile fascicles or cy- 
mose glomerules, (sometimes pedunculate and simple or 
branched), flowers sessile or subsessile, bracts small. 
Flowers bisexual (rarely unisexual and monoecious), ra- 
dially symmetrical, mostly small, sometimes heterosty- 
lous, hypanthium urceolate to ovoid or subglobose, calyx 
lobes 3-6, acute or rounded, persisting; corolla salver- 
form to funnelform, white, corolla tube densely hairy in 
the throat, corolla lobes 4-6, spreading or erect; stamens 
4-6, filaments very short and borne in the throat, corolla 
lobes 4-6, anthers dorsifixed near the base, included or 
slightly exserted; ovary 4-12-locular, ovules solitary in 
each locule, erect from the base of the locule or septum, 
style short or long, stigmas 4-10, linear or lobed. Fruits 
usually succulent, blue to purple, black, or red, with 4- 
1 2 pyrenes, the pyrenes 1 -seeded and 3-angled with flat 
sides, the dorsal side grooved, keeled or winged; seeds 
narrowly oblong. 

A genus of about 1 50 species, of southeastern 
Asia, Malaysia, tropical Australia, and Africa. Two 
species are found in the West Indies, and one spe- 
cies in our area and South America. Our species 
with eight-locular ovary and style with eight stig- 
mas is unique among Central American Rubi- 
aceae. Our species was originally described as a 
new genus, Dressleriopsis Dwyer. 



Lasianthus panamensis (Dwyer) Robbrecht, Ann. 
Missouri Bot. Gard. 69: 427. 1982. Dressleriop- 
sis panamensis Dwyer, Ann. Missouri Bot. Gard. 
67: 154. 1980. Figure 7. 

Herbaceous rhizomatous subshrubs to 0.6(-1) m tall, 
leafy stems 2-6 mm thick, hirsute with slender erect hairs 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



185 



to 2 mm long; stipules 4-7 mm long, 3-6 mm broad at 
the base, broadly triangular and hirsute along the midrib. 
Leaves with petioles 3-9 cm long, 1.5-2.5 mm thick, 
hirsute; leaf blades 1 0-1 9 cm long, 5-9 cm broad, oblong 
to elliptic-oblong, apex narrowed abruptly and short- 
acuminate, tip 0-8 mm long, base obtuse and rounded 
to subcordate-auriculate, often unequal at the petiole, 
drying thin-chartaceous, dark grayish brown, with scat- 
tered thin straight or crooked hairs 0.5-2 mm long on 
upper and lower surfaces, 2 veins 9-14/side and loop- 
connected near the margin to form an arcuate submar- 
ginal vein, often with a shorter (parallel) minor 2 vein 
between the major. Inflorescences axillary, dense fasci- 
cles of sessile flowers 1-2 cm broad, glomerulate or ver- 
ticellate, bracts ca. 2 mm long and difficult to see among 
the long hairs. Flowers monomorphic, with hypanthium 
ca. 2 mm long, subglobose, hirsute, calyx lobes 4, 3-4 
mm long, 2-3 mm broad, with hairs to 1 mm long; 
corolla white, tube 3-4 mm long, lobes 5 (4, 6), 1.5-4 
mm long, oblong; stamens 5 (4, 6), anthers ca. 1 mm 
long: ovary with 8 locules, style to 5 mm long, stigmas 
8, oblong and radiate. Fruits berry-like, to 1 1 mm diam., 
globose, purple-black, pyrenes 8 or fewer, 3 mm long, 
with an oblique scar on the concave side. 

Plants of wet evergreen forest formations on the 
Caribbean slope (at ca. 100 m elevation) and in 
the central highlands of Panama (to 1 000 m ele- 
vation). Flowering and fruiting in July. This spe- 
cies is known from the La Selva area, Heredia, 
and in Panama and Colombia. 

Lasianthns panamensis is recognized by its slen- 
der hirsute stems to 1 m tall, long-petiolate oblong 
leaves with many secondary veins forming an ar- 
cuate submarginal vein, sessile fasciculate inflo- 
rescences, and fleshy berries with up to eight py- 
renes. No other Central American species of 
Rubiaceae has ovaries with eight locules. Super- 
ficially, these plants resemble some species of 
Hqffmannia and a few species of Psychotria with 
axillary inflorescences. 



Lindenia Bentham 

REFERENCE S. Darwin, The genus Lindenia 
(Rubiacea). J. Arnold Arbor. 57: 426-449. 1976. 

Small shrubs growing along rivers and streams, 
branchlets terete, glabrous or puberulent; stipules inter- 
petiolar, borne above the petiole bases, short, usually 
persisting. Leaves opposite short-petiolate; leaf blades 
narrow, often drying dark, chartaceous, pinnately veined, 
domatia absent. Inflorescences terminal, short and few- 
flowered, flowers often in triads or solitary, bracts and 
bracteoles present, pedicels short. Flowers bisexual, ra- 
dially symmetrical, large and showy, hyanthium elon- 
gate-turbinate, with 5 longitudinal ribs or angles, calyx 
with 5 elongate lobes, lobes equal or unequal, persisting 
in fruit; corolla narrowly salverform with a narrow elon- 



gate tube, glabrous within, corolla lobes 5, convolute in 
bud and spreading at anthesis; stamens 5, sessile on the 
mouth of the tube, linear and exserted; ovary 2-locular, 
ovules very many, vertical, placentas longitudinally ad- 
nate to the septum, style slender and bifid. Fruits woody 
capsules, clavate to pyriform or obovoid, with persisting 
calyx lobes distally, 2-locular, splitting septicidally into 
2 valves from the top; seeds numerous, angulate/rhom- 
boidal. 

A genus of three species, with the other two 
species endemic in the western Pacific islands of 
Fiji and New Caledonia, respectively. The stream- 
side habitat, narrow leaves on thick stems, few 
terminal flowers, and very long corolla tubes make 
this genus quite distinctive. 



Lindenia rivalis Benth., PL Hartw. 84. 1841. Fig- 
ure 15. 

Small shrubs, 0.4-1. 2(-2) m tall, leafy branchlets 1.5- 
6 mm thick, glabrous or minutely puberulent with thin 
ascending hairs ca. 0.2 mm long, older stems drying 
black; stipules 3-5(-10) mm long, 1.5-4 mm broad, the 
broad basal portion 2-3 mm high and with a narrow 
awn to 2(-5) mm long, glabrous, thin and brown. Leaves 
clustered near the ends of branchlets, petioles 2-10(-16) 
mm long, 0.4-1 .5(-2) mm broad, little differentiated from 
the leaf base; leaf blades 3-12(-17) cm long, 0.8-3(-4) 
cm broad, oblanceolate to very narrowly elliptic-oblong 
or narrowly elliptic, apex tapering gradually and acute, 
sometimes with a short (0.5-1 mm) spine-like tip, base 
tapering gradually and acute, decurrent on petiole, drying 
chartaceous to coriaceous, the margin often revolute, 
glabrous above, glabrous or puberulent beneath with thin 
erect hairs 0.2-0.3 mm long, 2 veins 6-8/side, not loop- 
connected. Inflorescences of solitary flowers or short (2- 
1 mm) pedunculate triads (or clusters of up to 7 flowers), 
bracts difficult to see among the distal leaves 4-9 mm 
long, pedicels to 10 mm long, poorly differentiated from 
the flower base. Flowers nocturnal, hypanthium 5-9 mm 
long, 2-3.5 mm diam., puberulent, calyx tube minute, 
calyx lobes 10-1 7(-22) mm long, 0.7-2(-2.8) mm broad, 
narrowly oblong and acute, green; corolla white or white 
tinged with pink, tube 10-17 cm long, 2-3.5 mm diam.. 
puberulent externally with hairs 0.2-0.5 mm long, lobes 
1 5-27(-35) mm long, 5-14(-16) mm broad, narrowly to 
broadly elliptic, acute at the apex; stamens sessile, an- 
thers ca. 10 mm long, 1-1.4 mm broad; style slender 
equalling or slightly exceeding the tube, stigmatic area 
ca. 10m long. Fruits to 4 cm long, body of the fruit 15- 
25 mm long, 9-14 mm broad, with long persisting calyx 
lobes before dehiscence, broadly obovoid or pyriform, 
the woody valves twisting; seeds 1.5-2 mm long. 

Small woody shrubs of stream sides and often 
growing on rocks next to the water, in deciduous 
forest areas of Guanacaste Province, 0-700 m el- 
evation (to 1300 m in Honduras). Flowering in 
February-October in southern Central America 
(primarily April-July); probably fruiting through- 



186 



FIELDIANA: BOTANY 



out the year (mostly August-March). The species 
ranges from northeastern Mexico and along the 
Pacific slope of Central America to central Pana- 
ma. 

Lindenia rivalis is recognized by its small stiff 
narrow leaves on thick dark branches, short stat- 
ure and riverside habitat in seasonally deciduous 
areas, few terminal flowers, and extremely long 
narrow corolla tube. Opler's observation (Opler 
945 F) that the flower is nocturnal would be con- 
sistent with the long narrow tube being an adap- 
tation for Sphingid moth pollination. Our only 
other Rubiaceae with such long/narrow corolla 
tubes have much larger leaves. Also known asjaz- 
mincillo and lirio de aqua. 



Machaonia Humboldt & Bonpland 

Shrubs or small trees, stems often armed with spines 
(leafless short-shoots), glabrous or puberulent, terete; 
stipules interpetiolar, triangular or subulate and often 
with a distal awn, persisting. Leaves opposite (rarely ter- 
nate or verticillate), sessile or petiolate, thin-textured, 
pinnately veined. Inflorescences terminal, solitary, pa- 



niculate with opposite branching (or rarely umbeliform), 
bracteate, flowers in cymose or crowded distal clusters. 
Flowers bisexual and radially symmetrical, small, 4-5- 
parted, hypanthium turbinate or obovoid, slightly com- 
pressed laterally, calyx tube minute or cupulate, calyx 
lobes 4-5(-6), equal or unequal, persisting; corolla short- 
funnelform or short-salverform, white, corolla tube with 
long hairs in the throat, lobes imbricate in bud. rounded 
distally; stamens 4-5, filaments short or long, borne in 
the throat of the tube, anthers oblong, dorsi fixed and 
versatile, included or exserted; ovary 2-locular, with 1 
ovule pendulous from the apex of each locule. style slen- 
der, with 2 stigmas. Fruits small and dry, capsule-like 
and splitting from the bottom into 2 elongate cocci (mer- 
icarps), pendulous for a short period from the apex to 
the persisting stipe-like central axis; seeds elongate and 
cylindrical. 

A tropical American genus of about 30 species, 
found in Mexico, Central America, the West In- 
dies, and tropical South America. The genus is 
recognized by its small flowers with short corolla 
tubes, two-locular ovary with solitary pendulous 
ovules, and unusual capsule-like fruit. The sub- 
capitate clusters of distal flowers on an openly 
branching panicle, small thin leaves, and occa- 
sional presence of spines are also distinctive. 



Key to the Species of Machaonia 

la. Hypanthium/ovary of the flower with few small hairs; fruit 5-7 mm long (including calyx lobes), 
brownish and with few hairs often in rows; lowland evergreen Costa Rica M. martinicensis 

Ib. Hypanthium/ovary of the flower whitish with a dense covering of short whitish hairs; fruit 4-5 mm 
long, yellow or whitish with many hairs; Belize and central Panama M. acuminata 



Machaonia acuminata Humb. & Bonpl., PI. Ae- 
quin. 1: 101. 1806. 

Shrubs or small trees, occasionally with spines to 4 
cm long at the nodes, leafy branchlets 1.2-6 mm thick, 
glabrous to densely villose with hairs to 1 mm long, 
terete; stipules 2-4 mm long, triangular-subulate with a 
narrow awn apically, puberulent. Leaves often disti- 
chous, petioles 3-10 mm long, 0.6-1 mm broad, puber- 
ulent and often with the hairs restricted to adaxial side; 
leaf blades (3-)4-7(-10) cm long, 1.5^(-6) cm long, 
ovate to ovate-elliptic or broadly ovate, apex acute (ob- 
tuse) to short-acuminate and often slightly curved, base 
obtuse to rounded and subtruncate, drying thin-charta- 
ceous, glabrate above, with thin whitish hairs 0.3-0.7 
mm long on the lower surfaces, 2 veins 3-6/side, tufted 
hairs rarely present in the leaf axils. Inflorescences 4-20 
cm long, 3-10 cm broad, open panicles, peduncles 2-4 
cm long and 1.5-3 mm thick, puberulent, bracts to 7 
mm long and liner, peduncles of the lateral branches 5- 
40 mm long, flowers closely crowded in subcapitate dis- 
tal groups of 7 or more, flowers sessile or subsessile. 
bracteoles ca. 1 mm long. Flowers with hypanthium ca. 
2 mm long, turbinate or oblong, minutely puberulent 



with thin ascending whitish hairs, calyx lobes 0.5-1 mm 
long, erect, rounded distally, puberulent; corolla white, 
4-5 mm long, tube 1.5-3 mm long, sparsely puberulent 
or glabrous, lobes 1.5-2.5 mm long, ca. 1.1 mm broad, 
rounded at the tip; stamens exserted, anthers 0.3-0. 6 mm 
long; styles 1-3.5 mm long, stigmas 0.3-0.6 mm long. 
Fruits 4-5 mm long (including the persisting calyx), 1 .5- 
2 mm broad, narrowly obovoid or turbinate, minutely 
(0.1-0.2 mm) puberulent with ascending whitish hairs, 
splitting at the top into 2 valve-like parts and later sep- 
arating at the base, persisting central axis 2.5-3.2 mm 
long. 



Plants of lowland evergreen and partly decid- 
uous forest formations. Flowering in May-No- 
vember; fruiting in August-September and No- 
vember. This species is known from southern 
Mexico, northeastern Guatemala, Belize, and cen- 
tral Panama southward to Ecuador and Brazil. 

Machaonia acuminata is recognized by its oc- 
casional spines, thin small ovate leaves, small 
flowers with whitish puberulent ovary, and small 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



187 



fruit. This species has not been found in Honduras, 
El Salvador, Nicaragua, or Costa Rica. Its close 
similarity to M. martinicensis and peculiar distri- 
bution makes one wonder if the two are really 
different species. 



Machaonia martinicensis (EXT.) Standl., Publ. Field 
Mus. Nat. Hist., Bot. Ser. 22: 193. 1940. Tertrea 
martinicensis DC, Prodr. 4: 481. 1830. M. ro- 
tundata Griseb., Fl. Brit. W. Ind. 348. 1861. M 
rotundata var. dodgei Standl., Publ. Field Mus. 
Nat. Hist., Bot. Ser. 18: 1326. 1938. Figure 37. 

Shrubs or (rarely small trees or vines), l-4(-8) m tall, 
branches occasionally with spines to 35 mm long, leafy 
branchlets 1.6-5 mm thick, smooth glabrate; stipules 
2.5-4 mm long, rounded or triangular at the apex and 
with a short narrow awn. Leaves opposite (rarely alter- 
nate), petioles 4-12 mm long, 0.5-1.2 mm broad, gla- 
brous or puberulent on the adaxial side; leaf blades 4- 
9 cm long, 1.8-5 cm wide, ovate to ovate-oblong (ellip- 
tic-oblong), apex obtuse to acute or short-acuminate, 
base broadly obtuse or rounded and sometimes decur- 
rent on petiole, drying thin-chartaceous, glabrous above, 
glabrous below or with few short (0.2 mm) hairs beneath 
or with tufts of hairs (domatia) in the vein axils, 2 veins 
5-7/side. Inflorescences 6- 11 (-15) cm long, to 14 cm 
broad in fruit, peduncles 2-6 cm long, ca. 2 mm thick, 
with whitish hairs 0.2 mm long, bracts 3-5 mm long and 
linear (or sometimes leaf-like), secondary branches with 
(secondary) peduncles 1 0-20 mm long, with soft whitish 
hairs 0.2-0.3 mm long, flowers sessile or subsessile in 
distal subcapitate clusters of 5-1 5. Flowers with buds 5- 
6 mm long, hypanthium 1.5-2.5 mm long, sparsely pu- 
berulent, calyx lobes 0.7-1.2 mm long, broadly rounded 
distally; corolla white or pale greenish, tube 1.5-3 mm 
long, stiff, densely villous at the mouth within, sparsely 
puberulent externally, lobes 1-2 mm long; stamens 5, 
anthers ca. 0.6 mm long; style 3.5 mm long, stigmas 0.5- 
1 .2 mm long. Fruits 5-7 mm long, 2-3 mm broad, nar- 
rowly turbinate-oblong, with 2 longitudinal sulci and 
separating at the base, surface reddish brown or brown, 
with few hairs (often in lines), persisting axis 4-5 mm 
long. 

Shrubs usually growing near the ocean shore, in 
mangroves, or along rivers near the ocean in ev- 
ergreen forest areas of both the Caribbean and 
Pacific coasts. Flowering in March-April and June- 
August; fruiting in July-September. The species 
ranges from southeastern Nicaragua southward to 
Colombia and also in Jamaica. 

Machaonia martinicensis is recognized by its 
restriction to near-shore environments, small thin- 
ovate leaves, occasional spines, small flowers in 
close clusters in an open panicle, and the unusual 
capsule-like fruit splitting from the bottom. The 
species has been collected along the Caribbean 
shore and on the Osa Peninsula in Costa Rica. 



Macrocnemum P. Browne 

Trees or shrubs, branchlets terete, glabrous or puber- 
ulent; stipules interpetiolar, oblong to obovate, caducous 
or persisting with the leaves. Leaves opposite, petiolate; 
leaf blades with pinnate venation, domatia often present. 
Inflorescences terminal or axillary, paniculate with op- 
posite branches, bracteate, flowers in distal cymose 
groupings, sessile or pedicellate. Flowers bisexual and 
radially symmetrical, calyx tube very short and cupulate, 
calyx lobes 5, small, persisting; corolla funnelform or 
salverform, corolla lobes 5, broadly imbricate in bud 
and spreading at anthesis, often wider than long, mi- 
nutely puberulent within; stamens 5, filaments short and 
villous, anthers oblong; ovary 2-locular, ovules many, 
peltate and vertically imbricate on the central placenta. 
Fruits capsular, bisulcate and dehiscing loculicidally into 
2 valves; seeds flattened, narrowly elongate with thin 
wings. 

A small genus of about 20 species, ranging from 
Central America to Colombia and in the West 
Indies. The broad rounded stipules, broadly over- 
lapping corolla lobes (in bud), bilocular capsules, 
and many small imbricate winged seeds help to 
distinguish this genus. 



Macrocnemum glabrescens (Benth.) Wedd., Ann. 
Sci. Nat. Paris, ser. 4, 1: 76. 1854. Lasionema 
glabrescens Benth., Bot. voy. Sulph. 105. 1845. 
Figure 40. 

Trees (rarely shrubs) (4-)8-25 m tall, bark brown and 
the trunk deeply fluted, leafy branchlets 2-7 mm thick, 
glabrous or minutely (0.1 mm) appressed-puberulent, 
angular in early stages but becoming terete; stipules 
(8-) 10-20 mm long, 5-10 mm broad, apex oblong-ob- 
ovate and rounded, with thin appressed hairs 0.1-0.2 
mm long, leaving a scar around the stem above the node. 
Leaves often with the petioles unequal at the same node, 
petioles 5-20 mm long, 0.7-1 .8 mm broad, sulcate adax- 
ially; leaf blades 7-17(-21) cm long, 4-9 cm broad, ob- 
ovate to obovate-oblong, or ovate-oblong, apex abruptly 
narrowed and obtuse or short-acuminate apex, tip to 1 
cm long, base gradually narrowed and obtuse or acute, 
drying thin- to stiff-chartaceous, very dark above, gla- 
brous above, glabrous or minutely appressed-puberulent 
on the veins beneath, usually with tufts of hairs ca. 1 m 
long in the vein axils beneath (= domatia), 2 veins 5- 
9/side. Inflorescences terminal or axillary, solitary or 3 
when the first branching node is subtended by a pair of 
leaves, 8-28 cm long, to 20 cm broad, paniculate with 
few opposite lateral branches to 1 2 cm long, peduncles 
to 1 8 cm long, bracts 3-6 mm long or leaf-like, lanceo- 
late, flowers in distal cymes or clusters of more than 10, 
sessile or with pedicels to 3 mm long, bracteoles 0.5-1 .5 
mm long. Flowers with a hypanthium 3-4 mm long and 
ca. 2.4 mm diam., glabrous or minutely puberulent, ca- 
lyx tube minute, calyx lobes 0.3-1 mm long, broadly 
triangular; corolla bright rose pink, magenta or the tube 
becoming maroon, salverform, tube 6-10 mm long, 1 .3- 



188 



FIELDIANA: BOTANY 



2 mm diam., with 5 longitudinal ribs, glabrous or rarely 
sparsely and minutely puberulent, villous at the stamen 
attachment within, lobes 3-4 mm long, 4-5 mm broad, 
broadly ovate and rounded distally; stamens 5, filaments 
2-4 mm long, villous below the middle, anthers 0.8-1 
mm long; style 6-8 mm long, stigma lobes ca. 0.5 mm 
long, green. Fruits 9-16(-20) mm long, 2-4 mm broad 
(to 4 mm when split open), narrowly oblong-tubular, 
with obscure longitudinal ribs, valves opening but re- 
maining attached at both apex and base, persisting calyx 
ca. 2 mm broad and 1 mm high (together with disc); 
seeds 2-3 mm long, 0.5-1 mm broad, with a membra- 
naceous wing at opposite ends, body of the seed ca. 0.6 
mm long. 

Trees of evergreen or partly deciduous forest 
formations from near sea level to 400(-1000) m 
elevation. Flowering in December-April; fruiting 
in March-July in Costa Rica and Panama. The 
species ranges from the evergreen lowlands of 
southern Costa Rica, through Panama, to Colom- 
bia. 

Macrocnemum glabrescens is recognized by its 
broadly rounded stipules, usually thin-obovate 
leaves (often on petioles differing in length at the 
same node), large open panicles with small bright 
pink flowers, broadly overlapping corolla lobes, 
and narrow capsules with opened valves remain- 
ing attached at the base and apex. This species is 
only known from near the Hitoy Cerere reserve, 
Dominical, and the Golfo Dulce region in Costa 
Rica, but the original description mentions Nicoya 
and it is possible that it grows in moist forest on 
the southern part of the Nicoya Peninsula. This 
species appears to be common in central Panama; 
see flower description in Croat ( 1 978, p. 8 1 1 ). This 
species resembles Ferdinandusa panamensis. 
Standley (1938) listed palo cuadrado as a common 
name. 



Malanea Aublet 



Shrubs, woody lianas, or trees, stems glabrous or pu- 
berulent, terete; stipules interpetiolar, simple, caducous. 
Leaves opposite, petiolate, entire, pinnately veined, with 
domatia. Inflorescences axillary, paniculate with short 
opposite or subopposite lateral branches resembling spikes 
(or with fasciculate flowers on the central rachis or lateral 
branches), flowers sessile or subsessile, bracteolate. Flow- 
ers bisexual and radially symmetrical, small, hypanthi- 
um turbinate to campanulate, calyx lobes 4, short; co- 
rolla funnelform to rotate, pale green to white, tube short, 
throat villose within, corolla lobes valvate or slightly 
imbricate, villous on the adaxial surface; stamens 4, fil- 
ament borne on the throat of the corolla, anthers dor- 
si fixed and partly exserted; ovary 2-locular, ovules sol- 
itary in each locule and pendulous from the apex of the 



locule. Fruits fleshy, oblong or ovoid, 2-locular, exocarp 
thin-fleshy, endocarp woody or hard; seeds ellipsoid. 

A genus of 20-30 species in tropical South 
America and the West Indies. Malanea colom- 
biana has been discovered along the Caribbean 
coast in Belize, in northern Nicaragua, and in 
southeastern Costa Rica. Malanea erecia Seem, 
(photo F), said to have been collected by Seemann 
in Panama, was not treated in the Flora of Pan- 
ama. 



Malanea colombiana Standl., Publ. Field Columb. 
Mus., Bot. Ser. 7: 66. 1930. Chomelia coclensis 
Dwyer, Ann. Missouri Bot. Gard. 67: 97. 1980. 

Lianas or clambering shrubs with vining branches, 
leafy stems 1.7-8 mm thick, with closely appressed-as- 
cending straight hairs 0.4-0.8 mm long, glabrescent and 
lenticellate; stipules 6-18 mm long, 2.5-8 mm broad, 
oblong or obovate, with many ascending veins, ap- 
pressed-puberulent at the base and midrib, deciduous. 
Leaves with petioles (4-)7-22 mm long, 0.6-2 mm thick, 
appressed-puberulent; leaf blades 5-13 cm long, 2.5-7 
cm broad, ovate-elliptic to ovate or oblong-elliptic, apex 
obtuse to short-acuminate or acute, base obtuse to slight- 
ly rounded and subtruncate, drying stiffly chartaceous, 
dark brown above, glabrous above or with appressed 
hairs along the midvein and widely scattered elsewhere, 
with appressed thin whitish hairs 0.3-1 mm long be- 
neath, often with tufts of hairs .in the vein axils beneath. 
2 veins 5-7/side, distal secondaries to arcuate-ascend- 
ing, 3 veins numerous and closely parallel to form a 
straight or sinuous pattern between the secondaries. In- 
florescences 4-13 cm long, 3-8 cm broad, pyramidal 
with progressively shorter opposite branches, peduncles 
2-6 cm long, 0.6-1 .8 mm thick, sericeous, lateral branch- 
es l-3(-4) cm long, bracts 1.5-4 mm long and linear, 
bracteoles ca. 1 mm long, flowers sessile or with pedicels 
to 1 .5 mm long. Flowers 3-4 mm long, hypanthium 0.5- 
1 mm long, turbinate, calyx lobes 0.1-0.4 mm long, 
broadly triangular or poorly developed; corolla white or 
yellowish white, tube 1.5-2.5 mm long. ca. 1 mm diam. 
at mouth, lobes 4, 1-2 mm long, 1 mm broad at the 
base, villose within; anthers 0.5-0.8 mm long; style ex- 
serted, stigma lobes ca. 0.4 mm long. Fruits 5-6 mm 
long, 3-4 mm diam., oblong-cylindrical, becoming pur- 
plish red and drying black, persisting calyx very short 
(0.5 mm) and not elevated. 



Plants of evergreen or partly deciduous vege- 
tation, 2-200 m elevation. Flowering in June in 
Costa Rica (Herrera 3081 CR, MO); fruiting in Au- 
gust in northern Nicaragua (Molina 14926 & 
151125 EAP, F). The species ranges from Belize to 
Colombia. 

Malanea colombiana is recognized by its very 
small flowers with four-lobed perianth, corolla 
lobes conspicuously villous within, fleshy oblong 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



189 



fruit, unusual tertiary venation, and climbing hab- 
it. The tertiary veins of the leaf are many and 
parallel with inconspicuous transverse connec- 
tions. The Costa Rican collection has minor ve- 
nation that differs by having a more reticulate ar- 
rangement of 3 veins with sublineolate 4 veins. 
More material is necessary to confirm its place- 
ment under this name. The flowers resemble those 
of Elaeagia, Figure 39. 



Manettia Mutis ex Linnaeus 

Herbs or climbers, stems herbaceous or slightly woody, 
glabrous or puberulent; stipules interpetiolar, small, 
sometimes adnate to the petiole, triangular to laciniate, 
persistent. Leaves opposite and decussate, petiolate or 
subsessile; leaf blades often narrow, pinnately veined, 
domatia absent. Inflorescences axillary (rarely terminal), 
cymose to paniculate, fasciculate, umbellate or the flow- 
ers occasionally solitary, bracts and bracteoles present, 
flowers pedicellate. Flowers bisexual, monomorphic or 
distylous, radially symmetrical, small (in Central Amer- 
ica) to large, hypanthium turbinate, calyx lobes usually 



4 (5, 8), short or long, with glands or teeth in the sinuses; 
corolla funnelform or tubular, white, pink, red, lavender, 
blue, or yellow, corolla lobes 4(-5), valvate in bud; sta- 
mens 4, filaments attached in the throat or at the mouth, 
anthers versatile, exserted or included; ovary 2-locular, 
ovules numerous and imbricate, vertical on axile pla- 
centas, style filiform, stigmas 2 clavate or bifid. Fruits 
thin-walled capsules, obovoid or turbinate to subglo- 
bose, biloculate and bisulcate, splitting septicidally from 
the apex into 2 valves; seeds compressed -discoid, usually 
with a thin wing surrounding the central seed. 

A tropical American genus of about 100 species. 
The slender twining stems, smaller narrow leaves, 
small inflorescences, small flowers (in Central 
American species), small obovoid capsules break- 
ing into two valves, and winged or flattened seeds 
with erose margin help to distinguish these plants. 
This study has benefited from the herbarium an- 
notations of In-Cho Chung and David Lorence. 
Manettia luteo-rubra variety paraguariensis (Cho- 
dat) Chung (= M. inflata Sprague) is an ornamen- 
tal grown for its bright red flowers tipped with 
yellow. 



Key to the Species of Manettia 

la. Leaf blades narrowly lanceolate and often subsessile; seeds ca. 1 mm diam., orbicular with an erose 
margin (the wing minute or absent); flowers white, corolla tube 3-5 mm long M. barbata 

Ib. Leaf blades rarely consistently lanceolate, petioles usually well developed; seeds 2-3 mm long, 
oblong or obicular with a thin expanded marginal wing; flowers white, pink, red, or magenta, corolla 
tubes 4-13 mm long 2 

2a. Corollas white; capsules 4-6 mm long; leaf blades usually puberulent beneath; 1000-1600 m ele- 
vation M. flexilis 

2b. Corolla brilliant red to magenta or pink; capsules 6-10 mm long; leaf blades usually glabrous beneath; 
30-1 100 m elevation . . M. reclinata 



Manettia barbata Oersted, Vidensk. Meddel. 
Kjobenhavn 1852: 47. 1853. M. stenophylla J. 
D. Smith, Bot. Gaz. 56: 58. 1913. Figure 1. 

Slender vines to 3 m high, leafy internodes 0.7-2.5 
mm thick, glabrous or subglabrous; stipules 0.5-3 mm, 
long, broadly triangular, glabrous. Leaves with petioles 
1-5 mm long, 0.3-1.5 mm broad, glabrous; leaf blades 
3-10 cm long, 0.4-3 cm broad, lanceolate to narrowly 
elliptic-lanceolate or linear-lanceolate, apex tapering 
gradually and long-acuminate or acute, base obtuse or 
slightly rounded, drying chartaceous, pale grayish green 
or grayish brown, glabrous above and below, 2 veins 4- 
7/side. Inflorescences 10-15 mm long, axillary cymes of 
usually 3 flowers, peduncles only 1-4 mm long, bracts 
to 3 mm long, lanceolate, pedicels 2-5 mm long, brac- 
teoles 0.5-l(-2) mm long. Flowers with hypanthium 2- 
3 mm long, 1 .2-2 mm diam., calyx lobes 1 .2-2 mm long, 
0.7-1.2 mm broad, ovate; corolla white, tube 3-6 mm 
long, 2 mm diam., lobes 2-3 mm long, glabrous exter- 



nally and villous within; anthers ca. 1 mm long; stigma 
ca. 0.5 mm long, oblong to lanceolate. Fruits 4-8 mm 
long, 3-7 mm broad, persisting calyx to 1.5 mm long 
and usually recurved; seeds 0.7-1 mm diam., orbicular, 
wing reduced to a dentate rim around the seed, body of 
seed ca. 0.6 mm diam., discoid. 



Uncommon plants of evergreen forest forma- 
tions, from 500 to 1300(-2400?) m elevation. 
Flowering in November-January; fruiting in Jan- 
uary and March. This species is known only from 
central and southern Costa Rica. 

Manettia barbata is recogized by its narrowly 
lanceolate leaves, twining habit, few-flowered and 
short-pedunculate inflorescences, short corolla 
tubes, smaller capsules, and circular-discoid seeds 
with little evidence of a wing. 



190 



FIELDIANA: BOTANY 



Manettia flexilis Brandegee, Univ. Calif. Publ. 
Bot. 6: 196. 1915. M. estrellae Standl., J. Wash. 
Acad. Sci. 15: 6. 1925. Figure 1. 

Vines to 4 m high, leafy stems 0.7-2 mm long, pu- 
berulent with curled whitish hairs ca. 0.2 mm long; stip- 
ules 1-2 mm long, adnate to the petiole base, rounded 
distally or with short (0.3 mm) thick spike-like projec- 
tions. Leaves with petioles 2-10 mm long, 0.3-0.7 mm 
thick, puberulent; leaf blades (2.5-)3-7 cm long, 1-3 cm 
broad, oblong to ovate or lanceolate, apex acute to long- 
attenuate, base obtuse to acute, drying membranaceous 
or thin-chartaceous, dark green above, pale grayish green 
beneath, glabrous or puberulent above, short pilose be- 
neath with thin whitish hairs ca. 0.3 mm long, 2 veins 
4-6/side, the upper secondaries strongly arcuate-ascend- 
ing. Inflorescences axillary, cymose or umbelliform flow- 
ers few (rarely 1), peduncles 1-7 mm long, bracts ca. 2 
mm long, flowers closely crowded, pedicels 1-5 mm long. 
Flowers with hypanthium 2-3 mm long, densely puber- 
ulent with straight or curved multicellular hairs 0.2-0.5 
mm long, calyx cup to 0.4 mm long, calyx lobes (2-)3- 
4 mm long, 0.8-1.5 mm broad, lanceolate, with erect 
glands ca. 0.3 mm long in the sinsues; corolla white or 
rose red, tube 4-10 mm long, ca. 1.3 mm diam., short 
villose externally, glabrous within, lobes 4, 2-5 mm long, 
ca. 0.7 mm broad, ovate to triangular, densely villous 
externally; stamens 4, anthers ca. 0.7 mm long. Fruits 
5-7 mm long, 4-6 mm broad, broadly obovoid to subglo- 
bose, rounded and truncated at the apex, with few scat- 
tered hairs; seeds 2-3 mm long, oblong and with a thin 
translucent wing, body of the seed ca. 1 mm long and 
oblong. 

Plants of evergreen and partly deciduous forest 
formations in the central Cordilleras, from ca. 1 000 
to 1 600 m elevation. Probably flowering and fruit- 
ing throughout the year. The species ranges from 
southern Mexico to western Panama. 

Manettia flexilis is recognized by its smaller cap- 
sules, winged seeds, puberulent stems and leaves, 
and mid-elevation habitats. This species is infre- 
quently collected. 



Manettia reclinata Mutis in I ... Mant. PI. 2: 558. 
1771. Nacibea coccinea Aubl., Hist. pi. Guiane 
96, t. 37, f. 1. 1775. M. coccinea (Aubl.) Willd. 
in L., Sp. PI. ed. 4, 1: 624. 1797. M. cuspidata 
Bertero in Spreng., Syst. Veg. 1:415. 1825. M. 
panamensis Duchass. & Walp., Linnaea 23: 753. 
1850. M. costaricensis Wernham, J. Bot. 56, 
suppl. 1: 38. 1919. M. orbifera Wernham, loc. 
cit. 41. 1919. M, seleriana Loes., Verhand. Bot. 
Vereins Brandenb. 65: 107. 1923. Figure 1. 

Vines to 4 m high, stems often with 4 narrow longi- 
tudinal ridges, leafy stems 0.7-2.5 mm thick, sparsely 
puberulent (rarely glabrous), often with a row of minute 
(0. 1-0.2 mm) hairs along the longitudinal ridges; stipules 



1-2 mm long, broadly triangular, minutely puberulent. 
Leaves with petioles 2-14(-20) mm long, 0.6-1.2 mm 
broad, sparsely puberulent; leaf blades (2-)3- 10 cm long, 
l-3(-5) cm broad, narrowly ovate to ovate-elliptic or 
lanceolate, acute or acuminate at the apex, obtuse to 
acute at the base, leaves drying thinly chartaceous and 
dark green above, paler greenish gray beneath, usually 
glabrous above and below, with 4-6 major secondary 
veins strongly ascending on each side. Inflorescences ax- 
illary or terminal, usually solitary with 2-4 flowers per 
node, peduncles 0.5-2.5 cm long, puberulent, pedicels 
10-25 mm long (to 35 mm in fruit). Flowers with hy- 
panthium 3-5 mm long, oblong, calyx tube to 1 mm 
long, calyx lobes 4-8, unequal. 4-7 mm long, 0.3-1.3 
mm broad, linear to lanceolate; corolla red, rose red. 
deep pink, scarlet, or magenta, tube 6-13 mm long, 2- 
2.5 mm diam., glabrate to densely minutely puberulcnt 
externally, with yellowish hairs in throat, corolla lobes 
4 (5, 6), 2-4 mm long, 2.5-4.5 mm wide, ovate, puber- 
ulent externally and glabrous within; stamens 4 (5. 6), 
filaments ca. 1 mm long, anthers ca. 3 mm long; style 
ca. 1 1 mm long. Fruits 6-10 mm long, 6-9 mm broad, 
obovoid or subglobose, glabrous or pubescent with thin 
curled hairs to 0.5 mm long, with 8 longitudinal ribs (2 
in the sulci), calyx lobes persisting and recurved; seeds 
2-2.5 mm diam., thin-discoid and orbicular, with an 
erose translucent wing around the margin, body of the 
fruit 0.7-1 mm diam. 



Plants of wet evergreen forest formations, 0- 
1300 m elevation. Flowering in October-May; 
fruiting in December-May. The species ranges from 
southern Mexico and the West Indies into north- 
ern South America. 

Manettia reclinata is recognized by its larger red 
or pink flowers, few flowers per node, usually gla- 
brous leaves, larger rounded capsules borne on 
long pedicels, flat orbicular seeds with thin winged 
margin, and vining habit. This species appears to 
be much more common than its congeners in Cos- 
ta Rica. 



Mitracarpus Zuccarini 

Annual or perennial herbs, erect or decumbent, stems 
tctragonous in cross-section, glabrous or puberulent; 
stipules adnate to the petiole base to form a sheath with 
3-15 slender distal setae (awns), persistent. Leaves op- 
posite and decussate, subsessilc to short-petiolate, leaf 
blades usually narrow (linear to ovate), chartaceous, 
domatia absent. Inflorescences axillary or terminal, cap- 
itate or glomerulate with densely crowded flowers, the 
heads sometimes subtended by 4 leaf-like bracts, flowers 
sessile or subsessile. Flowers bisexual and monomor- 
phic, very small, hypanthium turbinate to subglobose. 
calyx tube short, calyx lobes 4(-5), unequal, persistent; 
corolla white, salverform or funnelform. tube usually 
with a ring of hairs within, glabrous or villous in the 
throat, corolla lobes 4 (3), valvate in bud; stamens 4, 
inserted on the throat, anthers oblong to linear, dorsi- 
fixed, included or exserted; ovary 2-3-locular, with 1 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



191 



ovule in each locule attached to a peltate placenta in the 
center of the septum, style short or long, with 2 short 
linear stigmas. Fruits thin-walled capsules, 2- or 3-locular 
with circumscissile or transverse dehiscence from below 
the middle (the upper portion breaking away with the 
persisting calyx and exposing the seeds), septum per- 
sisting with the basal part of the capsule; seeds ellipsoid 
to oblate or globose, ventral (adaxial) surface with 4 sulci 
radiating from a central area to give an X-like pattern 
demarking 4 broadly rounded areas, abaxial surface 
smooth. 

A Neotropical genus of 30-45 species, with most 
in Brazil. The genus is very similar to Crusea, 
Diodia, and Spermacoce with its weedy-herba- 
ceous habit, broad stipules with distal awns, nar- 
rowly elliptic leaves, small flowers in verticillate 
heads, and two-locular capsules with two seeds. 
Mitracarpus is unusual in the circumscissile de- 
hiscence of the capsule and the X-like (cruciform) 
sulcus on the inner face of the seed. 



Mitracarpus hirtus (L.) DC., Prodr. 4: 572. 1830. 
Spermacoce hirta L., Sp. PI. ed. 2: 148. 1762. 
S. villosa Sw., Prodr. 29. 1788. M. villosus (Sw.) 
Cham. & Schlend., Linnaea 3: 363. 1828. M. 
breviflorus Gray, PI. Wright. 2, 68. 1853. Fig- 
ure 4. 



persisting calyx and top of fruit 2-3.5 mm long, body of 
the yellowish brown capsule ca. 1 mm long, 0.6-1 mm 
broad; seeds 0.5-0.8 long and ca. 0.6 mm wide, oblong- 
oblate, yellowish brown, with 4 sulci forming an im- 
pressed X on the adaxial face, smooth or pitted on the 
abaxial face. 

Common weedy plants of open disturbed sites 
in both deciduous and evergreen formations, 0- 
1200(-1400) m elevation. Flowering primarily in 
June-December in Costa Rica. The species is found 
in the southwestern United States and throughout 
tropical America; it has become established 
through much of tropical Africa and has been found 
in India, Burma, and the western Pacific. 

Mitracarpus hirtus is recognized by its herba- 
ceous weedy habit, setose sheathing stipules, nar- 
row hispidulous leaves with few secondary veins, 
axillary heads of small congested flowers, and un- 
usual fruit and seed. It is found in both the Ca- 
ribbean and Pacific lowlands and the central up- 
lands. There has been considerable confusion 
regarding the nomenclature of this species and es- 
pecially the applicability of the names; we follow 
the recent annotations of C. Dennis Adams. These 
plants can be mistaken for species of Spermacoce, 
Diodia, and Hyptis (Labiatae). 



Herbs to 0.6(-1) m tall, erect or spreading, stems usu- 
ally simple distally and with few branches mostly near 
the base, often slightly woody at the base, leafy stems 
0.7-2.5 mm thick, with 4 longitudinal ridges and some- 
what quadrangular in cross-section, usually with curved 
thin hairs 0.2- 1.5 mm long and longer (1.5-2 mm) trans- 
lucent hairs at the nodes; stipules united to the petiole 
base to form a broad sheath l-3(-4) mm long, the straight 
distal margin of the sheath with 6-9(-l 3) linear setae 1- 
5 mm long. Leaves subsessile, petioles 0-3 mm long, not 
clearly differentiated from the base of the blade; leaf 
blades 2-5(-8) cm long, 0.5-1. 5(-2) cm broad, narrowly 
elliptic to lanceolate, or narrowly elliptic-oblong, apex 
tapering gradually and acute, base cuneate, drying stiffly 
chartaceous, glabrescent or sparsely hispidulous above, 
sparsely hispidulous beneath with hairs ca. 0.3 mm long 
and along the margin, 2 veins 2-3/side, strongly as- 
cending. Inflorescences axillary or terminal, flowers 
densely crowded in capitulae or verticils 4-10 mm high 
and 8-15(-20) mm broad, bracts 2-4 mm long but dif- 
ficult to distinguish among the sepal lobes in the tightly 
congested heads, flowers sessile or subsessile. Flowers 
with hypanthium 1-2 mm long, glabrous beneath and 
hispidulous distally, calyx lobes 4, unequal, the 2 larger 
1.5-2.5 mm long, lanceolate, thick centrally and with a 
hyaline margin; corolla white, glabrous or minutely hairy 
externally, tube 1.5-2 mm long, 0.3-0.9 mm diam., lobes 
4, 0.6-1.1 mm long. 0.4-1 mm broad at the base, ovate 
or triangular; stamens 4, filaments very short or absent, 
anthers 0.4-0.7 mm long, usually exserted; style 1-1.6 
mm long, stigma 0.3-0.5 mm long. Fruits with the top 
part coming off to expose the 2 locules and 2 seeds, 



Morinda Linnaeus 

Small to large trees or shrubs (lianas), stems terete or 
quadrangular in cross-section, glabrous or puberulent; 
stipules interpetiolar and sometimes slightly connate to 
form a short sheath above the petioles (intrapetiolar), 
entire to cuspidate or bifid, glabrous. Leaves opposite or 
3/node (sometimes only 1 at a flowering node), petiolate, 
pinnately veined, often slightly succulent in life, usually 
with tufts of hairs (domatia) in the vein axils beneath. 
Inflorescences terminal or axillary, with 1 or 2 (rarely 
more) capitate heads on a common peduncle (the heads 
rarely sessile), large or small bracts sometimes present, 
flowers sessile in the capitulum with the basal parts usu- 
ally united. Flowers bisexual (rarely unisexual), radially 
symmetrical, usually distylous, hypanthium free or unit- 
ed with other flowers, calyx tube urceolate or hemi- 
spheric, often entire distally, calyx lobes usually minute 
or absent; corolla funnelform or salverform, corolla tube 
glabrous or puberulent in the throat, puberulent at the 
base within, corolla lobes 4-7, narrow and valvate in 
bud; stamens 4-7, filaments short and inserted in the 
throat of the corolla, anthers dorsifixed and versatile, 
included or exserted, connective often prolonged distal- 
ly; ovary 2- or 4-locular 1 ovule in each locule, ovules 
erect and attached below the middle or at the base of 
the septum, style slender, with 2 short or long style 
branches. Fruits fused into a fleshy syncarp made up of 
the united ovaries (or their bases), often large and turning 
white, pyrenes 1 -seeded or part of a 2-4-locular woody 
structure; seeds obovoid or reniform. 



192 



FIELDIANA: BOTANY 



A pantropical genus of 50-80 species; most of 
the species are native to the Old World. The flow- 
ering capitula with the basal parts of the flowers 
united, and the latter forming a fleshy largely syn- 
carpous fruiting capitulum, make this genus very 
distinctive among our Rubiaceae. Three species 
have been recorded from southern Central Amer- 



ica; two additional native species (M. asperula 
Standl. and M. yucaianensis Greenm.) are found 
in northern Central America. There is consider- 
able variation within our species, and this can make 
their identification difficult. Current holdings of 
this genus are very limited; all the Costa Rican 
material is from near the seashore. 



Key to the Species of Morinda 

1 a. Leaf blades usually less than 5 cm broad, usually drying grayish or yellowish; branches often scandent 
M. royoc 

Ib. Leaf blades more than 5 cm broad, usually drying very dark in color; branches never clambering 
or scandent 2 

2a. Inflorescences usually solitary, with only 1 capitulum per peduncle, fruiting syncarps 3-1 2 cm diam.; 
largest leaf blades usually more than 20 cm long and 14 cm broad; only found near the ocean shore 
M. citrifolia 

2b. Inflorescences 1-3/node, with 1-3 capitula per primary peduncle, fruiting syncarp 0.5-3 cm diam.; 

largest leaf blades usually less than 20 cm long and 14 cm broad; lowland rain forests 

M. panamensis 



Morinda citrifolia L., Sp. PI. 176. 1753. Figure 19. 

Shrubs or small trees. (l-)2-8(-12) m tall, trunks to 
1 5 cm thick, wood yellow, branchlets quadrangular or 
terete, leafy stems 2.5-12 mm thick, glabrous; stipules 
6-20 mm long. 5-14 mm broad, oblong to suborbicular- 
triangular and rounded distally, glabrous. Leaves well 
spaced along the stem, petioles 12-20 mm long, ca. 2 
mm broad, glabrous, with slightly winged adaxial mar- 
gins; leaf blades 12-28(-40)cm long, 7-16(-24)cm broad, 
oblong to broadly elliptic-oblong or ovate-oblong, apex 
obtuse to acute (very short acuminate), base broadly 
obtuse to cuneate, drying chartaceous or thin-charta- 
ceous, often dark in color, glabrous above and below but 
with tufts of hairs in the vein axils beneath, 2 veins 6- 
8/side. Inflorescences axillary and drying black, solitary 
or 2-3/node, capitulae 9-20 mm long, to 20 mm diam., 
oblong to subglobose, peduncles 10-22(-30) mm long, 
1.2-2 mm thick, glabrous, flowers sessile and united at 
the base. Flowers united basally (hypanthia connate), 
calyx tube minute and truncated with a scarious margin: 
corolla white, slightly thickened, glabrous externally, tube 
6-10 mm long, cylindrical, corolla lobes usually 5, 3-8 
mm long, ca. 1.5 mm wide, obtuse, thick-fleshy; stamens 
4-6, filaments slightly unequal, anthers to 5 mm long, 
becoming twisted; stigma to 5 mm long and 0.8 mm 
wide, erose. Fruits united into a syncarp. 4-1 2 cm diam., 
fleshy succulent and irregularly globose to oblate, white 
with green "eyes" ca. 8 mm diam. formed by the calyx 
and disc of individual flowers, pyrenes to 10 mm diam.; 
seeds 3.5 mm long. 

Plants usually found near ocean beaches and 
lagoons along the Caribbean shore from Honduras 
to Panama, 0-20 m elevation. Flowering and fruit- 
ing throughout the year (Sanchez 1983). The orig- 



inal range of this species was from India to the 
East Indies and northern Australia. It has become 
naturalized in a number of areas around the Ca- 
ribbean. 

Morinda citrifolia is recognized by its large white 
Annona-\\ke syncarps, large opposite leaves on 
thick stems, and seaside Caribbean habitat. Her- 
barium specimens can be difficult to distinguish 
from M. panamensis. Yema de huevo is a common 
name; the fruits are edible. 



Morinda panamensis Seem., Bot. voy. Herald 1 36. 
1854. Figure 19. 

Shrubs or trees. 3-25 m tall, branchlets quadrangular 
in cross-section, leafy stems 1.5-6 mm thick, minutely 
(0.1 mm) farinose puberulent. glabrescent; stipules ap- 
parently with the distal part tearing off to leave a cupulate 
base 1-3 mm long forming a shallow cup around the 
stem, at first broadly oblong (to 10 mm long and 9 mm 
broad) and covering the shoot apex. I^eaves with petioles 
5-25 mm long. 1-2 m thick, glabrous or minutely (0.1 
mm) puberulent; leaf blades 9-21 cm long, 4-13 cm 
broad, broadly elliptic to elliptic-oblong or ovate-oblong, 
apex bluntly short-acuminate with tip to 10 mm long, 
base obtuse or cuneate, drying thin-chartaceous, dark, 
glabrous above, glabrous or minutely (0.2-0.3 mm) pu- 
berulent beneath and with dense tufts of hairs (domatia) 
in the vein axils. 2 veins 4-7/side. central 2 veins 1-4 
cm distant. Inflorescences 1-3/node, capitate, peduncles 
4-35(-55) mm long, 0.7-1.5 mm thick, the primary pe- 
duncle simple or with 3-4 equal or unequal branches to 
3 cm long, usually glabrous, each capitulum 4-10 mm 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



193 



long and 5-10 mm wide, oblong to globose, with 9 flow- 
ers or more, flowers sessile and united. Flowers united 
below, glabrous externally (rarely minutely puberulent) 
and drying black, hypanthium partly free distally for 1- 
2 mm, calyx tube ca. 0.3 mm long and subentire; corolla 
white, tube 5-10 mm long, lobes 4, 3-6 mm long, 1-2 
mm broad, narrowly oblong; stamens 4, anthers 2.5-3.5 
mm long; stigmas 3.5-5 mm long. Fruit a syncarp, 15- 
30 mm diam., subglobose, the calyx tube little (0-0.5 
mm) elevated above the surface of the fruit and 2.5-4 
mm diam. 

Plants of wet evergreen lowland rain forest for- 
mations, 0-600 m elevation. Flowering in March- 
May in Central America; fruiting in July-Septem- 
ber. This species ranges around the Gulf of Mexico 
and the Caribbean from Florida (U.S.A.), Mexico, 
Central America, and the West Indies to Panama. 

Morinda panamensis is recognized by its un- 
usual capitula of flowers with united ovaries, seeds 
imbedded within a globose syncarp, thin leaves 
drying very dark, and rain forest habitat. Some 
herbarium specimens can be very similar to spec- 
imens of M. citrifolia, and it is possible that there 
are intermediates in nature. An unusual collection 
from near Upala (Herrera 1783 CR, MO) is mi- 
nutely puberulent on all parts. 



Morinda royoc L., Sp. PI. 176. 1753. Figure 19. 

Shrubs, vines, or small trees, l-3(-7) m tall, often with 
scandent branches, leafy branchlets 1-5 mm thick, young 
stems with obscure longitudinal ridges and quadrangular 
or terete, minutely puberulent with thin hairs 0. 1 mm 
long; stipules l-2(-4) mm long, 3-5 mm wide at the 
base, triangular or with an awn 1-2 mm long, glabrous 
and drying yellowish brown. Leaves with petioles 4-14 
mm long, 0.5-1 mm broad, with adaxial margins, gla- 
brous or minutely puberulent; leaf blades 4-1 1(-13) cm 
long, 1-4.5 cm broad, narrowly elliptic-oblong to oblan- 
ceolate or linear-oblanceolate, usually 4 times longer than 
broad, apex acute to short-acuminate, base gradually 
narrowed and acute or attenuate, drying stiffly charta- 
ceous, grayish or yellowish, with the margin often in- 
volute, glabrous above and below but often with barbate 
hairs in the leaf axils and along the midvein, 2 veins 3- 
6/side, arcuate and weakly loop-connected near the mar- 
gin. Inflorescences solitary and axillary, the capitulae 4- 
12 mm diam., usually oblong, sessile or with peduncles 
0-7(-10) mm long, minutely puberulent, flowers sessile 
and united. Flowers united together in the lower half of 
their ovary, free portion of the hypanthium and calyx 
1-3 mm long, calyx entire distally (or obscurely 5-lobed), 
minutely puberulent; corolla white, 6-8 mm long, tube 
ca. 5 mm long and 2 mm diam., cylindrical, minutely 
(0. 1 mm) puberulent externally, lobes 1-2 mm long; sta- 
mens 5-6, anthers ca. 2 mm long; style 3-5 mm long, 
stigmas ca. 1.5 mm long. Fruit a syncarp, 8-25 mm 
diam., irregularly globose, pyrenes 5 mm long. 



Plants of the Caribbean lowlands (from pine 
savannas to evergreen forest formations) in Cen- 
tral America. Flowering in January-August. The 
species ranges from Florida (U.S.A.), Mexico, 
Central America, and the West Indies to northern 
South America. 

Morinda royoc is recognized by its vining 
branches, smaller narrow leaves that rarely dry 
very dark, small syncarpous capitulae, and small 
stipules. We have seen no specimens from Costa 
Rica, but the species has been collected near Blue- 
fields, Nicaragua, and in central Panama. Most 
Central American collections are described as 
vines. Appunia guatemalensis can appear quite 
similar, but the branches are not vining and the 
fruit develop separately. 



Mussaenda Linnaeus 

Shrubs, erect or climbing; stipules interpetiolar, united 
and solitary or separate and paired (4/node). Leaves op- 
posite, petiolate, pinnately veined and often acuminate 
at the apex, domatia absent. Inflorescences terminal, cy- 
mose, bracts and bracteoles deciduous; a few flowers of 
the inflorescence with a single greatly expanded colorful 
and leaf-like expanded sepal lobe. Flowers bisexual and 
radially symmetrical (except for those flowers where 1 
sepal lobe is greatly enlarged), calyx tube turbinate or 
ovoid, calyx lobes 5, 1 lobe greatly expanded in a few 
flowers of most species; corolla narrowly funnelform or 
salverform, glabrous or puberulent on the outer surface, 
corolla tube puberulent in the throat, corolla lobes 5; 
stamens 5, filaments very short, borne near the base or 
the upper part of the tube, anthers sagittate at the base; 
ovary 2-locular with many ovules, style 2-branched. Fruits 
usually fleshy and indehiscent (rarely dry and loculici- 
dally dehiscent); seeds small and ellipsoid. 

A tropical Old World genus of about 200 spe- 
cies. The greatly enlarged and colorful sepal lobes 
on a few flowers of each inflorescence are found 
in most species of the genus and account for the 
ornamental appeal of the following species. 



Mussaenda erythrophylla Schumach. & Thonn., 
Beskr. Guin. PI. 116. 1827. Figure 16. 

Shrubs or woody climbers, 1.5-3(-8) m tall, leafy 
branchlets 2-6 mm thick, densely velutinous with yel- 
lowish hairs ca. 1 mm long, terete; stipules 4/node, 6- 
1 1 mm long, ovate-triangular, glabrous on the inner face. 
Leaves with petioles 4-18 mm long, with hairs to 1.5 
mm long; leaf blades 4-12(-15) cm long, 3-7 cm long, 
ovate to broadly ovate-elliptic or ovate-orbicular, apex 
short-acuminate, base obtuse or rounded and subtrun- 
cate, densely puberulent on both surfaces with slender 



194 



FIELDIANA: BOTANY 



hairs 0.5-2 mm long, 2 veins 5-7/side, arcuate-ascend- 
ing. Inflorescences ca. 10 cm long and 20 cm broad, at 
the ends of distal un branched leafy stems, with opposite 
or trichotomous branches, densely velutinous. Flowers 
densely pubemlent externally, calyx lobes 6- 1 2 mm long, 
1-3 mm broad, red; the enlarged sepal lobe leaf-like, 
4.5-6.5 cm long, 3-6.5 cm broad, on petioles 2-8 mm 
long, with ca. 7 palmate veins, bright red; corolla yellow 
to pink, tube 15-30 mm long, lobes 3-9 mm long, 4-7 
mm broad, broadly ovate, whitish and papillate puber- 
ulent within. 



Popular small shrubs or climbers grown for or- 
nament in parks and gardens. The enlarged leaf- 
like sepal lobes are bright crimson to deep red and 
give a very colorful effect. They flower throughout 
the year but usually do not produce fruits. 



4 cm diam., peduncles 2-4 cm long, flowers many and 
closely congested. Flowers not conivent basally; corolla 
7-8 mm long, narrowly funnelform, lobes 5, ca. 1.5 mm 
long; style long-exserted. Fruits tightly congested in the 
spherical heads, splitting into 4 parts, seeds minute. 

The solitary pedunculate globose heads and 
larger drooping leaves distinguish this introduced 
species. It is fast-growing in open sites until it 
reaches about 10 m in height. Common names 
used in southeast Asia are kadam, kedam. and 
laran. The literature of this tree is to be found 
under Anthocephalus, now a synonym ofBreonia. 
See J. D. E. Fox, Anthocephalus chinensis, the La- 
ran Tree of Sabah. Econ. Bot. 25: 221-233. 1971. 



Neoiamarckia Bosser 



Nertera Banks & Solander 
Nomen conservandum 



Trees; stipules interpetiolar, triangular, deciduous. 
Leaves opposite, petiolate, entire, domatia absent. In- 
florescences terminal, usually solitary, capitate and pe- 
dunculate, flowers densely congested. Flowers bisexual 
and radially symmetrical, calyx lobes small; corolla fun- 
nelform, corolla lobes 45, imbricate in bud; ovary 2- 
locular at the base, placentation axile with many vertical 
ovules; stigma fusiform. Fruits capsules, loculicidally de- 
hiscent from the apex, thin-walled; seeds 1-5 in each 
locule, small and angular. 

A genus of two species of southeast Asia. One 
species is occasionally planted in Costa Rica. 

Neoiamarckia cadamba (Roxb.) Bosser, Bull. Mus. 
Hist. Nat. Paris 4 ser. sect. B. Adansonia 6: 247. 
1984. Nauclea cadamba Roxb., Fl. Ind., ed. 1, 
2: 121. 1824. Anthocephalus cadamba (Roxb.) 
Miq., Fl. Ind. Bat. 2: 135. 1856. A. morindae- 
folius Korth., Verh. Nat. Gesch. 1 54, t.. 48. 1 842. 
A. indicus A. Rich., Mem. Soc. Hist. Nat. Paris 
5: 238. 1834, nom. illeg. Cephalanthus chinensis 
a uctt.. non Lamarck; A. chinensis auctt. 

Trees, 5-15(-30) m tall, fast-growing in early stages, 
leafy stems 2.3-10 mm thick, glabrous, quadrangular; 
stipules oblong, covering the buds, caducous. Leaves 
usually somewhat pendant, deciduous, petioles 22-55 
mm long, 1-4 mm thick, glabrous or minutely puberu- 
lent, drying dark; leaf blades (7-) 13-3 4 cm long, (5-)6.5- 
18 cm broad, ovate to ovate-oblong or oblong, apex 
abruptly narrowed and blunt or short-acuminate, base 
obtuse to rounded and truncate or subcordate, drying 
brownish, glabrous above, glabrous or minutely (0.05 
mm) puberulent beneath, 2 veins (4-)8- 1 6/side, 3" veins 
subparallel. Inflorescences solitary, terminal on short lat- 
eral branchlets, each with a single globose capitulum 3- 



Perennial herbs, creeping and repent, glabrous or 
sparsely puberulent, much-branched and often rooting 
at the nodes (often forming mats); stipules interpetiolar, 
small, partly united to the petiole bases, entire or with 
2 teeth, persistent. Leaves opposite, very small, petiolate 
or sessile; leaf blades ovate to rounded, glabrous or pu- 
berulent, venation pinnate or subpalmate; domatia ab- 
sent. Inflorescences of solitary flowers, axillary or ter- 
minal, the flowers sessile or subsessile. Flowers bisexual, 
radially symmetrical, hypanthium ovoid to turbinate, 
calyx tube truncated and entire or slightly lobed; corolla 
broadly funnelform to tubular, corolla lobes 4-5, valvate 
in bud; stamens with thin filaments attached to the base 
of the tube, anthers basifixed, the connective apiculate. 
exserted; ovary 2-locular, with 1 ovule borne from the 
base of each locule, style deeply 2-branched and slender. 
Fruits fleshy drupes (rarely dry), rounded, with 2 plano- 
convex pyrenes. 

A genus of 6- 1 2 species found in Australia, New 
Zealand, Malaya, Indonesia, South China, and 
some Pacific islands; in addition, a single species 
ranges from Mexico through the higher moist 
mountains of Central and South America to Chile. 



Nertera granadensis (Mutis ex L.f.) Druce, Bot. 
Soc. Exch. Club Brit. Isles 1916: 637. 1917. Go- 
mozia granadensis Mutis ex L.f., Suppl. PI. 1 29. 
1781. Nertera depressa Banks & Solander ex 
Gaert.. Fruct. et Sem. PI. 1 : 1 24. 1 788. Figure 3. 

Creeping prostrate or pendant herbs to 5 cm tall and 
up to ca. 1 m long, terrestrial or low epiphytic, often 
forming mats, stems 0.3-1 mm thick and rooting at many 
nodes, often tetragonous in cross-section in life, glabrous 
or with a few scattered hairs; stipules 0.3-1 mm long, 
triangular, united at the base with the bases of the pet- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



195 



ioles, persisting. Leaves quite variable on different plants, 
petioles 0.7-6(-9) mm long, 0.2-0.4 mm broad, gla- 
brous, sometimes sulcate above; leaf blades (1.5-)2.5- 
8(-13) mm long, (1.2-)2-7(-12) mm broad, ovate-tri- 
angular to ovate-orbicular or ovate-deltoid, apex obtuse 
to rounded, sometimes with a minutely apiculate tip, 
base rounded and truncate to obtuse, often decurrent on 
petiole in larger leaves, drying thin-chartaceous but 
semisucculent in life, glabrous above and below (rarely 
puberulent), 2 veins 2-4/side, strongly ascending. Inflo- 
rescences absent, the solitary sessile flowers axillary to 
distal leaves, the flowers covered by surrounding leaves 
and very difficult to see in pressed herbarium material. 
Flowers minute (ca. 4 mm long), usually glabrous exter- 
nally, hypanthium 0.5-1 mm long, calyx lobes absent; 
corolla 1-3 mm long, greenish to yellow or white, tube 
0.6-1 mm long, widely funnelform (subcampanulate), 
lobes ca. 0.5 mm long, glabrous; anthers 0.3-0.5 mm 
long. Fruits sessile, 4-7 mm diam., globose, bright or- 
ange, orange-red, or deep red, fleshy part often translu- 
cent, pyrenes 2-2.5 mm long and 1.2-2 mm wide, ob- 
long. 

Plants of wet evergreen montane forest forma- 
tions, from (1000-)! 500 to 3400 m elevation in 
Central America. Fruiting throughout the year in 
Costa Rica. This species ranges from Mexico, 
through the high mountains of Central America 
and South America, as far south as Chile. This 
species, when interpreted in a broad sense, is said 
to be found throughout the range of the genus. 

Nertera granadensis is a very distinctive species 
with its low-creeping and mat-forming habit, very 
small paired leaves, lack of pubescence, solitary 
little flowers, bright orange-red berries, and re- 
striction to higher-elevation wet forest habitats. 
Collections with larger (10 mm) leaves can look 
very different from those with smaller (4 mm) 
leaves. It is common in some montane rain forests, 
covering mossy banks and old logs. In Costa Rica 
it seems to be restricted to the central Volcanic 
Highlands and the Cordillera de Talamanca. The 
brightly colored berries and mat-forming ability 
have made this species valuable as a greenhouse 
or moist-area ornamental. These plants resemble 
small collections of Didymaea. 



Oldenlandia Linnaeus 

REFERENCES E. E. Terrell, Synopsis of Olden- 
landia (Rubiaceae) in the United States. Phyto- 
logia 68: 125-133. 1990. E. E. Terrell and W. H. 



Lewis, Overview and annotated list of North 
American species of Hedyotis, Houstonia, Olden- 
landia (Rubiaceae) and related genera. Phytologia 
71: 221-243. 1991. D. A. Halford, Review of the 
genus Oldenlandia L. (Rubiaceae) and related gen- 
era in Australia. Austrobaileya 3: 683-722. 1992. 

Annual or perennial herbs (rarely subshrubs), stems 
erect or decumbent, simple or branched, glabrous or 
puberulent; stipules interpetiolar, small, acute to acu- 
minate, often united to the base of the petioles to form 
a short sheath, with 1 -several awns. Leaves opposite and 
decussate, sessile or short-petiolate; leaf blades usually 
narrow, often with a stiff mucronate tip, domatia absent. 
Inflorescences axillary or terminal, open and branched 
panicles and cymes or the flowers solitary or fasciculate 
in the leaf axils, flowers sessile to long-pedicellate. Flow- 
ers bisexual, monomorphic or distylous, usually small, 
hypanthium turbinate to hemispheric, calyx lobes 
(3-)4(-5-8), equal, narrowly to broadly triangular; co- 
rolla rotate or salverform to funnelform, white to lav- 
ender, pink, or purple, tube cylindrical, the throat often 
puberulent, corolla lobes (3-)4(-5), valvate in bud; sta- 
mens (3-)4(-5), anthers dorsifixed and sessile or on short 
filaments inserted on the throat, included or exserted; 
ovary 2-locular, usually with many horizontal ovules on 
peltate placentas attached near the base of the septum, 
style filiform, stigmas 2, linear to subglobose. Fruits cap- 
sular, often papery, globose to oblong, usually with a 
loculicidally dehiscent apex (beak), later also septicidally 
dehiscent; seeds usually many (50-100), angular to 
subglobose, smooth to reticulate or alveolate, often be- 
coming viscid when moistened. 

A pantropical and subtropical genus with prob- 
ably ca. 100 species but with problems regarding 
generic circumscription. Some authors have sug- 
gested placing this genus under Hedyotis or Hous- 
tonia, while others divide it into smaller genera; 
see the references cited above. Our plants of this 
genus are recognized by their delicate herbaceous 
habit, the very small or linear-lanceolate leaves, 
the axillary flowers or small few-flowered inflo- 
rescences, the minute flowers on filiform pedicels, 
and the broadly rounded thin-walled capsules with 
small seeds. 

Many Costa Rican collections formerly identi- 
fied as O. corymbosa, O. herbacea, and O. land- 
folia are probably the same species: O. corymbosa. 
This conclusion is based on an overview of the 
material and following the keys and descriptions 
of Verdcourt in the Flora of Tropical East Africa 
(1976). 



Key to the Species of Oldenlandia 

la. Leaf blades less than 5 mm long, usually broadly ovate; plants often forming small mats (not yet 
collected in Costa Rica) O. callitrichoides 



196 



FIELDIANA: BOTANY 



1 b. Leaf blades more than 5 mm long, narrowly elliptic-ovate to lanceolate or linear; plants diffuse or 
foming loose mats 2 

2a. Corolla 2-1 1 mm long; fruit with a beak 0.8-1 mm long (not yet known to occur in Costa Rica) 
O. herbacea 

2b. Corolla 0.5-2 mm long; fruit with a small or well-developed beak 3 

3a. Flowers pink or white marked with lavender, flowers often 2-3/peduncle; fruit globose to globose- 
oblate, rounded at the base but not saccate, with a beak 0.1-0.6 mm high O. corymbosa 

3b. Flowers white, flowers usually solitary; fruit broader than long and distinctly rounded (saccate) at 
the base, with a beak ca. 1 mm long O. lancifolia 



Oldenlandia callitrichoides Griseb., Mem. Am. 
Acad. 2, 8: 506. 1863. Oldenlandiopsis callitri- 
choides (Griseb.) Terrell & W. H. Lewis, Brit- 
tonia42: 185. 1990. 

Prostrate herbs to 10 cm tall, much-branched and of- 
ten forming mats, leafy stems 0.1-0.5 mm thick (when 
dried), glabrous; stipules adnate to petiole base and form- 
ing a short sheath to 0.5 mm long, sheath entire distally 
or with a small appendage, glabrous. Leaves very small, 
petiole 0.5-2 mm long; leaf blades 1-3.5 mm long, 1-3 
mm broad, rounded-triangular to ovate, apex bluntly 
obtuse or rounded, base broadly obtuse to rounded and 
subtruncate, somewhat decurrent on petiole, drying 
membranaceous, greenish, glabrous above and below (or 
with a few broad-based hairs ca. 0.2 mm long), 2 veins 
3-4/side, strongly ascending. Inflorescences of solitary 
flowers in distal leaf axils, usually only 1 flower per node, 
pedicels 1-9 mm long, filiform, glabrous. Flowers 2-3 
mm long, hypanthium 1-1.5 mm long, calyx lobes 0.2- 
0.6 mm long. Fruits 2-3 mm long (including the calyx 
lobes), ca. 1 mm diam., distally truncated, with a few 
whitish raphides on the greenish surface, calyx lobes ca. 
0.5 mm long. 

Distinctive little plants usually found on sandy 
soil in open sunny situations, in tropical lowland 
sites. This species has the smallest leaves found 
among Central American Rubiaceae. In general 
aspect, these plants resemble Pilea hernariarioides 
(Sw.) Weddell and P. microphylla (L.) Liebm. This 
species is known from the Yucatan peninsula and 
from central Panama, but it has not been collected 
elsewhere in Central America. Terell and Lewis 
based a new genus on this distinctive species (see 
synonomy). 



Oldenlandia corymbosa L., Sp. PI. 1 19. 1753. Fig- 
ure 3. 



liptic-oblong, apex abruptly obtuse to acute and with a 
minute (0.2 mm) apiculate tip, narrowed to the base, 
drying chartaceous and with the margins usually revo- 
lute, dark above, glabrous or minutely scabrid above and 
below (or with the cystoliths appearing like minute tri- 
chomes), 2 veins usually obscure. Inflorescences axil- 
lary, with single flowers on slender pedicels or cymose- 
umbellate with 3 (2-5) flowers on a common filiform 
peduncle 2-8 mm long, both singular and umbellate in- 
florescences often present on the same stem or node, 
pedicels 2-6(-12) mm long, filiform and glabrous. Flow- 
ers very small, hypanthium 0.5-1 mm long, obconical 
to cupular, glabrous, calyx lobes 4, 0.7-1.5 mm long, 
narrowly triangular, corolla rotate, white or white marked 
with lavender, blue, or purple, tube 0.6-1 mm long, lobes 
0.5-1.2 mm long; stamens 4, sessile on the distal half of 
the tube, anthers ca. 0.4 mm long. Fruits 1.7-2.7 mm 
diam., body of the fruit 1.2-2 mm long and truncated 
at the apex, raphides prominent, calyx lobes ca. 0.5 mm 
long, borne on slender pedicels 4-1 1 mm long; seeds ca. 
0.3 mm long, ellipsoid to depressed conic, reticulate. 

Plants of open weedy sites on sandy soils in 
evergreen and in seasonally deciduous formations, 
from near sea level to 1 500 m. Probably flowering 
and fruiting throughout the year. The species has 
been found in both the Caribbean and Pacific low- 
lands and in the Meseta Central. The species ap- 
pears to have originated in Africa but is now wide- 
spread in the tropics. 

Oldenlandia corymbosa is recognized by its short 
weedy habit, almost linear subsessile leaves, mi- 
nute flowers, and rounded capsules. The senior 
author believes that this is probably the correct 
name for all the Central American material for- 
merly called O. corymbosa, O. herbacea, and O. 
lancifolia. Separation of species on the basis of 
inflorescence differences or whether or not the stip- 
ules are bifid appears to be trivial. 



Herbs to 15(-30) cm tall, prostrate or erect, with few 
to many branches, leafy stems 0.2-1.3 mm thick, gla- 
brous or with a few hairs at the nodes; stipules with a 
sheath 0.3-2 mm long, with 2-5 slender unequal awns 
to 1.5 mm long. Leaves sessile or with petioles to 1 mm 
long; leaf blades (7-)15-30M5) mm long, 0.5-3.5(-6) 
mm broad, linear to linear-oblong or very narrowly el- 



Oldenlandia herbacea (L.) Roxb., Hort. Bengal. 
11. 1814. Hedyotis herbacea L., Sp. PI. 102. 
1753. 

Annual or perennial herbs, 5-60 cm tall, erect to 
spreading and decumbent, stems glabrous, with 4 lon- 



BURGER & TAYLOR: FLORA COSTAR1CENSIS. RUBIACEAE 



197 



gitudinal ridges; stipules forming a very short (0.1-0.5 
mm) sheath, truncate and with a few awns to 0.3 mm 
long. Leaves sessile; leaf blades 6-55 mm long, 1-4 mm 
wide, linear to linear-lanceolate, apex acute, base cu- 
neate, drying chartaceous, glabrous or with a few short 
hairs along the margins. Inflorescences axillary, flowers 
1 or 2/node, pedicels 3-30 mm long, filiform, glabrous. 
Flowers usually isostylous, hypanthium 0.5-1 mm long, 
ovoid, glabrous to papillate or puberulent, calyx lobes 
0.5-2.5 mm long, narrowly triangular, scabridulous on 
the margins; corolla white or lavender, or the tube green 
and the lobes with purple marks, corolla tube 2-1 1 mm 
long, cylindrical, lobes 1-3 mm long, ovate. Fruits 2.2- 
5 mm long, 1.5-2 mm diam.. subglobose to ovoid, drying 
pale yellowish, glabrous to puberulent, crowned by the 
dark calyx lobes, beak 0.8-1 mm long; seeds 0.2-0.4 mm 
long, ovoid to ellipsoid, reticulate, brown. 

Weeds of open sunny sites. Originally from Af- 
rica but now naturalized in Asia and parts of the 
Americas. The above description is based on 
Verdcourt (1976). This species has not yet been 
recorded from Costa Rica. Central American ma- 
terial earlier placed under this name is likely to be 
Oldenlandia corymbosa or O. lancifolia. 



Oldenlandia lancifolia (Schumach.) DC., Prodr. 4: 
425. 1830. Hedyotis lancifolia Schumach. in 
Schumach. & Thonn., Beskr. Guin. PI. 72. 1 827. 
Manettia bocaturensis Dwyer, Ann. Missouri 
Hot. Gard. 67: 278. 1980. 

Perennial (rarely annual) herbs to 90 cm long, pros- 
trate or creeping, usually much-branched near the base 
and with simple distal stems, leafy stems glabrous or 
rarely minutely scabridulous; stipules forming a sheath 
to 1 mm long, with 2-5 slender awns 0.5-1.5 mm long. 
Leaves sessile or subsessile; leaf blades 1 0-60 mm long, 
2-12 mm broad, linear or linear-lanceolate to narrowly 
elliptic, apex acute, base cuneate, drying chartaceous, 
dark green above, glabrous on both surfaces but often 
minutely scabrid along the revolute margins, 2 veins 3- 
5/side. strongly ascending but thin and difficult to see. 
Inflorescences axillary, of solitary flowers or several flow- 
ers on very short peduncles (or reduced lateral shoots), 
pedicels 5-30 mm long, filiform, glabrous or scabridu- 
lous. Flowers monomorphic, ca. 3 mm long, hypanthium 
ca. 0.8 mm long and 1.5 mm diam., cupulate, glabrous 
or with scattered short hairs, calyx lobes 1-1 .8 mm long, 
triangular, glabrous or scabridulous; corolla white (some- 
times tinged with pink or purple), tube ca. 1 mm long, 
lobes 1-2 mm long; stigma lobes 0.7-1 .4 mm long. Fruits 
2-3 mm long (including the 1 mm tall beak), 3.2-5 mm 
in diam., depressed subglobose; seeds 0.3-0.4 mm long. 

Herbs of both seasonally deciduous formations 
and lowland rain forest areas. The species is wide- 
spread in tropical Africa and has been introduced 
to parts of the South America and the West Indies. 
The preceding description follows that of Verd- 



court (1976). Central American material earlier 
ascribed to this species is probably O. corymbosa. 
Likewise, Steyermark's use of this name in the 
Flora de Venezuela (1974, pp. 408-41 1), may be 
incorrect. 



Osa Aiello 

REFERENCE A. Aiello, A re-examination of 
Portlandia and associated taxa. J. Arnold Arbor. 
60: 38-126. 1979. 

Small trees, stems slightly expanded at the nodes; stip- 
ules united (interpetiolar), small, with an acute central 
lobe. Leaves opposite, petiolate, attenuate at the apex, 
drying thin-chartaceous and grayish green, entire, pin- 
nately veined, domatia absent. Inflorescence of single 
axillary flowers, borne on pedicels continuous with the 
base of the hypanthium/ovary. Flowers large, radially 
symmetrical, glabrous, calyx with 6 long narrow lobes; 
corolla with a long tube and distally funnelform, corolla 
lobes 5, broadly triangular; stamens 5, anthers linear; 
ovary 2-locular, placentas borne on the septum, with ca. 
1 ovules in each locule, style filiform. Fruits thin-walled 
capsules, ellipsoid, with 6 longitudinal ribs, apparently 
opening septicidally, the calyx lobes persisting distally; 
seeds biseriate, slightly compressed, lacking wings, not 
imbricate, with persisting funicle. 

A monotypic genus known only from the Osa 
Peninsula of Costa Rica. No other species of Costa 
Rican Rubiaceae has such large flowers. This ge- 
nus is related to Portlandia and to a lesser extent 
to Hintonia. 



Osa pulchra (D. Simpson) Aiello, J. Arnold Arbor. 
60: 116. 1979. Hintonia pulchra D. Simpson, 
Phytologia 29: 277. 1974. Figure 15. 

A small tree or slender treelet, 2.5-15 m tall, leafy 
stem 1 .5-6 mm thick, glabrous, slightly expanded below 
the node; stipules 1-3 mm long, the broad base 1-2 mm 
high with a narrowed acute tip 0.5-1.5 mm long, per- 
sisting. Leaves with petioles 4-1 2 mm long, 0.7-1 .5 mm 
thick, glabrous, poorly differentiated from the lamina 
base; leaf blades 12-19 cm long, 3-6 cm broad, elliptic- 
oblong or narrowly oblong, apex gradually narrowed and 
acuminate, base acute to attenuate, drying thin-charta- 
ceous and grayish green, glabrous above and below, 2 
veins 6-8/side, 3 veins obscure, domatia lacking. Inflo- 
rescences of solitary flowers in the axils of distal leaves, 
pedicels ca. 1 5 mm long but merging imperceptibly into 
the flower base, ca. 1 mm diam., glabrous, drying black. 
Flowers glabrous externally, probably slightly pendu- 
lous, apparently homostylous, hypanthium ca. 6 mm 
long, obconic, calyx lobes 18-40 mm long, 0.5-2.5 mm 
broad; corolla long-tubular and funnelform distally 
(trumpet-shaped), white, tube 1 7-27 cm long, 4-5 mm 



198 



FIELDIANA: BOTANY 



diam. and gradually flaring to 80 mm diam. at the mouth, 
lobes ca. 1 5 mm long and 3040 mm broad at the base, 
broadly obtuse; anthers more than 20 mm long, 0.7 mm 
thick. Fruits ca. 3 cm long and 1.5 cm broad, oblong- 
ellipsoid, dark brown, the sepal lobes persisting but 
breaking off; seeds ca. 6 mm long, testa tuberculate. 

This species is known from only a few collec- 
tions in lowland rain forest near Rincon de Osa 
at ca. 50 m elevation on the Osa Peninsula. Flow- 
ering in January-February, with immature fruits 
in June and mature fruits in January. 

Osa pulchra is distinguished by its very large 
trumpet-shaped flowers that are thin in texture. 
No other Costa Rican member of the Rubiaceae 
has so long a flower. The long narrow basal tube 
suggests pollination by a long-tongued sphingid 
moth. The flowers are reminiscent of those of or- 
namental species ofBrugmansia (formerly includ- 
ed in Datura, Solanaceae). 



Palicourea Aublet 

REFERENCE C. M. Taylor, Revision of Pali- 
courea in Mexico and Central America. Syst. Bot. 
Monogr. 26: 1-102. 1989. 

Shrubs or small trees, glabrous or pubescent, stems 
terete, trigonous or quadrangular; stipules interpetiolar 
and often also intrapetiolar to form a sheathing tube with 
2 triangular lobes or awns on each side (4/node) or some- 
times elongated interpetiolarly into emarginate or bi- 
lobed apices with the intrapetiolar sheath poorly devel- 
oped, colleters present within at the base. Leaves opposite 
and decussate or rarely 3-4/node, petiolate or rarely ses- 
sile; leaf blades entire and often elliptic, apex acute to 
acuminate, base rounded to acute, drying chartaceous to 
subcoriaceous, pinnately veined, domatia present or ab- 
sent. Inflorescences terminal, paniculate with opposite 
or alternate branching, variable in form (from elongate 
and racemose to thyrsoid or broadly corymbose) with 
flowers usually in distal cymose groups, peduncles and 
branches of the inflorescences often colored red, orange, 
yellow, purple, or blue (rarely green), glabrous or pu- 
bescent, bracts and bracteoles usually present (in Costa 
Rica), flowers usually pedicellate. Flowers bisexual and 
usually distylous, radially symmetric or slightly bilat- 



erally symmetric when the corolla is curved or gibbous 
on 1 side, glabrous or pubescent externally, hypanthium 
usually turbinate, calyx tube with 5 calyx lobes (rarely 
truncated or spathaceous); corolla yellow, orange, red, 
purple, or blue (rarely white or cream), tubular to fun- 
nelform or salverform, membranaceous to camose. the 
corolla tube straight or curved, usually expanded (gib- 
bous) near the base, glabrous or puberulent externally, 
usually with a ring of hairs within the lower half of the 
tube (more rarely with the hairs in the upper half), corolla 
lobes 5, valvate in bud; stamens 5, inserted in the middle 
or on the throat of the tube, anthers dorsifixed and bifid 
at the base, included or exserted; ovary 2(-6)-Iocular, 
with 1 erect ovule from a basal placenta in each cell, 
stigmas 2-branched. Fruits fleshy, exocarp bluish to blu- 
ish black or purplish black; pyrenes usually 2. hemi- 
spheric with ca. 5 longitudinal ribs on the rounded back, 
usually with a longitudinal sulcus on the flattened inner 
face. 

A Neotropical genus of about 200 species, rang- 
ing throughout moist tropical vegetation from 
Mexico and the West Indies to southern Brazil 
and Paraguay. Most species are South American; 
there are 3 1 in Costa Rica. As in Hoffmannia. this 
genus has speciated profusely at middle and higher 
(1000-2800 m) elevations. These plants are rec- 
ognized by their colorful terminal inflorescences, 
corolla tubes often slightly bent or expanded at the 
base, ovaries with a single erect ovule in each loc- 
ule, and the fleshy fruit usually with longitudinal 
ribs when dried. The infructescences usually be- 
come purple regardless of their color during an- 
thesis. The stipules forming a short tube above the 
node and with two distal awns or lobes, and the 
leaves with many secondary veins are additional 
characteristics distinguishing many Costa Rican 
species. 

Palicourea is separated from the closely related 
genus Psychotria by the ring of hairs in the interior 
of the lower half of the corolla tube, a tendency 
for the corolla tube to be inflated or gibbous near 
the base, the more colorful inflorescences, and the 
generally larger more colorful corollas. Despite 
these differences, dried fruiting material may be 
very difficult to separate. 



Key to the Species of Palicourea 

la. Plants of lowland formations, rarely found from as high as 1000 m elevation 2a 

Ib. Plants of montane forest formations, from (900-)1000 to 3200 m elevation 5a 

2a. Flowers lavender or white marked with purple; stipule forming a sheath to 8 mm long [leaves 

with 9-15 pairs of secondary veins; from wet Caribbean slopes at 500-1000 m elevation] 

P. copensis 

2b. Flowers red, orange, or yellow; stipular sheath 0-2 mm long; wide-ranging species 3a 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



199 



3a. Leaves 3/node, stipular teeth 6/node, 8-1 2 mm long and persisting, leaf blades to 8 cm broad 

P. triphylla 

3b. Leaves 2/node, stipular teeth or lobes 4, 2-9 mm long, often deciduous, leaf blades to 16 cm 

broad 4a 

4a. Stipular lobes rounded; leaf blades 1 5-28 cm long, to 1 6 cm broad P. guianensis 

4b. Stipular teeth acute; leaf blades 7-14 cm long, to 7 cm broad P. crocea 

5a. Plants pilose with hairs 1.5-2 mm long; stipules united and bifid or emarginate on each side; 

flowers yellow and pilose P. standleyana 

5b. Plants glabrous to puberulent, the hairs not exceeding 1 mm in length; stipules 4-lobed, bifid or 

emarginate; flowers white, yellow, blue, or purplish 6a 

6a. Calyx teeth or calyx lobes regularly more than 1.5 mm long [often persisting in fruit and with 
slightly smaller dimensions; flowers usually blue, purple, or white (red to yellowish only in P. 

macrocalyx and P. orosiana)] 7a 

6b. Calyx teeth or calyx lobes 0.2-1 .5 mm long 1 7a 

7a. Calyx lobes more than 3.5 mm long 8a 

7b. Calyx lobes less than 3.5 mm long 1 5a 

8a. Calyx spathe-like and splitting down 1 side (to 20 mm long); corolla tube 3-4 cm long; 

leaves with 1 2-22 pairs of 2 veins P. spathaceae 

8b. Calyx 5-lobed (not spathe-like); corolla tubes 8-26 mm long; leaves with up to 19 pairs 

of 2 veins 9a 

9a. Corolla 20-26 mm long, ca. 4 mm diam. [calyx lobes 4-14 mm long, lanceolate; bracts 
of the inflorescence to 25 mm long and lanceolate; secondary veins 12-18 pairs; 1400- 

2200 m elevation] P. hammelii 

9b. Corolla tubes less than 20 mm long and 4 mm diam lOa 

lOa. Young stems minutely puberulent 1 la 

lOb. Young stems glabrous 1 2a 

1 la. Calyx lobes 10-14 mm long, corolla white; stipule lobes 7-20 mm long; leaves 

with 15-19 pairs of 2 veins; western Panama P. bella 

1 Ib. Calyx lobes 6-1 1 mm long, corolla yellow; stipule lobes 6-17 mm long; leaves 

with 8-12 pairs of 2 veins; central Costa Rica P. orosiana 

12a. Leaves with 17-20 pairs of 2 veins; stipule lobes 2-9 mm long [corolla white]; floral 

bracts 12-16 mm long; central Costa Rica P. bellula 

12b. Leaves with 7-14 pairs of 2 veins; stipule lobes 2-10 mm long 12a 

1 3a. Floral bracts 10-20 mm long, persisting and enclosing the flowers; corollas white; 

2600-2800 m elevation in the Talamanca mountains P. skotackii 

13b. Floral bracts 6-10 mm long, deciduous; corollas white with yellow or blue; Cor- 
dillera de Tilaran and central volcanic highlands, 1400-2200 m elevation . . 14a 
14a. Corolla white and blue; stipule lobes 2-4 mm long; inflorescence to 22 cm long and 

narrowly thyrsiform P . albocaerulea 

14b. Corolla white and yellow; stipule lobes 4-10 mm long; inflorescences to 15 cm long 

and paniculate P. macrocalyx 

1 5a. Young stems usually puberulent [flowers white; inflorescences narrowly thyrsiform; leaves 

2-7 cm broad; 800-1 600 m elevation] P. lancifera 

1 5b. Young stems glabrous 1 6a 

16a. Leaves l-2(-3) cm broad, blades to 8(-10) cm long; corolla blue and white (compare also 

1 7a below) P. salicifolia 

1 6b. Leaves becoming larger and broader; flowers variously colored 1 7a 

17a. Flowers purple to violet, blue, or white (if reddish then reddish purple or lavender) 18a 

1 7b. Flowers yellow to orange or red-orange 24a 

1 8a. Leaf blades not more than 9 cm long [0.7-3.5 cm broad]; young stems often slightly puberulent 

19a 

1 8b. Leaf blades usually becoming more than 9 cm long; young stems glabrous or puberulent 
beneath the inflorescence . . 20a 



FIELDIANA: BOTANY 



19a. Inflorescences blue, corolla white to violet or blue; leaves thick, petioles thick; 1800- 

3200 m elevation p. adusta 

19b. Inflorescences yellowish green, corolla white to yellow; leaves thin, petioles slender; 

1 300-2300 m elevation p, montivaga 

20a. Leaf blades l-3(-4) cm broad, to ca. 14 cm long 2 la 

20b. Leaf blades usually becoming more than 4 cm broad, to 24 cm long 22a 

2 la. Inflorescences thyrsiform and purple, to 16 cm long; calyx lobes ca. 0.4 mm long . . . 

P. angustifolia 

21b. Inflorescences paniculate and purple, to 8 cm long; calyx lobes ca. 1 m long 

P. purpurea 

22a. Corolla white, tube 3-6 mm long; calyx lobes ca. 0.3 mm long; leaves with 9-1 1 pairs of 2 

veins P. tilaranensis 

22b. Corolla blue to purple, tube 5-10 mm long; calyx lobes 1-3 mm long; leaves with 11-21 

pairs of 2 veins 23a 

23a. Inflorescence purple, paniculate; leaf blades to 26 x 13 cm, with 11-19 pairs of 2 veins 

P. discolor 

23b. Inflorescence blue, thyrsiform; leaf blades to 18 x 8 cm, with 15-21 pairs of 2 veins .... 

P. brenesii 

24a. Stems and veins on the lower leaf surfaces densely puberulent with short stiff scurfy or slender 

hairs; flowers puberulent P. vestita 

24b. Stems glabrous to sparsely puberulent; flowers glabrous or puberulent 25a 

25a. Stipule lobes/teeth 0.5-2 mm long; leaf blades 3-12 cm long and 1-3.5 cm broad 26a 

25b. Stipule lobes/teeth 2-8 mm long; leaf blades usually more than 7 cm long and 3 cm broad . . 27a 
26a. Flowers yellow or white with yellow and purple; leaves with 12-18 pairs of 2 veins; 1200- 

1 600 m elevation P. garciae 

26b. Flowers white and yellow; leaves with 6-9 pairs of 2 veins; wide ranging in Costa Rica; 

1 1 00-2000 m elevation P. montivaga 

27a. Inflorescence branches red to orange (to purple in fruit); corolla tube 6-10 mm long, orange; 

(800-) 1200- 1700 m elevation P. padifolia 

27b. Inflorescence branches yellow to greenish yellow; corolla tube 7-1 3 mm long, yellow; (800-)1000- 

2300 m elevation 28a 

28a. Corolla tube 7-1 1 mm long; stems glabrous; leaves with 8-13 pairs of 2 veins .... P. padifolia 
28b. Corolla tube 1 1-13 mm long; stems puberulent or glabrous; leaves with 1 1-19 pairs of 2 veins 

. P. lasiorhachis 



Palicourea adusta Standl., J. Wash. Acad. Sci. 18: 
279. 1928. Figure 53. 

Small shrubs or rarely little treelets, 0.5-3(-6) m tall, 
leafy branchlets 0.7-6 mm thick, glabrous or with minute 
(0.1-0.5 mm) appressed hairs, becoming terete; stipules 
2-6 mm long, united above the petiole and forming a 
very short (1-3 mm) tubular sheath, with 2 distal teeth 
1-3 mm long. Leaves opposite, petioles 4-12(-22) mm 
long, 0.5-1 mm thick, glabrous (rarely minutely puber- 
ulent); leaf blades 3-7(-10) cm long. 1-3.5 cm broad, 
narrowly to broadly elliptic, elliptic-oblong, apex acu- 
minate with tip 5-10(-l 5) mm long, base broadly obtuse 
to acute and slightly decurrent on the petiole, drying thin- 
chartaceous to stiffly chartaceous, brown to greenish, 
glabrous above and usually with linear cystoliths visible, 
glabrous or with a few minute appressed hairs on the 
veins beneath, 2 veins 6-1 I/side and usually joining a 
vein at the edge of the leaf, intersecondaries present. 
Inflorescences 6-12 cm long, 4-9 cm broad, open py- 



ramidal with opposite branching, yellowish to blue-gray, 
peduncles 1.5-5 cm long, 0.7-1.4 mm thick, puberulent 
with ascending hairs 0.2-0.4 mm long, bracts 1-4 mm 
long, linear-lanceolate, bracteoles 1-2.5 mm long, ped- 
icels 1-5 mm long. Flowers usually glabrous externally, 
hypanthium 1-1.5 mm long, turbinate, glabrous or 
sparsely puberulent (hairs to 1 mm long), calyx lobes 
0.5-0.8(-1.5) mm long. ca. 1 mm broad at the base; 
corolla tubular, bluish purple, violet, or white marked 
with purple or violet, usually glabrous, tube 6-10.5 mm 
long. 1-2 mm diam. and slightly expanded at the base, 
corolla lobes 5, 1.5-3 mm long, minutely whitish pa- 
pillate along the inner margin; anthers 1.5-2 mm long. 
Fruits 3-4.5(-6) mm long and 4 mm diam., obovoid, 
becoming dark purple to deep blue and translucent, with 
longitudinal ridges when dry, persisting calyx ca. 0.7 mm 
high. 

Plants of ridges and wet open areas in evergreen 
montane rain forest formations, ( 1 800-)2200-3 1 00 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



201 



m elevation. Probably flowering and fruiting 
throughout the year. These plants appear to prefer 
ridges and wet open areas. The species ranges from 
the Cordillera de Tilaran to western Panama. 

Palicourea adusta is recognized by its high-al- 
titude habitat, small leaves, small calyx lobes, usu- 
ally bluish flowers (less often yellowish or purple). 
This species is similar to P. montivaga with white 
flowers and P. salicifolia with yellowish inflores- 
cences. 



Palicourea albocaerulea C. M. Taylor, Syst. Bot. 
Monogr. 26: 15. 1989. Figure 53. 

Shrubs or small trees, 2-7 m tall, leafy stems 1.2-4 
mm thick, glabrous, quadrangular or rounded; stipules 
to 8 mm long, united at the base to form a sheath 1-4 
mm long, apex truncated with 2 teeth 2-4 mm long. 
Leaves opposite, petioles 8-22 mm long, 0.6-1.2 mm 
thick, glabrous, often drying reddish brown; leaf blades 
5-15 cm long, 2-5 cm broad, narrowly elliptic to nar- 
rowly oblong or elliptic-oblong, apex acuminate with tip 
6-15 mm long, base acute (obtuse) to attenuate and 
slightly decurrent on petiole, drying chartaceous, gla- 
brous above, glabrous beneath or with a few short (0. 1- 
0.3 mm) hairs along the midvein beneath, linear cys- 
toliths sometimes visible on the lower surface, 2 veins 
7-1 I/side and merging with a marginal vein at the leaf 
edge. Inflorescences 1 1-22 cm long, 3-8 cm broad, nar- 
rowly pyramidal (thyrsoid) with opposite or alternate 
lateral branches, peduncles 1.5-5 cm long, 1.6-2.2 mm 
thick, glabrous and drying dark brown, bracts 6-10 mm 
long, linear and acute, pedicels 3.5-10 mm long, brac- 
teoles 2-6(-8) mm long, glabrous. Flowers glabrous ex- 
ternally, hypanthium ca. 1 .5 mm long, ca. 0.5 mm diam. 
and funnelform, calyx lobes 5, 3.5-9 mm long, 1-1.2 
mm broad, narrowly oblong, often unequal; corolla tu- 
bular, white or white with blue, tube 8-1 1 mm long, 2- 
3 mm diam., slightly expanded near the base, corolla 
lobes 1-2 mm long and 1 .5 mm broad at the base; anthers 
ca. 2 mm long. Fruits 5-8 mm long (not including the 
calyx lobes) and 5-10 mm diam., ovoid or globose, blu- 
ish black, persisting calyx lobes ca. 7 mm long. 

Plants of lower montane cloud forest forma- 
tions, from 1450 to 1650 m elevation. Flowering 
in September-December; fruiting in June. This 
species is only known from the Cordillera de Ti- 
laran and adjacent areas in Costa Rica. 

Palicourea albocaerulea is recognized by its re- 
stricted range, generally glabrous parts, long inflo- 
rescences, long calyx lobes, and white corollas that 
turn blue with age. The narrow leaves and long 
slender petioles are also distinctive. This is our 
only species in which the flowers change from white 
to blue. Compare this species with P. lancifera 
with shorter calyx lobes and flowers that do not 
turn blue. 



Palicourea angustifolia H.B.K., Nov. gen. sp. 3: 
367. 1819. P. lanceolata Oersted ex Polakows- 
ky., Vidensk. Meddel. Dansk. Naturhist. Foren. 
Kjobenhavn 17. 1852. 

Shrubs or small trees, 2-5(-10) m tall, leafy branchlets 
1.3-4 mm thick, glabrous or rarely sparsely puberulent, 
terete; stipules to 10 mm long, the tubular sheath 2-4 
mm long and truncate but with slender awns 3-6(-8) 
mm long, glabrous, persisting. Leaves opposite, petioles 
2-9(-13) mm long, 0.6-1.2 mm thick, glabrous; leaf 
blades 5-15(-23) cm long, l-4(-5) cm broad, lanceolate 
to narrowly elliptic or very narrowly oblong, apex grad- 
ually narrowed and acute or acuminate, tip 4-14(-25) 
mm long, base acute and slightly decurrent on petiole, 
drying chartaceous, glabrous above, glabrous beneath or 
with short (0.4 mm) thin whitish hairs along the veins, 
2 veins 10-14/side. Inflorescences 3-14(-16) cm long, 
3-6(-8) cm broad, a narrowly pyramidal (thyrsoid) pan- 
icle, peduncles 1.5-3(-6) cm long, 1-2 mm, thick, gla- 
brous or minutely appressed-puberulent, lateral branch- 
es opposite, subopposite or alternate, purple to magenta, 
bracts 2-5 mm long, pedicels 0-3 mm long, bracteoles 
ca. 1 mm long. Flowers glabrous or minutely (0. 1 mm) 
papillate puberulent externally, hypanthium ca. 1 mm 
long, calyx lobes ca. 0.4 mm long, triangular; corolla 
slender funnelform, rose red to purple or fuchsia, tube 
9-14 mm long, 1.5-2.5 mm diam., expanded near the 
base, corolla lobes 5, 1.2-2.5 mm long; anthers 1.7-2.5 
mm long, included. Fruits 4-6 mm long and 5 mm diam., 
globose to ovoid, becoming purple and drying black, 
puberulent or glabrate, persisting calyx ca. 0.5 mm long. 

Plants of evergreen montane forest formations 
on both the Caribbean and Pacific slopes, from 
(1400-)! 600 to 2000(-2300) m elevation. Flow- 
ering and fruiting throughout the year but with 
most flowering collections made in January-Sep- 
tember. The species ranges from central Costa Rica 
to Peru. 

Palicourea angustifolia is recognized by the nar- 
row leaves, truncated stipule sheath with long awns, 
narrow inflorescences, and slender funnelform 
purple corollas with relatively short lobes. This 
species resembles P. padifolia and P. purpurea but 
differs in the relatively narrower inflorescences, 
shorter pedicels, and thinner, more funnelform co- 
rollas that are usually minutely puberulent at the 
base. In addition, P. padifolia has yellow corollas 
and red or orange inflorescence branches. 



Palicourea bella (Standl.) Dwyer, Ann. Missouri 
Bot. Gard. 67: 299. 1980. Psychotria bella 
Standl., J. Wash. Acad. Sci. 18: 185. 1928. 

Shrubs or small trees to 5 m tall, leafy stems ca. 4 mm 
thick and quadrangular, glabrous or with small (0. 1-0.3 
mm) hairs; stipules 10-20 mm long, to 15 mm broad. 



202 



FIELDIANA: BOTANY 



with thin leaf-like texture, united above the petioles, 
oblong, rounded and bilobed to emarginate. Leaves clus- 
tered distally, petioles 8-20 mm long, 1.2-2 mm thick, 
puberulent; leaf blades 9-22 cm long, 3-8.5 cm broad, 
elliptic to elliptic-obovate or oblong-elliptic, apex acu- 
minate, base gradually narrowed and acute, drying char- 
taceous, dark greenish brown, glabrous or with short (0.3 
mm) hairs on the veins and smaller hairs between the 
veins on both surfaces, 2 veins 12-19/side. Inflores- 
cences solitary, 8-20 cm long, 7-10 cm broad, broadly 
corymbiform, peduncles 5-13 cm long, to 2 mm thick 
and sparsely strigulose, bracts 10-26 mm long, ovate- 
oblong, translucent brown and sparsely hirtellous, sec- 
ondary peduncles 1 5-25 mm long, bright pink to purple, 
pedicels 2-8 mm long, obscured by the large bracts and 
bracteoles to 8 mm long. Flowers with hypanthium ca. 
2 mm long, glabrous or with conspicuous thin yellowish 
hairs ca. 0.7 mm long, the calyx lobes 6-14 mm long, 
3-6 mm broad, ovate to ovate-elliptic, drying thin trans- 
lucent with scattered short (0.2 mm) hairs; corolla tu- 
bular, white to pink, carnose, tube 14-20 mm long and 
2-3 mm diam., dilated at the base, corolla lobes 4-5 mm 
long, ovate; anthers ca. 3.5 mm long. Fruits ca. 6 mm 
long and 6 mm diam., ellipsoid, glabrous or sparsely 
puberulent. 

Plants of evergreen montane forest formations 
at (1500-)2000-2700 m elevation. Flowering in 
April. This species is only known from Chiriqui 
Province in Panama and a collection from the 
Caribbean slope of central Costa Rica at 1500 m. 

Palicourea bella is recognized by the broadly 
two-lobed stipules, corymbiform inflorescences 
with broad thin bracts, and the large thin calyx 
lobes. In life, both inflorescences and calyx are red- 
violet to bright pink in color. The similarity in 
morphology, texture, and vestiture of bracts and 
calyx lobes is an interesting example of heterotopy. 
This species is similar to P. bellula (q.v.) as well 
as to P. hammelii C. M. Taylor and P. ochnoides 
Dwyer of western Panama. Grayum 7046 (CR, MO) 
from Volcan Barva is tentatively placed here. 



Palicourea bellula C. M. Taylor, Syst. Bot. Mono- 
gr. 26: 24. 1989. Figure 49. 

Shrubs or slender treelets, to 2.5 m tall, leafy stems 
2.5-5 mm thick, glabrous, quadrangular and drying dark, 
grayish in age; stipules 5-1 1 mm long, 7-10 mm broad, 
united above the petioles for 2-3 mm to form a broad 
tube, with 2 rounded lobes 2-4 mm high, thin in texture 
and glabrous. Leaves opposite, petioles 4-18 mm long, 
1.5-2.5 mm thick, glabrous, sulcate adaxially near the 
base; leaf blades 7.5-14 cm long, 3.5-7 cm broad, broad- 
ly elliptic to slightly elliptic-obovate, apex acuminate, 
tip ca. 5- 1 0(- 1 5) mm long, base obtuse (acute) and slight- 
ly decurrent on petiole, drying stiffly chartaceous, green- 
ish, glabrous above, with thin whitish hairs 0.3-1 mm 
long along the sides of the midvein and on some 2 veins 
beneath, 2 veins ( 1 0-) 1 7-20/side. Inflorescences 1 or 3, 



9-14 cm long, 9-11 cm broad, rounded panicles, red- 
violet to fuchsia, peduncles 3-8 cm long, 1-3 mm thick, 
glabrous, bracts 12-16 mm long, 3-4 mm broad, lan- 
ceolate, bracteoles 5-12 mm long and 3-5 mm broad, 
narrowly ovate, pedicels 4-12 mm long. Flowers gla- 
brous externally, hypanthium ca. 1 mm long and 1.2 
mm diam.. drying dark and glabrous, calyx tube 2-3 mm 
long and broadly cupulate, calyx lobes 5. 4-7 mm long, 
2-3 mm broad, ovate, red-violet to fuchsia; corolla white, 
tubular, carnose, tube 12-15 mm long and 2.5 mm diam.. 
corolla lobes 2-3 mm long; anthers ca. 3 mm long. Fruits 
not seen at maturity. 

Plants of open sites in montane rain forest for- 
mations near the continental divide at 1 900-2200 
m elevation. Flowering in January-February, May, 
and November. This species is only known from 
between Volcan Viejo (Alajuela) and the south- 
eastern slope of Volcan Barva near the upper part 
of Rio Patria (Heredia Province) Costa Rica. 

Palicourea bellula is distinguished by its colorful 
glabrous inflorescences with conspicuous pink to 
purple bracts and calyx lobes, tubular white co- 
rollas, leaves with many closely parallel secondary 
veins, broadly rounded stipule lobes, and restrict- 
ed cloud forest habitat. This species is similar to 
P. bella, but the calyx and corolla are usually small- 
er and stiffer than in P. bella. Palicourea hammelii 
and P. ochnoides are similar but lack the broad 
imbricate calyx lobes. 



Palicourea brenesii Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 18: 1333. 1938. P. talamancana 
Standl. & L. O. Williams, Ceiba 1: 250. 1951. 
Figure 52. 

Shrubs, 1-4 m tall, erect or clambering, often with 
only a few distal branches 2.5-5 mm thick, glabrous or 
sparsely strigillose with hairs 0.5-1 mm long, drying dark, 
quadrangular; stipules united to form a tubular sheath 
3-6 mm long, truncated distally and with 2 prominent 
teeth (on each side) 4-7(-10) mm long and 1-2 mm 
broad at the base, usually glabrous, persisting. Leaves 
opposite, petioles 1.5-3 cm long, 1.2-2.2 mm thick, gla- 
brous; leaf blades 9-1 8 cm long, 3.5-8 cm broad, broadly 
elliptic to elliptic-obovate or elliptic-oblong, apex abruptly 
narrowed and short-acuminate, base obtuse to acute, 
drying stiffly chartaceous, brownish or yellowish, gla- 
brous above, glabrous or sparsely puberulent beneath, 
2 veins 15-21 /side. Inflorescences 8-15 cm long, 4-9 
cm broad (at broadest part), pyramidal or elongate thyr- 
soid panicles, often blue throughout, peduncles 2-5 cm 
long, 1.5-3 mm thick, sparsely to densely puberulent 
with thin hairs ca. 0.2 mm long, bracts 5-8 mm long, 
ca. 2 mm broad, lanceolate, pedicels 0-6(-8) mm long, 
bracteoles 1-5 mm long. Flowers minutely puberulent 
and glabrescent distally, hypanthium ca. 1 mm long and 
0.8 mm diam., calyx tube broadly cupulate, calyx lobes 
1-2.5 mm long, broadly triangular and obtuse; corolla 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



203 



blue, tubular, glabrous near the base and densely ap- 
pressed-puberulent distally or glabrous throughout, tube 
8-1 1 mm long, ca. 2 mm diam.. narrowed near the mid- 
dle, corolla lobes 5, 3-5 mm long; anthers ca. 2.5 mm 
long. Fruits 4-6 mm long, 4-6 mm diam., globose, gla- 
brate. 



Plants of evergreen montane rain forest for- 
mations, from 1800 to 2600(-3200) m elevation. 
Flowering and fruiting in February-June. The spe- 
cies has only been collected in the northern part 
of the Meseta Central (near Zarcero and Palmira) 
and along the Interamerican Highway in the west- 
ern part of the Cordillera de Talamanca in Costa 
Rica. 

Palicourea brenesii is distinguished by the larger 
leaves with many secondary veins, bright "china 
blue" thyrsoid panicels, and small blue to violet 
flowers with conspicuous calyx lobes. The usually 
glabrous stems contrast with the puberulent pe- 
duncle and inflorescence branches. The striking 
bright blue color of the inflorescence is similar to 
that found in species of Faramea. The disjunct 
collections from the Cordillera de Talamanca have 
smaller corollas and stipule lobes to 10 mm long. 
This species is very similar to P. discolor after the 
flowers have faded. 



Palicourea copensis (Dwyer) C. M. Taylor, comb, 
nov. Psychotria copensis Dwyer, Ann. Missouri 
Bot. Gard. 67: 365. 1980. Figure 52. 

Shrubs or small treelets, 2-5 m tall, leafy stems 2-5 
mm thick, with curved yellowish or whitish hairs 0.4- 
0.9 mm long, glabrescent; stipules 6-12 mm long, united 
to form a broad tube to 10 mm long, 4-8 mm broad (to 
14 mm broad below the inflorescences), bilobed with a 
distal sinus 1-6 mm deep, glabrous but with a ciliolate 
margin, persisting. Leaves with petioles 10-55 mm long, 
1.3-2.3 mm thick, glabrous or sparsely puberulent be- 
neath; leaf blades 9-23 cm long, 3.5-9 cm broad, elliptic- 
obovate to elliptic-oblong, apex narrowly acuminate and 
5-13 mm long, base cuneate and slightly decurrent on 
petiole, drying chartaceous, dark green above (pale green 
beneath), glabrous above or with a few hairs along the 
major veins, sparsely pubescent beneath with crooked 
whitish hairs 0.4-1.5 mm long, 2 veins 9-14/side and 
arcuate-ascending distally. Inflorescences terminal or 
pseudoaxillary, 1 or several per node, 2-7 (-1 1) cm long, 
3-6(-l 2) cm broad, pyramidal with few opposite branch- 
es, peduncles 8-26 mm long, ca. 1 .5 mm thick, pubescent 
or glabrous, bracts 3-7 mm long, flowers in distal cymose 
or irregular clusters. Flowers with hypanthium ca. 1 mm 
long, obconic, glabrous or pubescent, calyx yellowish, 
calyx lobes quite variable, 1-3 mm long, oblong and 
obtuse; corollas lavender or white marked with purple, 
salverform with a distinctly gibbous base, tube 7-12 mm 
long, 2-4 mm diam., puberulent, corolla lobes 3-5 mm 



long; anthers ca. 3 mm long. Fruits 6-7 mm long, 4-5 
mm diam., obovoid, becoming blue or violet. 

Plants of the very wet Caribbean escarpment, 
between 400 and 1 000 m elevation. Flowering and 
fruiting in March-May and November. The spe- 
cies is known from the Caribbean slopes of Volcan 
Barva and the P. N. Braulio Carrillo area in central 
Costa Rica. It was originally described from El 
Cope in Code Province, Panama. 

Palicourea copensis is recognized by its white or 
lavender flowers, variable sepal lobes, smaller 
thick-branched inflorescences, large persisting 
stipules, puberulence of usually curved hairs, and 
lower montane habitat. This is a rarely collected 
species that appears to prefer creek margins. 



Palicourea crocea (Sw.) Roem. & Schult., Syst. Veg. 
5: 193. 1819. Psychotria crocea Sw., Prodr. 44. 
1788. Figure 50. 

Shrubs or small trees, 1-5 m tall, leafy stems 1-4 mm 
thick, glabrous or minutely puberulent, terete; stipules 
united to form a short (0.5-2 mm) truncated tube with 
narrow distal teeth 1-4 mm long, glabrous, deciduous. 
Leaves opposite, petioles 6-12(-20) mm long, 1-1.5 mm 
thick, usually glabrous; leaf blades 7-14(-19) cm long, 
2.5-6(-7) cm broad, elliptic, narrowly elliptic, lanceolate, 
to elliptic-ovate or elliptic-oblong, apex acute to acu- 
minate, base obtuse to acute and slightly decurrent on 
the petiole, drying membranaceous to thin-chartaceous, 
greenish, usually glabrous above, with short (0.3 mm) 
thin hairs along the sides of the midvein beneath, 2 
veins 8-12/side. Inflorescences 6-15 cm long, 2.5-7 cm 
broad, nararow pyramidal to rounded with opposite or 
subopposite (alternate distally) branching, reddish to or- 
ange or pink, peduncles 1.5-1 1 cm long, 1-2 mm thick, 
glabrous or sparsely and minutely puberulent, bracts 1- 
10 mm long, linear-lanceolate and deciduous, pedicels 
4-8(-12) mm long, slender, bracteoles ca. 1 mm long 
and deciduous. Flowers distylous, glabrous or minutely 
puberulent externally, hypanthium ca. 1 mm long and 
0.7 mm diam., calyx lobes 0.3-0.5 mm long, broadly 
ovate; corolla tubular, dark red to yellow-orange or pink 
(rarely bluish green), tube 5-9 mm long, 1 .5-3 mm diam., 
inflated near the base, corolla lobes 1-2.5 mm long; an- 
thers ca. 2.7 mm long. Fruits 4-6 mm long and 4 mm 
diam., subglobose or ovoid, becoming dark blue or black, 
longitudinally ribbed when dry, persisting calyx minute. 



Plants of secondary growth in evergreen rain 
forest areas often along streams and paths in the 
Caribbean lowlands, from near sea level to 600 m 
elevation. Flowering and fruiting throughout the 
year (flowering mostly in April-August in Central 
America). The species ranges from central Mexico 
and the West Indies to Paraguay. 

Palicourea crocea is recognized by its smaller 



204 



FIELDIANA: BOTANY 



stature and lowland habitat, short stipular tube, 
colorful thyrsoid inflorescences, minute calyx lobes, 
and orange or red corolla tubes. This species is 
often found in swampy areas and other sites with 
poor drainage. This species is sometimes confused 
with P. padifolia, of higher elevations and with 
longer stipule sheaths, with P. guianensis, with 
longer stipule lobes, and with species ofHamelia. 



Palicourea discolor K. Krause, Bot. Jahrb. Syst. 
54: Beibl. 119: 40. 1916. P. macrosepala K. 
Krause, loc. cit. 41. 1916. P. panamaensis 
Standl., Ann. Missouri Bot. Gard. 25: 839. 1938. 
Figure 52. 

Shrubs or small trees, 1-4(1-6) m tall, leafy stems 3- 
6 mm thick, glabrous and quadrangular; stipules united 
to form a broadly tubular sheath 2-6 mm long, truncated 
apically and with 2 narrow teeth 4-7 mm long, glabrous 
and persisting. Leaves opposite, petioles l-3(-5) cm long, 
1.7-2.7 mm thick, glabrous; leaf blades 10-26 cm long, 
5-13 cm broad, elliptic to broadly elliptic-oblong, acute 
to short-acuminate, the tip 5-10(-15) mm long, obtuse 
or acute at the base, drying stiffly chartaceous, dark brown 
or greenish above, glabrous above, sparsely puberulent 
with inconspicuous (0.05-0.2 mm) thin whitish hairs on 
the veins beneath, 2 veins 11-19/side, united distally 
with a vein at the leaf edge. Inflorescences solitary or 
the basal branches subtended by leaves and apparently 
3, 12-22(-30) cm long, 10-1 5(-24) cm broad, pyramidal 
with opposite or subopposite branches, bright purple in 
life, peduncles 2-14 cm long, 2.5-4 mm thick, glabrous, 
bracts 5-8 mm long, 1 mm wide, linear-lanceolate, ped- 
icels 1-8 long, glabrous, bracteoles absent. Flowers gla- 
brous externally, hypanthium 0.5-1 .3 mm long and 0.3- 
1 mm diam.. calyx lobes 1.5-5 mm long, acute to Un- 
gulate, very variable in some collections; corolla purple 
or white flushed with purple, carnose, tube 7-13 mm 
long, 1-2 mm diam., corolla lobes 1-3.5 mm long. Fruits 
4-5 mm long and 4-5 mm diam., globose to globose- 
ellipsoid, with longitudinal ribs (dried), calyx persisting 
or deciduous. 

Plants of forest interiors and shaded thickets in 
evergreen montane forest formations, from 1 1 00 
to 1900(-2600) m elevation. Flowering and fruit- 
ing in February-August. The species ranges from 
the upper Rio Grande de Orosi and Moravia de 
Chirripo in the Cordillera de Talamanca of Costa 
Rica southward into the Chiriqui Highlands of 
western Panama. 

Palicourea discolor is recognized by its relatively 
large leaves and inflorescences, purple flowers and 
inflorescence branches, and variable calyx lobes 
to 5 mm long. This species is similar to P. brenesii 
with bright blue inflorescences and smaller corol- 
las and P. purpurea with smaller calyx lobes and 
smaller leaves. 



Palicourea garciae Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 22: 195. 1940, non P. garciae 
Steryerm. 1971. Psychotria copeyana Standl. & 
L. O. Williams, Ceiba 1: 251. 1940. Figure 53. 

Shrubs or small trees, 1.5-5 m tall, leafy branchlets 
1.5-3.5 mm thick, glabrous or with short (0.3 mm) as- 
cending yellowish hairs, quickly becoming glabrescent 
and terete, drying dark; stipules united to form a tubular 
sheath 2-4 mm long, each side with 2 short ( 1 mm) distal 
lobes and a small ( 1 mm) central sinus, glabrous or mi- 
nutely appressed-puberulent. Leaves opposite, petioles 
5-15 mm long, 0.6-1.2 mm thick, glabrous; leaf blades 
5-12(-15) cm long, 2-4(-4.5) cm broad, narrowly ellip- 
tic, narrowly elliptic-ovate to narrowly oblong or lan- 
ceolate, apex acuminate with tip 7-15 mm long, base 
acute to obtuse and slightly decurrent on petiole, drying 
stiffly chartaceous, much darker above than beneath, 
glabrous above or with hairs along the midvein, with 
few or many minute (0.05-0.2 mm) hairs along the major 
veins beneath and usually glabrescent, 2 veins 12-18/ 
side, arising from the midvein at near 90. Inflorescences 
solitary, 4-12 cm long, 6-12 cm broad, open pyramidal 
or rounded corymbiform with opposite (subopposite) 
branching, green or flushed with purple, peduncles 
(l-)2.5-4 cm long, very sparsely to densely puberulent 
with short (0.2 mm) thin hairs, bracts 2-5 mm long, 
linear-subulate, pedicels 1-5 mm long, bracteoles ca. 1 
mm long. Flowers glabrous externally, hypanthium 1- 
1.5 mm long, 0.8-1 mm diam., calyx lobes 5, 0.7-1.5 
mm long, acute; corolla funnelform and gibbous near the 
base, white or flushed with yellow or purple distally 
(grayish to purple near the base), tube 6-8 mm long, 
constricted in the middle (1-1..5 mm diam.) and gibbous 
near the base, corolla lobes 2-3 mm long and 1 mm 
broad at the base; anthers 2-2.5 mm long. Fruits 4-5 
mm long, 3-4 mm diam., globose to ellipsoid, becoming 
pale blue at maturity, with longitudinal ribs, persisting 
calyx ca. 1 mm long. 

Plants of montane rain forest formations, from 
1200 to 1600(-2000) m elevation. Flowering in 
March and May-August; fruiting in May-June and 
August. The species ranges from central Costa Rica 
to Colombia. 

Palicourea garciae is recognized by the small 
narrow leaves with many secondary veins, the small 
corolla tubes strongly bent below the middle and 
inflated at the base, and the small pale blue fruit. 
The very short tubular sheath and bifid interpetio- 
lar part of the stipule is noteworthy. Psychotria 
copeyana was based on a specimen ( Williams <&. 
Allen 16482 F) with immature inflorescences from 
El Copey. Vegetatively this species resembles sev- 
eral smaller-leaved species of Psychotria, but the 
strongly gibbous flowers are very different. 



Palicourea guianensis Aubl., Hist. pi. Guiane 1: 
173, t. 66. 1775. Psychotria palicourea Sw., Fl. 
Ind. Occ. 1797. Figure 50. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



205 



Shrubs or small trees, 2-6(-10) m tall, leafy stems 1.5- 
8 mm thick, glabrous or minutely puberulent, quadran- 
gular, often drying dark and contracted beneath the node; 
stipules free or united for 1-2 mm at the base, lobes, 5- 
8(-12) mm long, lobes 2-4 mm broad, blunt, glabrous 
and drying dark. Leaves opposite, petioles 8-24(-38) mm 
long, 1.3-3 mm thick, glabrous; leaf blades (12-) 15-28 
cm long, (5-) 7-1 8 cm broad, broadly ovate-elliptic to 
ovate-oblong, elliptic-oblong or ovate, apex acute or acu- 
minate, base obtuse to rounded and subtruncate, drying 
thin-chartaceous, concolorous, glabrous or minutely (0. 1 
mm) papillate-puberulent above and below, 2 veins 10- 
15/side. Inflorescences usually solitary, 7-18 cm long, 
4-12 cm broad, pyramidal or thyrsiform with many 
closely crowded alternate or subopposite branches, red- 
orange when flowering and later turning purple, pedun- 
cles 3-12 cm long, 1.8-3.2 mm thick, minutely puber- 
ulent with thin hairs ca. 0. 1 mm long, bracts absent or 
0.5-7 mm long and often adnate to the primary branches, 
pedicels 2-5(-8) mm long, minutely puberulent. Flowers 
monomorphic, minutely puberulent externally, hypan- 
thium ca. 1.5 mm long and 0.8 mm diam., tubular, calyx 
lobes 0.2-0.6 mm long, broadly obtuse; corolla yellow 
or orange, tube 8-25 mm long and 2-3 mm diam. at 
base, contracted in the middle and a slightly inflated 
distally, corolla lobes 1-2 mm long, triangular; ovary 2- 
6-locular. Fruits 4-7 mm long, 4-5 mm diam., ovoid to 
ellipsoid, becoming purple violet and drying black with 
prominent longitudinal ribs, persisting calyx and disc ca. 
1 mm high. 



mm thick, glabrous or minutely puberulent, drying dark; 
leaf blades 8.5-20 cm long, 2.5-8.5 cm broad, elliptic 
to elliptic-obovate, apex acuminate with tip 5-15 mm 
long, base gradually narrowed and acute or cuneate, 
slightly decurrent on petiole, drying stiffly chartaceous, 
dark olive green above, glabrous above or puberulent on 
the midvein, puberulent beneath with crooked hairs 0.05- 
0.3 mm long, 2 veins 12-18/side. Inflorescences 4-14 
cm long, to 1 3 cm broad, open pyramidal, pink to purple 
but drying dark, peduncle 15-35 mm long, 1.5-3 mm 
thick, glabrous or very sparsely minutely puberulent, 
proximal bracts 18-35 mm long, ca. 5 mm broad, lan- 
ceolate, distal bracteoles 6-20 mm long, pedicels 4-15 
mm long. Flowers glabrous externally, hypanthium ca. 
2 mm long and 1 mm diam., calyx 1 3-20 mm long, pink 
to purple, calyx lobes 4-1 2 mm long, lanceolate, unequal; 
corolla white, tubular, tube 20-27 mm long, ca. 4 mm 
diam., corolla lobes 3-6 mm long, triangular. Fruits 8- 
10 mm long, 7-8 diam., ellipsoid. 



Palicourea hammelii was originally described 
from collections made between 1 400 and 2200 m 
elevation in Chiriqui, Panama. It has been recently 
collected on the Fila Matama at 1 600 m elevation 
on the Caribbean slope of the Cordillera de Tala- 
manca (Herrera & Chacon 2781 CR, MO). This spe- 
cies resembles P. bella with ovate calyx lobes and 
P. bellula with a shorter corolla tube. 



Plants of open secondary formations in ever- 
green and partly deciduous forest areas, from near 
sea level to 900 m elevation. Flowering and fruit- 
ing occur throughout the year; flowering mostly in 
March-July in Costa Rica. The species ranges from 
southern Mexico and the West Indies southward 
to southern Brazil and Bolivia. 

Palicourea guianensis is recognized by its low- 
land habitat, the wide glabrous or very minutely 
puberulent thin-textured leaves, the large rounded 
stipule lobes, the conspicuous densely flowered 
yellow or orange inflorescences, and yellow or or- 
ange puberulent corollas. Most of our collections 
come from the Caribbean lowlands and from the 
Golfo Dulce area. Central American material has 
two-locular ovaries, but specimens from eastern 
South American may have three-or four-locular 
ovaries. Palicourea guianensis resembles P. padi- 
folia (q.v.). 



Palicourea hammelii Taylor, Syst. Bot. Monogr. 
26:43. 1989. 

Small trees to 5 m tall, leafy stems 3-7 mm thick, 
glabrous or minutely (0. 1 mm) appressed-puberulent, 
drying dark; stipules 6-12 mm long, with a sheathing 
base 3-5 mm long with 2 lobes (4/node) 3-7 mm long, 
persisting. Leaves with petioles 13-35 mm long, 1.3-2 



Palicourea lancifera Standl. & L. O. Williams, Cei- 
ba 1: 249. 1951. Figure 53. 

Shrubs, 1.5-3(-5) m tall, leafy stems 1.3-4 mm thick, 
glabrous to (less often) densely puberulent with crooked 
yellowish hairs 0.1-0.3 mm long, quadrangular at first, 
glabrescent; stipules united to form a tubular sheath 3- 
5 mm long, apex truncated with 2 narrow teeth 2-5 mm 
long on each side. Leaves opposite, petioles 7-25(-30) 
mm long, 0.4-1.2 mm thick, glabrous or minutely pu- 
berulent; leaf blades 5-1 5(-l 8) cm long, 2-5(-7) cm broad, 
elliptic to narrowly elliptic-oblong, apex acuminate with 
tip 6-20 mm long, base acute and decurrent on petiole, 
drying thin-chartaceous or chartaceous, dark greenish or 
brownish above, glabrous above, minutely (0. 1-0.4 mm) 
strigulose on the veins beneath, 2 veins 8-12/side and 
connecting with a vein along the leaf edge. Inflorescences 
solitary, 7-17(-21)cm long, 2-6(-9)cm broad, narrowly 
pyramidal with many short branches, green or marked 
with yellow, peduncles 1-4 cm long, 1-2 mm thick, gla- 
brous to densely puberulent, bracts 5-8 mm long, nar- 
rowly lanceolate, flowers often alternate along the lateral 
branches, bracteoles 1.5-4 mm long, pedicels 2-7(-17) 
mm long. Flowers glabrous or sparsely puberulent ex- 
ternally, hypanthium 1-1.5 mm long, ca. 1 mm diam., 
calyx lobes 1.5-3 mm long, often unequal, acute or 
rounded distally; corolla white or flushed with green or 
yellow, tubular, tube 7-1 1 mm long, ca. 2 mm diam. 
and expanded at the base, corolla lobes 1-2 mm long; 
anthers 2-2.5 mm long. Fruits 4-5 mm long and 4-5 
mm diam., ovoid or globose-oblate, persisting calyx ca. 
1.5 mm high. 



206 



FIELDIANA: BOTANY 



Plants of very wet lower montane cloud forest 
formations, from 800 to 1 700 m elevation. Flow- 
ering June-November; apparently fruiting 
throughout the year. This species ranges from the 
Reserva Forestal de San Ramon and Zapote de 
San Carlos (Alajuela) eastward to Moravia de 
Chirripo (Limon) and the western part of the Gen- 
eral Valley (San Jose). 

Palicourea lancifera is recognized by its narrow 
long-petiolate leaves, narrowly thyrsoid greenish 
inflorescences, slightly larger calyx lobes rounded 
distally, and white or pale yellow corolla tubes. 
This species resembles P. albocaerulea with longer 
calyx lobes and P. leucantha J. D. Smith of Mexico 
and northern Central America. 



Palicourea lasiorrhachis Oersted, Vidensk. Med- 
del. Dansk. Naturhist. Foren. Kjobenhavn 39. 
1 852. P. veraguensis Dwyer, Ann. Missouri Bot. 
Card. 67: 318. 1980. Figure 51. 

Shrubs or small treelets, l-5(-7) m tall, leafy stems 
1.8-5 mm thick, glabrous to pubescent with stiff yellow- 
ish curved hairs to 0.5 mm long; stipules united and 
forming a broad tube 2-9 mm long, truncated and with 
narrow teeth 2-5(-8) mm long, glabrous or minutely 
puberulent, persisting. Leaves opposite, petioles 7-24(-32) 
mm long, 1-2 mm thick, glabrous or puberulent; leaf 
blades 6-17(-26) cm long, 2-8(-10) cm broad, elliptic 
to narrowly ovate-elliptic or elliptic-oblong, apex taper- 
ing gradually and acuminate, tip 4-13(-20) mm long, 
base acute to obtuse, drying chartaceous, dark grayish 
green to brownish above, glabrous above, sparsely to 
densely puberulent along the major veins beneath with 
short (0.3 mm) yellowish or longer (1 mm) whitish as- 
cending hairs (rarely glabrous), 2 veins (8-)l 1-14(-18)/ 
side and joining a vein along the leaf edge distally. In- 
florescences 5-1 5(-20) cm long. 4-1 3 cm broad, narrow- 
ly pyramidal with 4-8 pairs of opposite or subopposite 
lateral branches from the central rachis, greenish to bright 
yellow, peduncles 1.5-7(-10)cm long, 0.7-2.8 mm thick, 
sparsely to densely puberulent with stiff multicellular 
hairs 0.3-1 mm long, bracts 2-9 mm long, linear-su- 
bulate, pedicels 1-6 mm long, minutely puberulent, yel- 
low to bright green, bracteoles 0.5-2 mm long. Flowers 
sparsely to densely pubescent externally (often glabres- 
cent), hypanthium 0.5-1.5 mm long, ca. 1 mm diam. 
distally, obconic, calyx lobes 0.3-1.2 mm long, obtuse: 
corolla yellow to greenish, tubular-funnelform, tube 6- 
15 mm long, narrowed in the middle, ca. 2 mm diam. 
at the base and 3 mm diam. distally, corolla lobes 1-2.5 
mm long, ca. 1.5 mm broad at the base, triangular; an- 
thers 1.5-2 mm long or 2.5-3 mm long in short-styled 
form. Fruits 4-5 mm long and 3-4 mm diam., obovoid, 
with prominent longitudinal ribs, puberulent or glabres- 
cent, blue or black. 



Understory plants of very wet montane rain for- 
est formations, from (1000-)! 300 to 2600 m el- 



evation. Flowering throughout the year (mostly in 
May-June); probably fruiting throughout the year. 
This species ranges from the Cordillera de Tilaran 
along the continental divide and Cordillera de Ta- 
lamanca to western Panama. 

Palicourea lasiorrhachis is recognized by the of- 
ten narrow leaves, usually puberulent beneath, the 
minutely puberulent inflorescences (rarely gla- 
brous) with yellow pedicels, the small calyx lobes, 
narrowly funnelform yellow corolla tubes, and 
small obovoid fruit. This common species is quite 
variable and includes individual collections that 
differ greatly from each other in regard to leaf size, 
leaf shape, puberulence, and aspects of flowers and 
inflorescences. A group of collections from near 
El Empalme have thicker stems; stifler, more ob- 
long leaves; secondary veins arising at almost 90 
angles; and dense puberulence. This species may 
be difficult to distinguish from P. vestita (q.v.). It 
is often confused with P. padifolia, which has red- 
orange inflorescence branches and more tubular 
corollas. Also compare P. montivaga (smaller 
plants with white flowers) and P. adusta (with blue 
flowers). 



Palicourea macrocalyx Stand!.. J. Wash. Acad. Sci. 
18: 278. 1928. Figure 49. 

Small shrub, subshrubs, or little treelets. 0.5-2(-6) m 
tall, leafy branchlets 1 .5^4(-10) mm thick, glabrous, flat- 
tened or quadrangular but quickly becoming terete: stip- 
ules with a broad thick sheath 2-6 mm long, truncated 
and with narrow acute teeth 4-10 mm long and 1-3 mm 
broad, glabrous and persisting. leaves opposite, petioles 
7-30(-45) mm long, 0.9-2.2 mm thick, glabrous; leaf 
blades 5-15(-18) cm long, 2-6(-9.5) cm broad, elliptic- 
obovate to obovate-oblong or elliptic (rarely broadly ob- 
long), apex acuminate with tip 4-13 mm long, base ob- 
tuse to acute (subtruncate). drying stiffly chartaceous to 
subcoriaceous. olive green or brownish above, glabrous 
above and below or with small (0.3-0.5 mm) hairs along 
the midvein, 2 veins 9-1 3/side. Inflorescences 3-8(-l 5) 
cm long. 2-10 cm broad, pyramidal, yellow or marked 
with purple, peduncles 1-8.5 cm long, 1-1.8 mm thick, 
glabrous, bracts ca. 6 mm long, pedicels 1-8 mm long, 
glabrous, bracteoles 1-4 mm long. Flowers glabrous ex- 
ternally, hypanthium ca. 1 mm long, narrowly obconic. 
calyx lobes 3-6 mm long, 1-2 mm broad, narrowly tri- 
angular to ovate, green or yellow and sometimes marked 
with purple: corolla tubular, yellow or greenish yellow, 
tube 8-13 mm long, ca. 2 mm diam.. slightly narrower 
in the center, corolla lobes 2-3 mm long, obtuse; anthers 
1-2 mm long. Fruits 5-6 mm long, 4-5 mm diam., el- 
lipsoid, becoming blue, longitudinally ribbed (when 
dried), with persisting calyx to 6 mm long. 

Plants of the very wet cloud forests along the 
continental divide, from 1300 to 2200 m eleva- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



207 



tion. Probably flowering throughout the year; 
fruiting in April-May and August-October. En- 
demic to Costa Rica, and collected primarily in 
the Cordillera de Tilaran, but ranging to near San 
Isidro de Heredia on the western slopes of Volcan 
Irazu. 

Palicourea macrocalyx is distinguished by the 
stiff slightly obovate leaves with many secondary 
veins, lack of pubescence, yellow inflorescences 
sometimes marked with purple, the larger imbri- 
cate greenish yellow calyx lobes, and yellow co- 
rollas. The name of this species should not be 
confused with P. macrosepala Krause, a synonym 
of P. discolor. This species is very similar to P. 
bellula, which has longer corolla tubes. 



Palicourea montivaga Standl.. J. Wash. Acad. Sci. 
18:279. 1928. Figure 53. 

Shrubs, 1-3 m tall (rarely slender treelets to 6 m), leafy 
stems 0.9-6 mm thick, glabrous or with a few thin hairs 
ca. 0.3 m long distally, terete; stipules with a tubular 
sheath 0.5-4 mm long, truncated distally and with tri- 
angular or linear teeth 0.5-2.5 mm long, glabrous and 
persisting. Leaves opposite, petioles 6-14(-25) mm long, 
0.4-1.1 mm thick, glabrous, slightly sulcate above; leaf 
blades 3-7(-9) cm long, l-2.5(-3) cm broad, lanceolate 
or elliptic-lanceolate to narrowly elliptic-oblong, apex 
tapering gradually and acute or acuminate, tip to 1 5 mm 
long, base acute (obtuse), drying chartaceous, dark green 
or brown above, glabrous above, glabrous or with a few 
thin hairs 0. 1 -0. 5 mm long on the veins beneath, 2 veins 
6-9/side. Inflorescences 2-10 cm long, 2-8 cm broad, 
pyramidal with open opposite branching, peduncles 8- 
45 mm long, 0.3-1.2 mm thick, glabrous or minutely 
puberulent, inflorescence branches usually bright yellow 
(green), bracts 2-3.5 mm long, linear, pedicels 1-6 mm 
long. Flowers usually glabrous externally, hypanthium 
0.5-1 mm long, ca. 1 mm diam. distally, obconic, calyx 
lobes 0.4-1 mm long, acute or obtuse; corolla white or 
yellow, tube 5-9 mm long, 1-1 .3 mm diam., corolla lobes 
1-2 mm long; anthers ca. 1.5 mm long. Fruits 4-5 mm 
long, 3-4 mm diam., obovoid, becoming longitudinally 
ribbed. 

Plants of montane rain forest formations, from 
(1 1 00-) 1300 to 2300(-2900?) m elevation. Prob- 
ably flowering throughout the year, but mostly in 
April-August; fruiting April-November. The spe- 
cies is endemic and ranges from Volcan Tenorio 
southward along the continental divide to the 
western slopes of Volcan Irazu. 

Palicourea montivaga is recognized by its small 
narrow leaves with long tips, glabrous parts, small 
yellowish inflorescences, small white or yellowish 
corollas, and montane habitat. This species is 
closely related to P. adusta with smaller bluish 
flowers and P. lasiorrhachis with relatively larger 



yellowish floral parts and shorter leaf tips. The 
plant attributed to this species from Panama 
(Dwyer, 1 980) is P. lasiorrhachis. Material of P. 
padifolia can also be confused with this species. 



Palicourea orosiana C. M. Taylor, Syst. Bot. 
Monogr. 26: 61. 1989. Figure 51. 

Shrubs, 2-8 m tall, leafy stems 1 .8-4 mm thick, stri- 
gulose with stiff straight hairs to 1 mm long; stipules with 
a short (1.5-3 mm) sheath (difficult to see under the 
strigulose hairs) and narrow distal teeth 6-17 mm long, 
covered with long (1-2 mm) thin hairs. Leaves opposite, 
petioles 4-12 mm long, 1-1.9 mm thick, puberulent in 
early stages with straight or curved ascending hairs; leaf 
blades 7-14 cm long, 2.5-4.5 cm broad, elliptic-oblong 
to narrowly elliptic, apex acuminate with tip 6-10(-13) 
mm long, base acute, drying stiffly chartaceous, yellow- 
ish, upper surface glabrous, glabrous on 2 veins and 
interveinal areas beneath but with straight stiffhairs 0.5- 
1.5 mm long on the sides of the midvein, 2 veins 8-1 3/ 
side. Inflorescences solitary, 7-1 1 cm long, 7-8 cm broad, 
pyramidal, yellowish, peduncles 3.5-6.5 cm long and 1 .5 
mm thick, strigulose, bracts and bracteoles 3-7 (-1 3) mm 
long, ca. 1 mm broad, yellowish and resembling the calyx 
lobes, pedicels 2-6 mm long. Flowers greenish yellow, 
hypanthium ca. 1.3 mm long and 1 mm diam., densely 
pubescent, calyx lobes 5-1 1 mm long, 0.8-1 . 1 mm broad 
near the base, narrowly triangular, acute, sparsely pu- 
bescent with thin whitish hairs ca. 0.5 mm long, drying 
yellow; corolla tubular, yellow, tube ca. 10 mm long, 
with whitish ascending hairs, corolla lobes ca. 2 mm 
long; anthers ca. 2.5 mm long. Fruits not seen. 

Plants of the wet cloud forests of the Caribbean 
slopes and continental divide, at 1200-2100 m 
elevation. Flowering in May-June. This species 
has only been collected above the upper Rio Gran- 
de de Orosi near Tapanti, Cartago, in Costa Rica; 
it is also known from western Panama. 

Palicourea orosiana is distinguished by its pu- 
berulent yellowish inflorescence and flowers, larg- 
er yellowish bracts and calyx lobes, and the long 
stipule lobes. The pubescence along the midvein 
on the underside of the leaf is also unusual. This 
species is similar to P. lasiorrhachis, which has 
much smaller calyx lobes, and to P. macrocalyx, 
which has glabrous inflorescences and broadly im- 
bricate calyx lobes. 



Palicourea padifolia (Willd. ex Roem. & Schult.) 
C. M. Taylor & Lorence, Taxon 34: 669. 1985. 
Psychotria padifolia Willd. ex Roem. & Schult., 
Syst. Veg. 5: 189. 1819. Psychotria mexicana 
Willd. ex Roem. & Schult., loc. cit. 192. 1819. 
Palicourea costaricensis Benth. ex Oerst., Vi- 
densk. Meddel. Dansk. Naturhist. Foren. Kjo- 



208 



FIELDIANA: BOTANY 



benhavn 1852: 38. 1853. Palicourea subrubra 
Polak., Linnaea 41: 571. 1877. P. galeottiana 
sensu Standley, not M. Martens (see Taylor & 
Lorence, 1985, cited above). Figure 51. 



Shrubs or small trees, (l-)2-7(-10) m tall, leafy stems 
1.5-5 mm thick, glabrous (rarely hirsutulous), quadran- 
gular but becoming terete; stipules with a tubular sheath 
1-4 mm long, truncated distally and with narrow awns 
2-10 mm long, glabrous. Leaves opposite (rarely 4/node), 
petioles 4-1 6(-22) mm long, 0.6-1 .8 mm thick, glabrous; 
leaf blades 6-16(-24) cm long, 2-6(-8.5) cm broad, nar- 
rowly elliptic-oblong to narrowly oblong, elliptic-ovate 
or lanceolate (rarely elliptic-obovate), apex acute to acu- 
minate with tip 4- 1 2(-20) mm long, base acute to obtuse, 
drying stiffly chartaceous, glabrous and with short (0.2 
mm) cystoliths often visible above, glabrous beneath 
except for thin whitish hairs 0.3-0.9 mm long on the 
major veins beneath or only along the midvein, 2 veins 
8-14/side. Inflorescences (5-)7-18 cm long, 4-14 cm 
broad, broadly pyramidal, reddish purple to salmon red 
or orange (yellow), peduncle 1.5-5 cm long, 1.2-2 mm 
thick, glabrous or sparsely puberulent, bracts ( l-)2-6(-8) 
mm long and linear-subulate, bracteoles 0.5-2 mm long, 
pedicels l-6(-10) mm long. Flowers glabrous or sparsely 
puberulent externally, hypanthium 0.7-1 mm long, 0.5- 
1 mm diam., calyx lobes 0.5-1 mm long, triangular, 
obtuse; corolla tubular to slightly funnelform, carnose, 
orange or yellow, tube 8-15(-18) mm long and 1.5-2.5 
mm diam., narrower in the middle, corolla lobes 2-3 
mm long, ca. 1.5 mm broad at the base, obtuse; anthers 
2.5-4 mm long. Fruits 4-6(-10) mm long, 4-6(-10) mm 
diam., ovoid to ellipsoid or globose, longitudinally ribbed, 
persisting calyx ca. 1 mm high. 



Plants of evergreen lower montane forest for- 
mations and in moist sites in deciduous forest for- 
mations, (800-)1000-2000(-2400) m elevation. 
Flowering and fruiting throughout the year (mostly 
flowering in December-August and fruiting in Jan- 
uary-August). The species is quite common in 
Costa Rica and ranges from the Cordilleras de 
Guanacaste and Tilaran, around the Caribbean 
side of the central volcanic chain, through the Cor- 
dillera de Talamanca. The species ranges from 
eastern Mexico to Panama. 

Palicourea padifolia is recognized by its usually 
narrow leaves, short-tubular stipule sheaths, col- 
orful inflorescences with (usually) reddish pedi- 
cels, minute calyx lobes, narrow yellowish (or or- 
ange) corolla tubes, and montane habitats with 
high rainfall. This is the most commonly collected 
species of Palicourea in Central America. This spe- 
cies is easily mistaken for P. angustifolia. P. crocea, 
and P. purpurea, but those species have purplish 
flowers. Palicourea lasiorrhachis is similar but has 
yellow inflorescence branches and corollas. Pali- 
courea padifolia is very closely related to P. thyr- 
siflora (Ruiz & Pavon) Roem. & Schult. of Ecuador 



and Peru and may be conspecific, but neither of 
the two species has been collected in Colombia or 
the eastern half of Panama. Fruiting plants of this 
species may be confused with species of Psy- 
chotria, but the inflorescences are more colorful 
in P. padifolia. 



Palicourea purpurea C. M. Taylor, Syst. Bot. 
Monogr. 26: 71. 1989. Figure 52. 

Shrubs or small trees, (l-)3-6(- 10) m tall, leafy stems 
1-4 mm thick, glabrous, stipules with sheaths 1-4 mm 
long, truncated distally and with slender teeth 2-5 mm 
long, glabrous and persisting. Leaves opposite, petioles 
6-25(-35) mm long, 0.6-1.3 mm thick, glabrous; leaf 
blades 7-19 cm long, 2-7 cm broad, narrowly elliptic to 
elliptic-oblong, apex acuminate with tip 5-15 mm long, 
base acute and slightly decurrent on petiole, drying char- 
taceous, dark brown above, glabrous above and below 
or with a few thin white hairs along the side of the mid- 
vein beneath, 2 veins 6-10/side. Inflorescences 5-1 2 cm 
long, 4-1 1 cm broad, pyramidal, purplish to deep lav- 
ender in color, peduncles 2-6 cm long, 1 .2-1 .8 mm thick, 
glabrous, bracts 2-7 (-10) mm long, linear-subulate, 
bracteoles 1-3 mm long, pedicels 0-1 1 mm long. Flowers 
glabrous externally, hypanthium 0.6-1 mm long and 1 
mm diam. at apex, obconic, calyx lobes 0.3-1 long and 
1 mm broad at the base, triangular; corolla tubular and 
carnose, purple to lavender, pink, or white, tube 11-18 
mm long and 2-4 mm diam.. slightly expanded at the 
base, corolla lobes 2-4 mm long; anthers 3.5-4 mm long. 
Fruits 4-6 mm long and 4-5 mm diam., globose, drying 
black and with longitudinal ridges. 

Plants of montane evergreen forest formations, 
(750-)1 200-2800 m elevation. Flowering in March, 
May-July, and December; fruiting in January- 
September and December. The species is found 
on the Caribbean slope and continental divide of 
the Central Volcanic chain and in the area above 
Rio Grande de Orosi and in western and central 
Panama. 

Palicourea purpurea is distinguished by its lon- 
ger purplish corollas, purple inflorescence branch- 
es, and globose fruit. It is very similar to P. padi- 
folia, but that species has yellow or orange corollas 
and red or orange inflorescence branches, and ovoid 
or slightly flattened fruit. Palicourea discolor is 
also similar but has larger leaves, inflorescences, 
and calyx lobes. Compare P. angustifolia also. 



Palicourea salicifolia Standl.. J. Wash. Acad. Sci. 
18: 280. 1928. P. austinsmithii Standl., Publ. 
Field Mus. Nat. Hist., Bot. Ser. 18: 1333. 1938. 
P. caerulescens Suesseng., Bot. Jahrb. Syst. 72: 
286. 1942. Figure 53. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



209 



Shrubs, 0.5-4 tall, leafy stems 1-4 mm thick, glabrous 
and quadrangular, young stems drying black, pale gray 
in age; stipules with short (1-2 mm) tubes truncated and 
with narrow teeth 1-2 mm long, glabrous. Leaves op- 
posite, petioles 3-9(-15) mm long, 0.4-0.8 mm thick, 
glabrous; leaf blades 4-8(-10) cm long, l-2(-3) cm broad, 
narrowly elliptic to lanceolate or lanceolate-oblong, apex 
usually tapering gradually and acute or acuminate, base 
acute and slightly decurrent on petiole, drying stiffly 
chartaceous, glabrous above and below (rarely minutely 
puberulent along the mid vein), 2 veins 9-13/side. In- 
florescences 2-6 cm long, 3-6(-8) cm broad, pyramidal, 
green or marked with blue or purple, peduncles 3-18 
mm long, glabrous and usually drying black, bracts 4-7 
mm long, 1 mm broad at the base, pedicels 0-8 mm 
long, bracteoles ca. 3 mm long. Flowers glabrous exter- 
nally, hypanthium ca. 1.3 mm long and 1 mm diam. 
distally, calyx lobes 2-3(-4) mm long, ca. 1 mm broad, 
oblong and rounded distally; corolla funnelform, car- 
nose, white and often flushed with blue or purple, tube 
6-13 mm long, ca. 2 mm diam., narrowed in the middle, 
corolla lobes 3-6 mm long, to 2 mm broad at the base; 
anthers 2-4 mm long. Fruits 6-8 mm long and 5-7 mm 
diam., obovoid or ellipsoid, flattened, becoming deep 
blue (pedicels also blue), with strong longitudinal ribs 
(dried). 



Plants of evergreen montane forest formations, 
from 1 500 to 2700 m elevation. Flowering in Jan- 
uary-September; fruiting in December-March and 
May-August. The species is known from Monte- 
verde and the northern part of the Meseta Central 
(near Zarcero), from the Cerro de Carpintera, and 
in the Cordillera de Talamanca. 

Palicourea salicifolia is recognized by the small 
narrow leaves tapering at both apex and base, small 
inflorescences that often dry black, slightly en- 
larged calyx lobes, and funnelform white corollas 
marked with blue or purple. The fruit have sharply 
denned longitudinal ribs. Two collections from be- 
low 2000 m (Dryer 768 & 965 from Monteverde 
and Stork 1 175 from Carpintera) have dried pale 
greenish and the inflorescences are little-branched. 
Because of its small leaves, this species resembles 
P. adusta and P. montivaga with shorter corolla 
tubes and calyx lobes. 



Palicourea skotakii C. M. Taylor, Syst. Bot. 
Monogr. 26: 76. 1989. Figure 49. 

Shrubs, 1.5-2 m tall, leafy stems 1.5-5 mm thick, 
glabrous, quadrangular or flattened; stipules 6-8 mm 
long, tubular sheath 3-8 mm long, broadly bilobed dis- 
tally with obtuse lobes ca. 2 mm long and 3 mm broad, 
glabrous, becoming coriaceous. Leaves opposite, petioles 
10-24 mm long, 1-1.6 mm thick, glabrous; leaf blades 
7-13(-17) cm long, 3-5-5. 5(-6.5) cm broad, elliptic to 
elliptic-oblong or elliptic-obovate, apex abruptly nar- 



rowed and acuminate, base acute and slightly decurrent 
on petiole, drying stiffly chartaceous to subcoriaceous, 
dark greenish above, glabrous above and below, 2 veins 
9-14/side and joining a vein along the leaf edge distally, 
parallel minor 2 veins usually present. Inflorescences 
4-10 cm long, 5-1 1 cm broad, usually 3-branched, 
rounded to pyramidal, peduncle 2.5-4 cm long and 3 
mm thick, glabrous, bracts 1 5-30 mm long, pale green 
(dried), bracteoles to 1 5 mm long, ovate and imbricate 
around the flowers, persisting. Flowers glabrous exter- 
nally, calyx lobes 2-6 mm long, lanceolate to ligulate, 
usually unequal; corolla funnelform, carnose, white, tube 
ca. 10 mm long, corolla lobes ca. 6 mm long; anthers 
ca. 2.5 mm long. Fruits 6-7 mm long, 5 mm diam., 
ellipsoid to slightly obovoid. 

Plants of high montane rain forest formations 
at 2700-2800 m elevation. Flowering in March 
and June. Endemic; like the type (Taylor & Skotak 
4756 DUKE), all collections are from along the In- 
teramerican Highway in the western part of the 
Cordillera de Talamanca. 

Palicourea skotakii is distinguished by its high- 
altitude habitat, stipules with broad lobes, lack of 
pubescence, large greenish ovate persisting bracts 
and bracteoles, and white corollas. This species 
resembles P. bella\ it also resembles Psychotria 
chlorochlamys and its allies. 



Palicourea spathacea C. M. Taylor, Syst. Bot. 9: 
226. 1984. Figure 49. 

Shrubs or small trees, 2-8 m tall, leafy stems 2-8 mm 
thick, quadrangular, with thin brownish hairs to 0.5 mm 
long but soon glabrescent; stipules 5-12 mm long, 4-8 
mm broad, bilobed, puberulent near the base and along 
the midrib abaxially. Leaves opposite, petioles 8-20 mm 
long, 1-2.2 mm thick, appressed pubescent with thin 
ascending hairs or glabrescent; leaf blades 12-28 cm 
long, 4.5-14 cm broad, elliptic-obovate to broadly ellip- 
tic, obovate or elliptic-oblong, apex short-acuminate with 
tip 8-15 mm long, base narrowed gradually and acute 
or cuneate, slightly decurrent on petiole, drying stiffly 
chartaceous, dark green or brown above (much paler 
beneath), glabrous above, with thin brownish ascending 
hairs to 1 mm long on the midvein beneath and shorter 
(0.1-0.3 mm) hairs on the minor venation and surfaces, 
2 veins 12-22/side (and usually with a weaker 2 be- 
tween them). Inflorescences solitary, drooping, 10-23 
cm long, 8-16(-20) cm broad, open broadly pyramidal 
or hemispheric, peduncles 3-9 cm long, 1.5-3.5 mm 
thick, puberulent near the base, bracts 1 5-20 mm long 
and 5 mm broad, bracteoles 8-15 mm long, ca. 4 mm 
broad, pink to purple (white), glabrous pedicels 12-26 
mm long, ca. 0.5 mm thick and black when dried. Flow- 
ers glabrous externally, hypanthium ca. 1.5 mm long and 
1.2 mm diam., cylindrical and drying black, calyx tube 
spathe-like and 1 5-20 mm long by 5-8 mm diam., split- 
ting open along 1 side, pink to purple; corolla tubular, 
camose, white, tube 3-4 cm long, 4-5 mm diam., straight 



210 



FIELDIANA: BOTANY 



or slightly curved, corolla lobes 4-7 mm long; anthers 
4.5-5 mm long. Fruits ca. 8 mm long (not including the 
persisting calyx), ca. 7 mm in diam., dark purple but 
drying black and with longitudinal ribs. 

Plants of the very wet lower montane rain forest 
formations on the Caribbean slope of the Cordil- 
lera de Talamanca, at 1300-1700 m elevation. 
Flowering in March, April, and November-De- 
cember; fruiting in April-May, July-August, and 
December. This species is only known from the 
upper drainage area of Rio Grande de Orosi near 
Tapanti, Cartago Province. 

Palicourea spathacea is a striking species; the 
bright pink or purplish bracts, bracteoles, and large 
spathaceous calyx make the inflorescences es- 
pecially conspicuous. The long white corollas, large 
leaves with many secondary veins and cuneate 
bases, and stipules with a single obtuse apex on 
each side are further distinctions. The calyx is in- 
flated before anthesis and splits for half to three- 
quarters of its length as the corolla emerges; there 
are no calyx lobes. 



Palicourea standleyana C. M. Taylor, Syst. Hot. 
Monogr. 26: 81. 1989. Psychotria brenesii 
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 
18: 1347. 1938, not Palicourea brenesii ; Standl., 
1938. Figure 54. 

Shrubs or small trees, 2-7 m tall, leafy branchlets 3- 
6 mm thick, densely pilose with thin yellowish hairs 1- 

3 mm long; stipules (6-)8-18 mm long, 3-8 mm broad, 
with a tubular sheath 4-8 mm long, acute or with 2 short 
(1-2 mm) lobes, sparsely pilose. Leaves with petioles 1- 

4 cm long, 0.7-1.7 mm thick, pilose with straight or 
slightly curved hairs to 2 mm long; leaf blades 9-23 cm 
long, 3-9 cm broad, broadly elliptic, elliptic-ovate or 
elliptic-obovate, apex acuminate to caudate-acuminate 
with 8-15(-20) mm long, base obtuse to acute, drying 
stiffly chartaceous, dark brown above, densely pilose 
above and below with hairs 1-1.5 mm long, 2 veins 
(8-) 10-1 7/side, connected distally to make a slightly ar- 
cuate submarginal vein. Inflorescences terminal, 3 or 1 
with 3 main branches from a short (0-1 5 mm) common 
peduncle, 7-12 cm long, to 14 cm broad, open trichot- 
omous panicles, primary (basal) lateral branches 3-7 cm 
long, pilose, bracts 4-1 1 mm long, ca. 1.5 mm broad, 
lanceolate, flowers in close distal clusters subtended by 
ovate pilose bracts, pedicels 0-5 long. Flowers puberu- 
lent externally, hypanthium ca. 2 mm long, calyx lobes 
2-4 mm long but difficult to see among the hairs; corolla 
tubular-funnelform, camose, deep lemon yellow to 
greenish yellow, tube 10-12 mm long, 1.5-2 mm diam., 
expanded at the base and with hairs to 2 mm long, corolla 
lobes 3-5 mm long; anthers 2.5-3 mm long. Fruits 6-8 
mm long (not including the 3-mm-long calyx), 5 mm 
diam., ellipsoid, pilose, becoming blue. 



Plants of very wet cloud forest formations along 
the Caribbean slope and continental divide, 1 100- 
1600 m elevation. Flowering in December-July; 
fruiting in January-June. The species has been col- 
lected in the Cordilleras de Guanacaste and Ti- 
laran, north of San Ramon (Alajuela), near Ta- 
panti (Cartago), in Chiriqui, Panama, and Narino 
Province, Colombia. 

Palicourea standleyana is recognized by the 
prominent pubescence on all parts, the large tu- 
bular stipules, stiff leaves with many secondary 
veins and submarginal veins, open three-branched 
inflorescences, and bright yellow corollas. This 
species is easily confused with Psychotria pilosa, 
but that species has much smaller corollas and 
smaller fruit and the corolla tubes lack the internal 
ring of the hairs that define Palicourea. 



Palicourea tilaranensis C. M. Taylor, Syst. Bot. 
Monogr. 26: 84. 1989. Figure 49. 

Small shrubs, ca. 1.5 m tall, leafy stems ca. 3 mm 
thick, quadrangular and glabrous, young stems drying 
black; stipules 5-8 mm long, with a very short (0.5-1.2 
mm) tubular sheath, stipule lobes to 4 mm long, broadly 
obtuse to deeply bilobed, persisting. Leaves opposite, 
petioles 13-40(-65) mm long, 0.7-1.7 mm thick, gla- 
brous; leaf blades 8-16 cm long, 3.5-6 cm thick, ovate 
to ovate-elliptic or ovate-oblong, apex acuminate with 
tip 7-14 mm long, base obtuse, drying stiffly chartaceous, 
dark greenish brown, glabrous above and below, 2 veins 
8-1 I/side. Inflorescences 12-15 cm long. 6-9 cm broad, 
pyramidal panicles with opposite branching, peduncles 
4-6 cm long, 1 .5-2 mm thick, glabrous, bracts 8-1 3 mm 
long, 2-4 mm broad, lanceolate, bracteoles ca. 5 mm 
long and 2 mm broad, flowers sessile or on pedicels to 
4 mm long within the persisting bracteoles. Flowers gla- 
brous externally, hypanthium 0.5-1 mm long, calyx lobes 
0.3-1.5 mm long, triangular; corolla tubular, carnose, 
white, tube 3-7 mm long, ca. 1.8 mm diam., slightly 
narrowed in the middle, corolla lobes ca. 2 mm long. 
Fruits not seen. 



Plants of evergreen lower montane rain forest 
formations, at about 15500-1800 m elevation. 
Flowering in February and June. This species is 
known only from the Monteverde Nature Reserve, 
in the Cordillera de Tilaran. 

Palicourea tilaranensis is recognized by its re- 
stricted range, large and persisting bracts and brac- 
teoles, small calyx lobes, and white corollas. It 
resembles Psychotria palicoureoides and its allies. 



Palicourea triphylla DC., Prodr. 4: 526. 1 830. Psy- 
chotria triphylla (DC.) Muell.-Arg. in Mart., Fl. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



21 



Bras. 6(5): 233. 1&&1. Palicourea longibracteata 
Bartlingex DC., Prodr. 4: 527. 1830. Palicourea 
parviflora Benth., Bot. voy. Sulph. 107. 1844. 
Figure 50. 



Shrubs or herbaceous subshrubs, l-3(-5) m tall, erect 
and usually unbranched (rarely clambering), leafy inter- 
nodes 2.3-7 mm thick, glabrous or sparsely pubescent 
on the new growth and nodes with short (0.1-0.3 mm) 
thin hairs, terete; stipules united for only 1-2 mm at the 
base or free, stipule lobes 6/node. 6-12(-15) mm long 
and ca. 2 mm broad at the base, narrowly triangular and 
acute, glabrous abaxially and ciliolate along the edge. 
Leaves 3/node (rarely 2 or 4), petioles 3-7(-l 5) mm long, 
1.4-2 mm broad, minutely pubescent but glabrescent; 
leaf blades 7-20 cm long, 3-8 cm broad, narrowly ellip- 
tic-oblong to lanceolate, narrowly elliptic or oblanceo- 
late, apex tapering gradually and acuminate with tip 5- 
1 5 mm long, base cuneate to acute (obtuse) and slightly 
decurrent on petiole, drying thinly to stiffly chartaceous, 
minute (0.05-0.2 mm) thin hairs along the major veins 
above, with thin whitish hairs 0.1-0.3 mm long on the 
veins and surfaces beneath, 2 veins 7-1 l(-15)/side. In- 
florescences 6-20 cm long (to 26 cm in fruit), 5-10 cm 
broad near the base, pyramidal to narrowly thyrsoid- 
cylindrical, red to orange or orange-yellow, peduncles 5- 
12(-18) cm long, 1.5-3 mm thick, puberulent with thin 
whitish hairs, bracts 5-20(-30) mm long linear-subulate, 
pedicels 1-5 mm long, puberulent. Flowers distylous, 
minutely puberulent, hypanthium ca. 1 mm long and 
0.5 mm diam., tubular, calyx lobes 0.2-1 mm long, 
broadly triangular; corolla tubular or slightly funnelform, 
yellow or reddish distally, tube 8-14 mm long, 1.5-3 
mm diam., inflated at the base, corolla lobes 1-2 mm 
long; anthers 2.5-3.5 mm long. Fruits 4-5 mm long, 3- 
5 mm diam., ovoid to globose with prominent longi- 
tudinal ridges (dried), becoming blue, purple, or black 
at maturity, glabrous or puberulent, calyx ca. 0.7 mm 
high. 



Plants of open wet or poorly drained sties in 
evergreen or partly deciduous forest formations in 
the Caribbean lowlands and on the Pacific slope 
of central and southern Costa Rica and in Panama, 
from near sea level to 700 m elevation, but to 1 300 
m on the semideciduous Pacific slope. Flowering 
and fruiting throughout the year (but flowering 
mostly in April-July). The species ranges from the 
Caribbean side of central Mexico and Cuba to 
Brazil and Bolivia. 

Palicourea triphylla is recognized by its ternate 
leaves, puberulence of short thin hairs, long per- 
sisting stipules with six teeth per node, orange- 
yellow inflorescences with persisting bracts, small 
corolla lobes, and gibbous corollas with multicel- 
lular hairs. The species is often found in open 
swampy sites. 



Palicourea vestita Standl., J. Wash. Acad. Sci. 18: 
277. 1928. Figure 51. 

Shrubs or small treelets, 1-4 m tall, leafy stems 1.5- 
5 mm thick, densely hirsutulous with yellowish hairs 
0.3-0.9 mm long; stipules with a tube 3-6 mm long, with 
narrow teeth 3-8 mm long, puberulent, persisting. Leaves 
opposite, petioles 6-1 8(-30) mm long, 0.8-1 .6(-2.2) thick, 
densely puberulent with yellowish hairs; leaf blades 
(6-)8.5-19 cm long, (1.7-)2.5-7 cm broad, elliptic to 
elliptic-oblong, elliptic-obovate or lanceolate, apex acu- 
minate (acute) with tip to 1 cm long, base acute to cuneate 
(obtuse), drying stiffly chartaceous, often yellowish green, 
glabrous above or with thin hairs along the midvein, 
pubescent along the major veins beneath with thin yel- 
lowish hairs 0.4-1.2 mm long, 2 veins 12-20/side and 
with minor short parallel 2 veins between them. Inflo- 
rescences 5-12 cm long, 5-8 cm broad near the base, 
thyrsiform pyramidal, yellowish, peduncles 2-8 cm long, 
1-3 mm thick, densely pubescent, bracts 4-10 mm long 
and 1.5 mm broad, bracteoles 1-4 mm long and per- 
sisting, pedicels 2-5 mm long. Flowers sparsely and mi- 
nutely puberulent externally, hypanthium 1-1.5 mm long, 
obconic, calyx lobes 0.7-1.5 mm long, acute; corolla 
bright yellow, tube 7-13 mm long, ca. 2 mm diam., 
slightly inflated at the base, corolla lobes 1.5-3 mm long; 
anthers 2.1-2.5 mm long. Fruits 5-8 mm long, 3.5-6 
mm diam., obovoid, persisting calyx ca. 0.7 mm high. 



Plants of evergreen montane wet forest forma- 
tions of the Cordillera de Talamanca, from 1 200 
to 2600 m elevation. Flowering in April-August; 
fruiting in June-August, October, and December. 
The species is known only from Costa Rica and 
western Panama. 

Palicourea vestita is recognized by the short yel- 
lowish hairs on most plant parts, the leaves with 
many secondary veins, the elongate thyrsiform in- 
florescences, and the bright yellow corolla tube. 
This species is distinguished from the very similar 
but more common P. lasiorrhachis by the more 
numerous secondary veins and longer calyx lobes. 



Pentagonia Bentham 
Nomen conservandum 

Shrubs or small trees (rarely large trees), main stems 
often unbranched or with few lateral branches, stems 
thick, flattened, quadrangular or terete in early stages, 
glabrous or puberulent, often with conspicuous rounded 
lenticels; stipules interpetiolar, free, large, triangular, 
usually early deciduous. Leaves opposite and decussate, 
usually very large, sessile or petiolate, the petioles some- 
times with auriculate (leafy) developments at the base; 
leaf blades entire or pinnatind, usually drying coriaceous, 
the minor venation lineolate-parallel and often with 2 
distinct orientations in the same leaf area (parallel with 
the tertiary veins and at right angles to the tertiary veins), 



212 



FIELDIANA: BOTANY 



domatia absent. Inflorescences axillary to distal leaves, 
usually short-pedunculate, cymose or corymbose to glo- 
merulate, generally with fewer than 25 flowers, bracts 
developed or minute, flowers subsessile or pedicellate. 
Flowers bisexual and monomorphic, radially symmet- 
rical, often large, usually densely pubescent externally, 
hypanthium conical to turbinate or campanulate, calyx 
tube well developed, calyx lobes 5-6 or spathaceous, 
lobes equal or unequal, often with glands inside at the 
base; corolla tubular to funnelform, carnose. white to 
yellow or red, glabrous within the throat, villous at the 
stamen attachment, corolla lobes 5-6, valvate in bud, 
short; stamens 5-6, filaments borne on the middle of the 
tube, equal or unequal, usually villous at the base, an- 
thers dorsifixed, included; ovary 2-locular, ovules many 
on expanded elongate placentas borne on the septum, 
stigmas subcapitate or branched. Fruits baccate, fleshy 
or becoming hard and nut-like when dry, usually globose 
and the surface often lenticellate-muricate, 2-locular, the 
large calyx often persisting on the mature fruit; seeds 
many, angular. 



A genus of about 20 species, ranging from Gua- 
temala into northern and western South America. 
The very large leaves with distinctive minor ve- 
nation, the small axillary inflorescences with larger 
crowded flowers, the stiff pubescent tubular co- 
rollas, and the globose indehiscent fruit with large 
persisting calyx distinguish this genus. The very 
large leaves are correlated with a lack of lateral 
branches on a solitary vertical trunk in many spe- 
cies. The leaves are often reddish or purplish be-. 
neath, making the plants even more striking. The 
genus is seriously in need of revision. The large 
leaves often very variable in form and size, the 
compact inflorescences resulting in crushed or hid- 
den flowers, and a paucity of collections account 
for the fact that the species are still poorly under- 
stood. 



Key to the Species of Pentagonia 

la. Leaf blades entire, never lobed distally; Caribbean lowlands and central cordilleras 2a 

Ib. Leaf blades conspicuously pinnatind or pinnately lobed; Pacific evergreen lowlands 6a 

2a. Leaves and stems with hairs ca. 1 .4 mm long, leaves sessile, the leaf blade slightly auriculate 
near the stem P. hirsula 

2b. Leaves and stems with hairs less than 0.5mm long, leaf blades petiolate and never with expanded 
auriculate tissue near the base 3a 

3a. Calyx lobes usually less than 3 mm long; inflorescences usually open and branched, with the 
branches easily seen at anthesis P. costaricensis 

3b. Calyx lobes usually more than 4 mm long; inflorescences compact, the branches of the inflo- 
rescence short and usually difficult to see (except in fruit) 4a 

4a. Bracts of the inflorescence minute or undeveloped; calyx often absent at the apex of the mature 

fruit; leaves obtuse to rounded/subtruncate at the base; Guatemala to Costa Rica 

P. donnell-smithii 

4b. Bracts of the inflorescence conspicuous in early stages, more than 10 m long and 4 mm broad; 
calyx persistent on the mature fruit; leaves rarely rounded or subtruncate at the base; south- 
ernmost Costa Rica and Panama 5a 

5a. Leaves subsessile with a thick petiole less than 2 cm long, lamina long-cuneate at the base; 
floral bracts 5-10 mm long P. wendlandii 

5b. Leaves with well-developed petioles more than 3 cm long, lamina obtuse to cuneate at the base; 

bracts 10-18 mm long P. macrophylla 

6a. Leaf blade long-decurrent (at its base) on the petiole . . P. linajita 

6b. Leaf blade not at all decurrent on the petiole P. gymnopoda 



Pentagonia costaricensis (Standley) W. Burger & 
C. M. Taylor, comb. nov. Nothophlebia costari- 
censis Standl., Contr. U.S. Natl. Herb. 17: 438. 
1914. Figure 14. 

Trees, 5-18 m tall, leaf stems 6-12(-22) mm thick, 
glabrous or subglabrous, flattened-quadrangular in early 
stages, often with dark round lenticels ca. 1 mm diam.; 



stipules 2-6(-8) cm long, lanceolate, glabrous or mi- 
nutely appressed-sericeous. Leaves with petioles (2.5-)4- 
7 cm long. 2.5-7 mm thick, glabrous or subglabrous and 
drying brown; leaf blades (2 l-)27-l 00 cm long, ( 1 2-)l 6- 
50 cm broad, broadly elliptic-obovate to very broadly 
obovate, apex abruptly narrowed and broadly obtuse, 
base obtuse and only slightly decurrent (rarely slightly 
auriculate at the petiole as in Gomez- Laurito 9321 CR, 
F), margin entire, drying coriaceous, glabrous above and 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



213 



below or with minute sericeous hairs on the veins be- 
neath, 2 veins 8- 11 /side, the minor venation closely 
parallel in 2 different directions (but both systems not 
always apparent). Inflorescences 3-12 cm long, 3-15 cm 
broad, often dichotomously branched, the primary pe- 
duncle 5-20 mm long, secondary branches 5-20 mm 
long, major branches lacking subtending bracts, ultimate 
branches usually with 3-flowered cymes, striate and gla- 
brous with elongate (0.4-2 mm) lenticels, pedicels 0-7 
mm long. Flowers with hypanthium 2-4 mm long and 
2-3 mm diam., obconic, minutely puberulent or gla- 
brous externally, calyx tube 47(-12) mm long, 4.5-6 
mm diam., cupular to short-tubular, with parallel ve- 
nation, rose-colored, calyx lobes 0.5-2 mm high, 2-4 
mm broad, broadly rounded to obtuse or obscure, vari- 
able and often unequal; corolla cream white or yellow, 
tube 12-20 mm long, 3-5 mm diam., minutely yellowish 
puberulent externally, corolla lobes 3-6 mm long; an- 
thers white. Fruits (based on Hammel& Grayum 14292) 
10-20 mm diam., globose with a persistent calyx tube 
4-5 mm high and 4 mm diam. at the top, calyx lobes 
often obscure. 

Trees of evergreen lowland rain forest forma- 
tions, 5-600(-900) m elevation. Flowering in 
April-May and November; fruiting in May, July- 
September, and November. The species is known 
only from the Caribbean lowlands, from northern 
Costa Rica to western Panama. 

Pentagonia costaricensis is recognized by the very 
large leaves, unusual minor venation, branched 
inflorescences, short calyx lobes, and smaller fruit. 
Standley erected the genus Nothophlebia for this 
unusual species but admitted (Standley 1938, p. 
1329) that it might be referred to Pentagonia. A 
collection from 1300-1400 m near Las Alturas 
(Almeda et al. 6699 F) with larger calyx lobes and 
glabrous corolla tube is tentatively placed here. 



Pentagonia donnell-smithii (Standl.) Standl., J. 
Wash. Acad. Sci. 17: 170. 1927. Watsonamra 
donnell-smithii Standl., Contr. U.S. Natl. Herb. 
17: 442. 1914. Figure 14. 

Small trees or treelets, 2-7(-12) m tall, often with a 
single stem, leafy stems 5-22 mm thick, at first ap- 
pressed-sericeous but quickly becoming glabrous; stip- 
ules 2.5-9 cm long, 6-25 mm broad, lanceolate to nar- 
rowly ovate, with appressed-ascending sericeous hairs 
0.2-0.5 mm long externally. Leaves often reddish be- 
neath when young, petioles (2-)3-l 2 cm long, 1 .5-6 mm 
thick, appressed-puberulent with thin ascending hairs ca. 
0.2 mm long; leaf blades (15-)26-90 cm long, (9-) 13- 
50 cm broad, broadly elliptic to broadly elliptic-oblong 
or broadly ovate, apex bluntly obtuse or rounded, base 
rounded and subtruncate to cuneate-obtuse, margin en- 
tire and without lobes, drying chartaceous to subcoria- 
ceous, pale to dark grayish green, glabrous above, sparse- 
ly to moderately puberulent with thin ascending hairs 
on the veins beneath, 2 veins 10-14/side, minor ve- 



nation closely parallel. Inflorescences 2.5-5 cm long, to 
8 cm broad (including both inflorescences of the node), 
primary peduncle 1-2 cm long and 2-3 mm thick branches 
of the inflorescence usually short and obscured by the 
flowers, bracts 0-2 mm long, pedicels 0-6 mm long. 
Flowers with hypanthium 3-7 mm long, ca. 3 mm diam. 
distally, calyx tube 6-10 mm long, ca. 6 mm diam., 
minutely ascending sericeous, calyx lobes 4-8 mm long, 
34 mm broad, broadly rounded distally, corolla yellow 
or white, corolla tube 1 5-30 mm long, 3-5(-6) mm diam. 
and broadest near the base, with short thin hairs ca. 0.2 
mm long, corolla lobes 5, 5-10 mm long, acute. Fruits 
16-40 mm diam., globose or globose-ovoid, outer wall 
hard and 2-3 mm thick, with small (0.5 mm) tuberculate 
lenticels, calyx deciduous or less often persisting on the 
mature fruit; seeds 3-5 mm long, ellipsoid, orange. 

Trees of evergreen rain forest formations on the 
Caribbean slope and lowlands of Costa Rica, from 
near sea level to 900 m elevation. Flowering in 
March-July and October-November in Costa Rica; 
probably fruiting throughout the year. The species 
ranges from Guatemala southward along the Ca- 
ribbean to central Costa Rica. 

Pentagonia donnell-smithii is recognized by the 
very large leaves with unusual minor venation, the 
small compact inflorescences lacking developed 
bracts, the prominent calyx lobes, stifFcorolla tubes, 
and usually globose fruit. The mature fruit in most 
Costa Rican collections are lacking the calyx, but 
the calyx persists regularly in Guatemala and Hon- 
duras. Standley (1938) suggested that this species 
be submerged in P. macrophylla. But while the 
two species appear to be closely related, P. macro- 
phylla has a red calyx, white corolla, and large 
distinctive bracts subtending the branches of the 
inflorescence in early stages. No such bracts are 
seen in P. donnell-smithii. Also, this species does 
not appear to occur in eastern Costa Rica, where 
intergradation might be expected. 

Sterile material of these species, and of P. cos- 
taricensis and P. wendlandii, may not be separable, 
since there is considerable variation within each 
species. For example, a dwarf treelet 1 m tall with 
very short (2 cm) petioles and slightly obovate 
leaves resembling the leaves off. wendlandii was 
collected near Puerto Viejo de Sarapiqui (A. Ji- 
menez 3424 CR). This collection lacks the bracts 
that are found in both P. macrophylla and P. wend- 
landii and is therefore placed here under P. don- 
nell-smithii. 



Pentagonia gymnopoda (Standl.) Standl., J. Wash. 
Acad. Sci. 17: 171. 1927. Watsonamra gym- 
nopoda Standl., Contr. U.S. Natl. Herb. 1 7: 444. 
1914. 



214 



FIELDIANA: BOTANY 



Small treelets, 1-6 m tall, usually with a vertical un- 
branched stem, leafy stems 6-16 mm thick, somewhat 
flattened in early stages but becoming terete, glabrous, 
grayish; stipules 4-6 cm long, 2-3.5 cm broad, broadly 
triangular to lanceolate, apparently glabrous or minutely 
appressed-puberulent. Leaves with petioles 5-12 cm long, 
2.7-5 mm thick, glabrous or minutely (0. 1 mm) puber- 
ulent; leaf blades 28-70(-100) cm long, 30-45(-70) cm 
broad, ovate to oblong in general outline but deeply 
pinnatitid to the distal part of the blade, with 5-7 major 
oblong lobes, the lobes 5-25(-35) cm long and 2-7(-8) 
cm broad, apex acute to acuminate, base truncate to 
acute, drying chartaceous, dull grayish green above, gla- 
brous above, minutely (0.1-0.2 mm) papillate-puberu- 
lent on the veins beneath, 2 veins 5-9/side. Inflores- 
cences closely crowded in the leaf axils, 2-5 cm long, 
subsessile and partly covered by the broad stipules or 
with a peduncle up to 1 cm long, bracts to 3 cm long, 
flowers closely crowded and apparently sessile. Flowers 
with hypanthium ca. 10 mm long, densely appressed- 
puberulent, calyx tube 6-10 mm long and ca. 6 mm 
diam., calyx lobes 5, 18-15 mm long, 5-8 mm broad, 
stiff and parallel veined; corolla white to yellow or pale 
green, tube ca. 20 mm long, puberulent externally, co- 
rolla lobes ca. 10 mm long; stamens 5, filaments 8-10 
mm long and unequal, anthers ca. 3 mm long; style ca. 
9 mm long. Fruits 15-20 mm diam., globose to ovate 
or pyriform, the persisting calyx to 22 mm long and 6 
mm diam. 

Poorly known plants of evergreen lowland rain 
forest formations. Flowering in January-Febru- 
ary; fruiting in January-February. The species 
ranges from the Carara Biological Reserve 
(8435'W) southeastward along the Pacific low- 
lands to central Panama. 

Pentagonia gymnopoda is recognized by the very 
large deeply pinnatifid leaves, small axillary inflo- 
rescences, congested flowers, and sessile fruit. The 
broad stipules often cover part of the inflorescence. 
Currently known from only about four collections 
in Costa Rica. Compare the closely similar P. ti- 
najita. 



Pentagonia hirsute Standl., J. Wash. Acad. Sci. 
17: 170. 1927. 

Probably small treelets, young branches ca. 10 mm 
thick, hirsute; stipules not seen. Leaves sessile and entire; 
leaf blades ca. 60 cm long and 27 cm broad, broadly 
obovate or broadly elliptic-obovate, apex short-acumi- 
nate, lamina gradually narrowed below the middle and 
cuneate but merging with a 3-cm-broad basal region that 
is slightly expanded (auriculate) and cordate-clasping at 
the stem, the auricles ca. 2 cm wide on each side, drying 
chartaceous, the upper and lower surfaces with numerous 
thin straight hairs 0.7-1 .8 mm long, 2 veins 14-1 7/side, 
with 1-3 branches distally and obscurely loop-connected 
near the margin. Inflorescences poorly known, small, 
with crowded sessile flowers in the leaf axils. Flowers 



poorly known, hypanthium densely hirsute, calyx ca. 24 
mm long (including both tube and lobes), membranous, 
hirsute with whitish hairs. Fruits unknown. 

Pentagonia hirsuta is a poorly known species 
from about 500 m elevation near Tsaki in the 
Talamanca valley, Limon Province, on the Carib- 
bean slope of easternmost Costa Rica. The type 
and only collection (Tonduz 9415 F, us) was made 
in March 1 895. The long hairs on stems and leaves 
are very unusual within Pentagonia. The leaf shape 
resembles that of P. wendlandii in the long base 
but not in its shape; P. wendlandii seems to have 
more narrowly obovate leaves not as auriculate at 
the base (and without the long hairs). 



Pentagonia macrophylla Ik-nth.. Hot. voy. Sulph. 
105, t. 39. 1845. Watsonamra macrophylla 
(Benth.) Kuntze, Rev. gen. pi. 302. 1891. W. 
pubescens Standl., Contr. U.S. Natl. Herb. 17: 
441. 1914. 

Trees or shrubs to 5(-10) m tall, often unbranched. 
leafy twigs 6-12 mm thick, minutely (0.1-0.3 mm) pu- 
berulent, quadrangular or becoming terete; stipules 3-7 
cm long, 12-15 mm broad at the base, triangular and 
acuminate, glabrous or puberulent. Leaves with petioles 
3-9(-12) cm long, 2.5-5 mm thick, minutely puberulent 
or glabrescent; leaf blades 22^45(-65) cm long 1 1-28(-37) 
cm broad, elliptic to elliptic-obovate or very broadly 
elliptic, apex gradually narrowed and acute, base grad- 
ually narrowed and obtuse or acute, drying subcoria- 
ceous, grayish, glabrous above, minutely puberulent be- 
neath, 2 veins 9-13/side, minor venation parallel. 
Inflorescences axillary, ca. 5 cm long (including the flow- 
ers), dense corymbs of 3-7 flowers, sessile or with short 
(ca. 1 cm) peduncles, bracts 10-18 mm long. 6-9 mm 
broad, oblong-ovate, apex obtuse, red, with many par- 
allel veins, minutely sericeous or glabrous with a ciliolate 
margin, pedicels 0-4 mm long. Flowers ca. 3 cm long, 
hypanthium 6-8 mm long, glabrous or minutely seri- 
ceous, calyx lobes 5, 6-12 mm long (becoming 20 mm 
long in fruit), to 7 mm broad, usually red; corollas yellow, 
sparsely to densely sericeous externally, tube 14-40 mm 
long, 3-10 mm diam., corolla lobes 5, 4-7 mm long, 
ovate with acute apices; stamens differing in size, anthers 
3.5-8 mm long; style 20-25 mm long. Fruits 1 5-28 mm 
diam., globose beneath the persisting calyx, red or or- 
ange; seeds ca. 4 mm long. 

Plants of lowland evergreen forest vegetation, 
at 10-900 m elevation in Panama. Flowering 
mostly in late April-September in central Panama 
(Croat, 1978). The species ranges from near the 
Panama border in Limon Province to Colombia. 

Pentagonia macrophylla is distinguished from 
its congeners by the larger bright red bracts and 
red calyx lobes. We have not seen material of this 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



215 



species collected in Costa Rica, but the species was 
studied near BriBri (Limon) by Lucinda McDade 
(see Oecologia 68: 218-223. 1986). 



Pentagonia tinajita Seem., Bot. voy. Herald. 134. 
1854. P. alfaroana Standl., J. Wash. Acad. Sci. 
17: 171. 1927. Figure 14. 

Small unbranched treelets, l-3(-4) m tall, stems at 
first with 4 prominent ridges or flattened soon becoming 
terete, leafy stems 5-12 mm thick, glabrous; stipules 3- 
5.5 long, 1.2-2 cm broad, ovate-oblong, obtuse or bifid 
with striate parallel venation, puberulent along the mid- 
rib. Leaves with petioles 0-1 1(-20) cm long (variable in 
length due to the decurrent lamina base), 2.5-4.5 mm 
thick; leaf blades 30-80(-100) cm long, 22^10 cm broad, 
pinnately lobed and ovate in general outline (from a 
cuneate base), the sinuses 5-15 cm deep near the base 
and becoming more shallow distally, pinnatifid in the 
lower VB of the lamina, sinuses usually rounded, usually 
obtuse at the apices, abruptly narrowed and cuneate at 
the base with long-decurrent lamina margins running 
down to the petiole or to the leaf base, leaves drying 
thin-chartaceous and dark green above (paler green be- 
neath), with few short (0.2 mm) hairs on the upper sur- 
faces, with more frequent short (0.3-0.4 mm) thin hairs 
on the lower surfaces, 2 veins 6-12/side, minor veins 
0.2-0.4 mm apart. Inflorescences sessile or subsessile, 
densely fasciculate, bracts to 2 cm long and 3-4 mm 
broad, with parallel longitudinal veins and difficult to 
distinguish from the calyx lobes. Flowers with hypan- 
thium ca. 5 mm long, calyx tube 10-15 mm long, calyx 
lobes 4-8 mm long, 3-4 mm broad, with the same stiff 
texture brown color and parallel venation as the bracts; 
corolla tube 2-2.5 cm long, 4 mm diam. at the base and 
8 mm distally, stiff, corolla lobes 6-8 mm long. Fruits 
10-20 mm long and 1 2-1 5 mm diam., globose or ovoid, 
the persisting calyx 10-20 mm long (tube and lobes). 

Poorly known plants of lowland rain forest for- 
mations, 0-200 m elevation. Flowering in Janu- 
ary; fruiting in February and April. The species 
ranges along the Pacific lowlands of central Costa 
Rica to western Panama. 

Pentagonia tinajita is recognized by the large 
thin pinnatifid leaves with decurrent lamina base, 
congested axillary inflorescences, and smaller fruit. 
Pentagonia alfaroana (based on Standley 40194 
F, us) was said to differ because the larger leaf lobes 
are also pinnatifid, but leaf lobing appears to vary 
greatly and does not appear to be a sound basis 
for separating a species. Compare P. gymnopoda, 
which may be conspecific. 



Pentagonia wendlandii Hook., Bot. Mag. pi. 5230. 
1861. Watsonamra wendlandii (Hook.) Kuntze, 
Rev.gen.pl. 320. 1891. 



Shrubs or small treelets to 3 m tall, main stems usually 
unbranched and with a cluster of leaves at the top, leafy 
stems 8-20 mm thick, with 4 prominent ridges, minutely 
appressed-puberulent or glabrous; stipules 2.5-6.5 cm 
long, lanceolate to ovate-oblong, minutely appressed- 
puberulent. Leaves entire and subsessile, petioles 5-15 
mm long, 6-10 m thick; leaf blades 50-100 cm long, 
25-50 cm broad, narrowly obovate to obovate-oblong, 
apex broadly obtuse or rounded, gradually narrowed be- 
low the middle, base cuneate and slightly auriculate, leaves 
drying stiffly chartaceous or subcoriaceous and brown, 
glabrous above, minutely (0.1 mm) puberulent on the 
veins beneath or glabrescent, 2 veins 14-16/side. Inflo- 
rescences to 5 cm long, with ca. 6-15 flowers, peduncles 
ca. 5 mm long, bracts 6-10 mm long, ca. 5 mm broad, 
oblong and brown, with whitish sericeous hairs along 
the midrib, flowers subsessile (or with pedicels to 1 mm 
long, fide Dwyer). Flowers (from Dwyer, 1980) with a 
calyx tube ca. 10 mm long, campanulate, purplish red, 
stiff, puberulent, with numerous glands at the base with- 
in, calyx lobes 5, slightly unequal, 5-10 mm long, 1 calyx 
lobe usually short and acute; corolla yellow, tube ca. 25 
mm long, narrowly cylindrical, glabrous to puberulent 
externally, corolla lobes 4-7 mm long, 2.2-2.6 mm wide, 
oblong to triangular; stamens 5, filaments to 1 6 mm long, 
unequal, anthers ca. 4 mm long, oblong; style ca. 18 mm 
long, stigmas 3.5 mm long. Fruits to 45 mm long, oblong 
rotund, drying black, the persisting calyx to 1 5 mm long. 

Pentagonia wendlandii occurs in Caribbean 
lowland rain forest formations, from near sea level 
to ca. 500 m elevation. The species ranges from 
Bocas del Toro Province to central Panama, but 
we have seen no flowering or fruiting material from 
Costa Rica. (A leaf associated with Gomez- Laurito 
9321 and collected near BriBri may be this species, 
but the flowering sheet appears to be P. costari- 
censis.) The species appears to be closely related 
to P. macrophylla Benth. of central Panama but 
differs in the long-cuneate leaf base with a very 
short petiole. See the discussion under P. donnell- 
smithii. 



Pentas Bentham 

Herbs or shrubs, stems erect or clambering, puberulent 
in most species, from a fibrous or woody rootstock; stip- 
ules interpetiolar with 2-many awns or narrow lobes. 
Leaves opposite or in whorls of 3-5, petiolate; leaf blades 
entire and usually narrowly ovate to lanceolate, pin- 
nately veined, without domatia. Inflorescences terminal 
or axillary to the distal leaf pair, usually much-branched 
and cymose. Flowers bisexual, radially symmetrical, 
monomorphic, dimorphic, or trimorphic, calyx lobes 5, 
equal or unequal with 1-3 longer than the others; corolla 
usually salverform with a narrow tube, throat pilose 
within, corolla lobes 5, ovate to oblong or lanceolate; 
stamens usually included, anthers usually linear and with 
short filaments; ovary 2- or 3-locular, with many ovules 
in each locule. Fruits capsules, usually ovoid and lon- 
gitudinally ribbed, beaked, splitting open at apex into 2 



216 



FIELDIANA: BOTANY 



or 4 valves; seeds minute, tetrahedral or subglobose. with 
reticulate testa. 

A genus of ca. 40 species of Africa, Arabia, and 
Madagascar. The following species is a popular 
garden ornamental with bright red, rose, lilac, or 
white flowers. 



Pentas lanceoiata (Forssk.) Deflers, Voy. Yemen, 
142. 1889. Ophiorhiza lanceoiata Forssk., F. 
Aegypt.-Arab. 42. 1775. Figure 31. 

Herbs or subshrubs with erect or clambering stems, 
0.5-2 m tall, stems 1.5-5 mm thick, puberulent with 
crooked whitish hairs ca. 0.5 mm long; stipules 2-9 mm 
long, with a short base and 3-9 slender setae, persisting. 
Leaves opposite, petioles 0-5(-l 5) mm long, puberulent; 
leaf blades (3-)4-13(-18) cm long, (l-)2-6 cm broad, 
ovate-lanceolate to lanceolate or elliptic, apex acute, base 
acute and slightly decurrent on petiole, drying stiffly 
chartaceous, sparsely to densely puberulent above and 
below, 2 veins 6-9/side and strongly ascending. Inflo- 
rescences terminal or with the distal leaf pair subtending 
the lateral branches, 3-9 cm long and equally broad, 
subcapitate or corymbose and hemispheric, puberulent. 
Flowers often trimorphic (style exserted and anthers in- 
cluded, anthers exserted and style included, both anthers 
and style included), calyx tube 1-3 mm long, calyx lobes 
5, usually unequal with the longer 4-12 mm long; corolla 
brilliant rose red to lilac or white, tube 14-25(-40) mm 
long, ca. 0.5 mm diam. but enlarged in the distal 2-6 
mm and 3-6 mm diam., throat filled with erect whitish 
hairs, corolla lobes 3-10 mm long, acute, ovate-lanceo- 
late to ovate-oblong, acute. 



A popular ornamental grown in gardens around 
the world. In Central America plants with deep 
red or pinkish red corollas are most common. It 
has become naturalized in Colombia. Compare 
Ixora coccinea, another garden favorite. 



Pentodon Hochsteter 

Annual or short-lived herbs, glabrous; stipules united 
to the petiole bases and forming a short truncated sheath 
with minute distal lobes. Leaves opposite, sessile or short- 
petiolate, entire with obscure pinnate venation, domatia 
absent. Inflorescences terminal or axillary and I/node, 
racemose with 1-4 distal nodes and 1-4 long- pedicellate 
flowers at each node or few flowered and irregular (as in 
ours). Flowers bisexual, distylous, glabrous externally, 
calyx with 5 small equal lobes; corolla funnelform, tube 
glabrous or with hairs at the throat within, corolla lobes 
5; stamens 5, included or exserted, subsessile from near 
the base of the tube or between the petal lobes; ovary 
2-locular, with many ovules in each locule borne on 
peltate placentas from the septum. Fruits capsules, lo- 
culicidally dehiscent; seeds many, small, black. 



A genus of probably two African species, with 
one now introduced into pans of the Americas. 
These plants are distinguished from the closely 
related Oldenlandia by the five-parted flowers. 



Pentodon pentandrus (Schumach. & Thonn.) 
Vatke, Oest. Bot. Zeitschr. 25: 231. 1875. Hed- 
yotis pentandra Schumach. & Thonn.. Kongel. 
Dansk. Vidensk. Selsk. Naturvidenske, Math. 
Am. 3: 71. 1827. Oldenlandia pentandra (Schu- 
mach. & Thonn.) DC., Prodr. 4: 427. 1830. H. 
halei Torr. & Gray, Fl. N. Amer. 2: 42. 1841. 
P. halei (Torr. & Gray) Gray, Syn. Fl. N. Amer. 
l.pt. 2: 28. 1884. 

Herbs with weakly erect or decumbent stems 10-30 
cm high, stems 0.8-1.8 mm thick, glabrous and slightly 
succulent; stipule sheath 0.5-1.5(-3) mm long, entire or 
with 1-3 short (0.3-2 mm) subulate lobes. Leaves with 
poorly defined petioles 0.5-10 mm long (in ours); leaf 
blades 1.5-4(-8) cm long, 0.4-2.5 cm broad, narrowly 
ovate-elliptic to elliptic (elliptic-lanceolate to linear-lan- 
ceolate), apex obtuse to acute, base obtuse to cuneate 
and decurrent on the petiole, drying membranaceous or 
thin-chartaceous (often translucent), greenish and gla- 
brous on both surfaces, 2 veins 3-4/side but difficult to 
see. Inflorescences 2-5 cm long, of (l-)3-5 irregularly 
arranged flowers in leaf axils or at apex (elongate race- 
mose inflorescences not seen in North American mate- 
rial), peduncles 0-8 mm long, pedicels 2-5 mm long. 
Flowers glabrous externally, hypanthium 2-3 mm long, 
calyx tube 0.3-1 mm long, calyx lobes 0.5-1 .5 mm long, 
acute; corolla white in ours, tube 1.5-4 mm long, 2-3 
mm diam., corolla lobes 5, 1-3 mm long, 0.7-2 mm 
broad, ovate-triangular. Fruits 2.5-4 mm long, 2-3 mm 
diam., obconic to oblong; seeds 0.3 mm long. 

Plants of moist depressions in open sunny sites, 
often along streams. Collected in central Nicara- 
gua as early as 1869 and recently found along Rio 
San Juan, and in Belize. Originally in tropical Af- 
rica, Texas to Florida, and Cuba. 

Pentodon pentandrus is recognized by its deli- 
cate habit with small thin ovate-elliptic leaves, 
small few-flowered and irregularly arranged inflo- 
rescences, small five-parted flowers, and small 
capsules with many seeds. The North American 
material (as represented by P. halei from the 
southern U.S.A. and illustrated in Correll & Cor- 
rell, 1982, fig. 615) differs greatly from typical Af- 
rican collections (Verdcourt, 1976, p. 264, fig. 38). 
Measurements in parenthesis in the description 
represent African material. Nevertheless, the spe- 
cies varies greatly in Africa, and the North Amer- 
ican material probably represents an early atypical 
introduction (Verdcourt, 1976, p. 263). Compare 
Sipanea biflora. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



217 



Pogonopus Klotzsch 

Trees or shrubs; stipules interpetiolar, free, small, del- 
toid to narrow, deciduous with the leaves. Leaves op- 
posite, petiolate; leaf blades usually thin, entire, pin- 
nately veined, domatia absent. Inflorescences terminal, 
paniculate and subcorymbose-cymose, pedunculate, 
bracteolate, 1 or 2 flowers of the inflorescence usually 
with a greatly enlarged sepal lobe forming a broad pet- 
iolate leaf-like and colorful "petal." Flowers bisexual, 
radially symmetrical (except for those with a single great- 
ly enlarged sepal lobe or where the tube is curved), hy- 
panthium turbinate, calyx tube short, calyx lobes 5, den- 
tate and deciduous (each inflorescence usually with 1 or 
2 flowers with a greatly expanded calyx lobe); corolla 
cylindrical, straight or curved, barbate in the throat, co- 
rolla lobes 5, short, valvate in bud; stamens 5, inserted 
near the base of the corolla tube, filaments slender and 
glabrous, anthers versatile dorsifixed above the middle, 
exserted; ovary 2-locular, ovules many on placentas lon- 
gitudinally adnate to the placenta, style slender with 2 
linear or oblong branches. Fruits capsular, subglobose 
to oblong-ovoid, 2-locular, areolate at apex, loculicidally 
bivalvate; seeds many, horizontal, crowded. 

A neotropical genus of three species; one species 
ranges from Mexico to South America and is grown 
as an ornamental. The greatly enlarged and bright- 
ly colored laminar sepal lobe of one or two flowers 
in each inflorescence is distinctive (but compare 
Mussaenda erythrophylld). 



Pogonopus speciosus (Jacq.) Schum. in Mart. Fl. 
Bras. 6, pt. 6: 265. 1889. Macrocnemum spe- 
ciousum Jacq., Hoit. Schoenbr. 1: 19,1.43. 1797. 
Macrocnemum exsertum Oersted, Vidensk. 
Meddel. Kjobenhavn 45. 1852. P. exsertusOer- 
sted, Amer. Centr. 17. 1863. Figure 16. 

Shrubs or small trees, 2-6(-l 2) m tall, leafy stems 1.5- 
5 mm thick, minutely (0.2-0.4 mm) appressed-pubes- 
cent and glabrescent in age, the older stems with con- 
spicuous rounded to linear whitish lenticels 0.5-1 (-2. 3) 
mm long; stipules 1-3.5 mm long, 3-4 mm broad and 
with a rounded or cuspidate tip to 1 m long. Leaves with 
petioles 2-15 mm long, 1-2 mm broad, appressed-pu- 
bescent, often with lateral wings continuous with the 
lamina margins; leaf blades 7-18(-25) cm long, 3- 
7.5(-9.5) cm broad, elliptic-obovate or elliptic-oblong 
(smaller leaves near the inflorescence usually ovate), apex 
acuminate with tip 5-15 mm long, base tapering grad- 
ually and cuneate-acute, decurrent on the petiole, drying 
thin-chartaceous, glabrous above, sparsely pubescent on 
the veins beneath with thin whitish hairs 0.2-0.4 mm 
long, 2 veins 4-10/side, the minor venation forming a 
small reticulum on both surfaces. Inflorescences termi- 
nal or axillary, 4-18 cm long, peduncles to 5 cm long, 
main axis usually with opposite branches, minutely ap- 
pressed-pubescent, flowers usually in compact distal cy- 
mose gruoups, bracteoles 2-4 mm long, lanceolate to 
linear, pedicels 0-3(-5) mm long. Flowers with hypan- 



thium 3 mm long, 2-3 mm diam., conical-turbinate, 
sparsely and minutely puberulent, drying dark, calyx lobes 
0.6-2 mm long, 0.2-0.3 mm broad, expanded calyx lobe 
with a petiole 8-20 mm long and lamina 2-4(-6) cm 
long and 1.5-3.5(-5) cm broad, broadly ovate or ovate- 
orbicular, bluntly obtuse to rounded, dark red to bright 
red or rose red, palmately veined; corolla tubular, rose 
red to dark red, tube 1 2-28 mm long, straight or curved, 
3-6 mm diam. distally, densely and minutely pubescent 
with ascending hairs, corolla lobes 3-5 mm long, 2-3 
mm broad at the base; stamens with filaments 20-23 
mm long, anthers ca. 2.5 mm long; style 30-35 mm long, 
stigmas 2-2.5 mm long. Fruits 6-9 mm long, 5-7 mm 
diam., oblong-urceolate, minutely (0. 1 mm) puberulent, 
with 2 longitudinal sulci, abruptly truncated at apex, the 
linear calyx lobes often persisting. 

Ornamental plants grown in both evergreen and 
deciduous environments in Central America, from 
near sea level to about 500 m elevation. Probably 
flowering principally in the rainy season: August- 
December. The species grows wild in southern 
Mexico, the Guanacaste lowlands, and central 
Panama and in Colombia and Venezuela. 

Pogonopus speciosus is recognized by the bright 
red or rose petaloid expansions of the calyx in a 
few flowers of each inflorescence. This species is 
not known from the wild in Nicaragua, El Sal- 
vador, or Honduras; it is possible that it was 
brought to Mexico as an ornamental in pre-Co- 
lumbian times. Compare Mussaenda erythrophyl- 
la, another ornamental with bright red petaloid 
sepal lobes. 



Posoqueria Aublet 

Trees, treelets, or shrubs, branches soon becoming 
terete, glabrous or pubescent, usually thick; stipules in- 
terpetiolar, free, large, triangular, usually early decidu- 
ous. Leaves opposite and decussate, petiolate; leaf blades 
usually large, entire, often coriaceous, usually with 
domatia. Inflorescences terminal, usually few-flowered 
and short (but the flowers very long), corymbose to cy- 
mose or umbellate, pedunculate, bracts absent or min- 
ute, flowers pedicellate. Flowers bisexual and mono- 
morphic, (4-)5(-6)-parted, radially symmetrical or 
bilaterally symmetrical because of the curved corolla tube, 
hypanthium little differentiated from the pedicel, calyx 
tube short, calyx lobes short-dentate, usually persisting, 
often auriculate at the base, with glands within; corolla 
long-salverform or long-tubular, white, corolla tube long 
and narrow, glabrous externally, glabrous or glandular- 
papillate within, corolla lobes imbricate or contorted in 
bud, rotate or reflexed; stamens unequal or subequal, 
short and inserted on the mouth of the tube, glabrous or 
pilose, anthers basifixed and sagitate, linear-oblong, ex- 
serted, (the pollen is said to be released explosively in 
some spp.); ovary 2-locular or incompletely 1-locular, 
ovules many in each locule and erect on stipitate bila- 
mellar placentas borne on the septum, style filiform, as 



218 



FIELDIANA: BOTANY 



long as the tube, stigmas short and bind. Fruits baccate 
or drying hard and indehiscent. globose to ovoid, 1 - or 
2-locular, often large; seeds large, hard, rounded and 
obtusely angled to flattened, imbedded in a gelatinous 
pulp. 

A Neotropical genus of 1 2- 1 6 species, with most 
of the species in South America; a single species 
ranges as far north as southern Mexico. The genus 
is distinctive because of the larger stiff leaves, large 
stipules, the long white flowers, and the globose 



indehiscent fruit with large seeds imbedded in a 
fleshy pulp. The corolla tubes are usually more 
than 10 cm long and less than 1 cm diam. The 
flowers are strongly fragrant at night when they 
open but yellowish and odorless by morning; these 
are probably pollinated by long-tongued moths. In 
some species the stamens are held asymmetrically 
and under tension over the mouth of the corolla 
tube until they are disturbed, when they snap for- 
ward. Compare Borojoa and Tocoyena. 



Key to the Species of Posoqueria 

la. Corolla tubes (20-)30-38 cm long; leaf blades (10-) 18-46 cm long leaf blades usually puberulent 
with very thin hairs beneath (but difficult to see, except along the larger veins), usually slightly rough 
and minor venation obscure beneath (uncommon) P. grandiflora 

Ib. Corolla tubes 8-20 cm long; leaf blades 10-20(-24) cm long, essentially glabrous beneath .... 2a 

2a. Leaf blades drying dull above with the tertiary veins not easily seen, the minor venation not raised; 

fruit dull yellow to orange, ellipsoid to ovoid, with pericarp leathery and 410 mm thick 

P. coriacea 

2b. Leaf blades lustrous above with tertiary veins easily seen and sometimes slightly elevated; fruit 
orange and globose, with pericarp hard and brittle, 1-3 mm thick P. latifolia 



Posoqueria coriacea M. Martens & Galeotti, Bull. 
Acad. Brux. 11: 240. 1844. 

Small trees or shrubs, 3-8(-20) m tall, branchlets 2- 
7 mm thick, glabrous; stipules 8-12(-28) mm long, ovate, 
ovate-oblong to ovate-lanceolate or suborbicular, acute, 
glabrous. Leaves with petioles 6-22 mm long, glabrous; 
leaf blades (6-)10-22(-27) cm long, (3-)5-12(-18) cm 
broad, narrowly elliptic to ovate or narrowly obovate, 
apex obtuse to abruptly acute, base cuneate to rounded 
and subtruncate, drying coriaceous, 2 veins 4-8/side. 
Inflorescences 2-4 cm long and 2-3 cm broad (not in- 
cluding the corolla lengths), cymose-corymbose, with 7- 
25 flowers, peduncles 1-2 cm long, glabrous or minutely 
puberulent, pedicels 5-10(-23) mm long, glabrate. Flow- 
ers with hypanthium + calyx 6-8 mm long, subulate 
glands present within the calyx, calyx lobes 5, unequal, 
0.5-1.2 mm long, rounded; corolla greenish white, gla- 
brous externally, long-tubular, tube 9-20 cm long, 1.5- 
3 mm diam. in the lower half, pilose within the throat 
and at the base of the lobes, corolla lobes 5, 15-25 mm 
long, unequal, rounded distally; anthers 7-10 mm long. 
Fruits 7-10 cm diam., ellipsoid to ovoid, brown, peri- 
carp leathery, 4-10 mm thick; seeds 10-12 mm diam., 
angular, aril white. 

Plants of lowland rain forest formations, from 
near sea level to 1 100 m elevation (to 1500 m in 
South America). Flowering January-March; fruit- 
ing in March-April, June-August, and October- 
November. The species ranges from Mexico to 
Brazil. 

Posoqueria coriacea is characterized by the long- 



tubular flowers, dull leaf surfaces, and fruit with 
thick leathery rind. The 2 veins are often some- 
what impressed in thick mature leaves when dried. 
The larger leaf dimensions noted above in paren- 
theses are from Steyermark's Flora of Venezuela 
treatment ( 1 974) and may not occur in Costa Rica. 
The breeding system of this species was studied 
by Bawa and Beach (1983). 



Posoqueria grandiflora Standl., J. Wash. Acad. Sci. 
18: 166. 1928. P. maxima Standl., Publ. Field 
Columb. Mus., Bot. Ser. 7: 57. 1930. Figure 15. 

Small trees, treelets, or shrubs, 2-7(-17) m tall, trunk 
to 18 cm dbh, leafy stems 2.5-12 mm thick, glabrous or 
minutely puberulent with thin erect hairs 0.1-0.2 mm 
long; stipules 8-18 mm long, 4-1 1 mm broad, triangular, 
glabrescent, deciduous. Leaves well spaced along the stem, 
petioles 7-20 mm long, 2-4 mm thick, glabrous or mi- 
nutely puberulent; leaf blades ( 1 3-)l 8-34(-46) cm long, 
6-1 7(-23) cm broad, elliptic to elliptic-obovate or ellip- 
tic-oblong, apex acuminate to short-acuminate or blunt- 
ly rounded, base tapering gradually to slightly rounded 
and obtuse or acute, drying subcoriaceous to coriaceous, 
pale grayish green beneath, glabrous above, with thin 
erect hairs 0.1-0.5 mm beneath or the hairs difficult to 
see and apparently glabrous, slightly rough to the touch 
beneath, 2 veins 4-8/side and prominent on both sur- 
faces, minor venation obscure on both surfaces. Inflo- 
rescences only 3-6 cm long (not including the corolla 
tubes), corymbose with ca. 10-20 flowers, peduncles 1- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



219 



2 cm long and 3-8 mm thick, pedicels 2-12 mm long, 
ca. 2 mm thick, glabrous or minutely puberulent. Flow- 
ers glabrous, hypanthium 3-6 mm long, little differen- 
tiated from the pedicel, calyx tube 3-6 mm long, 3-6 
mm diam., calyx lobes 5, 0.5-1.5 mm long, broadly 
rounded distally; corolla long-tubular with rotate lobes, 
white, tube ( 1 2.5-) 1 5-36 cm long 2.5-4 mm diam., lobes 
5, 20-38 mm long, 7-1 3 mm broad, oblong-obovate and 
rounded; stamens with unequal filaments to 1 5 mm long, 
0.5-1 mm thick, anthers 9-12 mm long, to 2 mm broad, 
linear-lanceolate. Fruits 7-12 cm long, 4-7 cm diam., 
ovoid to ellipsoid, with a slightly roughened bark-like 
(minutely lenticellate) brownish surface, pericarp 5-12 
mm thick; seeds 6-10 mm long, orange to white. 

Trees and treelets of evergreen lowland Carib- 
bean rain forest formations, from 40 to 300 m 
elevation. Flowering in February, April, July, and 
September-October; probably fruiting throughout 
the year. The species ranges from northeastern 
Costa Rica to Colombia. 

Posoqueria grandiflora is recognized by its very 
long flowers on small terminal inflorescences, the 
relatively large corolla lobes, and the large leaves 
with obscure minor venation and pubescence (when 
present), which makes the lower leaf surfaces 
slightly rough to the touch. This species appears 
to be less common than its congeners in Costa 
Rica. The species has been called "wild coffee" in 
the Caribbean lowlands. An unusual collection 
(Wilbur 207 '11 DUKE, F) from southern Costa Rica 
has large leaves with conspicuous long hairs, and 
flowers with very long corolla tubes, but is prob- 
ably no more than an extreme form of this species. 
The Colombian material ascribed to P. maxima 
appears to be conspecific; Steyermark erred in 
making it a subspecies of P. coriacea Mart. & Gal. 



Posoqueria latifolia (Rudge) Roem. & Schult., Syst. 
Veg. 5: 227. 1819. Solena latifolia Rudge, PI. 
Guian. 1: 26, t. 40. 1806. Stannia panamensis 
Walp. & Duchass., Linnaea 23: 755. 1850. P. 
panamensis (Walp. & Duchass.) Walp., Ann. 
Bot. Syst. 2: 797. 1852. Figure 15. 

Small trees to 9(-20) m tall, trunks to 25(-40) cm dbh, 
wood hard and reddish, leafy stems 2-6 mm thick, gla- 
brous; stipules 7-1 8 mm long, 3-8 mm broad at the base, 
triangular to ovate-oblong, sometimes slightly (1 mm) 
united above the petioles, apex obtuse to acuminate, stiff, 
glabrous. Leaves distant along the stem, petioles (4-) 7- 
20 mm long, 1 . 5-4 mm thick, glabrous; leaf blades (7-) 1 0- 
20(-24) cm long, (3-)4-10(-14) cm broad, elliptic-ob- 
long to elliptic-ovate or ovate, apex obtuse or short- 
acuminate, base abruptly narrowed and obtuse or round- 
ed, drying stiffly chartaceous to subcoriaceous, glabrous 



above and below, 2 veins 5-7/side, the minor venation 
visible on both surfaces. Inflorescences 2-5 cm long (not 
measuring the corolla tubes), corymbose with 7-1 8 flow- 
ers, peduncles 1-2 cm long, 2-3 mm thick, bracteoles 
ca. 0.5 mm long, pedicels 3-9 mm long, glabrous. Flow- 
ers glabrous externally, sweet aromatic, hypanthium 3- 
6 mm long, calyx lobes poorly developed, 0-0.5 mm 
long; corolla long-tubular with usually reflexed lobes, 
white, tube 8-14(-16) cm long, 2-3.3 mm diam., lobes 
5 (4), 12-20(-26) mm long, 4.5-5.5 mm broad, narrowly 
oblong, rounded; stamens 5 (4), filaments to 6 mm long, 
ca. 0.4 mm thick, anthers 6-7.5 mm long, ca. 1 mm 
broad, linear-oblong, with basal lobes ca. 1 mm long; 
style ca. 9 cm long. Fruits 4-6 cm diam., globose to 
ovoid, yellow or orange at maturity, the pericarp only 
1-3 mm thick, surface becoming wrinkled; seeds 6-12 
mm long, often triangular, translucent in life, arils white 
to yellow-orange, in a fleshy sweet pulp. 

Trees and treelets found in rain forests, partly 
deciduous forests, and moist sites in deciduous 
forests, from 2 to 700(-1200) m elevation. Prob- 
ably flowering and fruiting throughout the year 
(but mostly flowering in March-October and fruit- 
ing in October-April). This common species oc- 
curs in all the lowland evergreen areas of Costa 
Rica. The species ranges from southern Mexico to 
the Amazon basin of Brazil and Bolivia. 

Posoqueria latifolia is recognized by the larger 
stiff glabrous leaves, the small terminal inflores- 
cences with very long tubular flowers, and the glo- 
bose fruit. The barely visible minor (3 and 4) 
venation appears to be a consistent way of differ- 
entiating the leaves of P. latifolia from those of P. 
coriacea. Croat (1978, p. 814) remarks that the 
anthers are united at anthesis along one side of the 
tube and burst apart when contacted. He also notes 
that the flowers open late in the day and do not 
persist on the following day. Common names re- 
corded in Costa Rica are boca de vieja, carica,fruta 
de mono, guayaba de mono, guayaba mica, man- 
zana de mico, picarito, and querica. The names 
jicarillo and querica are used in southeastern Nic- 
aragua. The fruit's pulp is edible and sweet. 



Psychotria Linnaeus 
Nomen conservandum 

REFERENCES C. Hamilton, A revision of Me- 
soamerican Psychotria subgenus Psychotria (Ru- 
biaceae), part I: Introduction and species 1-16. 
Ann. Missouri Bot. Gard. 76: 67-1 1 1 . Part II: Spe- 
cies 17-47. Ann. Missouri Bot. Gard. 76: 386- 
429. Part III: Species 48-6 1 and appendices. Ann. 
Missouri Bot. Gard. 76: 886-916. 1989. A. Mo- 



220 



FIELDIANA: BOTANY 



lina, Revision de las Especies de Cephaelis en 
Mexico, Centre America y las Antilllas. Ceiba 4: 
1-38. 1953. (Other references are listed under spe- 
cific species below, under the family description, 
and at the end of the text.) 



Shrubs or small treelets, less often medium-sized trees 
or herbaceous subshrubs (rarely lianas), terrestrial or rarely 
epiphytic, stems often slightly thickened at the nodes, 
glabrous or puberulent; stipules usually united and in- 
terpetiolar with 1 or 2(-5) apices or lobes on each side, 
sometimes also united around the stem above the pet- 
ioles (intrapetiolar) and forming a short tube or sheath, 
rarely separate to the base and appearing free, deciduous 
or persisting, colleters often present on the stems at the 
adaxial base of the stipules (and drying reddish in sub- 
genus Psychotria). Leaves opposite and decussate (rarely 
3 or 4/node), petiolate or rarely subsessile, usually acu- 
minate at the apex; leaf blades entire and pinnately veined, 
raphides (cystoliths) present and obscure or conspicuous, 
some species with domatia in the vein axils beneath 
(some African species with bacterial nodules in the leaves). 
Inflorescences mostly terminal and occasionally becom- 
ing pseudoaxillary by further growth of an axillary branch, 
less often consistently axillary, usually solitary at a node, 
usually pedunculate and paniculate with opposite 
branching, bracts large to small or rarely undeveloped, 
bracts forming an involucre in some species formerly 
placed in Cephaelis, flowers often borne in distal brac- 
teolate cymes, sessile or pedicellate. Flowers bisexual (in 
Central America), radially symmetrical, usually small, 
often distylous with long-styled (pin) and short-styled 
(thrum) forms within the same species, calyx tube usually 
short and cupulate, with 4 or 5(-6) short calyx lobes or 
without lobes and entire; corolla tubular to funnelform 
or campanulate, white to pink or yellowish, corolla tube 
usually short or narrow, often with tufted hairs at the 
throat within, glabrous or puberulent externally, corolla 
lobes 4-5(-6), always valvate in bud, often thickened at 
the tips; stamens 4-5(-6), borne from the middle or up- 
per part of the tube, filaments slender, anthers narrow, 
included or exserted, disc ring-shaped and encircling the 
base of the style; ovary usually 2-locular (rarely with 4- 
6 locules), 1 erect ovule borne from the base of each 
locule or from the base of the thick septum, style long 
or short, with 2 (rarely 4-6) linear stigmas. Fruits fleshy 
drupes, red, yellow, blue, purple, black, or white when 
ripe, often with spongy arenchymatous tissue, usually 
with 2 (4-6) hard pyrenes, pyrenes plano-convex with a 
flattened interior (adaxial) face and a rounded exterior 
(abaxial) surface, often with longitudinal ridges on the 
exterior surface (rarely with transverse ribs or projec- 
tions), usually with a median longitudinal sulcus on the 
inner face. 



Psychotria is the largest genus of Rubiaceae and 
one of the largest genera of Angiosperms, with an 
estimated 1,500-1,600 species in the tropics and 
subtropics of both hemispheres. Together with 
Piper (Piperaceae) and Miconia (Melastomaceae), 



Psychotria is one of Costa Rica's three most spe- 
ciose woody genera, especially common in the un- 
derstory of evergreen forests and forest edges. Some 
species are difficult to separate from similar spe- 
cies in Coussarea, Faramea. and Palicourea (q.v.). 
While Cephaelis appeared to be a very distinctive 
genus in Central America, a great number of in- 
termediate species have necessitated the inclusion 
of its species in Psychotria. Our treatment has ben- 
efited greatly from Hammers work on Psychotria 
at La Selva (in Taylor, 1991). Likewise, Hamil- 
ton's recent revision (see references above) of the 
species in subgenus Psychotria has been especially 
helpful, as these species are often difficult to dif- 
ferentiate; we have not deviated from his species 
concepts in this treatment. 

Psychotria is generally characterized by the sol- 
itary terminal inflorescences with opposite 
branching (in most species), the 1 - or 2-lobed in- 
terpetiolar stipules, entire and pinnately veined 
opposite leaves, smaller flowers with minute calyx 
lobes, short corolla tubes, valvate corolla lobes, 
basal solitary ovules, and fleshy fruit usually with 
2 hard seed-like pyrenes. The short shrubby habit, 
preferences for areas of high rainfall, fruit often 
with longitudinal ridges, and white or yellowish 
flowers are additional characteristics. Those spe- 
cies with axillary inflorescences tend to have un- 
branched succulent stems or few-branched woody 
stems (see below). All the fruits appear to be bird- 
dispersed; they are fleshy and bright red, blue to 
purplish and black, or arenchymatous and whitish. 
Two species of Psychotria found in Costa Rica, P. 
emetica and P. ipecacuanha, are used medicinally. 

Most species of Psychotria in Costa Rica are 
readily recognizable but the differences between 
closely related species can be subtle. There appear 
to be real problems of intcrgradation in the epi- 
phytic species, and the succulent-stemmed her- 
baceous species with axillary inflorescences pre- 
sent problems of species delimitation. Except for 
the groups just mentioned, the majority of species 
stand well apart and give little evidence of hy- 
bridization or intergradation. The difficulty in 
identifying a specimen to species often is due to 
the large number of species in the genus, and the 
wide range of variation found within some species. 
Below we provide keys to four groups within the 
genus; groups 2 and 3 appear to be largely mono- 
phyietic. The illustrations are an additional ave- 
nue for identification and are grouped by inflo- 
rescence position, subgeneric placement, and 
general similarity. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



221 



Key to Four Species Groups of Psychotria in Costa Rica 

1 a. Plants nearly always epiphytic and with subcoriaceous leaves drying grayish; ovary usually with 4 
locules; fruits red and often with 4 seeds; or plants of Cocos Island Group 1 

Ib. Plants not epiphytic, leaves various; ovary with 2 (rarely 5) locules; fruit usually with 2 (rarely 5) 
seeds; and plants not known from Cocos Island 3 

2a. Inflorescences regularly axillary; main stems usually short (ca. 1 m) and succulent in a majority of 
species, both succulent and woody stems often with hollow sections within when dried . . . Group 2 

2b. Inflorescences terminal, only occasionally axillary where lateral shoots have continued to develop; 
main stems usually woody and rarely with hollow sections when dried 3 

3a. Leaves drying greenish, gray-green, yellow-green, yellowish brown, or brown (note that leaves treated 
with isopropyl alcohol may turn reddish brown and that some species have pinkish venation in 
life), domatia rarely present; stipules often persisting, the stipules not subtending and enclosing a 
ring of reddish hairs (colleters) in early stages; fruit usually becoming blue, purple, or black (red 
only in P. haematocarpa with very small inflorescences, or orange in later stages in P. racemosa 
with 5 locules and 5 -seeded fruit); subgenus Heteropsychotria Group 3 

3b. Leaves drying grayish to black, pinkish brown or dark reddish brown, domatia present in a few 
species; stipules usually falling off as the leaves expand (caducous), and usually enclosing a short 
ring of reddish hairs (these often persisting just above the stipule scar); fruit always red at maturity 
(never with more than 2 seeds); subgenus Psychotria Group 4 



Key to Group 1: Epiphytic Species of Psychotria in Costa Rica and Three Species of Cocos Island 

The epiphytic species appear to represent a very natural (probably monophyletic) group. However, 
there is the strong likelihood that a large number of our collections belong to a single polymorphic entity, 
for which P. guadalupensis is the earliest name. Psychotria maxonii appears to be quite distinct, but 
there are a significant percentage of collections that appear to be intermediate between P. guadalupensis 
and P. pithecobia. See the discussion under P. guadalupensis. 

la. Epiphytic plants of continental Costa Rica 2 

1 b. Trees and shrubs of Cocos Island 4 

2a. Leaves to 5(-7) mm wide, 10-28 mm long, narrowly lanceolate to narrowly oblong; inflores- 
cences to 3 cm long P. maxonii 

2b. Leaves 8-40 mm wide, 20-120 mm long, more than 40 mm long if lanceolate or narrowly 

oblong; inflorescences 3-14 cm long 3 

3a. Leaves with 6 or more pairs of secondary veins, the veins clearly elevated and visible on the 
dried leaf surfaces, leaves more than 4 cm long and 2 cm broad; inflorescences with peduncles 

2-7 cm long P. pithecobia 

3b. Leaves with fewer than 6 pairs of major secondary veins, the veins often obscure on the surfaces 

of the dried leaves, leaves 3-10 cm long and 1-5 cm broad, very variable (on different plants) 

as regards size, shape, and texture; peduncles 0.5-3 (rarely to 5) cm long . . .P. guadalupensis 

4a. Leaves to 25 cm long, drying subcoriaceous and dark reddish brown; stipules forming an acute 

F/CMs-like cap over the shoot apex and early deciduous P. cocosensis 

4b. Leaves to 1 5 cm long, drying thin-chartaceous and greenish or grayish; stipules with 2 acute lobes 

on each side and persisting 5a 

5a. Inflorescences less than 3 cm long, with thick (1 mm) lateral branches less than 5 mm long; stipule 

lobes 24 mm long P. brachybotrya 

5b. Inflorescences usually becoming more than 3 cm long, with slender (0.3 mm) lateral branches 5- 
10 mm long; stipule lobes ca. 1 mm long Psychotria sp. A 

Key to Group 2: Species of Psychotria with Axillary Inflorescences 

la. Inflorescences axillary to both leaves of the node, usually with 2 or more inflorescences at distal 
nodes, inflorescences usually less than 5 cm long; fruits becoming blue, black, or purple 2 

222 FIELDIANA: BOTANY 



Ib. Inflorescences axillary to only 1 leaf at each node, usually with 1 inflorescence at distal nodes, 

inflorescences often more than 5 cm long; fruits white to yellow or red .8 

2a. Inflorescences subsessile and forming a dense verticillate cluster at the node 3 

2b. Inflorescences short-pedunculate, not forming dense verticillate clusters around the node 

5 

3a. Inflorescences lacking an involucre of bracts, calyx lobes to 3 mm long (not Psychoiria 

spp.) Hoffmannia spp. 

3b. Inflorescences subtended by an involucre of broad bracts, calyx lobes to 2 mm long 

4 

4a. Bracts subtending the inflorescences pale green; stipules to 10 mm long and 5 mm 

broad; widely distributed at 1 200-2800 m elevation P. aubletiana 

4b. Bracts reddish; stipules 12-22 mm long and almost as broad; 700-1500 m near Orosi 

and Muneco P. cartagoensis 

5a. Leaves drying greenish to greenish brown, shrubs or small trees with many branches, young 
stems glabrous; stipules forming a short sheath; inflorescences with many branches, often 

with more than 1 fruits P. cooperi 

5b. Leaves drying grayish or dark brown, subshrubs or few-branched shrubs, young stems mi- 
nutely puberulent; stipules not forming a sheath; inflorescences few-branched, rarely with 

more than 5 fruits 6 

6a. Inflorescences usually 4/node; plants growing to 2(-5) m tall, with lateral branches; leaf blades 

drying stiffly chartaceous [often obtuse at the base; peduncles to 1 5 mm long] . . . P. erecta 

6b. Inflorescences 2/node; plants growing to 1 m tall, unbranched; leaf blades drying thin-char- 

taceous 7 

7a. Leaf blades drying brownish, often cuneate basally; peduncle to 5 mm long, inflorescence 

less than 2.5 cm long P. emetica 

7b. Leaf blades drying greenish, often decurrent basally; peduncle to 70 mm long, inflorescences 

more than 3 cm long P. aggregata 

8a. (from Ib) Stipules lobed or fimbriate distally, to 3 cm long, translucent to opaque 9 

8b. Stipules entire distally, bluntly obtuse to rounded, to 1 cm long, thick and opaque 11 

9a. Stipules to 20 mm broad, drying yellowish and translucent, fimbriate to bluntly lobed [in- 
florescences neither capitate nor hirsute; fruit becoming white] P. cartagoensis 

9b. Stipules to 7 mm broad, drying dark and opaque, with stiff narrow or filiform teeth; fruit 

becoming red, purple, or blue 10 

lOa. Leaves pilose with hairs to 2 mm long; inflorescences to 15 cm long, much-branched and 

lacking an involucre; fruit becoming purple or blue P. pilosa 

lOb. Leaves glabrous or with short (0.5 mm) hairs; inflorescences to 5 cm long, capitate and 

involucrate; fruit red P- ipecacuanha 

1 la. Leaves with more than 18 pairs of closely parallel secondary veins, veins becoming prominently 
raised and the leaf corrugated in age, bluntly obtuse to rounded distally; plants rarely exceeding 

40 cm in height [fruits red] - P- polyphlebia 

1 Ib. Leaves with fewer than 18 pairs of secondary veins, not becoming corrugated, usually acuminate 

at apex; plants usually more than 50 cm tall 

1 2a. Fruits becoming orange or red 

12b. Fruits becoming white or yellowish green 

13a. Leaf blades drying dark above and much paler beneath, young leaves glabrous; pyrenes with 

a longitudinal costa on the back; commonly collected . P. uliginosa 

13b. Leaf blades drying greenish to dark brown above and only slightly paler beneath; young 
leaves densely hirsute beneath with hairs to 1 mm long; pyrenes lacking dorsal costae; rare 

P. siggersiana 

14a. Young leaves villose beneath with hairs to 1 mm long, a definite arcuate submarginal vein present 
2-3 mm from the leaf edge, with 16-22 pairs of major secondary veins; fruit becoming greenish 

yellow P capacifolia 

14b. Young leaves minutely puberulent or glabrous, a definite submarginal vein absent, with 10-16 
pairs of secondary veins (and often loop-connected in the distal part of the leaf); fruit becoming 
white and spongy. (Note: This is a variable group of plants that may intergrade.) 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 223 



1 5a. Inflorescences with thin widely spreading distal branches, bracts subtending the major branches 
less than 4 mm long, flowers borne separately along the distal branches P. macrophylla 

15b. Inflorescences with thick (1 mm) distal branches or the inflorescences compact and short (5 cm), 
bracts subtending the basal branches or basal flowers more than 4 mm long 16 

1 6a. Peduncles 0-3 cm long, flowers loosely clustered to capitate; leaves usually elliptic (rarely consis- 
tently narrow); widespread in Costa Rica P. aggregata 

1 6b. Peduncles to 1 5 cm long, flowers usually capitate on the peduncle or on 3 short branches on the 

peduncle; leaves usually elliptic-lanceolate; eastern part of the Cordillera de Talamanca 

P. aggregata (sensu stricto) 



Key to Group 3: Species of Subgenus Heteropsychotria and Similar Species in Costa Rica 
(Including Species Formerly in Cephaelis) 

la. Flowers borne in dense capitate or subcapitate inflorescences with closely clustered flowers, bracts 
usually conspicuous and enclosing the flowers; inflorescences terminal and solitary or 3 closely 

grouped together (species formerly placed in Cephaelis and others) 2 

Ib. Flowers in open or congested inflorescences but not capitate or subcapitate (flowers sometimes in 
small distal capitula or glomerules on secondary branches within branched inflorescences); bracts 

small to conspicuous or absent, bracts rarely enclosing and obscuring the flowers 16 

2a. Inflorescences subtended by 2(-3) large bracts forming a basal cupulate involucre, capitulum 

solitary, 3-10 cm broad 3 

2b. Inflorescences usually with more than 4 basal bracts, lacking a single cupulate involucre at 

the apex of the peduncle; inflorescence of 1 or 3 capitula, 1-5 cm broad 6 

3a. Bracts reddish (rarely yellow), inflorescences erect; very common shrubs in lowland 

evergreen formations 4 

3b. Bracts purple to lilac, inflorescences erect or pendant; rarely collected species 5 

4a. Stems and leaves glabrous; stipules with 2 rounded lobes on each side . . P. elata 
4b. Stems and leaves densely tomentulose; stipules with 2 sharp teeth on each side 

P. poeppegiana 

5a. Montane (800-1200 m) plants; bracts purple, inflorescences pendant; leaf blades with 

ca. 1 4 pairs of 2 veins P. correae 

5b. Lowland (0-500 m) plants; bracts lilac, inflorescences erect; leaf blades with ca. 7 pairs 

of 2 veins P. borucana 

6a. Stipules with more than 2 slender teeth on each side (more than 6 teeth per node); unbranched 
subshrubs less than 1 m tall; fruit red [medicinal plants rarely collected in Costa Rica] . . . 

P. ipecacuanha 

6b. Stipules with 2 slender teeth on each side, or with 2 rounded lobes or unlobed (with 4 or 
fewer stipule lobes per node); plants mostly shrubby and more than 1 m tall (except P. 

guapilensis); fruit blue or purple 7 

7a. Corollas more than 4 cm long; fruit becoming more than 1 2 mm long; bracts usually bluntly 

obtuse and green P. chiapensis 

7b. Corollas less than 3 cm long; fruit less than 1 2 mm long; bracts acute to rounded and green 

to purple 8 

8a. Basal bracts of the inflorescences rounded to bluntly obtuse at the apex 9 

8b. Basal bracts acute to acuminate 14 

9a. Bracts white to pale green or bluish, glabrous, broadly rounded distally forming a tight 

cupulate involucre beneath the congested capitulum [0-600 m elevation] 

P. glomerulata 

9b. Bracts green to deep purple, glabrous to puberulent, bluntly obtuse distally (rounded 

in P. platypoda), not forming a definite cup at the base of the inflorescence 10 

1 Oa. Bracts deep purple or reddish purple, inflorescences often densely compacted and spher- 
ical; plants usually less than 1 m tall P. guapilensis 

1 Ob. Bracts greenish to purple, inflorescences never spherical, often loosely compacted; plants 
usually more than 1 m tall 11 



224 FIELDIANA: BOTANY 



1 la. Inflorescences usually of 3 pedunculate capitula; bracts white to pink (purple in fruit); 
stipular tube 2-5 mm long; 2 veins 8-12/side [1000-2500 m elevation] ...P. dichroa 

lib. Inflorescences usually of a single pedunculate capitulum; bracts whitish to green or 
purple; stipular tube ca. 1 mm long; 2 veins 1 1-22/side .12 

12a. Bracts broadly ovate and rounded at the base, purple; stipule lobes short and bluntly 
rounded; 1200-2300 m elevation p, molinianum 

1 2b. Bracts usually narrowed at the base, whitish to green or purple; stipules lobes short to 
long, acute; 0-800 m elevation 13 

13a. Bracts elliptic-obovate to oblanceolate; flowers usually puberulent, corolla tube 9-13 
mm long; commonly collected p. suerrensis 

1 3b. Bracts broadly rounded; flowers glabrous, corolla tube 3-6 mm long; rare in Costa Rica 

P. platypodd 

14a. (from 8b) Bracts orange or reddish orange, ovate and broadly overlapping; plants confined 

to the southern Pacific slope, 600-1300 m elevation P. aura nti bract ea 

1 4b. Bracts green or greenish with white, blue, or purple, linear lanceolate to ovate but not broadly 

overlapping; plants wide ranging 15 

1 5a. Bracts narrowly ovate, often marked with blue; inflorescences ca. 3 cm long and 2 cm broad 

with erect bracts; stipule lobes thin and translucent, to 6 mm long; Caribbean slope, 900- 

1 600 m elevation P. hazenii 

15b. Bracts often linear-lanceolate, usually marked with white or purple; inflorescences short (1 

cm) with broadly spreading bracts; stipule lobes stiff and opaque, to 4 mm long; widely 

ranging in moist evergreen formations, 0-800 m elevation P. hoffmannseggiana 

1 6a. (from 1 b) Young stems densely and conspicuously puberulent with hairs 0.3-2 mm long; peduncles 

and branches of the inflorescence usually densely puberulent 17 

16b. Young stems glabrous or sparsely puberulent with inconspicuous hairs less than 0.3 mm long; 

peduncles and branches of the inflorescences glabrous or puberulent 25 

17a. Hairs usually becoming more than 0.8 mm long, leaf blades drying dark above and 12-30 

cm long, often with more than 14 pairs of major 2 veins; flowers in dense distal clusters, 

subtended by conspicuous bracts 18 

17b. Hairs rarely exceeding 0.8 mm in length; leaf blades usually drying greenish (rarely dark 

brownish above), usually less than 20 cm long, with less than 14 pairs of major 2 veins; 

flowers in open or small distal clusters, bracts various 19 

1 8a. Leaf blades with 9-18 pairs of major 2 veins; flowers subtended by densely puberulent 
lanceolate bracts ca. 2 mm broad; wide-ranging P. pilosa 

18b. Leaf blades with (14-) 17-23 pairs of major 2 veins; flowers subtended by sparsely 

puberulent ovate bracts 2-3 mm broad; southwestern Costa Rica .... P. mortoniana 

19a. Ovary with 5 locules, fruits often with 5 seeds; stipules with 2 long (6-14 mm) stiff persisting 

awns on each side; leaf blades with conspicuous subparallel 3 veins [9-21 (-26) cm long] 

P. racemosa 

19b. Ovary with 2 locules, fruits never with more than 2 seeds; stipules rarely with 2 long stiff 

awns (P. umbelliformis); leaf blades rarely with conspicuous 3 veins 20 

20a. Inflorescence umbelliform, flowers in 3 small glomerules on equal primary branches at the 

apex of a long peduncle; leaves glabrous above [rare] P. umbelliformis 

20b. Inflorescences not as above; leaves puberulent or glabrescent above .21 

2 la. Leaf blades cuneate at the base and long-decurrent on the petiole, thin in texture; flowering 

portion of the inflorescences often broader than long; deciduous and evergreen formations 

of the Pacific slope P pubescens 

21b. Leaf blades not cuneate and long-decurrent at the base (sometimes decurrent in P. steyer- 

markii), mostly stiff-chartaceous when dried; flowering portion of the inflorescence rarely 

broader than long; plants of evergreen formations 

22a. Pubescence of young stems usually in narrow longitudinal lines; leaf blades 3-12 cm long 

and 1-3 cm broad; stipules with narrow awns 2-5 mm long [plants of lower montane (800- 

1 800 m) cloud forests; inflorescences few-branched and racemose] P. steyermarkii 

22b. Pubescence not in narrow longitudinal lines; leaf blades 6-20 cm long, 2-5 cm broad; lines; 

plants rarely collected above 800 m elevation 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 225 



23a. Plants of the Osa Peninsula; inflorescences with hairs to 1.5 mm long, branches and flower 
clusters distant and subtended by linear bracts to 8 mm long [leaf blades 5-12 cm long and 
with 10-12 pairs of major 2 veins] P. acicularis 

23b. Plants not known from the Osa Peninsula or from the Pacific slope below 400 m elevation; 
inflorescences with hairs usually less than 1 mm long, flower clusters congested or distant, 
subtended by linear bracts usually less than 4 mm long 24 

24a. Inflorescences with crowded flowers and branches, usually less than 4 cm broad, narrowly 
pyramidal, erect, calyx lobes 0.3-1.5 mm long; leaf blades 7-20 cm long, with 8-14 pairs of 
major 2 veins; awns of the stipules to 3 mm long P. hebeclada 

24b. Inflorescences usually with open branching and separate flower clusters, usually more than 
3 cm broad, broadly pyramidal, often pendant; calyx lobes 0.2-0.5 mm long; leaf blades 6- 
1 5 cm long and with 8-1 1 pairs of major secondary veins; awns of the stipules to 7 mm long 

P. pittieri 

25a. (from 16b) Inflorescences large and many-branched, usually becoming more than 12 cm long and 

1 cm broad, broadly paniculate; leaf blades usually large (often to more than 1 8 cm long); fruit 

rarely more than 4 mm diam. when dried (to 5 mm in P. solitudinum) 26 

25b. Inflorescences smaller, rarely more than 12 cm long (and if so narrowly racemiform) and usually 

less than 10 cm broad, paniculate to subcapitate or racemiform; leaf blades often less than 15 cm 

long; fruit 3-12 mm diam. when dried 31 

26a. Stipule lobes to 15 mm long and 5 mm broad at the base, conspicuous; flowers distant in 
small (1-2) alternate sessile groups along the slender distal branches of the inflorescence; 
fruits grayish blue to white P. microbotrys 

26b. Stipule lobes rarely more than 5 mm long, usually less than 2 mm broad at the base; flowers 
distant in small sessile groups only in P. solitudinum; fruit blue to black 27 

27a. Stipules at first acute distally but splitting and developing 2 acute teeth or awns separated 
by a broad U-shaped sinus; leaf blades often drying dark brown above; rarely collected below 
400 m elevation P. berteriana 

27b. Stipules at first rounded or obtuse at the apex, splitting into 2 broad lobes separated by a 
narrow V-shaped sinus; leaf blades drying dark greenish brown to yellowish green or grayish 
green; plants growing from to 1 700 m elevation 28 

2 8a. Flowers in small (1-2) separate groups along the (usually dichotomous) slender distal branches 
of the inflorescence [corolla tube 4-5 mm long, narrowed at the base and urceolate distally]; 
southwest Pacific slope of Costa Rica P. solitudinum 

28b. Flowers in distal small cymes on multiple-branched inflorescences with opposite and cymose 

branching; corolla various; plants not collected from the southwest Pacific area of Costa Rica 

29 

29a. Plants only known from the upper Rio Grande de Orosi and Tapanti above 1 300 m elevation; 
floral bracts ca. 3 mm long, often persisting with the flowers and obtuse distally; corolla tube 
ca. 6 mm long and tubular P. tapantiensis 

29b. Plants not collected above 1200 m elevation; floral bracts to 3 mm long, early deciduous; 
corolla tube 2-7 mm long 30 

30a. Corolla tube 2-3 mm long, funnelform or tubular; floral bracts usually rounded at the apex; 
leaf blades with 8-14 pairs of major 2 veins; fruit blue P. luxurians 

30b. Corolla tube 6-7 mm long, narrowed at the base and expanded distally; floral bracts usually 
acute at the apex; leaf blades with ca. 8-10 pairs of major 2 veins; fruit becoming blue or 

black P. angustiflora 

3 la. (from 25b) Inflorescences 12-20 cm long and only 2-5 cm broad, racemiform or thyrse-like, 

peduncles 6-12 cm long; leaf blades with a distinct marginal vein within ca. 0.3 mm of the leaf 

edge and connecting the distal ends of the major secondary veins; stipules narrowly 2-lobed . . . 

P. cincta 

31b. Inflorescences rarely over 14 cm long, usually more than 4 cm broad when over 10 cm long, not 

racemiform or narrowly thyrse-like; leaf blades lacking a distinct marginal vein along the leaf edge; 

stipules various 32 

32a. Inflorescences with conspicuous bracts and bracteoles to 14 mm long and 3 mm broad, the bracts 

persisting with the flowers . 33 



FIELDIANA: BOTANY 



32b. Inflorescences without conspicuous bracts, bracts rarely more than 3 mm long or more than 1 mm 

broad, bracts often deciduous before the flowers mature 37 

33a. Inflorescence small and compact, less than 3 cm long but expanding in fruit; stipules with 

narrow awns 2-6 mm long [0-900 m elevation] p. brachybotrya 

33b. Inflorescences larger and more open, to 10 cm long; stipule lobes less than 2 mm long or 

triangular (not narrowly awned) 34 

34a. Stipules very shortly (1-3 mm) bilobed with a U-shaped sinus between them [lateral branches 

of the inflorescences not subtended by bracts; stipules 2-5 mm long] P. officinalis 

34b. Stipules bilobed distally with a short or long V-shaped sinus between the lobes 35 

35a. Flowers subtended by ovate bracteoles 2-3 mm long, lateral branches of the inflorescences 

subtended by adnate bracts to 8 mm long; stipules 3-7 mm long P. brachiata 

35b. Flowers subtended by lanceolate to oblanceolate bracts 4-14 mm long, lateral branches of 

the inflorescence usually without subtending bracts; stipules 8-20 mm long 36 

36a. Flowers subtended by lanceolate bracts 4-9 mm long; inflorescences with 4 or more nodes 

with lateral branches; corolla tube 3-7 mm long P. capitata 

36b. Flowers subtended by oblanceolate bracts 5-14 mm long; inflorescences with 1-3 nodes with 

lateral branches; corolla tube 7-10 mm long P. calochlamys 

37a. (from 32b) Stipules with 1 narrow lobe (usually 2/node), the awn-like lobe acute (rarely bifid) 

distally and with an inner (adaxial) tooth; 500-1600 m elevation P. valeriana 

37b. Stipules with 2 lobes on each side (4/node), the lobes separate laterally and without a tooth-like 

adaxial (inner) appendage, or the lobes/teeth not developed; 0-2000 m elevation 38 

38a. Inflorescences less than 1.5 cm long, less than 12 mm wide; fruit becoming red at maturity .... 

P. haematocarpa 

38b. Inflorescences becoming more than 2 cm long and more than 2 cm broad; fruit blue to purple or 

black at maturity (orange during development in P. racemosd) 39 

39a. Leaf blades linear-lanceolate to narrowly elliptic-oblong, largest blades less than 3 cm broad; 1000- 

2000 m elevation P. goldmanii 

39b. Leaf blades rarely linear-lanceolate, larger leaves over 3 cm broad; 0-1500 m elevation 40 

40a. Inflorescence branches 0.7-1.5 mm thick when dried, flowering part of the inflorescence (beyond 

the peduncle) broader than long 41 

40b. Inflorescence branches slender (ca. 0.4 mm thick) when dried, flowering portion of the inflorescences 

usually longer than broad [fruit less than 8 mm long and 7 mm diam. when dried] 43 

4 la. Corolla tubes less than 3 mm long; inflorescences consistently with 4 lateral branches at the 

first node; rare in Costa Rica at 1 500 m elevation P. allenii 

41b. Corolla tubes 10-15 mm long; inflorescences with 2 or 4 opposite lateral branches at the first 

node; from below 1 200 m elevation in Costa Rica 42 

42a. Fruit 8-12 mm long and 7-10 mm diam., without transverse projections; leaf blades stiffly 

chartaceous; calyx lobes ca. 0.2 mm long P. eurycarpa 

42b. Fruit 4-5 mm long and ca. 6 mm diam., with transverse projections when dried; leaf blades 

thinly chartaceous; calyx lobes 0.2-1 mm long P. domingensis 

43a. Leaf blades drying membranaceous or thin-chartaceous, usually cuneate at the base and decurrent 

on the petiole; plants often found in deciduous forest formations .44 

43b. Leaf blades usually drying chartaceous, rarely cuneate and conspicuously decurrent on the petiole; 

never collected in deciduous areas 

44a. Leaf blades with 3-7 pairs of major 2 veins, often obovate; corolla tube 6-10 mm long; fruit 

7-10 mm long P microdon 

44b. Leaf blades with 9-15 pairs of major 2 veins, elliptic to ovate; corolla tube 2.5-4 mm long; 

fruit 5-6 mm long P pubescens 

45a. Fruits often with 5 seeds, ovary with 5 locules; stipules with stiff sharp awns 6-14 mm long and 

persisting; leaf blades with 3 veins subparallel [with 7-12 pairs of major 2 veins] P. racemosa 

45b. Fruit never with more than 2 seeds, ovary with 2 locules; stipules with awns or lobes 2-8 mm 

long; 3 8 veins not subparallel 46 

46a. Stipules with lobes or teeth absent or rarely 1 mm long, corolla tube 5-8 mm long (rare in Costa 

Rica) P. phanerandra 

46b. Stipules with lobes or teeth 2-8 mm long; corolla tubes 0.7-5 mm long 47 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 227 



47a. Stipules with narrow lobes 2-3 mm long; fruits ca. 4 mm long and 6 mm with smooth rounded 
surfaces (dried); leaf blades with long (1-3 cm) drip tips; inflorescences often small (to 6 cm long) 
P. acuminata 

47b. Stipules with narrow awns 3-8 mm long; fruits ca. 3 mm long and 4 mm diam. with longitudinal 
ribs and transverse depressions; leaf blades with narrowed tips 1-2 cm long; inflorescences to 12 
cm long P. deflexa 



Key to Group 4: Species of Subgenus Psychotria and Similar Species in Costa Rica 

la. Leaf blades 3-9 cm long, very rarely more than 7 cm long; small shrubs of evergreen forests ... 2 
Ib. Leaf blades 5-30 cm long, the largest leaf blades usually more than 9 cm long; plants of evergreen 

and deciduous habitats 6 

2a. Young stems densely puberulent with short (0. 1-0.2 mm) reddish brown hairs; stipules acute 
at the apex and caducous; inflorescences less than 1 6 mm long [ 1 200-2200 m elevation] . . 

P. parvifolia 

2b. Young stems glabrous (except for a ring of reddish hairs above the stipular scar); stipules 
bilobed or bifurcate distally; inflorescences usually more than 20 mm long (except in P. 

chagrensis) 3 

3a. Inflorescences capitate and sessile, less than 1 7 mm long and subtended by large bracts to 1 
cm long, corolla tube 4-8 mm long; stipules bifurcate distally with a V-shaped sinus [wet 

lowlands, 0-300 m elevation] P. chagrensis 

3b. Inflorescences paniculate and pedunculate usually more than 20 mm long and subtended by 
inconspicuous bracts, corolla tubes 2-4 mm long; stipules with 2 distal lobes separated by a 

broad U-shaped sinus 4 

4a. Stipule lobes glabrous or minutely (< 0.05 mm) papillate puberulent; inflorescence branches 

ca. 0.4 mm thick when dried [0-500 m elevation in Nicaragua and Panama] 

P. fruticetorum 

4b. Stipule lobes minutely (ca. 0.1 mm) ciliolate; inflorescence branches 0.2-0.3 mm thick when 

dried; plants known to occur in Costa Rica 5 

5a. Leaf blades rarely more than 5 cm long; petioles 2-10 mm long (the leaves not sessile); 

inflorescences not exceeding 5 cm in length [0-1800 m elevation] P. graciliflora 

5b. Leaf blades (at least the larger blades) usually exceeding 1 cm in length, petioles 0-50 mm 

long (leaves sessile in some spp.); inflorescences usually exceeding 5 cm in length 21 

6a. (from Ib) Young stems conspicuously puberulent with reddish hairs 0.2-2 mm long, internodes 

puberulent above the ring of reddish hairs (colleters) at the stipule scar 7 

6b. Young stems glabrous or inconspicuously puberulent with minute (0.03-0. 1 mm) hairs, but a ring 

of reddish hairs (colleters) often present just above the stipule scars 14 

7a. Inflorescences small (1-5 cm), flowers usually congested distally; peduncles to 2 cm long . . 

8 

7b. Inflorescences larger (4-15 cm long), flowers separate or congested, peduncles 1.5-15 cm long 

10 

8a. Leaf blades with an arcuate submarginal vein; stipules with a tube 4-14 mm long and 

narrow awns; fruits 5-6 mm long [0-200(-800) m elevation] P. psychotriifolia 

8b. Leaf blades usually lacking an arcuate submarginal vein but the distal veins often loop- 
connected; stipules with a tube 0-5 mm long, usually lacking narrow awns; fruits 6-14 

mm long 9 

9a. Leaf blades usually elliptic-obovate or elliptic; stipules 3-1 1 mm long; fruits 6-8 mm 

long; widespread at 20-800(-1200) m elevation P. nervosa 

9b. Leaf blades usually oblanceolate to narrowly elliptic-oblong; stipules 8-14 mm long; 

fruits unknown; Chiriqui Highlands ca. 1 200 m elevation P. boquetensis 

lOa. Leaf blades with 14-33 pairs of major 2 veins, leaf blades 12-30 cm long; peduncles 4-14 

cm long 11 

lOb. Leaf blades with 8-13 pairs of major 2 veins, leaf blades 6-30 cm long; peduncles 1.5-8 cm 
long 12 



228 FIELDIANA: BOTANY 



1 la. Leaf blades with 14-22 pairs of major 2 veins; lacking a definite submarginal vein, 3 

veins subparallel; floral bracts ca. 3 mm long, narrowly triangular P. micrantha 

1 Ib. Leaf blades with 20-34 pairs of major 2 veins, with definite submarginal veins and 

subparallel 3 veins; floral bracts ca. 7 mm long and ovate P. sixaolensis 

12a. Leaf blades rounded at the base and subcordate at the petiole, to 30 cm long, petioles 6-12 
cm long; stipules ca. 2 cm long; rarely collected plants of the Caribbean escarpment [200- 

500 m elevation] p. insignis 

1 2b. Leaf blades not rounded and subcordate at the base (or if so not regularly exceeding 1 8 cm 

in length), petioles usually less than 6 cm long; widespread 13 

1 3a. Corolla tubes 2-2.5 mm long; fruit 4-6 mm long; peduncles 1.5-5 cm long; leaf blades usually 
drying reddish brown or brown, major veins not impressed on the upper surfaces; (200-)400r 

1600 m elevation P. jimenezii 

13b. Corolla tubes 1.5-2 mm long; fruit 5-7 mm long; peduncles 3-8 cm long; leaf blades usually 
drying grayish, major veins impressed on the upper leaf surfaces; 20-400 m elevation .... 

P. neillii 

1 4a. (from 6b) Larger leaf blades usually more than 1 cm broad and usually more than 20 cm long 

15 

14b. largest leaf blades less than 10 cm broad, rarely more than 20 cm long 21 

15a. Inflorescences short (3-1 1 cm long) and compact (2-5 cm broad), with lateral branches to 1 

cm long and often difficult to see; plants 0.5-2 m tall 16 

1 5b. Inflorescences to 30 cm long and 4-15 cm broad, with lateral branches more than 1 cm long 

and clearly visible; plants 1-10 m tall 18 

16a. Leaf blades with 16-19 pairs of major 2 veins, usually more than 25 cm long, rounded 

to obtuse at the apex; bracts ca. 5 mm long P. chitarriana 

16b. Leaf blades with 8-14 pairs of major 2 veins, usually less than 25 cm long, short- 
acuminate to acute at the apex; bracts 0.2-2 mm long 17 

17a. Stipules with a basal tubular sheath and 2 narrow distal lobes; leaf blades with 10-14 
major 2 veins on each side, usually obovate; inflorescences to 7 cm long, compact 

panicles of cymes : P. alfaroana 

1 7b. Stipules not tubular at base, broadly ovate and bluntly obtuse at the apex; leaf blades 
with 8-1 1 pairs of major 2 veins, usually broadly elliptic; inflorescences to 3 cm long, 

with short lateral branches and appearing subumbellate P. lamarinensis 

18a. Stipules united over the shoot apex (as in Ficus) and tearing off as the leaves begin to grow, 

narrowly conical in early stages; fruit 5-9 mm long (var. compressicaulis of) 

P. panamensis 

18b. Stipules united only at their base, open distally and often with the base persisting, ovate- 
triangular in form; fruit 4-6 mm long 

19a. Leaf blades subsessile and subcordate at base, usually 3-4 times longer than wide, 2 veins 

arising from the mid vein at angles of 80-100 P. rosulatifolia 

19b. Leaf blades with prominent petioles, never rounded at the base, blades 2-3 times longer than 

wide, 2 veins usually arising at angles of 60-70 .20 

20a. Secondary veins not usually loop-connected; stipules 8-30 mm long and acute to acuminate 

at the apex; inflorescences to 30 cm long .P. grandis 

20b. Secondary veins loop-connected distally to form an arcuate submarginal vein; stipules 5-12 

mm long and rounded to acute; inflorescences to 10 cm long . P. limonensis 

2 la. (from 5b and 14b) Flowers in dense sessile verticils on a single elongated (4-9 cm) rachis or in 

dense clusters in a compact often globose arrangement, inflorescences solitary and apical, spiciform 

or capitate/subcapitate, primary branches short (< 1.5 cm) or absent 

21b. Flowers usually in open cymes, or on open spreading branches of the inflorescences when closely 
clustered, never spiciform or globose-capitate, primary branches of inflorescences usually more 

than 1 cm long 

22a. Flowers in dense sessile verticillate clusters along the slender elongated (3-9 cm) rachis, 
inflorescences spiciform with a single pair of proximal opposite branches or none; leaf blades 
usually with domatia in the distal vein axils P- viridis 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 229 



22b. Flowers in dense subcapitate globose of ellipsoid inflorescences with a rachis less than 4 cm 

long; leaf blades lacking large domatia P. alfaroana 

23a. Stipules united over the shoot apex and forming a cap with a single acute apex as in Ficus (usually 
not split or open distally during development), narrowly conical to cylindrical and elongate, usually 
becoming more than 5 mm long before splitting, caducous as the leaves begin to expand (note that 
new lateral branches beneath the inflorescences often are enclosed within F/cws-like stipules after 
other stipules have fallen); inflorescences with opposite branches; leaves lacking a submarginal 

vein 24 

23b. Stipules united over the shoot apex only in very early stages (if present the united cap usually 
splitting before reaching 5 mm in length and open distally in later development), caducous or 
tardily deciduous, larger F/cs-like stipules not present; inflorescences with opposite or whorled 

branches; leaves with or without a submarginal vein 27 

24a. Leaf blades usually with pit domatia or tufted hairs in the vein axils beneath, drying grayish 

or pinkish gray above, with 10-13 pairs of major 2 veins; ca. 600 m elevation in the Cordillera 

de Tilaran P. mexiae 

24b. Leaf blades lacking pit domatia or tufted hairs in the vein axils, drying grayish to dark reddish 

brown, with 5-16 pairs of major 2 veins; 400-2100 m elevation 25 

25a. Stipules 2-6 mm long, central 2 veins arising at angles of 60-90, leaf blades 4-12 x 1.5- 

5 cm [drying dark grayish above, with 5-7 pairs of major 2 veins; fruits 4-7 mm long] . . 

P. orosiana 

25b. Stipules usually becoming more than 6 mm long; central 2 veins arising at angles of 50-70, 

leaf blades 6-22 x 2-10 cm 26 

26a. Leaf blades usually drying grayish above, with 6-8 pairs of major 2, usually narrowly elliptic 

and with long petioles; stipules becoming 5-45 mm long; dried fruits 7-10 mm long 

P. sarapiquensis 

26b. Leaf blades usually drying reddish brown to dark brown (less often dark grayish), with 6-16 

pairs of major 2 veins, very variable in shape and texture; stipules becoming 8-80 mm long; 

fruits 5-9 mm long P. panamensis 

27a. (from 23b) Inflorescences only 2-4 cm broad, with short (< 2 cm) lateral branches (sometimes 
becoming larger as the fruit develop) [flowers often closely congested distally]; leaf blades with 2 
veins often weakly loop-connected near the margin; plants usually found in deciduous and partly 

deciduous formations 28 

27b. Inflorescences usually 4-8 cm broad, the lateral branches usually more than 2 cm long; leaf blades 

with or without loop-connected 2 veins; plants not found in deciduous formations 31 

28a. Corolla tube 4-5 mm long; fruit 6-9 mm long; inflorescences 1-3 cm long, with opposite 

branches or the flowers subsessile at the end of the peduncle [broad ovate stipules with an 

obtuse apex often subtending the young inflorescence] P. quinqueradiata 

28b. Corolla tube 1-3.5 mm long; fruit 4-8 mm long; inflorescences 1 .5-10 cm long, with opposite 

or whorled branches 29 

29a. Inflorescences with only 2 branches at each node, often umbelliform; corolla tube 1.5-2 mm 

long; stipules 6-14 mm long, acute and becoming narrowly 2-lobed at the apex; leaf blades 

narrowly elliptic, to 20 cm long P. tenuifolia 

29b. Inflorescences usually with 4 branches at the first node (2 longer and 2 shorter), rarely 

umbelliform; corolla tube 2.2-3.5 mm long; stipules 2-8 mm long and bluntly obtuse; leaf 

blades usually obovate to elliptic, to 1 5 cm long 30 

30a. Pit domatia rarely present in distal vein axils beneath; leaves usually drying gray to pinkish 

gray; calyx not usually persisting on the fruit; deciduous formations (in Costa Rica) 

P. carthagenensis 

30b. Domatia or tufts of hairs often present along the midvein beneath; leaf blades usually drying 

gray to greenish gray [calyx often persisting on the fruit with lobes 0.5-3 mm long]; in both 

evergreen and deciduous areas P. horizontalis 

3 la. (from 27b) Inflorescences usually with 4 lateral branches at the first node (2 smaller descending 

and 2 larger ascending); leaf blades usually narrowly elliptic to oblanceolate and with an arcuate 

submarginal vein formed by the loop-connected secondaries (compare dichotomy 28 also) . . 32 

31b. Inflorescences with 2 opposite lateral branches at the first node; leaf blades various 33 

230 FIELDIANA: BOTANY 



32a. Corolla tube 2-2.5 mm long; leaves usually drying grayish; wet lowlands (0-800 m elevation) 

P. clivorum 

32b. Corolla tube 3-5 mm long; leaves usually drying reddish or grayish brown; montane (2000- 

2600 m elevation) forests p. sylvivaga 

33a. Leaf blades usually with large (1-3 mm) domatia in the vein axils beneath, often opening by 
ellipsoid slits [blades 1 1-24 cm long, usually elliptic-oblong and drying dark reddish brown; 
inflorescence axes winged; peduncles ca. 2 mm thick; fruit 7-9 mm long; rarely collected in Costa 

Rica] P. remota 

33b. Leaf blades without large webbed domatia in the vein axils, smaller domatia present or absent 

34 

34a. Calyx lobes well developed (0.5-1 mm long); peduncles 1.5-2.5 mm thick when dried, primary 
branches of the flowering inflorescences 1-1.5 mm thick when dried; leaf blades lacking domatia'. 

often oblanceolate, to 22 cm long; montane (1300-2600 m elevation) forest species 35 

34b. Calyx lobes minute (0.3 mm) or absent; peduncles 0.5-1 .5 mm thick when dried, primary branches 
of the flowering inflorescences usually less than 1 mm thick when dried; leaf blades often with 
domatia on the lower surfaces, not usually oblanceolate or more than 1 5 cm long (except in P. 

sarapiquensis); 0-2500 m elevation 37 

35a. Primary branches of the inflorescence usually diverging at more than 100 from the more 
distal rachis ( reflexed); leaf blades with 10-15 pairs of major 2 veins, with or without an 
arcuate submarginal vein [flowers on open cymes on tertiary branches of a much-branched 

inflorescence, peduncles 5-9 cm long; 1 700-2000 m elevation] P. stockwellii 

35b. Primary branches of the inflorescence usually diverging at 90 or less from the rachis, per- 
pendicular or ascending; leaf blades with 8-1 1 pairs of major 2 veins; usually with a definite 

submarginal vein 36 

36a. Flowers in dense glomerules at the ends of the few (ca. 5) 1 or (ca. 9) 1 and 2 branches; 

peduncles 1.5-5.5 cm long; 900-1500 m elevation P. monteverdensis 

36b. Flowers in open cymes or pedicellate on 2 or 3 branches of the inflorescences; peduncles 

4-1 2 cm long; 1 300-2600 m elevation P. sylvivaga 

37a. Stipules developing 2 small (1-3 mm) slender lobes at the apex (4 acute tips per node) and usually 

united to form a short tube at the base; corolla tube usually more than 3 rtim long 38 

37b. Stipules acute at the apex (without small lobes), a short tube not apparent at the base; corolla tubes 

rarely more than 3 mm long (except in P. chiriquina) 

38a. Leaves drying dark reddish brown, petioles 7-18 mm long; stipules 4-10 mm long; 1500- 

2500 m elevation P. chiriquina 

38b. Leaves drying dark to pale grayish, petioles 0-5 mm long; stipules 2-5 mm long; 400-1900 

m elevation P- orosiana 

39a. Fruits 7-10 mm long, obovid-oblong; stipules conical, united and early caducous; leaf blades 

glabrous beneath and without domatia [600-1600 m elevation] P. sarapiquensis 

39b. Fruits 4-8 mm long (unknown in P. laselvensis), subglobose; stipules free distally, not united into 

a conical cap; leaf blades glabrous or minutely puberulent and often with domatia beneath 40 

40a. Highland (1500-2500 m) plants of easternmost Costa Rica and Chiriqui, Panama; leaves usually 

drying reddish brown; corolla tube 3-6 mm long P. chiriquina 

40b. Lower (0-1000 m) elevation plants; leaves usually drying grayish; corolla tube 1-3 mm long 41 
4 la. Leaf blades with domatia rarely present in distal vein axils, blades to 13(-16) cm long; stipules 

triangular; La Selva area, 50-300 m elevation P laselvensis 

41b. Leaf blades with domatia often present along the central and proximal part of the midvein, blades 
to 17 cm long; stipules ovate-triangular; widespread, 0-1000 m elevation P. marginata 

Psychotria acicularis C. M. Taylor, sp. nov. Figure TYPUS-/. Utley A K. Utley 1036 (holotypus CR. >- 

typi F, MO!), from region to southwest of airstrip, 20-60 

:>:> - m alt., 20 July 1974, Rincon de Osa, Puntarenas, Costa 

Rica. 

Species Psychotriae pittieri Standley affinis. sed ab ea 

bracteismajoribus(3-10mmlongis)etangustoribusdif- Small shrubs, 0.5-2 m tall, leafy stems 

f ert thick, densely hirsutulous with thin hairs 0.4-1.5 mm 

BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 231 



long, older stems glabrescent; stipules 3-8 mm long, with 
a short (1-3 mm) tube and 2 slender teeth 3-7 mm long 
on each side, densely hirsutulous in early stages, per- 
sisting or deciduous. Leaves with petioles 2-9(-13) mm 
long, ca. 0.7 mm thick, densely hirsutulous, often some- 
what unequal at the same node; leaf blades 5-12 cm 
long, 2-4.5 cm broad, narrowly ovate-elliptic to elliptic 
or elliptic-oblong, apex acuminate with a tip 4-12 mm 
long, base usually obtuse, drying thinly chartaceous or 
membranaceous, dark green above, with slender hairs 
0.5-1.2 mm long on the upper surface, especially along 
midvein, becoming glabrescent, more densely hirsutu- 
lous beneath with slightly shorter more persistent hairs, 
2 10-15/side. Inflorescences terminal and solitary, 5- 
12 cm long, open or elongate panicles with usually 4 
branches at apex of the peduncle and 2 additional pairs 
of opposite or alternate branches, peduncles 3-8 cm long, 
ca. 0.7 mm thick (dried), hirsutulous with hairs to 1.5 
mm long, bracts 3-10 mm long and linear, flowers sub- 
sessile in condensed distal cymes, bracteoles 2-4 m long, 
linear. Flowers densely puberulent externally, hypanthi- 
um ca. 1 mm long, sericeous with ascending thin whitish 
hairs ca. 0.5 mm long, calyx lobes 0.5-1 mm long, lan- 
ceolate; corolla white, narrowly salverform, tube ca. 3 
mm long and 0.5 mm diam., corolla lobes ca. 1.2 mm 
long; anthers ca. 1 mm long. Fruits 3-3.5 mm long, 2.5- 
3 mm diam., broadly ellipsoid, bluish with thin ascend- 
ing hairs ca. 0.5 mm long, persisting calyx lobes ca. 0.5 
mm long. 

Plants of the rain forest interior on the Osa Pen- 
insula, at elevations of 20-80 m. Flowering in May 
and July; fruiting in May-August. This species is 
only known from near Rincon de Osa on the Osa 
Peninsula in southwestern Costa Rica. 

Psychotria acicularis is recognized by its hir- 
sutulous indumentum on all parts, small stature, 
small leaves, open inflorescences with conspicuous 
linear bracts, sessile flowers borne in distal clus- 
ters, small blue fruit, and limited geographic range. 
This species, a member of subgenus Heteropsy- 
chotria, resembles P. pittieri vegetatively but dif- 
fers in its linear bracts and inflorescences with four 
branches at the first (basal) node. Additional spec- 
imens seen were Aguilar 6423 (CR), Burch 4413 
(DUKE, MO), Burger & Stolze5449(cR, F, MO), Duke 
16111 (MO), Kennedy 1 934 (MO), Raven 21532 (CR, 
F), and Utley & Utley 1174 (CR, F). 



Psychotria acuminate Benth., Bot. voy. Sulph. 107. 
1 845. P. cuspidata sensu Standl. et auctores, non 
Bredem. ex Roem. & Schult. fide Steyermark 
1974. Figure 56. 

Shrubs or subshrubs, l-2(-3) m tall, much-branched, 
leafy stems 1 .5-5 mm thick, glabrous, quadrate to terete; 
stipules with a short (0.5-2 mm) truncated tube with 2 
narrow awns 1.5-3 mm long on each side, these decid- 
uous. Leaves with petioles 5-15(-18) mm long, 0.7-1.4 



mm thick, minutely (0.05 mm) papillate-puberulent; leaf 
blades 7-15(-19) cm long, 3-8(-10) cm broad, elliptic 
to oblong-elliptic or ovate-elliptic, apex short- to long- 
acuminate or caudate-acuminate with tip 1-3 cm long, 
base acute to obtuse, drying thin-chartaceous and green- 
ish or brownish, glabrous above and below, or with tuft- 
ed domatia in the vein axils beneath, 2 veins 7-12/side, 
midveins sometimes reddish beneath. Inflorescences 
solitary and terminal, 2-6 cm long (to 7 cm in fruit); 2- 
4(-5) cm broad, paniculate, peduncles 1-4 cm long, 0.5- 
1 mm thick, minutely papillate-puberulent (or appar- 
ently glabrous), primary branches opposite or alternate, 
bracts subtending the inflorescence branches absent or 
small (1 mm) and adnate to branches, floral bracts ab- 
sent, pedicels 0-2 mm long. Flowers distylous, minutely 
(0.05 mm) papillate-puberulent externally, hypanthium 
ca. 0.5 mm long and 0.5 mm diam., calyx 0.3-0.7 mm 
long with weakly denned lobes; corolla white or pale 
yellow, tube 2-4(1-5) mm long and 0.5-1 mm diam., 
lobes 5, 1.5-2.5 mm long, rounded; anthers 1-1.2 mm 
long, linear. Fruits 4 mm long, 5-7 mm diam., 2-lobed 
and rounded, smooth, becoming blue, bluish gray, or 
purple; pyrenes 2.5 mm long and 3 mm broad, hemi- 
spheric with a longitudinal sulcus on the inner face. 

Plants of evergreen lowland rain forest forma- 
tions on both the Caribbean and Pacific slopes, 
from near sea level to 900(-1 100) m elevation in 
Costa Rica. Flowering in January-August in 
southern Central America (usually May-June at 
La Selva). Fruiting is primarily in June-Decem- 
ber. The species ranges from Mexico and the West 
Indies to Venezuela and Peru. 

Psychotria acuminata is characterized by its gla- 
brous or minutely papillate-puberulent parts, lus- 
trous acuminate leaves with pronounced narrow 
drip tips, small flowers and inflorescences, 2-awned 
stipules, and smooth rounded bilobed blue fruit. 
In some specimens, the stipules appear to be un- 
developed. The small stature, leaf size, and foliage 
drying greenish are similar to many other species, 
but the small yellowish papillate-puberulent inflo- 
rescences and smooth bilobed fruit are distinctive. 
Also, the veins on the undersides of the leaves are 
sometimes pink. The presence of domatia-like tufts 
of hairs in the vein axils is uncommon among our 
species of Psychotria subgenus Heteropsychotria. 
The breeding biology was studied by Bawa and 
Beach (1983). Psychotria valeriana of higher ele- 
vations is similar but has a single stipular awn 
with an adaxial tooth on each side of the stem. 
[The name P. cuspidata deserves more careful re- 
view; it may be that Steyermark failed to recognize 
a specific distinction between the Caribbean ma- 
terial and his Venezuelan specimens. C.M.T.] 



Psychotria aggregata Standl., Contr. U.S. Natl. 
Herb. 18: 128. 19 16. P. tonduziiSiandl, J. Wash. 



232 



FIELDIANA: BOTANY 



Acad. Sci. 15: 287. 1925. Montamans pana- 
mensis Dwyer, Ann. Missouri Bot. Gard. 67: 
286. 1980. Figure 12. 

Herbacous subshrubs with succulent stems, 0.5-1.5(-2) 
m tall, leafy stems 1 .6-6 mm thick (dried), quadrangular, 
glabrous; stipules 2-9(-12) mm long, to 7 mm broad, 
triangular with a short (2-3 mm) cupular base and bifid 
conical appendages 4-7 mm long, base persisting. Leaves 
with petioles 3-1 0(- 1 3) cm long, 1 -2 mm thick, glabrous; 
leaf blades 12-30(-38) cm long, (3-)5-17 cm broad, el- 
liptic-oblong to elliptic-obovate or elliptic-oblanceolate, 
apex tapering gradually and acute or acuminate, tip 5- 
1 5(-22) mm long, base obtuse to cuneate or acute and 
decurrent on petiole, drying chartaceous and dark to pale 
grayish green, glabrous above, glabrous or minutely pa- 
pillate-puberulent beneath, 2 veins 10-19/side. Inflo- 
rescences axillary, solitary, 4-1 5 cm long, 2-4 cm broad, 
congested to capitate or with 1-2 pairs of lateral branch- 
es, peduncles l-5(-8) cm long, 1-2 mm thick, glabrous 
or minutely and sparsely puberulent, bracts at the lateral 
branches 2-8 mm long, flowers subsessile in closely 
crowded distal glomerules on the 1 or 2 branches, brac- 
teoles 1-4 mm long. Flowers with hypanthium 1-2 mm 
long, glabrous or sparsely and minutely puberulent, calyx 
0.5-2 mm long, lobes 5, 0.5-4 mm long, obtuse to linear; 
corolla white, funnelform glabrous or minutely puber- 
ulent externally, tube 2-4 mm long, ca. 1 mm diam., 
corolla lobes 5, ca. 1 mm long, obtuse and galeate; an- 
thers ca. 1.2 mm long. Fruits 5-7 mm long (dried), 4-6 
mm diam. (to 10 mm in life), ovoid to subglobose, spongy, 
white; pyrenes ca. 6 mm long, with margin and median 
keel thickened dorsally. 

Plants of evergreen montane rain forest and cloud 
forest formations, from 10 to 2300 m elevation 
(but most common in the 400-1 700-m range). Ap- 
parently flowering and fruiting throughout the year. 
The species ranges from northern Costa Rica into 
western Panama. 

Psychotria aggregata is recognized by its short 
succulent unbranched main stems, large essen- 
tially glabrous leaves, axillary inflorescences with 
flowers in dense bracteolate clusters, and spongy 
white fruits. This species is quite similar to P. 
macrophylla but differs in the more condensed 
inflorescences and the pyrenes. (This and related 
species have recently been studied by Molly Ne- 
pokroeff, wis, 1 992.) The type of P. aggregata (Pit- 
tier 3264 us from Horqueta, Chiriqui, Panama) is 
a very poor specimen, but it is an early name for 
this variable taxon, commonly called P. tonduzii 
by previous authors. 



Psychotria alfaroana Standl., J. Wash. Acad. Sci. 
18:273. 1928. Figure 66. 

Herbaceous erect subshrubs, 0.2-0.6(-1) m tall, usu- 
ally with a single erect unbranched stem, rhizomatous, 



leafy stems 2-4 mm thick, glabrous, terete; stipules to 
20 mm long, basal sheath 5-8 mm long and 4 mm broad, 
with 2 narrow distal lobes 3-12 mm long, glabrous or 
sparsely puberulent, caducous. Leaves with petioles (2-)6- 
25(-30) mm long. 1.5-2 mm thick, sulcate above, gla- 
brous; leaf blades 1 1-22(-30) cm long, 4-1 1 cm broad, 
obovate to elliptic-obovate or elliptic-oblanceolate. apex 
short-acuminate or subacutc. base gradually narrowed 
and cuneate, decurrent on petiole, drying thin- to stiffly 
chartaceous, grayish or grayish green, glabrous above. 
with minute (0. 1-0.3 mm) thin hairs along major veins 
beneath, 2 veins 10-14/side, weakly connected by an 
arcuate submarginal vein. Inflorescences terminal, sol- 
itary, 3-8 cm long, 2-4 cm broad, compact globose or 
elongate dense panicles of cymes, peduncle 1-4 cm long, 
1.2-2 mm thick, glabrous, bracts 0.5-1 mm long, ped- 
icels 1-2 mm long. Flowers glabrous externally, hypan- 
thium ca. 1.5 mm long, calyx tube ca. 1 mm long, cu- 
pulate, calyx lobes 5, 0.5-2 mm long, narrowly oblong; 
corolla white, tube 2.5-4 mm long, ca. 2 mm diam.. 
lobes 5, 1 .5-3 mm long, ca. 1 mm broad at base; stamens 
5, anthers 0.8-1.3 mm long. Fruits 8-12 mm long, 4-6 
mm diam., ellipsoid, red at maturity, becoming black 
when dried, persisting calyx to 3 mm long; pyrenes ca. 
7 mm long, usually with 5 dorsal ribs. 

Plants of wet rain forest formations of the Ca- 
ribbean slope and continental divide, 20- 
900(-1100) m elevation. Flowering in January- 
September; fruiting in January-February and June- 
August. The species ranges from the Cordillera de 
Guanacaste in Costa Rica to Bocas del Toro Prov- 
ince in Panama. 

Psychotria alfaroana is recognized by its very 
short stature, usually obovate leaves, compact 
subglobose inflorescences, and larger fruit. The 
bright red fruit and leaves drying grayish are char- 
acteristics of subg. Psychotria. Standley and Vale- 
ric made a number of collections of this species 
near Tilaran, including the holotype (41579 us). 
A collection from along Rio Corobici (Opler 138 
F) is provisionally placed here; this is the only 
collection from the Pacific lowlands. Several col- 
lections with exceptionally large (35 x 15 cm) 
leaves attenuate almost to the base and then 
abruptly subauriculate are tentatively placed here: 
Gdmez-Laurito 11570, Hammelet al. 16895, and 
Herrera & Chacdn 2355 (all at CR). 



Psychotria allenii Standl., Ann. Missouri Bot. 
Gard. 27: 342. 1940. 

Small trees or shrubs, (2-)4-10 m tall, leafy stems 1- 
4 mm thick, glabrous or minutely and sparsely puber- 
ulent; stipules united around the stem for 0.5-3 mm, 
with 2 acute lobes 1-5 mm long, separated by a V- or 
U-shaped sinus, persisting. Leaves with petioles 6-18 
mm long, 0.8-1.3 mm thick, glabrous; leaf blades 5-16 
cm long, 3-7(-8) cm broad, ovate-elliptic or ovate-ob- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



233 



long to narrowly elliptic-oblong, apex acuminate with 
tip ca. 5 mm long, base obtuse to slightly rounded, drying 
dark olive green or grayish green and chartaceous, gla- 
brous above, glabrous or minutely puberulent beneath, 
2 veins 5-7/side, distal veins strongly arcuate-ascend- 
ing. Inflorescences 8-10 cm long, 5-7 cm broad, pyra- 
midal paniculate with distal branches progressively 
shorter, peduncles 3.5-6.5 cm long, 0.9-2.3 mm thick, 
glabrous or puberulent, the first node with 4 lateral 
branches 2-3.5 cm long subtended by linear bracts 1-3 
mm long, flowers mostly sessile and crowded in distal 
cymes. Flowers with hypanthium 0.4-0.8 mm long, gla- 
brous or sparsely papillate-puberulent, calyx ca. 0.7 mm 
long with glabrous lobes ca. 0.3 mm long; corolla white, 
tube 0.7-2 mm long and 1 mm diam., lobes 5, ca. 1.7 
mm long; stamens 5. Fruits 6-1 5 mm long and ca. 6 mm 
diam. when dried, bright blue; pyrenes smooth, subglo- 
bose. 

Plants of moist evergreen lower montane for- 
ests, 50-900(-1500) m elevation. Flowering and 
fruiting in February-August in Panama. The spe- 
cies is known only from Monteverde in Costa Rica 
(Koptur 74 MO with flowers and early fruits in 
October). The species ranges to central Panama. 

Psychotria allenii is recognized by its distinctive 
pyramidal inflorescences with relatively thick pe- 
duncles, four stout lateral branches at the first few 
nodes, and small sessile flowers. This species is 
quite similar to P. officinalis, but that species has 
floral bracts and larger flowers and lacks the robust 
lateral branches of the inflorescence. The Costa 
Rican collection has narrower elliptic-oblong leaves 
than Panamanian material, and it was a 2-m shrub 
rather than a tree. 



Psychotria angustiflora K. Krause, Bot. Jahrb. 54, 
Beibl. 119: 43. 1916. P. mima Standl., Publ. 
Field Mus. Nat. Hist., Bot. Ser. 22: 386. 3 1 Oct. 
1 940 (holotype: Skutch 4589 F from Costa Rica), 
not P. mima Standl., loc. cit. 22: 204. 10 Sept. 
1 940 (holotype: L. B. Smith 1878 F from Brazil). 
Figure 57. 

Shrubs or small trees to 5 m tall, leafy stems 1.5-5 
mm thick, glabrous, terete; stipules 3-8 mm long, 2-6 
mm broad, triangular to ovate and rounded at the nar- 
rowed tip, with a short (1-3 mm) tube, obtuse or bilobed 
with a small ( 1 mm) sinus, glabrous, persisting. Leaves 
with petioles 15-35(-60) mm long, 1.2-3 mm thick, gla- 
brous, with 2 adaxial ridges; leaf blades (9-) 1 5-24(-30) 
cm long, (4-)7-14(-16) cm broad, oblong to ovate-ob- 
long, broadly elliptic or broadly elliptic-obovate, apex 
short-acuminate with a tip 5-10 mm long, base broadly 
obtuse (rarely acute), drying thin-chartaceous and green- 
ish gray or greenish brown above, glabrous above and 
below, 2 veins 8-1 I/side, central 2 veins ca. 2 cm dis- 
tant in larger leaves. Inflorescences terminal, solitary 
(3-) 17-45 cm long, (8-) 15-25 cm broad, pyramidal pa- 



niculate with distant opposite lateral branches, peduncles 
6-15 cm long, 1.5-2 mm thick, glabrous or minutely 
puberulent, bracts to 7 mm long and 1.5 mm broad, 
flowers subsessile in distal cymes or along slender distal 
branches, subtended by bracteoles ca. 1 mm long. Flow- 
ers minutely papillate-puberulent externally, hypanthi- 
um ca. 0.7 mm long, calyx ca. 0.4 mm long, calyx lobes 
ca. 0.2 mm long, acute; corolla pale yellow or greenish 
yellow (rarely white), tubular-obovoid or funnelform, 
tube 5-7.5 mm long, 0.9-1 .3 mm diam. in the lower half 
and 2-3 mm distally, lobes 5, 1-2.5 mm long, triangular 
and acute, erect. Fruits 4-5 mm long and 3-5 mm diam. 
(to 8 mm in life), subglobose with ca. 6 longitudinal ribs, 
becoming blue or black; pyrenes 1.5-3.5 mm long. 

Plants of evergreen rain forest formations, from 
near sea level to 900 m elevation on the Caribbean 
slope. Flowering in January-August; fruiting in 
June and October-November. The species ranges 
from near La Selva, Heredia, to Brazil. 

Psychotria angustiflora is recognized by its large 
open inflorescences with small distal bracteoles, 
narrowly obovoid yellowish corolla, larger leaves, 
and ovate stipules that are obtuse or minutely two- 
lobed at apex. The stipules may become two-part- 
ed as the expanding stem breaks them apart. The 
flower buds are quite narrow before the corolla is 
fully expanded. The isotype of P. angustiflora 
(Tonduz 12996 us) fits in well with the material 
placed here. This species should be compared to 
P. berteriana (smaller flowers, different stipules, 
narrower leaves) and P. luxurians (smaller flowers, 
larger bracteoles). 



Psychotria aubletiana Steyerm., Mem. New York 
Bot. Card. 23: 694. 1972. Cephaelis axillaris 
Sw., Prodr. 45. 1788, not P. axillaris Willd. P. 
aubletiana var. centro-americana Steyerm., loc. 
cit. 698. 1972. 

Herbs or subshrubs, 0.5-1 .5(-4) m tall, erect and few- 
branched, stems slightly succulent, leafy stems 1-5 mm 
thick, glabrous or rarely densely puberulent with soft 
yellowish hairs ca. 0.2 mm long, quadrangular, drying 
dark; stipules 5-10 mm long, to 6 mm broad, basal sheath 
3-5 mm long, ovate with a short (2-4 mm) sinus and 2 
irregular rounded lobes per side. Leaves with petioles 6- 
20(-40) mm long, 0.7-1.8 mm thick, glabrous or mi- 
nutely puberulent; leaf blades 4-15(-18) cm long, 1- 
4(-6.5) cm broad, elliptic-oblong, narrowly oblong-ob- 
ovate to narrowly elliptic-obovate, oblong-lanceolate or 
narrowly elliptic, apex acuminate (rarely acute) with tip 
4-1 2 mm long, acute at the base (obtuse in larger leaves), 
drying stiffly chartaceous, dark greenish above, much 
paler beneath, glabrous above, glabrous beneath or with 
short (0.1-0.2 mm) hairs on major veins, 2 veins 6-87 
side (with lesser parallel intermediate 2 veins). Inflo- 
rescences axillary and sessile, 1.3-2 cm diam., densely 
compact, subglobose, with distal 2-lobed stipules resem- 



234 



FIELDIANA: BOTANY 



bling bracts, bracts obovate, purple or green, 2-7 mm 
long, flower 5-1 5/head, sessile. Flowers with 5 calyx lobes 
1-2 mm long; corolla white, glabrous, tube ca. 10 mm 
long, lobes 5, 3-3.5 mm long, triangular, stamens 5, 
anthers 1-2 mm long. Fruits 5-7 mm diam., ellipsoid, 
bright blue; pyrenes 3-5 mm long and 2-4 mm broad. 

Understory plants of evergreen montane rain 
forest formations, from (600-)1 200 to 2300(-2800) 
m elevation in Costa Rica. Probably flowering and 
fruiting throughout the year. The species ranges 
from Guatemala through Central America to Co- 
lombia, Venezuela, and the lesser Antilles. 

Psychotria aubletiana is recognized by its small 
stature, sessile rounded inflorescences often encir- 
cling distal nodes and subtended by an involucre 
of broad bracts, short white corolla tubes, and 
bright blue fruit. This species is not easily confused 
with other species of Psychotria. 



Psychotria aurantibractea C. M. Taylor, nom. nov. 
Cephaelis pittieri K. Krause, Bot. Jahrb. Syst. 
54, Beibl. 1 19: 45. 1916, not Psychotria pittieri 
Stand i. Figure 18. 

Shrubs, 1-3 m tall, leafy stems 1.3-4.5 mm thick, 
glabrous, often narrowed below the node when dried; 
stipules united for only 0.5-2 mm around stem, with 2 
narrowly triangular or linear teeth 2-8 mm long, sepa- 
rated by 2-3 mm, persisting. Leaves with petioles 5-40 
mm long, 0.8-1.5 mm thick, glabrous; leaf blades 8- 
21 (-28) cm long, 2-6 cm broad, narrowly oblong to nar- 
rowly elliptic-oblong or lanceolate, apex gradually ta- 
pering and acute or acuminate, tip to 1 5 mm long, base 
cuneate to acute, drying membranaceous or thin-char- 
taceous and dark brown, glabrous above, glabrous or 
sparsely and minutely (0. 1 mm) puberulent on the veins 
beneath, 2 veins 10-16/side. Inflorescences terminal, 1 
or 3, 2.5-6 cm long, 4-6 cm broad (to 8 cm when tri- 
partite), usually with 3 capitate or cymose clusters of 
flowers on short (1-4 mm) primary branches, peduncles 
5-25 mm long, 1.3-2.2 mm thick, minutely puberulent, 
bracts 8-15 mm long, 4-9 mm broad, ovate to lanceo- 
late, acuminate, orange (rarely red), sparsely puberulent 
and ciliolate along margins, flowers sessile. Flowers en- 
closed by bracts and bracteoles, hypanthium ca. 1 .5 mm 
long, densely pubescent with straight ascending hairs, 
calyx lobes 5, ca. 1.3 mm long, dentate with narrrow 
lobes; corolla yellow or yellow-rose, tube ca. 5 mm long, 
lobes ca. 2 mm long. Fruits 5-7 mm long, 2.5-5 mm 
diam., ovoid-ellipsoid, with prominent longitudinal cos- 
tae and short thin hairs, color unknown, persisting calyx 
1 .3-2 mm long, conical; pyrenes ridged. 

Plants of continually wet or seasonally dry ev- 
ergreen forest formations, from 200 to 1200 m 
elevation. Flowering in May-July; fruiting in Au- 
gust-September and January. This species is en- 
demic to southern Costa Rica, from San Isidro del 



General to San Vito and the Osa Peninsula, and 
on the adjacent Caribbean slope (at 83W). 

Psychotria aurantibractea is recognized by its 
terminal inflorescences composed of large ovate, 
orange or reddish bracts, narrow leaves, and re- 
stricted geographical range. The colorful bracteate 
inflorescences made this and other species for- 
merly placed in Cephaelis so distinctive in Central 
America. 



Psychotria berteriana DC., Prodr. 4: 515. 1830. 
Figure 58. 

Shrubs or small trees, 1.5-6(-8) m tall, leafy stems 
(1.5-)3-5 mm thick, glabrous or minutely puberulent 
with hairs 0.1-0.2 mm long; stipules at first triangular 
and acute with a small distal cleft, becoming 2-lobed or 
2-awned (on each side), basal sheath 0.5-2 mm long with 
acute lobes 1-3.5 mm long. Leaves with petioles 10- 
30(-52) mm long, 0.6-1.8 mm thick, glabrous or mi- 
nutely puberulent; leaf blades 8-20(-26) cm long, 3- 
9(-ll) cm broad, narrowly ovate-elliptic to elliptic or 
elliptic-oblong, apex acuminate with 5-15 mm long, base 
acute and cuneate to obtuse (rarely rounded and sub- 
truncate in large leaves), drying membranaceous to thin- 
chartaceous, dark green or brown, glabrous or with min- 
ute (0.1-0.2 mm) hairs along the veins above, glabrous 
or with thin whitish hairs ca. 0.2 mm long beneath, 2 
veins 7-15/side. Inflorescences terminal and solitary. 5- 
16(-22) cm long, 5-15 cm broad, oblong but becoming 
broadly pyramidal paniculate with opposite or alternate 
branches, yellowish in life, peduncles 5-10 cm long. 1 .3- 
2.2 mm thick, sparsely puberulent with thin hairs ca. 0.2 
mm long, bracts 3-8 mm long, ca. 0.6 mm broad, flowers 
sessile in distal cymes, bracteoles 1-2 mm long. Flowers 
minutely puberulent externally (rarely glabrous), hypan- 
thium ca. 0.5 mm long, calyx lobes ca. 0.2 mm long; 
corolla tubular-salverform, yellow to white, tube 1.5-4 
mm long, 1-2 mm diam., glabrous proximally, lobes 5. 
1-1.5 mm long; anthers 1-1.4 mm long. Fruits 34 mm 
long (5 mm including the persisting calyx), 3-4 mm diam.. 
subglobose, fleshy, lustrous black; pyrenes with 3-5 dor- 
sal ridges. 



Plants of evergreen rain forest formations, from 
near sea level to 1 700 m elevation (to 1 200 m on 
the Pacific slope of southern Costa Rica). Flow- 
ering throughout the year; probably fruiting 
throughout the year. The species ranges from Mex- 
ico to South America. 

Psychotria berteriana is characterized by its large, 
many-branched, pyramidal inflorescences, thin 
leaves often with long slender petioles and drying 
dark, two-lobed stipules, small flowers, and small 
rounded fruit. Compare P. luxurians and P. an- 
gustiflora (with longer thinner corolla tubes). A 
broader interpretation of P. berteriana might re- 
quire the inclusion of P. luxurians. The greenish 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUB1ACEAE 



235 



yellow inflorescences resemble those found in Pal- 
icourea tilarensis. 



Psychotria boquetensis Dwyer, Ann. Missouri Bot. 
Gard. 67: 349. 1980. 

Shrubs, 0.5-2 m tall, leafy stems 1.2-3.5 mm thick, 
tomentulous with reddish hairs 0.4-1 mm long; stipules 
8-14 mm long, 2-6 mm broad, oblanceolate to ovate, 
obtuse to acuminate at apex (not bilobed in the type as 
described by Hamilton, 1989), caducous. Leaves with 
petioles 1-6 mm long, 0.7-1.2 mm broad, with curved 
thin reddish hairs ca. 0.3 mm long; leaf blades 4-16 cm 
long, 1-4 cm broad, narrowly elliptic-oblong to oblan- 
ceolate, apex acuminate with tip 4-15 mm long, base 
gradually narrowed and acute or cuneate, decurrent on 
petiole, drying thinly chartaceous and dark reddish brown, 
puberulent along the primary and secondary veins above 
and below, with hairs 0.2-0.7 mm long along the margin, 
2 veins 10-13/side, with minute tufted domatia in the 
axils beneath. Inflorescences terminal or pseudoaxillary, 
several at a node, 2-4 cm long, small panicles, peduncles 
ca. 20 mm long and 0.6 mm thick, bracts represented 
by dense whitish hairs 0.5-1 mm long, flowers sessile or 
subsessile in compact cymes. Flowers glabrous on the 
distal surfaces, hypanthium ca. 1 mm long and puber- 
ulent, calyx lobes 1-1.5 mm long, narrowly triangular, 
corolla white, urceolate, tube 1.5-2.5 mm long and 1 
mm diam., lobes 5, 1-1.5 mm long; stamens 5, anthers 
0.5-0.7 mm long. Fruits unknown. 

Plants of the Boquete region of the Chiriqui 
Highlands at about 1 200 m elevation. Flowering 
material was collected in May-June. The species 
is only known from western Panama. 

Psychotria boquetensis is recognized by the nar- 
row leaves with conspicuous reddish hairs, mem- 
branaceous broad caducous stipules, small inflo- 
rescences, and narrow calyx lobes to 1.5 m long. 
The species is closely related to P. nervosa but 
differs in the large calyx lobes, narrower leaves, 
and smaller inflorescences. This distinctive species 
is a member of subgenus Psychotria. 



Psychotria borucana (A. Molina) C. M. Taylor & 
W. Burger, comb, nov., Cat. fl. Peru (in press). 
Cephaelis affinis Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 8: 1 84. 1 930, not P. affinis Baker. 
C. borucana A. Molina, Ceiba 4: 31. 1951. Fig- 
ure 17. 

Shrubs or woody herbs, 1.5-4 m tall, leafy stems 2.5- 
8 mm thick, glabrous and drying dark, quadrangular or 
flattened; stipules 9-20 mm long and 5-10(-15) mm 
broad, basal sheath 1-2 mm long, ovate-triangular, ob- 
tuse, with parallel venation, glabrous or minutely brown- 
ish puberulent. Leaves with petioles 12-50 mm long, 



1 .7-3(-5) mm thick, glabrous (as in the type) or minutely 
papillate-puberulent with erect brownish hairs 0. 1 mm 
long; leaf blades 1 5-28(-32) cm long, 7- 14(- 16) cm broad, 
obovate to elliptic obovate (as in the type) to broadly 
elliptic, apex short-acuminate (acute) with tip 5-15 mm 
long, base gradually narrowed and cuneate to obtuse, 
drying stiffly chartaceous, dark grayish above and mark- 
edly paler beneath, glabrous above, glabrous beneath (as 
in the type) or with minute hairs 0.1-0.2 mm long 
throughout, 2 veins 6-10/side. Inflorescences terminal, 
capitulae solitary, 2-4 cm broad, peduncle 5-20 cm long, 
2.5-4 mm diam., glabrous or minutely puberulent, basal 
bracts mostly 2, 4-15 mm long, broadly ovate to reni- 
form, interior bracteoles 4-8 mm long, spatulate, pale 
purple, lavender, or green marked with purple, flowers 
enclosed by bracts. Flowers with hypanthium ca. 2 mm 
long, calyx 1-3 mm long and 3 mm broad; corolla fun- 
nelform, white, glabrous, 20-22 mm long, tube 14-19 
mm long and 2-4 mm diam., lobes 5(-6), 2.5-3 mm long 
1.5- 2 mm broad at the base, acute; stamens 5(-6), an- 
thers subsessile, ca. 2.8 mm long; style ca. 16 mm long. 
Fruits ca. 8 mm long and 8 mm diam., rounded-oblong, 
becoming blue or purple, 1 0-ribbed when dried; pyrenes 
with 4-6 ribs. 

Plants of evergreen forest formations of the Pa- 
cific slope, from around 50 to 500 m elevation in 
southern and south-central Costa Rica. Flowering 
in May-July; fruiting in late August and Novem- 
ber. The species is also known from Peru. 

Psychotria borucana is recognized by its large 
leaves, long-pedunculate purple heads, two broad 
inflorescence bracts with reticulate venation, long 
corolla tube with short lobes, unusual stipules, and 
restricted range (in Costa Rica). The leaves are 
quite variable in shape. Compare P. correae with 
pendant heads and more secondary veins. 



Psychotria brachiata Sw., Prodr. 45. 1788. Figure 
58. 

Shrubs or small trees, (0.5-)1.5-3(-5) m tall, leafy 
branchlets 1.2-5 mm thick, glabrous, quadrangular be- 
coming terete; stipules 3-7 mm long, 2-5 mm broad, 
rounded and bilobed with a sinus 0.5-2 mm deep and 
2 rounded lobes, glabrous and persisting. Leaves with 
petioles (1-) 1.5-3. 5 cm long, 1-1.5 mm thick, glabrous; 
leaf blades 9-17(-21) cm long, 3-7(-ll) cm broad, el- 
liptic to elliptic-oblong or elliptic-obovate, apex usually 
short-acuminate with tip 5-10(-l 5) mm long, base acute 
to attenuate (obtuse), drying thin-chartaceous or char- 
taceous and greenish or brownish, glabrous above, gla- 
brous beneath except for a line of short (0.5 mm) hairs 
along sides of midvein, 2 veins 7-10/side. Inflorescences 
terminal and solitary or 3 (4, 5), 10-20 cm long, 3-10 
cm broad, open pyramidal panicles with opposite sep- 
arated (6-30 mm) lateral branches mostly arising at 90 
angles, peduncles 1.5-6 cm long, with thin hairs ca. 0.4 
mm long (mostly in 2 longitudinal rows), often with 3 
2 branches bearing the distal flower clusters, bracts 3- 
8 mm long and 1.5-3 mm broad, lanceolate, floral bracts 



236 



FIELDIANA: BOTANY 



2-3 mm long, broadly ovate, ciliolate along the margins, 
pedicels 0-2 mm long, flowers borne in capitulae 5-10 
mm broad. Flowers with hypanthium 1-1.5 mm long, 
calyx 1-1.5 mm long, yellowish, calyx truncate or slightly 
dentate; corolla yellow or white (often with bluish tips 
and yellow throat), funnelform, tube 3-5 mm long and 
0.7-2 mm diam., usually glabrous externally on the low- 
er half, lobes 5, 1-1.5 mm long. Fruits 4-5 mm long 
(including calyx) and 3-6 mm diam., obovoid to ellip- 
soid but becoming subglobose and spongy in life (to 10 
mm diam.), bright blue to dark blue or purple, with 
prominent longitudinal ribs, persisting calyx ca. 1 mm 
long; pyrenes 3-4 mm long, with 3-5 ribs. 

Plants of secondary vegetation in evergreen 
tropical and premontane wet forest formations on 
the Caribbean slope and in the General Valley, 
from 5 to 800 m elevation in Costa Rica. This 
species appears to flower and fruit throughout the 
year in Central America. At La Selva flowering is 
primarily in May-July and fruiting is mostly in 
July-September. The species ranges from the West 
Indies and Mexico through Central America to 
Peru. 

Psychotria brachiata is recognized by its open 
inflorescences with well separated short opposite 
branches bearing closely clustered yellowish flow- 
ers in distal groups subtended by prominent per- 
sisting bracts. The small stature, nearly glabrous 
leaves usually drying greenish, spongy bright blue 
fruit with longitudinal ribs, and persisting stipules 
are additional features. Immature inflorescences 
of P. brachiata are racemose with the congested 
flowers in unexpanded capitula at the ends of the 
opposite lateral branches. 

Steyermark (1972, pp. 585-586) distinguishes 
P. caerulea Ruiz & Pavon from P. brachiata by 
its larger corollas (11-12 mm long), external co- 
rolla surface that is densely puberulent through- 
out, more prominent calyx lobes, larger clusters 
of flowers, and larger floral bracts forming a more 
definite involucre. Specimens that fit Steyermark's 
definition of P. caerulea appear to occur in Bocas 
del Toro, Panama (D. Simpson in herb. 1977), 
and it is possible that they also occur in nearby 
Costa Rica. The question of whether or not these 
two species intergrade and are conspecific might 
make a worthwhile study. 



Psychotria brachybotrya Muell.-Arg. in Mart., Fl. 
Bras. 6(5): 327. 1881. P. iquitosensis Standl., 
Publ. Field Columb. Mus., Bot. Ser. 8: 195. 1930. 
Figure 56. 

Shrubs or subshrubs, l-2(-3) m tall, leafy branchlets 
0.8-4 mm thick, glabrous; stipules separate or with a 



short (0.5-1 mm) sheath, lobes 2/side, 2-7 mm long, 
0.5-1 mm broad at base, linear to narrowly triangular, 
usually separated by a U-shaped sinus, glabrous, per- 
sisting. Leaves with petioles 3-10 mm long (poorly dif- 
ferentiated from the decurrent leaf base), glabrous; leaf 
blades 9-16(-18) cm long, 3-7(-8) cm broad, ovate-el- 
liptic to ovate or elliptic, apex usually tapering gradually 
and short-acuminate or acute, tip 5-10 mm long, taper- 
ing gradually or abruptly to the attenuate base (acute to 
broadly obtuse above the narrowed base), drying thinly 
chartaceous and greenish, glabrous above and below, 2 
veins 4-8/side and arcuate ascending, distal 3 veins of- 
ten subparallel. Inflorescences terminal and solitary, 1 .2- 
3 cm long but enlarging in fruit, 1-2 cm wide, narrowly 
paniculate with short opposite lateral branches or some- 
times subcapitate, peduncle 3-18 mm long, ca. 1 mm 
thick, with thin whitish hairs to 0.5 mm long, 2 branches 
0-5 mm long, cymes congested, usually subtended by 3 
involucrate narrowly ovate to lanceolate bracts, median 
bracts 3-6 mm long and 2-3 mm broad, with 2 shorter 
lateral bracts, all 3 united at base, flowers sessile. Flowers 
with hypanthium ca. 0.5 mm long, puberulent, calyx 
lobes 5, 0.2-0.3 mm long; corolla white, salverform, tube 
1.5-4 mm long, ca. 1 mm diam. distally, glabrous or 
puberulent, lobes 5, 0.8-1.5 mm long. Fruits ca. 4 mm 
long and 4 mm diam., globose to oblong, becoming pur- 
ple-black at maturity (blue-black on Cocos Island), with 
prominent longitudinal ribs (transverse ridges some- 
times visible when dried), calyx minute (0.3 mm high) 
or obscure; pyrenes 3-4 mm long, with 4-5 prominent 
ridges. 

Plants of the lowland rain forest formations on 
both the Caribbean and Pacific slopes, from near 
sea level to 400(-900) m elevation and usually 
found in shaded, poorly drained sites. Flowering 
in July-August; fruiting in August-October and 
December. In addition to the mainland plants, a 
distinctive population of this species is found on 
Cocos Island (see below). The species ranges from 
Costa Rica and Panama to Brazil and Bolivia. 

Psychotria brachybotrya is distinguished by its 
well-separated stipule lobes, glabrous leaves dry- 
ing green, small compact inflorescences, flowers 
subtended by an involucre of bracts, small white 
flowers, and purplish black fruit. Inflorescences 
change in shape as they grow and expand. Collec- 
tions from Cocos Island placed under this name 
differ from the mainland material in having small- 
er leaves with blades (4-)6-9 cm long and 1.5-3.5 
cm broad; the stipules, flowers, and fruit are very 
similar. Compare this species to P. hoffmannseg- 
giana, P. platypoda, and P. officinalis. Hammel (in 
Taylor, 1991) noted that P. brachybotrya is usually 
a larger plant than P. officinalis and the latter has 
a linear ventral sulcus on the pyrene. 



Psychotria ca loch lam ys Standl., Publ. Field Co- 
lumb. Mus., Bot. Ser. 8: 199. 1930. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



237 



Small shrubs or treelets, 1.5-4.5 m tall, leafy stems 
2.5-4.5 mm thick, glabrous or sparsely and minutely 
puberulent; stipules 1 3-28 mm long and 5-9 mm broad 
at the base, ovate-triangular with 2 attenuate-acute lobes 
6-10 mm long, greenish to wine red or violet, with fine 
parallel venation, chartaceous. Leaves with petioles 3- 
16(-25) mm long, 0.9-2 mm thick, glabrous; leaf blades 
8-23 cm long, 2-8.5 cm broad, elliptic-ovate to oblong- 
lanceolate or lanceolate, apex gradually tapering and acu- 
minate, tip 5-10 mm long, base acute to obtuse, drying 
chartaceous and greenish, glabrous above and below, 2 
veins 1 1-17/side, often with lesser 2 veins between the 
major. Inflorescences solitary and terminal, 12-14 cm 
long and 6-7 cm broad, hemispheric panicles with 2- 
3(-4) branches at the first and second nodes, peduncles 
3-8 cm long and 1.5-2 mm thick, white, minutely (0.1 
mm) puberulent, bracts subtending the first whorl of 
branches 1 .5-2 cm long, 3-4 mm wide, narrowly elliptic, 
resembling the stipules in texture, distal bracteoles 5-14 
mm long, 2-4 mm broad, lanceolate, persisting, white 
to violet, enclosing the sessile flowers. Flowers ca. 15 
mm long, hypanthium 0.6-1 mm long, calyx 0.6-0.8 mm 
long, lobes 5, triangular; corolla white to purple or pale 
violet, yellow in throat, tubular-salverform, tube 7-10 
mm long, lobes 5, 2-3 mm long; anthers ca. 1.5 mm 
long and curved. Fruits ca. 6 mm long and 4 mm diam., 
oblong, dark purple or black; pyrenes with 4-5 ribs. 



In Costa Rica this disjunct species is only known 
from the hills near Golfito, Puntarenas, at about 
1 00 m elevation, flowering in December-January. 
The species also occurs in Choco, Colombia, and 
Amazonian Peru. 

Psychotria calochlamys is distinguished by its 
large deciduous stipules, long and persisting white 
or lilac bracts and bracteoles, and large flowers. 
The unusual form and size of the stipules is similar 
in P. capitata, a close relative. The inflorescence 
with large white (or purplish) bracts and flowers 
is unlike any other Costa Rican species of Psy- 
chotria and reminiscent of some species of Pali- 
courea. The disjunct occurrence in Costa Rica is 
similar to that of some other species in the Golfo 
Dulce region. This species may prove to be con- 
specific with P. stipulosa Muell. Arg. of Amazo- 
nian South America, but that species (interpreted 
in a broad sense) includes variation not seen in 
our material. 



The leaves drying grayish and the bright red fruit 
are characteristics of the subgenus Psychotria. This 
species has a disjunct distribution and is not known 
from Costa Rica. It is found in southern Mexico 
and Guatemala and again in Colon and Code 
provinces in Panama. 



Psychotria camponutans (Dwyer & Hayden) Ham- 
mel, Selbyana 12: 139. 1991. Cephaelis cam- 
ponutans Dwyer & Hayden, Ann. Missouri Bot. 
Gard. 55: 35. 1968. 

Semisucculent herbs or subshrubs 0.5-1.2 m tall, rhi- 
zomatous, stems unbranched, leafy stems 24 mm thick, 
quadrangular or terete, glabrous, drying black and con- 
tracted below the nodes; stipules 6-8(-15) mm long, 5- 
6(-10) mm broad, broadly ovate with a serrated margin 
and 2 long lobes, glabrous and thin, distal parts decid- 
uous but with a persisting truncated sheath 1-1.5 mm 
long. Leaves with petioles 18-55 mm long, 1.2-2 mm 
thick, glabrous, drying black; leaf blades (1 1-)18-25 cm 
long, (4-)7-9 cm broad, narrowly elliptic-oblong to ob- 
long or oblong-obovate, apex acuminate with tip 6-13 
mm long, acute at base and slightly decurrent on petiole, 
drying thinly chartaceous, dark or grayish above, gla- 
brous above and below, 2 veins 8-15/side but these 
difficult to see. Inflorescences axillary and I/node, 1-2 
cm long, to 2 cm diam., capitate, peduncle 4-10 mm 
long, with involucrate bracts 8-15 mm long, 5 mm broad, 
ovate and acuminate, glabrous and green, flowers sessile 
and congested, each enclosed by 2 lanceolate bracteoles 
6-8 mm long. Flowers glabrous, calyx lobes 5-6, ca. 1 
mm long; corolla white, funnelform, tube ca. 3 mm long, 
corolla lobes 1-1.5 mm long. Fruits 7-8 mm long, 4-5 
mm diam., ellipsoid to obovoid, crowned by a persisting 
calyx, deep maroon red or pale rose red, bracts red to 
purple; pyrenes 6-7 mm long, ribbed. 

Plants of the wet evergreen forests of the Carib- 
bean slope, from 300 to 900 m elevation. Flow- 
ering in April and July; immature fruit were col- 
lected in September. Costa Rica to central Panama. 

Psychotria camponutans is distinguished by its 
short semisucculent erect stems with small axillary 
capitate inflorescences, reddish fruit, and restric- 
tion to the Caribbean slope. Compare P. uliginosa 
and P. cartagoensis. 



Psychotria calophylla Standl., Contr. U.S. Natl. 
Herb. 18: 129. 1916. 

Small trees, 3-18 m tall, with larger (13-26 x 
5-12 cm) elliptic to obovate leaves usually reddish 
tomentulose beneath, with an arcuate submarginal 
vein; the inflorescences paniculate with usually 3 
ranks of branching, the flowers in distal glomer- 
ules, and larger (10-14 x 6-9 mm) ellipsoid fruit. 



Psychotria capacifolia Dwyer, Ann. Missouri. Bot. 
Gard. 67: 353. 1980. Figure 54. 

Herbs or subshrubs, 0.4-1 (-2?) m tall, usually suc- 
culent and unbranched, leafy stems 3-12 mm thick, gla- 
brous or with hairs to 1 mm long at nodes, terete, drying 
dark; stipules 3-6 mm long, to 8 mm broad, broadly 
obtuse to truncate and entire, with a conical caducous 
lateral appendage to 3 mm long, densely hirtellous. Leaves 
densely hirtellous in early stages but usually glabrescent 



238 



FIELDIANA: BOTANY 



in age, petioles 3-10 cm long, 2-4 mm thick, with hairs 
ca. 1 mm long in early stages; leaf blades 2040 cm long, 
9-1 6(-20) cm broad, oblong to elliptic-oblong or elliptic- 
obovate, apex abruptly narrowed and short-acuminate, 
base cuneate to obtuse, drying thinly chartaceous and 
grayish green, usually glabrous above, glabrescent or with 
thin hairs to 1 .4 mm long beneath, 2 veins 1 7-22/side, 
connected by a distinct slightly arcuate submarginal vein 
2-3 mm from the margin. Inflorescences axillary, I/node, 
(8-) 15-3 5 cm long, 5-12 cm wide, open panicles with 
opposite branching, peduncles 7-22 cm long, 1.5-2.2 
thick, with thin yellowish to reddish brown hairs 0.2- 
0.7 mm long, bracts 1-2 mm long, flowers sessile on 
distal dichotomous branches or distal clusters, bracteoles 
ca. 0.5 mm long. Flowers puberulent externally, hypan- 
thium ca. 0.7 mm long, calyx to 1 mm long, calyx lobes 
5, 0.3-0.5 mm long, triangular; corolla white or pale 
yellow, funnelform, tube 2.5-4 mm long, 0.5-1 mm diam. 
basally and 2 mm distally, lobes 5, ca. 1 mm long. Fruits 
5-6 mm long, 3-4 mm broad, ovoid, yellow-green or 
whitish, with ca. 10 longitudinal ribs; pyrenes ca. 5 mm 
long. 

Plants of evergreen rain forest formations on the 
Caribbean slope, ranging from 5 to 1300 m ele- 
vation. Flowering in January-September; fruiting 
in March-November. The species ranges from 
southern Nicaragua to western Panama. 

Psychotria capacifolia is recognized by its short 
succulent stems, pubescence on younger parts, large 
leaves, axillary often large puberulent inflores- 
cences, well-developed (only slightly arcuate) sub- 
marginal vein, and the yellowish green fruit. The 
corolla interior is said to have a "ball" of hairs 
within (Dwyer, 1980, p. 354). This species closely 
resembles some material placed under P. macro- 
phylla, P. siggersiana, and P. aggregate, but the 
differences used in the keys appear to reflect a real 
discontinuity of gene flow. 



Psychotria capitata Ruiz & Pav., Fl. Peruv. 2: 59, 
pi. 206. 1799. 

Shrubs or small treelets l-3(-4) m tall, leafy stems 2- 
4 mm thick, glabrous; stipules 8-18 mm long, 4-7 mm 
broad, ovate-lanceolate, acute at apex, with 2 triangular 
lobes 1-10 mm long, glabrous, deciduous. Leaves with 
petioles 4-20 mm long, 1-2 mm thick, glabrous, often 
drying yellowish green; leaf blades 6-16(-22) cm long, 
2.5-6(-8) cm broad, elliptic-oblong to elliptic, ovate- 
elliptic or elliptic-lanceolate, apex short-acuminate with 
tip 5-12 mm long, base acute or obtuse and slightly 
decurrent on petiole, drying stiffly chartaceous, greenish, 
glabrous above and below (except for short hairs along 
the sides of the midvein beneath), 2 veins 9-14/side, 
arising at almost 90 from midvein. Inflorescences ter- 
minal and solitary, 5-1 3 cm long, 3-6 cm broad, an open 
racemose or congested panicle, white at anthesis, pe- 
duncles (l-)2.5-7 cm long, ca. 1.5 mm thick, glabrous 
(rarely puberulent), branches opposite or ternate (alter- 



nate) and usually without subtending bracts, flowers ses- 
sile in distal cymose groupings of 2-4 subtended by white 
lanceolate bracteoles 4-9 mm long and 1-2 mm broad. 
Flowers glabrous externally, hypanthium ca. 1 mm long, 
calyx ca. 0.5 mm long, calyx lobes 4-5, ca. 0.3 mm long, 
triangular; corolla white or cream (yellow in throat), tube 
3-7 mm long, 1 mm diam. near base, lobes 4-5, 1.5-4 
mm long; stamens 4 or 5 anthers 1-2 mm long. Fruits 
5-7 mm long, ca. 5 mm diam., subglobose, dark blue- 
black to purple-black or black; pyrenes ribbed. 



Plants of lowland evergreen forest formations, 
0-200 m elevation. Flowering and fruiting 
throughout the year (mostly in December-July). 
The species ranges from Belize (Croat, 1978), the 
Caribbean lowlands of Nicaragua and Panama to 
Peru and Brazil; it is not known from Costa Rica. 

Psychotria capitata is recognized by its white 
flowers and inflorescences, conspicuous bracts on 
the distal parts of the inflorescences, leaves usually 
glabrous and drying greenish, and the large bilobed 
stipules. This species is similar to P. calochlamys, 
which has larger bracts and stipules and less prom- 
inent secondary venation. Compare also Psycho- 
tria officinalis and Palicourea tilaranensis. This 
species appears to be less variable in Central 
America than in South America (cf. Steyermark, 
1974). 



Psychotria cartagoensis Nepokroeff, nom. nov. 
Cephaelis latistipula Standl., J. Wash. Acad. 
Sci.18: 281. 1928, not Psychotria latistipula 
Benth. Figure 13. 



Herbaceous subshrubs, 0.3-1 m tall, stems erect and 
usually unbranched, leafy stems 2-6 mm thick, glabrous, 
quadrangular or terete; stipules (5-) 1 2-22 mm long, 1 2- 
18 mm broad, ovate in general outline, bilobed with a 
narrow sinus 2-7 mm deep, glabrous, stiffand persisting. 
Leaves well separated along the stem, petioles 2-6(-7) 
cm long, 1.2-3.7 mm thick, glabrous, drying dark; leaf 
blades 13-27 cm long, 5-9 cm broad, elliptic-oblong, 
oblong-lanceolate, elliptic-obovate, or ovate-elliptic, apex 
abruptly narrowed or rounded and shortly acute or short- 
acuminate, base acute to cuneate, drying stiffly charta- 
ceous, dark gray-green above, much paler beneath, 2 
veins 9-13/side, arising at 60-80 and arcuate distally. 
Infloresences axillary and sessile or subsessile, broadly 
capitate, 1-2 cm long and 24 cm broad, becoming dark 
magenta or purplish, bracts 7-8 mm long and 2-4 mm 
broad, innermost bracteoles lanceolate, flowers enclosed 
by many bracteoles. Flowers glabrous externally, hypan- 
thium ca. 2 mm long, turbinate, calyx cup 0.3-1 mm 
long, calyx lobes 4-5(-6), 1-4 mm long, lanceolate; co- 
rolla 4-5 mm long, funnelform, white, lobes 5, 1.2-3 
mm long, acute; anthers 2.5 mm long. Fruits ca. 6 mm 
long and 4 mm broad (not including the 2 mm long 
persisting calyx), reddish purple. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



239 



Understory plants of evergreen wet forests of 
the Caribbean slope, from 800 to 1 500 m eleva- 
tion. Flowering in March (Standley 39695 us the 
type of C. latistipuld) and May-June; fruiting in 
June and November. The species has only been 
collected in the valley of Rio Sarapiqui (Heredia 
Province) and near Orosi and Mufieco (Cartago) 
in central Costa Rica. 

Psychotria cartagoensis is recognized by its short 
unbranched habit, broad bilobed stipules, axillary 
and subsessile heads, and absence of pubescence. 
Among our species, Hoffmannia congesta is most 
likely to be confused with this species, but that 
species lacks the involucrate bracts and has many- 
seeded fruits. This species and its allies have been 
studied by Molly Nepokroeff (wis, 1992); P. \vil- 
buriana Dwyer and P. dukei Dwyer of Panama are 
closely related. 



Psychotria carthagenensis Jacq., Enum. PL Carib. 
16. 1760. Figure 65. 

Shrubs, subshrubs, or small treelets, 0.3-3(-4) m tall, 
leafy branches 1.5-4 mm thick, glabrous and terete, be- 
coming gray; stipules 3-8 mm long, 1.5-5 mm broad, 
ovate to oblong or slightly obovate, apex rounded or 
bluntly acute, glabrous, drying dark reddish brown, ca- 
ducous. Leaves with petioles 3-10 mm long, ca. 1 mm 
thick, with decurrent lamina base; leaf blades (4-)6- 
13(-15) cm long, (1.7-)2-5(-7) cm broad, oblanceolate 
to narrowly elliptic-obovate or narrowly elliptic, apex 
obtuse to very shortly acuminate (tip ca. 4 mm long), 
base gradually narrowed and cuneate or acute, decurrent 
on petiole, drying chartaceous, gray or reddish gray, gla- 
brous on both surfaces, 2 veins 6-8/side, domatia rarely 
present in distal vein axils. Inflorescences terminal and 
solitary, 3-8(-10) cm long, 2.5-4 cm broad, a compact 
panicle with usually 3 nodes of opposite branches di- 
minishing in size toward apex (also verticillate or um- 
bellate), peduncle to 4.5(-6) cm long, 0.6-1 .3 mm thick, 
glabrous, bracts subtending the lateral branches 0.5-1.5 
mm long, subulate to broadly ovate, flowers in distal 
cymes, pedicels 0-2 mm long, bracteoles to 0.5 mm long. 
Flowers glabrous or minutely (0.05 mm) papillate-pu- 
berulent externally, hypanthium ca. 1 mm long, calyx 
ca. 0.5 mm long, subentire or with 5 short triangular 
lobes; corolla white, funnelform-salverfbrm, tube 2.5-3 
mm long, ca. 1 mm diam. near base, lobes 5, 1.2-2 mm 
long; anthers 1-1.5 mm long. Fruits ca. 5 mm long and 
4 mm diam., oblong or ellipsoid, becoming red or or- 
ange, with 10 vertical ribs, glabrous; pyrenes with 2 sulci 
on the inner face and 5 ribs on the exterior surface. 



Plants of shaded sites in seasonally dry decid- 
uous forest formations of Guanacaste and north- 
ernmost Alajuela Province, from 5 to 200(-600) 
m elevation (rarely to 1400 m elsewhere). Flow- 
ering in November-August (primarily February- 



July); fruiting throughout the year. The species 
ranges from Mexico and the West Indies to Bolivia 
and Argentina. 

Psychotria carthagenensis is recognized by its 
deciduous forest habitat, smaller oblanceolate or 
obovate leaves, lack of conspicuous pubescence, 
small flowers and inflorescences drying reddish, 
and inflorescences often with nodes bearing two 
larger and two smaller lateral branches. The ten- 
dency of the leaves to dry gray or reddish and the 
red or orange fruit are characteristics of members 
of subgenus Psychotria. The correct spelling of this 
species is carthagenensis (not carthaginensis}. This 
species is similar to P. tenuifolia of similar habitats 
but with bilobed stipules. 



Psychotria chagrensis Standl., J. Wash. Acad. Sci. 
15: 105. 1925. Figure 60. 

Shrubs, 1-3 m tall, many-branched and often flat- 
topped, leafy branchlets 1-3 mm thick, glabrous, reddish 
brown or dark; stipules 3-10 mm long, 1 -3 mm broad 
near base (broader when subtending the inflorescence), 
narrowly tubular to narrowly ovate and acute at apex 
with 1 or 2 slender awns 1-3 mm long, glabrous and 
drying dark reddish brown, caducous. Leaves usually 
closely clustered distally, petioles 1-12(-17) mm long, 
ca. 0.7 mm thick, glabrous; leaf blades 2.5-8(-10) cm 
long, 0.9-3(-3.5) cm broad, obovate to oblanceo- 
late, elliptic-obovate or elliptic, apex acuminate or cau- 
date-acuminate to bluntly obtuse, tip ca. 5 mm long, 
base cuneate to acute and decurrent on petiole, drying 
stiffly chartaceous, gray or reddish brown, glabrous above 
and below, 2 veins 6-9/side (but sometimes difficult to 
see), joined by an arcuate submarginal vein near margin. 
Inflorescences terminal (rarely pseudoaxillary), 6-16 mm 
long, to 7 mm broad, sessile, subtended by broad stipule- 
like dark reddish brown glabrous bracts to 1 cm long, 
fasciculate or capitate, with few sessile flowers, bracteoles 
4-5 mm long. Flowers distylous, glabrous externally, hy- 
panthium ca. 1 mm long, calyx 2-4 mm long, lobes 1- 
2 mm long, narrowly triangular, corolla funnelform, white, 
tube 49 mm long and 1-1.3 m diam., lobes 5, 2-3 mm 
long; stamens 5, anthers ca. 1 m long. Fruits becoming 
8 mm long (not including persisting calyx), 3-6 mm 
diam., oblong-ellipsoid to ovoid, persisting calyx 2-3 
mm long, becoming red or purple pyrenes ca. 6 mm long, 
weakly ribbed. 



Understory plants often found in low wet de- 
pressions in evergreen rain forest formations, from 
near sea level to 700 m elevation on the Caribbean 
slope and Osa Peninsula. Probably flowering and 
fruiting throughout the year (flowering mostly in 
March-August). At La Selva flowering is most 
common in May-June and fruiting in November- 
December. The species ranges from southern Nic- 
aragua to Colombia and Peru, but with disjunct 



240 



FIELDIANA: BOTANY 



populations in Veracruz, Mexico, and Izabal, 
Guatemala. 

Psychotria chagrensis is recognized by its short 
(often flat-topped) habit, very small obovate leaves, 
distinctive stipules, small sessile fasciculate or cap- 
itate inflorescences, and unusual calyx lobes. The 
leaves drying grayish or reddish and the red fruit 
are characteristic of subgenus Psychotria. The 
slender secondary veins often have point inden- 
tations on the lower surface after the leaves have 
been dried. The very small leaves are unusual 
among the woody plants of the lowland rain forest 
floor; compare P. graciliflora, P. parvifolia, and 
Randia loniceroides. 



from near sea level to 700 m elevation. Flowering 
in May-September and January; fruiting in July- 
March. The species ranges from southern Mexico 
and Belize to central Panama. 

Psychotria chiapensis has the longest corolla 
tubes among Central American Psychotria species; 
it is pollinated by long-tongued sphingid moths 
(cf. Bawa & Beach, 1983). The terminal subcapi- 
tate inflorescences (often in groups of three) with 
broad bracts and the relatlively large four-edged 
fruit are also distinctive. The paucity of puberu- 
lence, and leaves usually broadest at or above the 
middle are additional characters that make this 
species stand apart. It is called cocobolito in Pan- 
ama. 



Psychotria chiapensis Standl., Contr. U.S. Natl. 
Herb. 23: 1390. 1926. Cephaelis tetragona J. D. 
Smith, Bot. Gaz. 61: 376. 1916, not Psychotria 
tetragona Seem., 1867. Figure 18. 



Psychotria chiriquiensis (Standl.) C. M. Taylor, 
comb. nov. Cephaelis chiriquiensis Standl., Ann. 
Missouri Bot. Gard. 28: 469. 1941. Figure 17. 



Shrubs, treelets, or small trees, 2.5-8(-10) m tall, much- 
branched with rounded crown, leafy stems 2-6(-8) mm 
thick, terete and often conspicuously contracted for 5- 
10 mm below the node after drying, glabrous or with 
short (0.2 mm) hairs in new growth, center with white 
spongy pith; stipules 2-5 mm long, 2 4(-8) mm broad, 
broadly triangular, rounded with a small apical sinus, 
with 2 triangular acute lobes to 2 mm long. Leaves with 
petioles 7-35 mm long, 0.8-2.3 mm thick, glabrous 
(sometimes puberulent near inflorescences); leaf blades 
9-20(-23) cm long, 3-8(-9) cm broad, elliptic-obovate 
to elliptic-oblong or narrowly oblong-obovate, apex short- 
acuminate or acute with tip 5-10 mm long, base usually 
gradually narrowed and cuneate base, slightly decurrent 
on petiole, drying thin-chartaceous and grayish green to 
dark brown above, distinctly paler beneath, glabrous 
above and below except for short (0.2 mm hairs) along 
sides of midvein beneath, 2 veins 9-12/side. Inflores- 
cences terminal, 1 (or 3 when basal lateral branches are 
axillary to distal leaves), 4-12 cm long, 5-8 cm broad, 
subcapitate or corymbiform with 3-7 dense bracteate 
flower clusters, peduncles l-8(-15) cm long, 1.3-3 mm 
thick, minutely puberulent with hairs 0.2 mm long, bracts 
6-10 mm long, 4-8 mm broad, ovate to ovate-elliptic, 
obtuse to rounded, minutely ciliolate along margins, with 
6-many flowers closely clustered and sessile. Flowers 
distylous, calyx resembling the bracts in texture and col- 
or, 3-5 mm long, calyx lobes 5, ca. 1.5 mm long, tri- 
angular; corolla salverform, white, glabrous externally in 
ours, tube 20-45 mm long, 1-3 mm diam. for most of 
its length, lobes 5, 9-15 mm long and 3 mm broad; 
anthers ca. 4 mm long. Fruits 12-16 mm long (not in- 
cluding the calyx) and 9-1 3 mm diam., ellipsoid to ovoid, 
purple-black, strongly 4-angled, persisting calyx 2-3 mm 
long; pyrenes 10-13 mm long, 7 mm broad and 3 mm 
thick, bony, with 3 dorsal ribs and concave areas be- 
tween. 

Plants of evergreen lowland rain forest forma- 
tions on both the Caribbean and Pacific slopes, 



Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla- 
brous; stipules 3-5 mm long, with a short (1 mm) tube 
and 4 rounded ovate lobes 2-4 mm long at each node, 
these separate on the internode but overlapping above 
petioles, persisting. Leaves well separated along stems, 
petioles 15-40(-57) mm long, 0.5-1.5 mm thick, gla- 
brous; leaf blades 8-16(-20) cm long, 3-7 (-9) cm broad, 
elliptic to elliptic-oblong, apex acuminate with tip 5-10 
mm long, base obtuse to acute, drying stiffly chartaceous, 
dark green to yellowish brown or grayish brown, glabrous 
above and beneath, 2 veins 1 1-22/side, loop-connected 
distally to form an arcuate submarginal vein. Inflores- 
cences terminal (pseudoaxillary), solitary or 3, 3-12 cm 
long, 3-8 cm broad, capitulum often with 3 short (2-5 
mm) primary branches (especially in fruit), involucral 
bracts 12-20 mm long, 20-50 mm broad, broadly ovate, 
purple to reddish violet, peduncle l-6(-8) cm long, 2- 
3.5 mm thick, glabrous, flowers subsessile within many 
imbricate bracteoles 8- 12 x 5-1 Omm, elliptic to oblong. 
Flowers glabrous externally, hypanthium ca. 1 mm long, 
0.5 mm diam., turbinate, calyx tube ca. 1 mm long, lobes 
ca. 0.5 mm long; corolla tubular-funnelform, purplish, 
tube 9-12 mm long and 1.7 mm diam., lobes 5-6, 1.5- 
2 mm long; anthers to 2 mm long. Fruits elliptic, blue, 
enclosed within the inflorescence; pyrenes with 4-5 ridg- 



Plants of lower montane rain forest formations, 
from 1 200 to 1 800 m elevation. Flowering in Jan- 
uary-September; fruiting in October-December. 
This species ranges from Volcan Tenorio to south- 
ward along the Caribbean slope and continental 
divide to the Chiriqui Highlands. 

Psychotria chiriquiensis is recognized by its in- 
volucrate capitate inflorescences with broad pur- 
ple bracts, long-petiolate leaves with many prom- 
inent secondary veins, lack of pubescence, and 
rounded stipule lobes usually overlapping above 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



241 



the petiole. This species is named in honor of An- 
tonio Molina R., who monographed the Mexican 
and Central American species ofCephaelis in 1953 
and has made many important collections in Cen- 
tral America. This species is very similar to P. 
data, with larger red bracts, heads not three- 
branched, and white corollas. It is also similar to 
P. dichroa. 



Psychatria chiriquina Standl., Contr. U.S. Natl. 
Herb. 18: 129. 1916. Figure 61. 

Shrubs or small trees, 2-6 m tall, leafy stems 1-4 mm 
thick, glabrous, dark reddish brown; stipules 4-8(-10) 
mm long, 3-5 mm broad, ovate to elliptic, apex acute, 
ciliolate to erose or shortly bilobed, united around stem 
at base, glabrous or puberulent, caducous. Leaves with 
petioles 7-18(-35) mm long (sometimes variable on the 
same stem), 0.4-1.3 mm broad, glabrous; leaf blade 5- 
15 cm long, 2-6 cm broad, narrowly elliptic, elliptic- 
oblanceolate to oblanceolate or oblong-obovate, apex 
gradually tapering and acute or acuminate, tip ca. 10 
mm long, gradually narrowed to an acute base and de- 
current on petiole, drying stiffly chartaceous, dark red- 
dish brown above, glabrous above and below except for 
hairs along sides of midvein, occasional pit domatia in 
vein axils beneath, 2 veins 6-12/side. Inflorescences 
terminal, usually solitary or 3, 3-10(-l 7) cm long, a few- 
branched panicle, often trichotomous, peduncles 1.5-5 
cm long, 0.5-2 mm thick, essentially glabrous, pedicels 
0-3 mm long. Flowers glabrous externally, calyx ca. 1 
mm long, truncate or with 5 small (0.6 mm) lobes; corolla 
white, salverform or funnelform, tube 3-6 mm long, 1.5- 
2 mm diam., glabrous or puberulent externally, lobes 5, 
1.5-2 mm long; stamens 5, anthers ca. 1 mm long. Fruits 
5-8 mm long, 4-6 mm diam., oblong to subglobose, 
becoming red, persisting calyx 0.5-1 mm long; pyrenes 
with 45 rounded ribs. 



Psychotria chitariana Dwyer & C. Hamilton, Phy- 
tologia64: 221. 1988. 

Subshrubs, 30-100 cm tall, leafy stems 49 mm thick, 
glabrous, drying dark; stipules 15-18 mm long, ovate, 
with 2 awns ca. 3 mm long, caducous. Leaves with pet- 
ioles 0-27 mm long, ca. 3 mm thick, glabrous, with 
lateral margins and flattened above; leaf blades 25-40 
cm long, 10-17 cm broad, elliptic-obovate to obovate- 
oblong or oblanceolate, apex obtuse to acute, base grad- 
ually narrowed and cuneate or expanded and auriculate 
at the petiole, drying thinly chartaceous, grayish brown 
to reddish brown, glabrous above and below, 2 veins 
12-17/side, loop-connected 2-3 mm from the margin. 
Inflorescences terminal and solitary, dense panicles of 
congested cymes, globose or hemispheric, 3-1 1 cm long, 
3-5 cm broad, peduncles l-2.5(-^.5?) cm long, bracts 
ca. 5-6 x 1-3 mm, glabrous, flowers closely congested, 
pedicels 3-5 mm long, bracteoles to 0.5 mm long. Flow- 
ers glabrous externally, calyx ca. 1 mm long, cupulate, 
lobes 5, short; corolla funnelform, greenish white, tube 
4. 5-5 mm long, 1-1.5 mm diam., lobes 1.5-2. 3 mm long, 
triangular or oblong; anthers ca. 1.5 mm long. Fruits not 
seen. 

Known only from the Caribbean slope of the 
Cordillera de Talamanca in Costa Rica, from 200 
to 800 m elevation. Flowering in April-May; im- 
mature fruits were collected in September. The 
species ranges from the valley of Rio Chitaria east 
of Turrialba (Liesner et al. 15400 CR, MO holotype) 
to the valley of Rio Estrella. 

Psychotria chitariana is distinguished by its short 
stature, large obovate leaves with rounded and 
obtuse apex, and unusual inflorescence. The leaves 
drying dark grayish (or reddish brown) is a char- 
acteristic of subgenus Psychotria. Compare P. al- 
faroana with more open inflorescences and acute 
leaf bases. 



Plants of evergreen montane cloud forest for- 
mations, from (900-)1500 to 2500 m elevation. 
Flowering in January-May (July-August in Nic- 
aragua); fruiting in August-April (November-May 
in Nicaragua). The species is known from northern 
and central Nicaragua and from the easternmost 
part of the Cordillera de Talamanca in Costa Rica 
and the Chiriqui Highlands in Panama. 

Psychotria chiriquina is recognized by its higher 
elevation habitats, stipules often with two small 
distal lobes, narrow leaves tapering gradually to 
apex and base, pit domatia, small flowers, and 
poorly developed calyx lobes. The red fruit and 
leaves/branchlets drying dark reddish brown are 
characters of subgenus Psychotria. Compare P. 
sarapiquensis, with a shorter calyx, and P. sylvi- 
vaga. This species is also similar to P. panamensis 
and with very different stipules. 



Psychotria cincta Standl., Publ. Field Columb. 
Mus., Hot. Ser. 7: 90. 1930. Figure 58. 

Shrubs, l-2.5(-4) m tall, leafy branches 1.5-4 mm 
thick, glabrous and drying greenish or dark; stipules 
forming a small basal (1-3 mm) sheath, with 2 awns (4/ 
node), awns 6-14 mm long, ca. 0.5 mm broad, glabrous, 
persisting. Leaves with petioles 8-20 mm long, 1-1.5 
mm thick, glabrous; leaf blades 12. 5-21 (-28) cm long, 
3. 5-7 (-9) cm broad, elliptic-oblong to oblong or ovate- 
oblong, apex tapering gradually and acuminate, tip 8- 
18 mm long, base acute to obtuse, drying stiffly char- 
taceous, green or grayish green, glabrous above and be- 
low, 2 veins 9-1 1 (- 1 4)/side, leaf margin with a distinctly 
thickened vein along the edge. Inflorescences terminal 
and solitary, usually pendant, 12-20 cm long, 2-4 cm 
broad, racemiform thyrsoid panicles with short (1-2 cm) 
usually alternate lateral cymose branches, peduncles 6- 
14 cm long, 0.6-1.2 mm thick, sparsely and minutely 
puberulent with short (0.1-0.2 mm) thin hairs, bracts 



242 



FIELDIANA: BOTANY 



linear, adnate to lateral branches or absent, bracteole 0- 
0.5 mm long, pedicels 0-2 mm long. Flowers minutely 
puberulent externally, hypanthium 0.5 mm long, calyx 
0.2-0.4 mm long, truncate or with minute lobes; corolla 
funnelform or salverform, white or greenish white, tube 
2-3 mm long, ca. 0.5 mm diam., lobes 5, 1-1.5 mm long. 
Fruits 4-6 mm long, subglobose, obscurely ribbed, gla- 
brous, red becoming black; pyrenes shallowly ribbed. 



Plants of evergreen lowland Caribbean rain for- 
est formations, from near sea level to 500 m ele- 
vation. Flowering in February-July; fruiting in 
August. The species ranges from Nicaragua to Co- 
lombia. 

Psychotria cincta is recognized by its persisting 
stipules with long narrow awns, the often oblong 
leaves usually drying yellowish green, the long nar- 
row pendant inflorescences with unusually short 
lateral branches, and the small puberulent flowers. 
In addition, the thickened leaf margins are nota- 
ble. This distinctive species is presently known 
only from La Selva and nearby areas in Costa Rica. 
Compare P. deflexa, which lacks the thickened leaf 
margins and has pyrenes with transverse ribbing. 



Psychotria clivorum Standl. & Steyerm., Publ. Field 
Mus. Nat. Hist., Bot. Ser. 23: 87. 1944, based 
on P. limonensis var. angustifolia Standl., loc. 
cit. 17: 282. 1937. Figure 61. 



Shrubs or small trees, 1.5-6 m tall, leafy stems 2-4 
mm thick, glabrous; stipules 10-16 mm long, 2-5 mm 
broad (broader beneath the inflorescences), with a sheath 
1-4 mm long, ovate to elliptic, usually bilobed with 2 
acute lobes and a short (1-3 mm) sinus at apex, glabrous, 
dark reddish brown, caducous. Leaves variable in shape 
(on different plants), petioles 6-35 mm long, 1-1.7 mm 
thick, glabrous; leaf blades 10-22 cm long, 3.5-8 cm 
broad, narrowly elliptic-oblong to elliptic or oblanceo- 
late, apex acuminate, tip 7-15 mm long, base acute to 
cuneate and often long-decurrent on petiole, drying 
membranaceous to thinly chartaceous, grayish or gray 
tinted with red, glabrous above and below (midvein rare- 
ly sparsely puberulent beneath), 2 veins 8-12/side, loop 
connected 1-3 mm from margin, small pit domatia usu- 
ally present. Inflorescences terminal and solitary, 6-10 
cm long, 3-7 cm broad, open panicles, usually with 4 
lateral branches from 2 proximal nodes, peduncles 1-5 
cm long and 2.2 mm thick, minutely puberulent or gla- 
brous, reddish brown, bracts to 2(-3) mm long, pedicels 
0-2 mm long. Flowers glabrous externally, hypanthium 
ca. 0.6 mm long, conical, calyx tube ca. 0.5 mm long 
with minute (0.2 mm) broadly triangular lobes; corolla 
funnelform, white, tube 2-2.5 mm long, 1.2-1.5 mm 
diam., lobes 1-2 mm long; anthers ca. 1 mm long. Fruits 
5-6 mm long, 4-5 mm diam., ellipsoid or obovoid, lon- 
gitudinally ribbed, red; pyrenes with 3-5 longitudinal 
ridges. 



Plants of evergreen forest formations of the Ca- 
ribbean slope; from near sea level to 800 m ele- 
vation (to 1400 m in Guatemala). Most flowering 
collections have been made in March-May. A lit- 
tle-collected species ranging from Veracruz, Mex- 
ico, to central Panama. 

Psychotria clivorum is recognized by its larger 
narrow leaves gradually narrowed to the base with 
numerous pit domatia and distinct submarginal 
vein, stipules tubular at base and sometimes bi- 
lobed distally, inflorescences with whorled lateral 
branches at first two nodes (usually of two unequal 
pairs), small flowers, and reddish fruit. The ten- 
dency of the leaves to dry grayish and the stipules 
to dry reddish is characteristic of subgenus Psy- 
chotria. Specimens of this species may be difficult 
to separate from material of P. orosiana and P. 
sarapiquensis. 



Psychotria cocosensis C. Hamilton, Phytologia 64: 
222. 1988. Figure 62. 

Shrubs or small trees to 5 m tall, leafy stems ca. 2-7 
mm thick, glabrous; stipules (7-) 1 2-35 mm long, ( 1 .5-)2- 
5 mm broad, lanceolate, glabrous, drying dark reddish 
brown, caducous. Leaves with petioles 5-16(-23) mm 
long, 1-2.5 mm thick, glabrous; leaf blades (8-) 12-1 7 
cm long, (3-)5-7 cm broad, narrowly elliptic to elliptic- 
oblong or broadly elliptic-obovate, apex acuminate or 
caudate-acuminate with tip 4-12 mm long, base acute 
to attenuate and decurrent on petiole, drying subcoria- 
ceous, reddish brown to grayish brown, glabrous above 
and beneath, 2 veins 9-1 2/side, small pit domatia often 
present in the vein axils beneath. Inflorescences terminal 
or pseudoaxillary, solitary or 3-parted, 1.3-3 cm long, 
few-branched panicles with distal 3-flowered cymules, 
peduncles 0-7 mm long, ca. 1 mm thick, glabrous, bracts 
0.5-1.5 mm long, triangular, glabrous, pedicels 2-3 mm 
long. Flowers puberulent externally, hypanthium ca. 0.5 
mm long, glabrous, calyx tube ca. 1 mm long and 2 mm 
broad, cupulate, slightly denatate; corolla funnelform, 
white, tube ca. 3 mm long and 2 mm diam., lobes 5, ca. 
4 mm long, 2 mm wide; stamens 5, anthers ca. 2 mm 
long. Fruits ca. 6 mm long, 5-6 mm diam., ellipsoid to 
globose, orange then red and finally purple, drying dark 
reddish brown, persisting calyx 0.8-1.5 m long. 

Plants known only from Cocos Island, from near 
sea level to 50 m elevation. We have seen four 
collections: Foster 4132 F, Holdridge 5169 us, Pit- 
tier 12375 us (the holotype), and Pittier 16279 GH, 
us. Flowering in April-June; fruiting in January, 
April, and June. 

Psychotria cocosensis is similar to P. panamen- 
sis but differs in its subcoriaceous leaves and larger 
fruits with persisting cupulate calyx. The very small 
domatia, short inflorescences, and flowers with long 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



243 



corolla lobes are distinctive. The red fruit and ten- 
dency of the leaves to dry reddish brown are char- 
acteristics of subgenus Psychotria. The Holdridge 
collection is unusual in having broadly elliptic- 
obovate leaves that have dried grayish and sig- 
nificantly paler beneath. 



Psychotria cooperi Standl., Publ. Field Columb. 
Mus., Bot. Ser. 4: 296. 1929. Figure 13. 

Small trees or shrubs, 3-8 m tall, to 10 cm dbh, leafy 
stems 1 .4-5 thick, glabrous, terete or slightly tetragonal; 
stipules 5-9(-l 1) mm long, to 8 mm broad, with a broad 
tube, truncate to rounded or bilobed with a small ( 1 mm) 
sinus at apex, glabrous, persisting or deciduous. Leaves 
opposite or 3 at a node, petioles 7-25 mm long, 0.7-1.7 
mm thick, glabrous; leaf blades 8-21 cm long, 3.5-7 cm 
broad, elliptic-oblong to elliptic or elliptic-obovate, apex 
acuminate with tip 3-10 mm long, base gradually nar- 
rowed and acute or cuneate base, slightly decurrent on 
petiole, drying chartaceous, dark yellowish green to dark 
brown above, distinctly paler beneath, glabrous except 
for short (0.2-0.3 mm) stiff hairs along the sides of the 
midvein above and below (sometimes with thin hairs 
along the secondary veins beneath), 2 veins 7-1 I/side, 
arcuate-ascending near the margin. Inflorescences ter- 
minal or pseudoaxillary and becoming verticilate, sessile 
or subsessile, 1-3.5 cm long, to 3.5 cm broad, a compact 
globose irregularly branched panicle, peduncle 1-7 mm 
long, bracts to 10 mm long, bracteoles ca. 1 mm long 
and digitate-glandular, pedicels to 6 mm long in fruit. 
Flowers glabrous externally, distylous, hypanthium ca. 
0.7 mm long, calyx lobes 5, ca. 1 mm long; corolla fun- 
nelform, white, tube 4-5 mm long, 1.3 mm diam., lobes 
reflexed at anthesis, ca. 2 mm long. Fruits 4-5(-10?) mm 
long, 2.5-3.5 mm diam., obovoid or turbinate and trun- 
cated distally, dark blue or purple, persisting calyx ca. 1 
mm long; pyrenes 3-4 mm long, with 4-5 longitudinal 
ribs. 



Plants of lowland rain forest formations from 
20 to 300(-600) m elevation, often found in light 
gaps and stream sides. Flowering and fruiting 
throughout the year with most collections from 
January-March. The species ranges from north- 
eastern Costa Rica to Colombia. 

Psychotria cooperi is recognized by its small con- 
gested axillary inflorescences (sometimes appear- 
ing verticillate), the broad short-tubular stipules, 
leaves often drying yellowish green, small white 
flowers, and blue or purple obovoid fruit. The 
flowers are mostly sessile in terminal inflores- 
cences but become axillary as the stems continue 
to grow. These small treelets are called cocobolito 
in Bocas del Toro, Panama. Fruiting material may 
resemble Palicourea copensis. 



Psychotria correae (Dwyer & Hay den) C. M. Tay- 
lor, comb. nov. Cephaelis correae Dwyer & 
Hayden, Ann. Missouri Bot. Gard. 55: 36. 1 968. 
Figure 17. 

Shrubs or small trees, 2-3.5(-5) m tall, leafy stems 3- 
6 mm thick, glabrous, quadrangular; stipules with a very 
short (1-3 mm) sheath with 2 broadly rounded often 
overlapping lobes ca. 4 mm long and 4 mm broad sep- 
arated by a U-shaped sinus, glabrous. Leaves with pet- 
ioles 1 5-80 mm long (opposing petioles often unequal), 
1.2-2.5 mm thick, glabrous; leaf blades 1 1-28 cm long, 
5-13 cm broad, broadly elliptic-oblong, ovate-oblong, 
apex abruptly narrowed and short-acuminate with tip 
ca. 4 mm long, base broadly obtuse, drying chartaceous, 
green, glabrous above and below, 2 veins 13-17(-20)/ 
side. Inflorescences solitary and terminal or pseudoax- 
illary, capitulae 1-2 cm long, 2-3 cm diam., involucre 
5-10 cm wide, peduncles (3-)5-12(-22) cm long, ca. 2 
mm thick, glabrous, erect and becoming pendant, the 2 
large involucral bracts 2.5-5 cm long and 4-5 cm broad, 
ovate to reniform, rose red to magenta or purple, gla- 
brous, bracteoles 8-15 mm long, obovate. Flowers gla- 
brous externally, calyx lobes 5, ca. 1 mm long; corolla 
narrowly funnelform, white, tube to 10 mm long, cylin- 
drical, lobes 5, ca. 3 mm long; anthers ca. 3 mm long. 
Fruits 10-15 mm long, ca. 6 mm diam., obovoid-oblong, 
blue; pyrenes with 4-5 slightly elevated ridges 

Plants of evergreen cloud forest formations along 
the central Cordilleras at 900-1100 m elevation. 
Flowering in March-August; fruiting in October. 
This species is only known from the Cordilleras 
de Guanacaste and Tilaran and the Province of 
Code in Panama. 

Psychotria correae is recognized by the larger 
leaves with many secondary veins, the unusual 
stipules, the long-pedunculate inflorescences be- 
coming pendant in later stages, and the large pur- 
plish bracts subtending the capitulum. Specimens 
of this species can be confused with P. elata, which 
has erect capitulae with smaller involucral bracts 
and shorter peduncles, and to P. chiriquiensis, 
which lacks involucral bracts. This attractive spe- 
cies was named in honor of our respected Pana- 
manian colleague Mireya Correa. 



Psychotria deflexa DC., Prodr. 4: 510. 1830. P. 
patens auct. non Sw. fide Steyermark 1972. Fig- 
ure 56. 

Shrubs, (0.5-)l-3 m tall, leafy stems 0.5-5 mm thick, 
glabrous or sparsely puberulent in early stages; stipules 
united around stem for 0.5-2 mm and with 2 subulate 
or linear awns 3-8 long on each side, glabrous, persisting. 
Leaves with petioles 3-9(-12) mm long, 0.5-1.3 mm 
thick, glabrous; leaf blades 7-15(-18) cm long, (1.5-)2- 



244 



FIELDIANA: BOTANY 



5(-7) cm broad, narrowly ovate-elliptic to narrowly el- 
liptic, elliptic-lanceolate or rarely lanceolate, apex ta- 
pering gradually and acute or acuminate, tip 10-20 mm 
long, base broadly obtuse to acute, drying chartaceous, 
greenish or brown, glabrous above and below, 2 veins 
4-8/side. Inflorescences terminal (rarely pseudoaxillary) 
and solitary, 4-1 1 cm long and 2-4 cm broad, narrowly 
pyramidal or thyrsiform panicles with slender central 
rachis and short (5-14 mm) opposite or subopposite 
lateral branches (expanding somewhat in fruit), pedun- 
cles 2.5-5 cm long, 0.4-1.1 mm thick, glabrous or mi- 
nutely puberulent, lateral branches without subtending 
bracts, pedicels 1-2 mm long, bracteoles ca. 1 mm long. 
Flowers glabrous externally, hypanthium ca. 1 mm long, 
calyx ca. 0.5 mm long, dentate; corolla white, funnel- 
form, tube 1.6-3 mm long, 0.3-1 mm diam. (in Costa 
Rica), lobes 4-5, ca. 1 mm long; stamens 4, anthers ca. 
1 mm long. Fruits 2-3 mm long and 3-4 mm diam. (8- 
10 mm when aerenchymatous), oblate and somewhat 
bilobed, longitudinally ridged, becoming purple, violet, 
or blue (pulpy white within); pyrenes with 3-4 ridges and 
transverse ribs. 



sheath, with 2 short (1-2 mm) obtuse or rounded lobes 
separated by a U-shaped sinus, glabrous. Leaves with 
petioles 4-25 mm long, 0.7-1 .2 thick, glabrous; leaf blades 
4-1 1(-13) cm long, 1.8-4.3 cm broad, narrowly elliptic 
to lanceolate, apex tapering gradually and acuminate, tip 
5-10 mm long, base acute and slightly decurrent on pet- 
iole, drying subcoriaceous, dark brownish green above, 
glabrous above and below, 2 veins (6-)8-12/side. Inflo- 
rescences 1-3 and terminal, 3-10 cm long, 3-8 cm wide, 
usually subtended by a pair of smaller (15 mm) leaves, 
capitula ca. 1 5-25 mm long and equally broad (enlarging 
in fruit), subtended by 2 involucrate bracts 12-20 mm 
long and 8-15 mm broad, white becoming pinkish (dark 
purple in fruit), peduncles 3-9 cm long and 1.5-3.5 mm 
thick, glabrous, flowers sessile within bracteoles. Flowers 
glabrous externally, hypanthium ca. 1.5 mm long, calyx 
ca. 3 mm long, lobes ca. 1.5 mm long and triangular; 
corolla funnelform, white to pink, tube 5-10 mm long, 
ca. 2 mm diam., corolla lobes 2-3 mm long, bluntly 
acute. Fruits 8-9 mm long (including the 1 mm high 
calyx), 4 mm diam., narrowly obovoid, blue-black, dry- 
ing dark reddish brown; pyrenes usually smooth. 



Understory plants of wet evergreen or partly 
deciduous forest formations from ca. 500 to 1200 
m elevation on the Caribbean slope of the cordil- 
leras and near sea level on the Osa Peninsula. 
Flowering in June-August; fruiting in July-March. 
This species ranges from Mexico to Peru and Bo- 
livia. 

Psychotria deflexa is recognized by its persisting 
narrowly awned stipules, narrow leaves, general 
lack of pubescence (only the slender narrow inflo- 
rescence is minutely puberulent), small flowers, 
and purple or blue fruit with transverse ribs. Flow- 
ers in Costa Rican material appear to be much 
smaller than those described by Steyermark (1974, 
p. 1284). This species may resemble P. cincta and 
P. microbotrys. There are collections (Kernan & 
Phillips 661 & 789 CR) from Corcovado National 
Park that have unusually thin pedicels and inflo- 
rescences. However, these aberrant samples are 
bridged by other collections (Liesner 2842 & 3198 
CR, also from the Osa Peninsula) that appear to 
be intermediate with the more normal inflores- 
cences. These populations are worthy of further 
study. 



Psychotria dichroa (Standl.) C. M. Taylor, comb, 
nov. Evea dichroa Standl., Contr. U.S. Natl. 
Herb. 18: 124. 1916. Cephaelis dichroa (Standl.) 
Standl. Publ. Field Columb. Mus., Bot. Ser. 4: 
296. 1929. 

Small shrubs, 0.6-2 m tall, leafy stems 1.3-4 mm thick, 
glabrous; stipules 2.5-6 mm long, with a short (2-3 mm) 



Plants of evergreen montane forest formations, 
1 200-2600 m elevation. Flowering January-Sep- 
tember. This species is only known from the Chi- 
riqui Highlands of western Panama and the Ca- 
ribbean slope of the Cordillera de Talamanca. 

Psychotria dichroa is recognized by the leaves 
with prominent 2 veins, one to three terminal 
small involucrate heads, white flowers, and high- 
land habitat. In Costa Rica, it is rarely collected 
and appears to be restricted to the understory of 
high-elevation (2000-2500 m) Quercus forests. 
This species is similar to P. chiriquiensis with red- 
dish purple capitula. 



Psychotria domingensisJacq.,Enum. PI. 16. 1760. 
Psychotria pavetta Sw., Prodr. 45. 1788. Pali- 
courea domingensis (Jacq.) DC., Prodr. 4: 529. 
1 830. Palicourea pavetta (Sw.) DC., loc. cit. 525. 
1830. Psychotria mombachensis Standl., Publ. 
Field Columb. Mus., Bot. Ser. 8: 188. 1930. 

Shrubs, 1-2 m tall, leafy stems 2-5 mm thick, gla- 
brous, often with a shrunken area below the node after 
drying; stipules 2-5 mm long, with 2 triangular lobes 
separate almost to base by a broad (1-2 mm) U-shaped 
sinus, acute or awned, glabrous, deciduous or persisting. 
Leaves with petioles 5-20 mm long, 0.7-1.7 mm thick, 
glabrous, often drying yellowish; leaf blades 8-19 cm 
long, 3-7 cm broad, elliptic-oblong to narrowly elliptic- 
obovate, apex acuminate or gradually tapering and acute, 
base acute to obtuse and slightly decurrent on petiole, 
drying membranaceous to thin-chartaceous, yellowish 
green to greenish brown or brown, glabrous above and 
below, 2 veins 8-1 I/side. Inflorescences solitary or 3, 
terminal or pseudoaxillary, 4-9 cm long, 3-6 cm broad, 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



245 



a rounded open panicle with opposite branching, pe- 
duncle 3-20 mm long, ca. 1.5 mm thick and yellowish 
when dried, glabrous or minutely papillate-puberulent, 
bracts absent or 1-6 mm long and linear, flowers in distal 
cymules of 3, pedicels 0-3 mm long, ca. 1 mm thick, 
bracteoles minute or absent. Flowers glabrous or mi- 
nutely puberulent externally, hypanthium ca. 1.2 mm 
long, calyx tube ca. 0.5 mm long, lobes 0.2-1 mm long, 
narrowly acute; corolla white, tubular-salverform, tube 
ca. 12 mm long, 0.8-1.7 mm diam., lobes ca. 4-5 mm 
long and 2 mm broad. Fruits 4-5 mm long, ca. 6 mm 
diam., subglobose to ellipsoid, purple-black, with 4-5 
longitudinal costae and smaller transverse ribs when 
dried; pyrenes angled. 

Plants of evergreen or seasonally deciduous for- 
mations, from 20 to 900 m elevation. Flowering 
in April and June (the Nicaraguan type of P. mom- 
bachensis (Maxon et al. 7818 F) was flowering in 
July); a Costa Rican collection from near Tilaran 
(Standley & Valeria 45222 us) was fruiting in Jan- 
uary. The species is frequent in the West Indies 
and Guatemala-southern Mexico but is known 
from only a few collections in southern Central 
America. 

Psychotria domingensis is recognized by the in- 
florescences with thick branches and relatively few 
flowers, larger fruit with longitudinal and trans- 
verse ridges, thin glabrous leaves that often dry 
yellowish green, two-lobed stipules, and dried stems 
often conspicuously contracted beneath the node. 
The few collections from Costa Rica come from 
both evergreen and deciduous areas. This species 
is rather similar to P. eurycarpa of wet forests, but 
that species has stiffer leaves lustrous above, poor- 
ly developed calyx lobes, and larger fruit without 
transverse ribs. It is also similar to P. microdon 
and some species of Coussarea (see fig. 47). 



Psychotria dukei Dwyer, Ann. Missouri Bot. Gard. 
67: 371. 1980. 

Shrubs or herbaceous subshrubs, 0.5-1. 5(-2) m tall, 
leafy stems succulent in life, 1.8-7 mm thick (dried), 
glabrous; stipules 7-20 mm long, 8-1 8 mm broad, ovate- 
triangular, bilobed or fimbriate, yellowish and membra- 
naceous, lobes 1-4 mm long, deciduous. Leaves with 
petioles 1-5 cm long, 1-2.3 mm broad, glabrous; leaf 
blades (1 1-) 13-27 cm long, 4-1 1 cm broad, elliptic-ob- 
long to oblong or elliptic-ovate, apex short-acuminate 
with tip 5-10 mm long, base obtuse to cuneate and de- 
current on petiole, drying chartaceous, dark green or 
greenish brown, glabrous above, minutely (0. 1 mm) pu- 
berulent along the major veins beneath, 2 veins 10-18/ 
side. Inflorescences axillary, solitary at each node, 2.5- 
9 cm long, pyramidal, often with the flowers crowded in 
3 capitula on 3 primary branches 3-20 mm long (rarely 
with further secondary branches), peduncles 8-35(-60) 



mm long, ca. 1.2 mm thick, minutely puberulent or gla- 
brous, bracts 2-10 mm long, triangular to lanceolate, 
bracteoles 2-5 mm long, often forming a small involucre 
beneath the sessile flowers. Flowers glabrous externally, 
hypanthium ca. 1 mm long, calyx tube ca. 0.5 mm long, 
entire; corolla tubular-funnelform, white, tube 3-4 mm 
long, ca. 0.5 mm diam. basally, lobes ca. 1.5 mm long. 
Fruits 5-6 mm long, 3-4 mm diam., oblong, white; py- 
renes longitudinally ridged. 

Plants of lower montane rain forest formations 
from (800-)1 100 to 1800 m elevation. Flowering 
in March and June-August; fruiting in March- 
November. The species ranges from the Cordillera 
de Tilaran to Choco, Colombia. 

Psychotria dukei is recognized by its usually un- 
branched succulent stems, axillary inflorescences 
with flowers in capitate clusters, white fruit, and 
unusual stipules. This species is easily confused 
with P. aggregata or P. macrophylla, but the large 
thin stipules and inflorescence bracts are distinc- 
tive. This species is rarely collected in Costa Rica. 
Compare P. dukei with P. wilburiana Dwyer of 
Panama. 



Psychotria elata (Sw.) Hammel, Selbyana 1 2: 1 39. 
1991. Cephaelis elata Sw., Prodr. 45. 1788. C. 
punicea Vahl, Eclog. Amer. 1:19. 1796. C. cos- 
taricensis Schlechtend., Linnaea 28: 546. 1856. 
Evea elata (Sw.) Standl., Contr. U.S. Natl. Herb. 
18: 123. 1916. C. elata forma lutea Standl., Publ. 
Field Mus. Nat. Hist., Bot. Ser. 18: 1279. 1938. 
Figure 17. 

Shrubs, subshrubs, or small trees, 0.5-5(-8) m tall, 
leafy stems 1.3-5 mm thick, glabrous, terete or qua- 
drangular; stipules united around stem for 1-2 mm, with 
2 ovate lobes on each side, 2-5 mm long and ca. 3 mm 
broad, obtuse to broadly rounded and separated by a U- 
or V-shaped sinus, glabrous, persisting. Leaves with pet- 
ioles 4-22(-30) mm long, 1-2 mm wide, glabrous; leaf 
blades 6-25 cm long, 2.5-7(-8.5) cm broad, oblong to 
elliptic-oblong, oblong-obovate, or oblanceolate, apex 
acute to short-acuminate, tip 4-10 mm long, base cu- 
neate to acute or obtuse, drying thinly to stiffly charta- 
ceous, green, glabrous above and below, 2 veins (9-)l 3- 
20(-23)/side. Inflorescences terminal and solitary (rarely 
2-3), capitate and globose-hemispherical, capitula 1.5- 
4 cm long, involucre 2-7 cm broad, peduncles 2-13 cm 
long, 1.5-3 mm thick, erect or becoming pendant, gla- 
brous, the 2 involucrate basal bracts 15-55 mm long, 
1 5-45 mm broad, deep red to orange-red (yellow), ovate 
to reniform, rounded to acuminate, glabrous, bracteoles 
5-10 mm long, flowers sessile. Flowers glabrous exter- 
nally (rarely puberulent), calyx ca. 1 mm long, dentate; 
corolla white (rarely pink or yellow), narrowly funnel- 
form, tube ca. 16 mm long, 1.5-2.5 mm diam., lobes 5, 
2.5-4 mm long, narrowly triangular; stamens 5, anthers 
2-3 mm long. Fruits 5-10 mm long, 2-5 mm diam., 



246 



FIELDIANA: BOTANY 



ovoid, blue becoming black; pyrenes 4-7 mm long and 
3-4 mm broad, 5-ribbed. 

Common shrubs of evergreen forest edges and 
forest interiors, from 30 to 1700 m elevation. 
Flowering and fruiting throughout the year (pri- 
marily flowering in January-August). Fruiting at 
La Selva in July-September. The species ranges 
from Mexico and the West Indies, through Central 
America to Colombia. 

Psychotria elata is recognized by its glabrous 
parts, usually narrowly oblong leaves with many 
secondary veins, solitary long-pedunculate heads, 
large brilliant red (rarely yellow) bracts subtending 
the capitulum, and white corollas. The yellow- 
bracted form appears to be most common in the 
General Valley and Golfo Dulce area. This is one 
of Central America's most distinctive species of 
Rubiaceae, common in light gaps and on the edges 
of forests. The flowers are visited by butterflies 
and hummingbirds (Freeman & Stiles, 1990); 
breeding biology was studied by Bawa and Beach 
(1983). This is one of the species that made Ce- 
phaelis so distinctive a genus in Central America. 
Compare P. poeppigiana with conspicuous pubes- 
cence, P. chiriquiensis with purple capitula lacking 
the large involucral bracts, P. correae with longer 
peduncles and larger bracts, and P. dichroa with 
several smaller heads. 



Psychotria emetica L.f, Suppl. PI. 144. 1781. Fig- 
ure 13. 

Small herbs and subshrubs, 0.2-1 m tall, rhizomatous, 
stems usually unbranched, terete, leafy internodes 1-4 
mm thick, appressed-puberulent with stiff crooked 
brownish hairs 0.2-0.4 mm long; stipules 2-4 mm long, 
1-2 mm broad at base, narrowly triangular and ap- 
pressed-puberulent, drying dark, usually caducous. Leaves 
with petioles 3-12(-20) mm long, 1-2 mm broad, ap- 
pressed-puberulent with stiff brownish hairs; leaf blades 
(7-)8.5-13(-17) cm long, 2-5(-6.5) cm broad, elliptic to 
narrowly elliptic-oblong, elliptic-obovate or oblanceo- 
late, apex acute or short-acuminate, tip ca. 5 mm long, 
base cuneate to acute and slightly decurrent on petiole, 
drying thinly chartaceous, dark grayish green or dark 
grayish brown, glabrous above, sparsely appressed-pu- 
berulent with short (0. 1-0.3 mm) hairs beneath, 2 veins 
5-9/side. Inflorescences axillary, usually solitary in each 
axil (2/node), 1-2 cm long, condensed cymose or sub- 
capitate, with 3-10 flowers, peduncles 3-12 mm long, 
ca. 0.7 mm thick, puberulent, bracts 1-2 mm long, tri- 
angular-acute, pedicels 0-5 mm long. Flowers with hy- 
panthium ca. 1 mm long, conical and appressed-puber- 
ulent, calyx lobes 1-1.5 mm long, narrowly triangular; 
corolla white, funnelform, glabrous externally, tube 2-4 
mm long, 1-1.5 mm diam. at mouth, lobes 5, 1.5-2 mm 
long, acute. Fruits 8-10 mm long and 4-6 mm diam., 



oblong or ellipsoid, persisting calyx 1-1.5 mm long, be- 
coming blue; pyrenes ca. 5 mm long, smooth. 



Plants of shaded understory in lowland ever- 
green rain forest formations, from 10 to 300 m 
elevation. In Costa Rica and Panama flowering in 
February-August; fruiting mostly in June-No- 
vember. This species ranges from Guatemala to 
Bolivia. 

Psychotria emetica is recognized by its short, 
usually unbranched stems, small condensed axil- 
lary inflorescences, and bright blue fruit. These 
plants are occasionally cultivated and have been 
called raicilla and raicilla macho in Costa Rica, 
Nicaragua, and Panama. This species differs from 
the closely similar P. erecta by its smaller stature, 
puberulent stems, and floral details. The roots con- 
tain the drug ipecac but it is of inferior quality to 
that found in P. ipecacuanha. 



Psychotria erecta (Aubl.) Standl. & Steyerm., Publ. 
Field Mus. Nat. Hist., Bot. Ser. 23: 24. 1943. 
Ronabea erecta Aubl., Hist. pi. Guiane 1: 156. 
1775. Figure 13. 

Shrubs or treelets, (0.5-)l-3(-8) m tall, leafy stems 
1 .5-7 mm thick, sparsely appressed-puberulent with thin 
ascending hairs ca. 0.3 mm long, becoming glabrescent 
and drying dark; stipules with a persistent short (0.5 mm) 
base and single slender awn 2-6 mm long, ca. 0.5 mm 
wide, triangular to subulate, appressed-puberulent. Leaves 
with petioles 8-18(-35) mm long, 0.7-2(-2.5) mm thick, 
appressed-puberulent with hairs 0.2-0.3 mm long; leaf 
blades 8-20 cm long, 3-9 cm broad, elliptic-oblong to 
oblong or slightly obovate, apex abruptly narrowed and 
short-acuminate, tip 4-1 1 mm long, base obtuse to acute, 
drying chartaceous to subcoriaceous, dark olive green or 
brownish green, glabrous above, glabrous or sparsely pu- 
berulent beneath with thin appressed hairs, 2 veins 5- 
8/side. Inflorescences 1-3 in each axil (2-6/node), 1-3 
cm long, 5-10 mm broad, subcapitate condensed cymose 
with 3-7 closely crowded flowers, peduncles 0-8(-18) 
mm long, ca. 0.7 mm thick, appressed-puberulent with 
ascending yellowish hairs, flowers subsessile or sessile, 
bracteoles to 1 mm long. Flowers with hypanthium 1-2 
mm long, sparsely appressed puberulent near the base, 
calyx ca. 1 mm long, glabrous, lobes ca. 0.2 mm high; 
corolla white, tubular-salverform, glabrous, tube 3^ mm 
long, 1 mm diam., glabrous externally, lobes 5(-6), 1.5- 
3 mm long. Fruits 8-10 mm long, 5-8 mm diam., ellip- 
soid or oblong, glabrous, becoming blue-black; pyrenes 
5-7 mm long, smooth. 

Plants of evergreen rain forest formations, from 
near sea level to 700(-1000) m elevation on both 
the Caribbean and Pacific coasts. At La Selva this 
species is usually found on ridges near light gaps. 
Flowering in May-October; probably fruiting 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



247 



throughout the year. This species ranges from 
Mexico and the West Indies to Bolivia. 

Psychotria erecta is recognized by its axillary 
small subcapitate inflorescences with few small 
flowers, blue fruit, and oblong leaves often drying 
dark. This species is similar to P. emetica but grows 
to a larger size with broader and thicker leaves and 
glabrescent stems. Both species have axillary, not 
pseudoaxillary, inflorescences. Compare Appunia 
guatemalensis. 



Psychotria eurycarpa Standl., J. Wash. Acad. Sci. 
18: 275. 1928. Figure 47. 

Shrubs or small trees, 1.5-5(-7) m tall, leafy stems 1- 
3 mm thick, glabrous or rarely minutely papillate-pu- 
berulent; stipules 1-3 mm high and 3-4 mm broad, trun- 
cate to broadly rounded with a small (0.2 mm deep) 
sinus, persisting or deciduous. Leaves with petioles 12- 
28(-35) mm long, 0.7-1.5 mm thick, glabrous and often 
drying yellowish; leaf blades 6-14(-21) cm long, 2.7- 
7(-l 1) cm broad, ovate-elliptic to elliptic or broadly el- 
liptic, apex short-acuminate with a narrowed tip 3-10 
mm long, base obtuse to acute, drying stiffly chartaceous, 
yellowish green, glabrous and lustrous above, glabrous 
beneath, 2 veins 4-7/side. Inflorescences terminal and 
solitary, 5-1 1 cm long, 5-9 cm broad, broadly pyramidal 
panicles with opposite branching or umbellate, the first 
node often with 4 lateral branches, peduncles 1 5-55 mm 
long, 1-2 mm thick, yellowish, glabrous, bracts ca. 1 mm 
long, flowers sessile or subsessile in glomerules of 2-5. 
Flowers nocturnal, glabrous or very minutely papillate- 
puberulent externally, hypanthium ca. 1 mm long and 
0.6 mm diam., obconic, calyx tube 0.3-0.7 mm high 
with minute (0.2 mm) lobes; corolla narrowly funnel- 
form, white, tube 6-1 5 mm long and 0.8 mm diam. (1.7 
mm distally), lobes 5 (6, 7), 3-8 mm long, 0.5-1 mm 
broad, narrowly oblong. Fruits 8-20 mm long (to 25 mm 
when succulent), 7-1 5 mm diam., oblong to subglobose, 
blue-black or dark purple (often drying pale yellowish 
brown), longitudinal ridges present but not prominent 
when dried; pyrenes 12-17 mm long, obscurely 3-5- 
angled. 

Plants of lowland rain forest formations and 
lower montane cloud forest formations, from 20 
to 900(-1100) m elevation. Flowering in Febru- 
ary-May; fruiting in October-April and June. This 
species is only known from the Caribbean coastal 
plain, Cordillera de Guanacaste, Cordillera de Ti- 
laran, and the Caribbean slopes of the Central 
Highlands as far east as Rio Reventazon, and in 
western Panama. 

Psychotria eurycarpa is recognized by the usu- 
ally glabrous parts often drying yellowish green, 
the short stipules (truncated or with minute apical 
sinus), few-branched inflorescences with thick pe- 
duncles and thick lateral branches and inconspic- 
uous bracts, longer corolla tubes, and the large 



fruit. Breeding biology was studied by Bawa and 
Beach ( 1 983). A collection (Taylor 3546) from Las 
Cruces (San Vito) is tentatively placed here; it is 
the highest elevation ( 1 1 00 m) and easternmost 
collection in Costa Rica. Folsom 9188 DUKE, from 
La Selva with corolla tubes to 18 mm long and 
corolla lobes to 1 2 mm long, is provisionally placed 
here. Individuals of this species may be difficult 
to distinguish from P. domingensis (with bilobed 
stipules and flowers in distal triads) and Coussarea 
psychotrioides (q.v.). 

Psychotria fruticetorum Standl., J. Arnold Arbor. 
11:42. 1930. Figure 60. 

Shrubs, subshrubs, or small treelets, 0.3-3(-5) m tall, 
leafy stems 0.7-3.5 mm thick, glabrous; stipules 24 mm 
long, 1-3.5 mm broad at base, triangular to ovate, trun- 
cate to acute, with 2 awns 1-1.5 mm long, glabrous or 
minutely (0.05 mm) papillate, drying reddish brown, 
caducous. Leaves with petioles 0-6(-8) mm long, 0.7- 
1.2 mm thick, glabrous; leaf blades 3-8.5(-l 1?) cm long, 
1 4(-5?) cm broad, elliptic to obovate or oblanceolate, 
apex acute to subacuminate, base cuneate to acute, dry- 
ing stiffly chartaceous to subcoriaceous, grayish to dark 
reddish brown, glabrous above and below, 2 veins 4 
7/side, often with minute pit domatia or tufts of hairs 
in the axils of 2 veins beneath. Inflorescences terminal 
and solitary, 3-4.5 cm long, 3-4 cm wide, rounded pan- 
icles with opposite or whorled branches, peduncles 5- 
20 mm long, 0.5-0.8 mm thick, glabrous, bracts ca. 1 
mm long, flowers subsessile in distal cymes, pedicels 0.5- 
1 mm long. Flowers glabrous externally, hypanthium ca. 
0.8 mm long, obconic, calyx lobes ca. 0.5 mm long; 
corolla funnelform to salverform, white, tube ca. 2 mm 
long, 1-1.2 mm diam., lobes 5, ca. 1.5 mm long and 0.7 
mm broad at the base; stamens 5, anthers ca. 0.8 mm 
long. Fruits 4-5 mm long, 3-4 mm diam., ellipsoid with 
ca. 10 longitudinal ridges, bright red; pyrenes ridged. 

Plants of wet evergreen lowland forest forma- 
tions (and in open pine savannas in Nicaragua), 
from near sea level to 200 m elevation (to over 
1 300 m in Mexico and Guatemala). Flowering in 
southern Central America in March-May and Sep- 
tember; fruiting in July-December. The species 
ranges from Mexico, mostly along the Caribbean, 
to Panama but is disjunct from Nicaragua to cen- 
tral Panama. 

Psychotria fruticetorum is recognized by the 
small, often cuneate and subsessile leaves, cadu- 
cous stipules with two distal awns, general lack of 
pubescence, small domatia, small inflorescences, 
and small flowers and fruit. The tendency of the 
leaves to dry grayish or reddish and the bright red 
berries are characteristics of subgenus Psychotria. 
Presently, this species is not known from Costa 
Rica, though it grows near Bluefields, Nicaragua, 



248 



FIELDIANA: BOTANY 



and in central Panama. It may be restricted to 
limestone soils. 



Psychotria glomerulata (J. D. Smith) Steyerm., 
Mem. New York Bot. Gard. 23: 670. 1972. Ce- 
phaelis glomerulata J. D. Smith, Bot. Gaz. 16: 
12, pi. 1. 1891. Figure 18. 

Small shrubs or treelets, 0.6-2(-4?) m tall, much- 
branched, leafy stems 1-5 mm thick, quadrangular, gla- 
brous; stipules with a short (0.5-2 mm) sheath, truncated 
and entire at apex, glabrous, persisting. Leaves with pet- 
ioles 3-1 1(-15) mm long, 0.8-1.3 mm thick, glabrous, 
yellowish green; leaf blades 5-14(-17) cm long, 1.5- 
5(-6.5) cm broad, narrowly oblong to narrowly elliptic- 
oblong, oblong or lanceolate, apex acuminate with tip 
4-1 2 mm long, base obtuse to acute or cuneate and slightly 
decurrent on petiole, drying chartaceous to stiffly char- 
taceous, glabrous above and below, 2 veins 9-13/side 
and loop-connected near the the margin. Inflorescences 
terminal and solitary, capitula 1.2-2.5 cm long, 1.5-3.5 
cm broad, oblate to subglobose, subtended by 4 decus- 
sate involucrate bracts 10-16 mm long and 1 2 mm broad, 
oblong to broadly obovate and rounded distally, greenish 
yellow or whitish and sometimes edged with blue or 
purple, glabrous, peduncle to 5(-15) mm long, flowers 
tightly enclosed within the bracts and spatulate bracte- 
oles. Flowers glabrous externally, hypanthium ca. 1 mm 
long, calyx lobes 5, 0.5-1 .5 mm long; corolla funnelform, 
white, tube 10-15 mm long, 1-2 mm diam.. lobes 5, 
1.5-3 mm long; stamens 5, anthers 2-3 mm long. Fruits 
6-13 mm long and 5-10 mm diam., ellipsoid, bright 
blue; pyrenes 4-6 mm long and 3-4 mm diam., appar- 
ently smooth. 

Understory shrubs of evergreen rain forest for- 
mations, from near sea level to 600 m elevation 
along the Caribbean slope and coastal plain. Prob- 
ably flowering and fruiting throughout the year but 
flowering mostly In March-April and August-Oc- 
tober. This species ranges from southern Mexico 
and Belize along the Caribbean lowlands to Pan- 
ama. 

Psychotria glomerulata is recognized by its short 
stature, narrow leaves, lack of pubescence, solitary 
sessile terminal heads with four (to eight) white or 
yellow-green involucrate bracts, and lowland Ca- 
ribbean habitat. Our figure is based on the original 
illustration. This species is similar to P. guapilensis 
with long stipule lobes; P. apoda Steyerm. of South 
America may be synonymous. 



Shrubs or small treelets, 1.5-3(-5) m tall, leafy stems 
0.6-4 mm thick, glabrous, flattened or quadrangular at 
first, soon terete and with thickened nodes; stipules 1-3 
mm long, 1.5-2.5 mm broad, with 2 acute lobes and 
distal sinus 0.3-2 mm deep, glabrous, deciduous or per- 
sisting. Leaves with petioles 3-22 mm long, 0.3-1 mm 
thick, glabrous; leaf blades (2-)3-10(-13) cm long, 
(0.8-)l-2.5(-3) cm broad, narrowly elliptic-oblong to 
lanceolate or linear-lanceolate, apex long-acuminate with 
tip 7-15 mm long, base gradually narrowed or abruptly 
acute and slightly decurrent on petiole, drying charta- 
ceous, dark green or brown above, glabrous above and 
below, 2 vein 8-1 3/side, often with parallel intermediate 
2 veins. Inflorescences terminal and solitary (pseu- 
doaxillary), 4-14 cm long, to 1 1 cm broad, open pyra- 
midal panicles with distant slender opposite or subop- 
posite branches, peduncles 15-52 mm long, 0.7-1.2 mm 
thick and glabrous, bracts 3-10 mm long and linear to 
triangular (or leaf-like), distal branching cymose (often 
minutely puberulent), flowers usually sessile in triads. 
Flowers glabrous on the exterior, hypanthium ca. 1 mm 
long, obconic, calyx tube ca. 1 mm long, subentire or 
with 5 acute lobes to 0.5 mm long; corolla funnelform, 
greenish white, tube 4-6 mm long, 0.6-1 mm diam., 
lobes 5, 1-2 mm long. Fruits 3-5 mm long, 2.5-4 mm 
diam., ellipsoid-oblong, blue, purple, or white, persisting 
calyx 0.3-0.7 mm long; pyrenes 3-5-angled or -ridged. 

Plants of evergreen montane forest formations, 
from 1000 to 2000 m elevation. Flowering in 
March, April, and October; fruiting in January and 
March-August. The species ranges from the Cor- 
dillera de Guanacaste eastward along the high- 
lands to Darien, Panama. 

Psychotria goldmanii is recognized by its higher- 
elevation habitat, smaller narrow long-acuminate 
glabrous leaves, calyx with minute lobes, longer 
corolla tube, and small blue fruit. The leaves often 
dry with the lower surfaces markedly paler in color 
and obscure 3 venation. The stipules with two 
small lobes are also distinctive. Few other species 
of Psychotria in southern Central America develop 
such narrow linear-lanceolate leaves (an extreme 
form of which is represented by the type of P. 
torresiana, Standley 39769 us). Compare P. stey- 
ermarkii with more numerous 2 veins and P. val- 
eriana with one stipule appendage. Smaller-leaved 
specimens of this species may be difficult to dis- 
tinguish from Palicourea montivaga, but that spe- 
cies has shorter leaves, very slender petioles, a 
tendency to develop a short stipular tube, inflo- 
rescences that dry yellowish, and corollas that are 
swollen at the base. 



Psychotria goldmanii Standl., Contr. U.S. Natl. 
Herb. 18: 130. 1916. P. torresiana Standl., J. 
Wash. Acad. Sci. 15: 288. 1925. P. eugenifolia 
Dwyer, Ann. Missouri Bot. Gard. 67: 375. 1980. 
Figure 55. 



Psychotria graciliflora Benth. in Oerst, Vidensk. 
Meddel. Dansk. Naturhist. Foren. Kjobenhavn 
1852: 35. 1853. P. vallensis Dwyer, Ann. Mis- 
souri Bot. Gard. 67: 438. 1980 (fide Hamilton 
1989). Figure 60. 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



249 



Shrubs, 0.6-2(-3) mm tall, much-branched and often 
flat-topped, leafy stems 0.7-2(-3) mm thick, glabrous 
(except for reddish colleters at the node), usually with 2 
(opposite) longitudinal ridges between nodes; stipules 
with a narrowly triangular to ovate base 1-2 mm long 
and 2 awns 1-2 mm long, with ascending hairs 0.1-0.2 
mm long, reddish brown, deciduous. Leaves mostly clus- 
tered near the ends of branchlets, petioles 2-10 mm long, 
0.5-1 mm broad, glabrous; leaf blades 1 .5-6(-8) cm long, 
0.7-3 cm broad, elliptic to ovate-elliptic, apex bluntly 
obtuse to acute or subacuminate, base cuneate to acute 
and decurrent on petiole, drying chartaceous, dark gray 
or reddish gray above, glabrous above and below, 2 
veins 3-6/side, pit domatia rarely present. Inflorescences 
terminal and solitary, 2-6 cm long, 2-5 cm broad, py- 
ramidal rounded panicles with 1-2 pairs of opposite 
branches or umbellate, peduncles l-2(-3.5) cm long, ca. 
0.5 mm thick, glabrous and drying dark, bracts and brac- 
teoles to 1 mm long, flowers often sessile in distal cymes 
or triads. Flowers glabrous externally, hypanthium ca. 
0.7 mm long, obconic, calyx tube ca. 0.5 mm long, lobes 
0.2-1 mm high; corolla white, salverform, tube 2.5-3.5 
mm long, 0.5-0.8 mm diam., lobes 5, 1-2 mm long; 
stamens 5, anthers ca. 0.9 mm long. Fruits 4-6 mm long, 
3-5 mm diam., globose to ellipsoid, with ca. 10 longi- 
tudinal ridges, bright red or orange; pyrenes ca. 4 mm 
diam. 

Understory plants of wet evergreen forest for- 
mations on both the Caribbean and Pacific slopes, 
from near sea level to 1700 elevation (to 2500 m 
elsewhere, and not found below 1000 m on the 
Pacific slope of Costa Rica). Flowering in Febru- 
ary-August (mostly in March-May); fruiting 
throughout the year. The species ranges from 
southern Mexico to Colombia. 

Psychotria graciliflora is recognized by its very 
small leaves, usually horizontal glabrous branches, 
caducous stipules with two slender pubescent awns, 
small open inflorescences with slender peduncles, 
and small flowers with minute calyx lobes. The 
bright red fruit and tendency for leaves to dry 
grayish or reddish are characteristics of subgenus 
Psychotria. Costa Rican material of this species 
has smaller leaves with fewer secondary veins than 
material from Honduras and Nicaragua (Hamil- 
ton, 1989, cited under the genus). This species 
resembles P. carthagenensis, P. chagrensis, and P. 
parvifolia (see fig. 60). 



Psychotria grandis Sw., Prodr. 43. 1788. Figure 
64. 

Shrubs or small trees, (1.5-)4-8(-10) m tall, trunks to 
20 cm diam., leafy stems 4-10 mm thick, terete, glabrous 
or minutely papillate-puberulent in early stages; stipules 
8-30 mm long, 5-20 mm broad, broadly ovate-trian- 
gular with the margins often revolute, acute to acuminate 



and often minutely (1 mm) 2-lobed, usually glabrous, 
often inflated at the base, deciduous. Leaves with poorly 
denned petioles 0.5-3.5 cm long, 1.3-3 mm broad, gla- 
brous; leaf blades ( 1 1-) 1 8-40 cm long, (3-)6-l 6 cm broad, 
obovate to elliptic-obovate or oblanceolate, apex obtuse 
to short-acuminate with tip ca. 5 mm long, gradually 
narrowed to the cuneate base and long-decurrent on pet- 
iole, drying chartaceous to subcoriaceous, grayish green 
to pinkish gray or dark grayish above, glabrous above 
and below, 2 veins 12-16/side. Inflorescences terminal 
or pseudoaxillary and solitary, 12-25(-30) cm long, 10- 
18 cm broad, open pyramidal panicules with opposite 
branching or with 4 branches (2 larger and 2 smaller) at 
lower nodes, peduncles 11-18 cm long, 2-4 mm thick, 
glabrous or minutely (0.05 mm) papillate-puberulent, 
flowers 2-5 in distal cymules, bracts ca. 2 mm long and 
triangular, pedicels 1-3 mm long, bracteoles ca. 0.5 mm 
long. Flowers mostly glabrous externally, hypanthium 
ca. 0.8 mm long, obconic, calyx tube ca. 0.6 mm long, 
lobes 5, 0-0.3 mm high, triangular; corolla white, fun- 
nelform, tube 2-4 mm long, 1.5-2 mm diam., lobes 5, 
1.5-2 mm long and 1.1 mm broad at the base; stamens 
5, anthers 0.8 mm long. Fruits 5-7 mm long and 5 mm 
diam., broadly ellipsoid or subglobose, orange or red, 
surfaces smooth and rounded; pyrenes ca. 5 mm long, 
with 4-5 rounded dorsal ribs. 

Trees of wet evergreen forest formations on both 
the Caribbean and Pacific slopes of Costa Rica, 
from near the seashore to 800 m elevation. Flow- 
ering in February-July; fruiting mostly in July- 
February. The species ranges from southern Mex- 
ico and the Greater Antilles to northern South 
America. 

Psychotria grandis is recognized by its often larger 
(tree) habit, large glabrous-obovate leaf blades de- 
current on the petioles, large ovate-acuminate 
stipules with reflexed lateral margins, large inflo- 
rescences, and small flowers. The red fruit and 
vegetative parts drying grayish or reddish are char- 
acteristics of the subgenus Psychotria. Guatemalan 
material is often puberulent and with larger inflo- 
rescences; Nicaraguan collections have larger fruit. 
This species is closely related to P. costavenia Gri- 
seb., which ranges from Mexico and Cuba into 
southern Nicaragua and is smaller in its parts. 



Psychotria guadalupensis (DC.) Howard, J. Ar- 
nold Arbor. 47: 139.1 966, sensu lato. Loranthus 
guadalupensis DC., Prodr. 4: 294. 1830. Vis- 
coides pendulum Jacq., Select. Stirp. Amer. 73, 
pi. 51, f. 1. 1763. P. parasitica Sw., Prodr. 44. 
1 788, nomen illeg. P. pendula (Jacq.) Urb., Symb. 
Ant. 1: 445. 1900, non P. pendula Hooker f., 
1880. P. peperomiae Standl., Contr. U.S. Natl. 
Herb. 18: 132. 1916. P. orchidearumSiandl.J. 
Wash. Acad. Sci. 18: 276. 1928. Figure 60. 



250 



FIELDIANA: BOTANY 



Small epiphytic shrubs, stems 0.2-1 m long, leafy stems 
1-4 mm thick, glabrous, often slightly succulent; stipules 
with a short (0.5-2.5 mm) usually membranaceous trans- 
lucent sheath, breaking apart and becoming thickened 
and whitish at the base. Leaves with petioles 1-6 mm 
long, 0.7-1.2 mm thick, glabrous; leaf blades 12-40 mm 
long, 7-22 mm broad, ovate or ovate-elliptic to obovate, 
apex bluntly obtuse to rounded, base obtuse to acute and 
slightly decurrent on petiole, drying stiffly chartaceous 
to subcoriaceous, gray to dark gray above, paler beneath, 
glabrous above and below, with 2-5 obscure 2 veins per 
side arising at angles of 20-40 (veins rarely prominent). 
Inflorescences terminal and solitary, 1.5-5 cm long, 
equally broad, few-flowered cymes to pyramidal panicles 
with opposite or 3 -partite branching, peduncles 3-30 
mm long, 0.4-1 . 1 mm thick, glabrous and reddish in life 
but often drying black, bracts 2-6 mm long, longer bracts 
linear, flowers in distal cymes or triads, pedicels 0.5-5 
mm long. Flowers glabrous externally, hypanthium 1- 
1.2 mm long, 0.5-0.7 mm diam., calyx tube ca. 0.3 mm 
long, calyx lobes 0.4-1 mm long, triangular and acute, 
often reddish; corolla white or rose, salverform, tube 
(4-)6-8 mm long, ca. 1 mm diam., lobes 4, 1.5-2 mm 
long. Fruits 3-6 m long, 3-5 mm diam., globose to ob- 
late, usually with 4 pyrenes, red or purple and finally 
black. 

Epiphytes of lower montane cloud forest for- 
mations of the Caribbean slope and continental 
divide, from (200-)1000 to 2300 m elevation. 
Flowering in January-August; fruiting in Febru- 
ary-September and December. The species ranges 
from Mexico and the West Indies to the Guianas. 

Psychotria guadalupensis is recognized by its 
epiphytic habit, unusual stipular tube, thick leaves 
often with obscure venation, variable inflores- 
cences with slender peduncles, longer-tubed flow- 
ers, and four-locular ovary. The leaves drying 
grayish and red fruit are characters shared with 
subgenus Psychotria. The differentiation between 
this species and material placed under P. pithe- 
cobia may be artificial, though it does separate a 
large portion of the specimens effectively. Costa 
Rican material placed here has longer corolla tubes, 
and leaf secondary veins are less numerous and 
less prominent when dried as in P. pithecobia, and 
they arise from the midvein at a smaller angle. But 
there do appear to be a few intermediates, and 
paucity of collections makes it difficult to tell 
whether or not the longer corolla tubes of P. guad- 
alupensis are a consistent difference. (See Ho- 
ward's discussion of variation in this species, J. 
Arnold Arbor. 47: 139-142. 1966.) 

The preceding description refers to Costa Rican 
material (excluding specimens assigned to P. pi- 
thecobia) and differs from that given by Steyer- 
mark (1974). For example, corolla tubes are ca. 3 
mm long in Venezuela (and in P. pithecobia), much 



shorter than usually seen in this material. It seems 
probable tbat the length of the corolla tube varies 
as greatly (between different plants) as do so many 
other characters of inflorescences and leaves in this 
complex. 

The type of P. orchidearum Standl. (Standley & 
Valeria 50863 us) has rather small (2 cm long) 
leaves and small inflorescences, but the leaves are 
not as narrow as those in P. maxonii. The type of 
P. peperomiae Standl. (Pittier 3235 us from Pan- 
ama) has somewhat larger leaves and is interme- 
diate between the larger-leaved forms of this spe- 
cies and the type of P. orchidearum. The fact that 
all these plants share the same habitats and alti- 
tudinal range suggests that they may be part of a 
single polymorphic complex. 



Psychotria guapilensis (Standl.) Hammel, Selby- 
ana 12: 139. 1991. Cephaelis discolor Polak., 
Linnaea 41: 572. 1877, not/*, discolor (Griseb.) 
Rolfe. C. tonduzii Krause, Bot. Jahrb. Syst. 54, 
Beibl. 1 19: 45. 1916, not P. tonduzii Standl. C. 
nana Standl., J. Wash. Acad. Sci. 17: 171. 1917, 
not P. nana K. Krause. Evea guapilensis Standl., 
J. Wash. Acad. Sci. 15: 104. 1925. C. nicara- 
guensis Standl., Trop. Woods 16: 46. 1928. C. 
guapilensis (Standl.) Standl., Publ. Field Co- 
lumb. Mus., Bot. Ser. 4: 295. 1929. Figure 7. 

Herbs or subshrubs, 25-90 cm tall, with only 1 or 2 
unbranched terete stems 2-4 mm thick, glabrous; stip- 
ules with basal portion 2-4 mm long, with 2 narrow 
triangular lobes 3-8 mm long, glabrous, often deciduous. 
Leaves with petioles 2-9.5 cm long, 0.8-2 mm thick, 
glabrous; leaf blades 9-24 cm long 3.5-10 cm wide, el- 
liptic-oblong to oblong or elliptic-obovate, apex abruptly 
narrowed and acuminate, tip 6-1 5 mm long, base obtuse 
to acute and slightly decurrent on petiole, drying thinly 
chartaceous, dark green or brown, glabrous above and 
below, 2 veins 10-14/side. Inflorescences terminal and 
solitary, ca. 5-8 cm long, the dense capitula 2-3 cm long 
and 2.5-5 cm broad, globose to oblate, peduncles 4-40 
mm long, ca. 1.5 mm thick, glabrous, with many ovate- 
involucrate bracts 6-10 x 6-10 mm, covering the outer 
surface and becoming dark reddish purple in color, brac- 
teoles spatulate and with purple hairs, flowers sessile. 
Flowers glabrous externally, hypanthium ca. 1.5 mm 
long, calyx short-cupulate, lobes 4-5(-6), 0.8-1.5 mm 
long, triangular, corolla pale pink to purple, funnelform, 
tube ca. 2.5 mm long, lobes 5(-6), 1.5-2 mm long. Fruits 
ca. 10 mm long, pyriform to ellipsoid, bright blue; py- 
renes 5-6 mm long and 4 mm broad, smooth to slightly 
angled on the dorsal surface. 

Understory plants in primary rain forest for- 
mations of the Caribbean slope in Costa Rica, 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



251 



from 30 to 900 m elevation. Probably flowering 
and fruiting throughout the year but most collec- 
tions have been made in July-September. The spe- 
cies ranges from southeastern Nicaragua to the 
Choco region of Colombia. 

Psychotria guapilensis is recognized by its small 
herbaceous habit, long and slender petioles, dense 
rounded short-pedunculate terminal heads cov- 
ered by many purplish to maroon bracts, and bright 
blue fruit. This species resembles P. glomerulata. 



Psychotria haematocarpa Standl., J. Wash. Acad. 
Sci. 18: 274. 1928. Figure 56. 

Shrubs, 1-2.5 m tall, leafy stems 0.8-2.5 mm thick, 
glabrous and terete, with 2 opposite longitudinal ridges 
in early stages; stipules with a short (ca. 1 mm) truncate 
sheath with 2 filiform awns per side 2-5 mm long, early 
deciduous or the base of the stipule becoming thickened. 
Leaves with petioles 2-8(-14) mm long, ca. 1 mm thick, 
glabrous, drying yellowish; leaf blades 6-15 cm long, 2- 
5.5 cm broad, elliptic to oblong-elliptic, apex acuminate 
with tip 5-13 mm long, base attenuate or acute and 
decurrent on petiole, drying membranaceous or thin- 
chartaceous, green or grayish green, glabrous above and 
below, 2 veins 6- 1 0/side and loop-connected near (1.5- 
3 mm) the margin. Inflorescences terminal (pseudoax- 
illary), solitary, rounded to subcapitate 4-10 mm long, 
to 10 mm broad, condensed cymes with 5-9 flowers, 
peduncles 2-5 mm long, ca. 0.5 mm thick, glabrous (or 
minutely puberulent), bracts 0.5-2 mm long, green, gla- 
brous, pedicels 0-2 mm long. Flowers minutely papil- 
late-puberulent externally, hypanthium ca. 1 mm long, 
calyx 1-2 mm long, thin, lobes 5, 0.8-1.3 mm long, 
triangular; corolla short-salverform, pale greenish or 
white, tube 2-2.5 mm long, ca. 0.5 mm diam. at base, 
lobes 0.5-1 mm long, acute. Fruits 5-6 mm long, 4-5 
mm diam., elliptic-oblong to globose, orange and finally 
bright red (dark purple?) at maturity; pyrenes with 45 
rounded ridges. 

Plants of evergreen forests on the Caribbean slope 
from 50 to 1000 m elevation and on the Pacific 
slope at 500-1000 m. Flowering in July-August 
near La Selva; fruiting in October-February. This 
species ranges from Nicaragua to Colombia. 

Psychotria haematocarpa is recognized by its 
small stipules with deciduous filiform awns, small- 
er green dried leaves with loop-connected second- 
ary veins, very small condensed inflorescences, and 
red fruit. The calyx resembles the corolla in size 
and texture. This is one of only a few species of 
subgenus Heteropsychotria with red fruit, rather 
than blue or black. These are the smallest inflo- 
rescences of any of our species of Psychotria (or 
any of our shrubby species of Rubiaceae), and this 
small size may explain the paucity of collections, 
as the flowers and fruit are very difficult to see 



among the leaves. Compare P. brachybotrya, with 
larger bracts and purple fruits, and smaller spec- 
imens of P. hoffmannseggiana, with larger bracts. 



Psychotria hazenii Standl., Publ. Field Columb. 
Mus., Dot. Ser. 7: 96. 1930. P. ramonensis 
Standl., Publ. Field Mus. Nat. Hist., Bot. Ser. 
18: 1360. l938.CephaelischlorochlamysSlandl, 
loc. cit. 1278. 1938. Figure 55. 

Small shrubs, 1-2 m tall, leafy stems 0.9-3 mm thick, 
glabrous, drying yellowish green; stipules with a tubular 
sheath 3-7 mm long, with 2 narrow lobes 4-1 1 mm long 
and 0.5-1.5 mm broad, linear to narrowly lanceolate, 
glabrous. Leaves with petioles 6-20(-40) mm long, 0.3- 
0.8(-1.3) mm broad, glabrous; leaf blades (3.5-)6-l 1(-17) 
cm long, (l-)1.8-3.5(-4) cm broad, narrowly elliptic to 
elliptic-oblong, apex tapering gradually or abruptly and 
acuminate, tip 5-12 mm long, base acute to obtuse and 
slightly decurrent on petiole, drying chartaceous, dark 
green or brownish, glabrous above and below (or sparsely 
and minutely puberulent along the midvein beneath), 2 
veins 11-17/side. Inflorescences terminal and solitary 
(or 3), 2-4 cm long, 1 .5-2.5 cm broad, loosely condensed 
bracteate heads, peduncles 10-25 mm long (to 30 mm 
in fruit), ca. 1 mm thick and glabrous, bracts subtending 
the inflorescence branches 1-2 cm long, 4-7 mm broad, 
ovate and acute or acuminate, glabrous externally (pu- 
berulent within), chartaceous, green with bluish tips, 
flowers sessile within enclosing bracts. Flowers with hy- 
panthium ca. 1.8 mm long and 1.4 mm diam., obovoid- 
oblong, white, glabrous, calyx tube ca. 0.7 mm long, 
lobes 5, 1-2 mm long, ca. 1.2 mm broad at the base, 
triangular, bluish distally; corolla not seen. Fruits 4-8 
mm long, oblong, becoming bright blue with a spongy 
texture, persisting calyx 1-2 mm long. 

Plants of lower montane rain forest formations 
along the Caribbean slope and continental divide, 
from 900 to 1 600 m elevation. Probably flowering 
in May-July; fruiting in August-November. In 
Costa Rica this species is only known from the 
Caribbean slopes of the Cordillera de Tilaran and 
central Volcanic Highlands; it is also found in Co- 
lombia and Ecuador. 

Psychotria hazenii is recognized by the tubular 
stipules with two long narrow awns, the slender 
petioles, the condensed inflorescences with large 
broadly imbricate bracts, and bright blue fruit. 
Our inability to find material with corollas sug- 
gests that flowering is nocturnal. This species re- 
sembles P. steyermarkii with smaller bracts and 
P. pittieri; compare also Palicourea skotakii. 



Psychotria hebeclada DC., Prodr. 4: 513. 1830. 
Probably including P. molliramis (K. Schum. & 
Krarse.) Steyerm., Mem. New York Bot. Gard. 



252 



FIELDIANA: BOTANY 



23: 529. 1972. Palicourea molliramisK. Schum. 
& Krause, Bot. Jahrb. Syst., 40: 331 1910. Fig- 
ure 55. 

Shrubs or subshrubs, 0.7-2(-3) m tall, leafy stems 1- 
4 mm thick, terete, sparsely to densely puberulent or 
hispidulous with curved whitish hairs 0.2-0.8 mm long, 
often contracted beneath the node when dried; stipules 
with a short truncate tube 0-2 mm long, with 2 distal 
linear lobes or spines 1-3 mm long, puberulent, persist- 
ing or breaking apart. Leaves with petioles 4-15(-30) 
mm long, 0.8-1.2 mm thick, puberulent; leaf blades 7- 
20 cm long, 3-8 cm broad, elliptic to ovate-elliptic or 
elliptic-obovate, apex acuminate with tip 4-14 mm long, 
base cuneate to acute (obtuse), drying membranaceous 
to thin-chartaceous, dark green to brownish green above, 
much paler beneath, sparsely puberulent or subglabrous 
above, puberulent beneath (especially along the midvein) 
with thin straight hairs 0. 1-0.4 mm long, 2 veins (6-)8- 
14/side. Inflorescences terminal and solitary (or 3), 
(2.5-)3-6(-12) cm long, 1.5-4(-8) cm broad, narrowly 
pyramidal or subcylindrical open panicles of closely 
spaced flowers (in early stages), expanding in fruit, pri- 
mary branches opposite or alternate, peduncles to 45(-60) 
mm long, 1-1.8 mm thick, with curved or crooked thin 
hairs 0.2-0.6 mm long, bracts 2-4 mm long, linear (not 
apparent beneath the proximal branches), pedicels 1- 
2(-5) mm long, often purplish, bracteoles 1-2 mm long. 
Flowers distylous, puberulent externally, hypanthium ca. 
0.7 mm long and 0.5 mm diam., obconic, calyx lobes 5, 
0.3-1.5 mm long, ca. 0.3 mm broad, ligulate; corolla 
white or tinged with pink or purple near the mouth, 
salverform, tube 2-4 mm long, 1-2 mm diam. and often 
constricted in middle, lobes 5, 1.5-2 mm long; anthers 
1.2-1.7 mm long. Fruits 4-5 long, 3-5 mm diam., sub- 
globose, with 8 or 1 sharply denned longitudinal ridges, 
glabrous, purple or blue-black; pyrenes 2.5-4.5 mm long, 
ridged. 



Plants of evergreen forest formations on the Ca- 
ribbean slope (20-600 m elevation) and on the 
Pacific slope of southern Costa Rica from 40 to 
700(-1000?) m. Flowering in March-August in 
Costa Rica and Panama; fruiting in May-January. 
The species ranges from southern Mexico to Ec- 
uador and Venezuela. 

Psychotria hebeclada is recognized by its small 
stature, distinctive pubescence on most parts, two- 
awned stipules, relatively compact pyramidal to 
subcylindrical conical inflorescences (in early 
stages), flowers with well-developed calyx lobes, 
and small fruit finally becoming black. These plants 
are often found along rivers, streams, and forest 
edges. This species was called P. pubescens in the 
Flora of Barro Colorado Island, but P. pubescens 
is found in deciduous forests and has smaller calyx 
lobes. It appears that P. hebeclada is synonymous 
with P. molliramis as used by Steyermark (1974). 
Poor material of this species may resemble P. ra- 
cemosa. Liesner 14126 CR with glabrous inflores- 



cences, large calyx lobes, and minutely papillate- 
puberulent corolla tube is tentatively placed here; 
it may be a closely related South American species. 



Psychotria hoffmannseggiana (Willd. ex Roem. & 
Schult.) Muell. Arg. in Mart., Fl. Bras. 6(5): 356. 
1881. P. furcata DC., Prodr. 4: 512. 1830. P. 
involucrata sensu Standley and others, in part, 
not Sw. (cf. Steyermark, 1974). Figure 56. 

Shrubs, 0.7-2(-4) m tall, leafy stems 0.7^4.5 mm thick, 
minutely (0.1-0.2 mm) glabrous to papillate-puberulent 
(conspicuously pubescent with thin straight hairs in a 
few collections); stipules 2-4 mm long, basal sheath 0.3- 
2 mm long, with 2 awns 1-3 mm long (4/node) separated 
by a U-shaped sinus, glabrous (pubescent), usually per- 
sisting. Leaves with petioles 2-8 mm long, ca. 0.5 mm 
thick, glabrous or with hairs 0. 1-0.2(-0.5) mm long; leaf 
blades 5-14(-17) cm long, 1.5-6(-8) cm broad, ovate- 
elliptic, narrowly ovate, elliptic or elliptic-oblong (lan- 
ceolate), apex tapering gradually and acuminate with tip 
3-14 mm long, base cuneate to acute and often decurrent 
on petiole, drying thinly chartaceous and usually green- 
ish, glabrous above or puberulent only on midvein, gla- 
brous or puberulent beneath with thin hairs 0. 1-0.3 mm 
long (more rarely with hairs ca. 0.5 mm long), 2 veins 
5-9/side. Inflorescences terminal and solitary, 6-25 mm 
long, involucre up to 6 cm broad, capitate or condensed- 
cymose with (3-)7-30 congested flowers, peduncles 3- 
20 mm long, 0.7-1 .8 mm thick, pubescent or glabrescent, 
bracts 3-22(-35) mm long, 1-6 mm broad, linear-lan- 
ceolate, conspicuous and persisting, green to purple, 
flowers sessile. Flowers glabrous or puberulent exter- 
nally, hypanthium 1-2 mm long, calyx lobes 0.2-0.8 mm 
long, acute to rounded; corolla white, funnelform, tube 
2-6 mm long, 0.8-2.5 mm diam., lobes 1.3-2 mm long; 
stamens 4 or 5, anthers ca. 1 mm long. Fruits 3-6 mm 
long, 4-5 mm diam., subglobose to slightly oblong, with 
8 longitudinal ridges, becoming purple to dark maroon 
or black, glabrous or puberulent; pyrenes ridged. 



Plants of wet evergreen lowland rain forest for- 
mations, from near sea level to 700 m elevation, 
both on the Caribbean slope and in the General 
Valley and Golfo Dulce area. Flowering primarily 
in May-August; fruiting in July-March. The spe- 
cies ranges from Veracruz, Mexico, into Venezuela 
and Brazilian Amazonia. 

Psychotria hoffmannseggiana is recognized by 
its persisting two-awned stipules, small capitate 
inflorescences with conspicuous narrow spreading 
or recurved persisting bracts, small flowers, and 
purple fruit. The subcapitate inflorescences can 
expand after anthesis and become slightly 
branched. The leaves can vary greatly in size in 
different plants and, while most of our plants have 
little pubescence, a few are densely pubescent. 
Plants of southwestern Costa Rica often have con- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



253 



spicuously larger bracts than specimens from other 
areas. This species is very similar to P. officinalis, 
which has larger branched inflorescences. The ma- 
terial placed here was called P. involucrata Sw. by 
Standley (1938 and in herb.), but Steyermark (1972, 
p. 603) showed that the Swartz name is a synonym 
of P. officinalis. Some authors may prefer to retain 
P. furcata as a distinct species, but the South 
American material of P. hoffmannseggiana in- 
cludes a wide range of variation, some of which 
is similar to that seen in the type of P. furcata. 



cences, and the small flowers often with large calyx 
lobes. The leaves are often undulate along the edge 
in life and may have an arcuate submarginal vein. 
The red fruits and tendency for the leaves to dry 
grayish are characteristics of subgenus Psychotria. 
There seems to be great variation in flowers and 
inflorescence, with thicker puberulent peduncles 
being correlated with more congested flowers. The 
calyx lobes also seem to vary greatly in their de- 
velopment. Compare P. quinqueradiata. 



Psychotria horizontalis Sw., Prodr. 44. 1788. P. 
longicollis Benth. in Oerst., Vidensk. Meddel. 
Dansk. Naturhist. Foren. Kjobenhavn 1852: 33. 
1853. Figure 65. 

Small shrubs to little treelets, 1-3 m tall, leafy stems 
14 mm thick, glabrous (rarely minutely puberulent), 
usually becoming pale grayish; stipules 2-7 mm long, 1- 
4 mm broad at base, triangular to narrowly ovate, obtuse 
to acute, usually glabrous, drying reddish brown, cadu- 
cous. Leaves with petioles l-7(-ll) mm long, 0.5-1.3 
mm thick, glabrous or minutely puberulent; leaf blades 
3 9(-l 3) cm long, 1 .5-4.5(-6) cm broad, elliptic to ovate- 
elliptic, obovate, lanceolate or oblanceolate, apex acute 
to short-acuminate, base obtuse or slightly auriculate, 
drying thin-chartaceous or chartaceous, dark gray to 
pinkish gray or brown, glabrous above and below (rarely 
with thin hairs ca. 0.2 mm long on the veins beneath), 
2 veins 5-9/side, occasionally with tufted hairs or pit 
domatia in vein axils beneath. Inflorescences terminal 
and solitary, 2-6 cm long (to 13 cm in fruit), l.5-4(-7) 
cm broad, rounded open panicles with opposite or 
whorled branching (often globose-umbellate), peduncles 
l-3(-7) cm long, 0.5-1.3 mm thick, glabrous or puber- 
ulent, with 2-5 flowers separate or closely clustered in 
distal cymes, pedicels 0-2 mm long, bracteoles 0.2-1 
mm long. Flowers glabrous externally, hypanthium 0.7- 
1 mm long, obconic, calyx tube ca. 0.6 mm long, lobes 
5, 0.5-3 mm long, linear to ligulate; corolla funnelform, 
white, tube 2.3-3.5 mm long and 0.7-1.4 mm diam., 
lobes 5, ca. 1.5 mm long; anthers 0.7-1 .2 mm long. Fruits 
4-8 mm long, 3-6 mm diam., ellipsoid to ovoid, orange 
becoming bright or dark red; pyrenes ca. 4 x 3 mm, 
with 3-5 rounded ridges. 

Plants of both evergreen and seasonally decid- 
uous forest formations, most often collected from 
the seasonally dry Pacific slope, 20-1600 m ele- 
vation. Flowering most often in May-August; 
fruiting throughout the year (June-December in 
Costa Rica). This species ranges from Mexico to 
Ecuador and Brazil; it is also found in Cuba and 
Hispaniola. 

Psychotria horizontalis is recognized by its 
smaller often subsessile leaves often with domatia, 
unlobed stipules, often umbelliform inflores- 



Psychotria insignis Standl., Contr. U.S. Natl. Herb. 
18: 130. 1916. Figure 66. 

Small treelets or shrubs, 2.5-6 m tall, leafy stems 3- 
8 mm thick, distinctly flattened in early stages, densely 
dark reddish brown pubescent with hairs 0.1-0.4 mm 
long; stipules 12-27 mm long, 4-10 mm broad, ovate- 
lanceolate with an acute bifid tip, awns 2-4 mm long, 
reddish pubescent with longer hairs along the midrib and 
margins, usually caducous. Leaves with petioles 3-5.5 
cm long, 1.5-3 mm thick, densely pubescent with dark 
reddish brown hairs 0.1-0.5 mm long; leaf blades 14- 
30 cm long, 7-18 cm broad, broadly elliptic to broadly 
elliptic-ovate or elliptic-obovate, apex abruptly nar- 
rowed and acuminate with tip 7-18 mm long, rounded 
at the subcordate based, basal lobes often unequal, form- 
ing a sinus 0-10 mm deep, drying chartaceous, dark 
grayish brown or dark reddish brown, glabrous above 
(except the midvein), densely pubescent beneath, 2 veins 
14-18/side and loop-connected in distal part of the blade. 
Inflorescences solitary and terminal, 5-9 cm long, 4-6 
cm broad, short pyramidal panicles with 4 lateral branches 
(2 short and 2 longer) at the first node and short dense 
branches at the closely congested second and third nodes, 
peduncles 1 .8-4 cm long, 1 .4-2 mm thick densely pilose, 
bracts of the first node 6-8 mm long, flowers sessile in 
dense glomerules of 3-10, bracteoles 1-3 mm long. Flow- 
ers reddish puberulent externally, hypanthium 0.5-1 mm 
long, obconic, calyx tube 0.5 mm long, lobes 5, 0.5-1 
mm long, triangular; corolla white, tube 2-3 mm long, 
with prominent hairs 0.3-0.4 mm long, lobes 5, ca. 1 
mm long. Fruits 5-6 mm long, 3-4 mm diam., ellipsoid 
with longitudinal sulci (dried), puberulent, calyx per- 
sisting. 



Plants of the wet evergreen Caribbean slope, 
from 20 to 400 m elevation. Flowering in Septem- 
ber-October; fruiting in September. We have seen 
only three collections: Herrera 2248 (CR) and Gra- 
yum et al. 8929 (CR, MO) from central Costa Rica 
and the type (Pittier 4410 us holotype) from San 
Bias, Panama. 

Psychotria insignis is recognized by the densely 
pubescent parts, large long-petiolate leaves often 
subcordate at the base, and unusual compact in- 
florescence. The upper surface of the leaf blade is 
sometimes vaginate at the juncture with the pet- 



254 



FIELDIANA: BOTANY 



iole. This species is a member of subgenus Psy- 
chotria. Our material differs from the type in that 
the 2 veins do not arise at 90 angles from the 
midvein and the submarginal is not as well de- 
veloped. This name is used provisionally for the 
Costa Rican collections. 



Psychotria ipecacuanha (Brotero) Stokes, Bot. mat. 
med. 1: 365. 1812. Callicocca ipecacuanha Bro- 
tero, Memoria sobre a Ipecacuanha do Brasil 
27. 1 801, Trans. Linn. Soc. London 6: 137. 1802. 
Cephaelis ipecacuanha (Brotero) A. Rich., Bull. 
Fac. Med. 4: 92. 1818. Figure 13. 

Herbaceous subshrubs, 25-50 cm tall, usually with a 
single erect unbranched stem, rhizomatous, leafy stems 
1.5-4 mm thick, glabrous, terete; stipules with a short 
(2 mm) sheath, truncate with 4-8 setae per side 3-6 mm 
long, persisting. Leaves crowded distally, petioles 3-8 
mm long, 1-2 mm thick, glabrous; leaf blades 7-17 cm 
long, 4-9 cm broad, obovate to oblong or elliptic-ob- 
ovate, apex acute or short-acuminate, base cuneate to 
rounded-obtuse, drying membranaceous to thin-char- 
taceous, glabrous or minutely puberulent above and be- 
low, 2 veins 5-7/side. Inflorescences terminal or axil- 
lary, solitary, capitulae to 2 cm long and 1-3 cm broad, 
subglobose, peduncles 1-4 cm long, deflexed, involucrate 
bracts 5-10 mm long, ovate, acute, flowers sessile. Flow- 
ers distylous, glabrous externally, hypanthium ca. 2 mm 
long, ellipsoid, calyx teeth 5, ca. 0.5 mm long; corolla 
funnelform, white, tube 3-4 mm long, cylindrical, lobes 
5, 1.5-2.5 mm long; stamens 5, anthers ca. 1.6 mm long. 
Fruits ca. 10 mm long, becoming red then black; pyrenes 
6-7 mm long, ridged. 

Plants of the lowland Caribbean rain forest for- 
mations (0-600 m elevation), and probably the 
result of introduction for cultivation in Central 
America. The species ranges from southeastern 
Nicaragua to the Amazon basin of Brazil. 

Psychotria ipecacuanha is recognized by its short 
unbranched rhizomatous habit, unusual fimbriate 
stipules, leaves with broad blades and short peti- 
oles, small involucrate heads, and red to black 
fruit. The roots and rhizomes are the source of the 
alkaloid emetin and are used in folk medicine or 
cultivated for pharmaceutical use as ipecac. The 
species is referred to as "ipecac" and raicilla or 
ipecacuana. 



Psychotria jimenezii Standl., J. Wash. Acad. Sci. 
1 5: 288. 1 925. P. wendlandiana Oerst. ex Standl., 
loc. cit. 18: 9. 1928. Figure 63. 

Shrubs and small treelets, 2-5(-7) m tall, to 15 cm 
dbh, leafy stems 1.2-4.5 mm thick, pubescent with as- 



cending reddish hairs ca. 0.3 mm long (rarely glabrous), 
terete; stipules 5-12 mm long, 2-5 mm broad at the base, 
triangular to ovate, acute to acuminate (or shortly bifid), 
puberulent, margins ciliolate, drying reddish brown. 
Leaves with petioles variable among plants, 0-20(-35) 
mm long, 0.8-1.5 mm thick, hirsutulous to papillate- 
puberulent; leaf blades 6-16 cm long, 1.7-5 cm broad, 
oblanceolate to narrowly obovate or narrowly elliptic- 
oblong, apex acute to acuminate with tip 3-10 mm long, 
base gradually narrowed and cuneate or rounded-auric- 
ulate, drying membranaceous to thin-chartaceous, dark 
brown or almost black above, glabrous above, with short 
(0. 1-0.2 mm) reddish hairs on the veins beneath, 2 veins 
8-1 2/side. Inflorescences terminal and solitary, 4-10 cm 
long, 3-5 cm broad, open pyramidal panicles, peduncles 
1 5-50 mm long, ca. 1 mm thick, yellowish green in life, 
densely reddish brown hirsutulous when dried, first node 
with 4 branches (2 long and 2 short), bracts 2-6 mm 
long, triangular, flowers subsessile in crowded distal clus- 
ters of 3-8, bracteoles 0.5-1 mm long. Flowers usually 
glabrous externally, hypanthium ca. 0.7 mm long, ob- 
conic, calyx ca. 0.5 mm long, calyx lobes obscure; corolla 
tubular, cream white to yellowish or greenish yellow, 
tube 1.5-2.5 mm long and 1.4 mm diam., lobes 5, ca. 
1.2 mm long, 0.9 mm broad at base; stamens 5, anthers 
0.7 mm long. Fruits 4-6 mm long, 4-5 mm diam., sub- 
globose, bright red; pyrenes with 4-5 rounded longitu- 
dinal ridges. 

Understory plants of evergreen lower montane 
cloud forest formations along the Caribbean slope 
and continental divide, from (200-)400 to 1600 
m elevation. Flowering in January-June; fruiting 
in February- April, August-October, and Decem- 
ber. The species is found in the Cordilleras de 
Guanacaste and Tilaran and on the Caribbean side 
of the Meseta Central in central and north-central 
Costa Rica. 

Psychotria jimenezii is recognized by its reddish 
pubescence (sometimes glabrous), usually oblan- 
ceolate leaves, triangular stipules acute to acu- 
minate, inflorescences with four branches at the 
first node and second node (two short basal and 
two longer ascending branches), and small gla- 
brous flowers. The tendency of the leaves to dry 
grayish and the fruit becoming red are character- 
istics of the subgenus Psychotria. The length of 
petioles and shape of the leaf (especially the base) 
can differ greatly in different collections of this 
species. The narrowed lamina base is subauricu- 
late in the type of P. wendlandii (Wendland 781 
us). Compare P. horizontalis, P. orosiana, and P. 
laselvensis. 



Psychotria lamarinensis C. Hamilton, Phytologia 
64: 227. 1988. Figure 66. 

Shrubs, ca. 2 m tall, leafy stems 1.7-4.5 mm thick, 
glabrous, grayish, terete; stipules 8-12(-20) mm long, 6- 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



255 



1 2 mm broad, broadly ovate, apex obtuse, reddish brown, 
glabrous, caducous. Leaves with petioles 2-8 mm long, 
1.4-2.7 mm thick, glabrous, slightly sulcate above; leaf 
blades 1 3-22 cm long, 5-13 cm broad, elliptic to broadly 
elliptic or elliptic-obovate, apex acute or short-acumi- 
nate with tip ca. 5 mm long, base obtuse to cuneate or 
slightly truncated (rounded), drying chartaceous, grayish 
or reddish brown, 2 veins 8-1 I/side, minute pit domatia 
sometimes present in the vein axils. Inflorescences ter- 
minal or pseudoaxillary, 2-5, 1.5-3 cm long, to 2 cm 
broad, condensed globose panicles of cymes with short 
lateral branches (or appearing umbellate), peduncles 4- 
20 mm long, 0.8-1.2 mm thick, glabrous, dark red, usu- 
ally with 4 subequal branches at first node, bracts 0.2- 
2 mm long, triangular, bracteoles not apparent, pedicels 
0.5-1 .5 mm long. Flowers glabrous externally, calyx tube 
ca. 0.5 mm long, cupulate, truncate or with very short 
teeth; corolla tubular, white, tube 2.5-3 mm long and 1 
mm diam., lobes 5, 1-2 mm long and 1 mm broad, with 
a linear (1.5 mm) extension from near the apex; stamens 
5, anthers ca. 0.7 mm long. Fruits 7-8 mm long, 4-5 
mm diam., oblong-ellipsoid; pyrenes smooth. 

Plants of deeply shaded sites in evergreen rain 
forest formations on the Caribbean slope, at 20- 
500 m elevation. Flowering in March-May; fruit- 
ing in November. This endemic species ranges from 
Canalete (Alajuela) to Cahuita (Limon). 

Psychotria lamarinensis is recognized by its larg- 
er broad glabrous short-petiolate leaves (cuneate 
basally but slightly rounded at the petiole), small 
short-branched globose panicles, and broadly ovate 
stipules. The leaves drying grayish or reddish brown 
are characteristic of species in subgenus Psy- 
chotria. The extensions on the corolla lobes are 
separate in bud. This species is closely related to 
P. quinqueradiata, but P. lamarinensis has shorter 
corolla tubes and larger leaves with the veins di- 
verging at angles of 70-80. 



Psychotria laselvensis C. Hamilton, Phytologia 64: 
228. 1988. Figure 61. 

Shrubs, 1.2-4 m tall, leafy stems 1.3-3 mm thick, 
erect, glabrous, bark smooth; stipules 2-4(-6) mm long, 
1-3 mm broad, triangular to ovate, glabrous or puber- 
ulent on the midrib externally, caducous. Leaves sub- 
sessile or with petioles 1-7 mm long, 0.4-0.8 mm thick, 
glabrous; leaf blades 5-13(-16) cm long, 1.5-4(-6) cm 
broad, elliptic to narrowly elliptic-oblong or elliptic-lan- 
ceolate, apex acute or short-acuminate with tip to 1 cm 
long, base acute to cuneate and often slightly auriculate 
at the petiole, drying thin-chartaceous, dark grayish above, 
glabrous above and below, 2 veins 7-10/side, pit dom- 
atia often present in distal vein axils beneath. Inflores- 
cences terminal and solitary, 6-1 5 cm long, 4-7 cm broad, 
open pyramidal, panicles with thin opposite branches, 
peduncles 4-9 cm long, 0.4-0.7 mm thick, glabrous, with 
2 or 4 lateral branches at the first node, bracts 0.5-1 .5(-3?) 
mm long, flowers often in distal pairs or triads, pedicels 



ca. 0.5 mm long, bracteoles ca. 0.5 mm long. Flowers 
distylous, glabrous externally, hypanthium ca. 0.7 mm 
long, obconic, calyx lobes poorly developed (to 0.2 mm 
long); corolla narrowly funnelform, white or pale yellow, 
tube 1.5-3 mm long, 1-2 mm diam., lobes 5, 1.5-2 mm 
long; anthers 1-1.2 mm long. Fruits 4-5 mm diam., 
globose, red; pyrenes with 4-5 rounded ridges. 



Plants of lowland Caribbean rain forest under- 
story, at 50-300 m elevation on the Caribbean 
slope. Flowering in February-June (mostly May); 
fruiting in July. The species is only known from 
the La Selva research station and nearby areas, 
Heredia, and northern Limon provinces. 

Psychotria laselvensis is recognized by its small- 
er elliptic to oblanceolate leaves, larger inflores- 
cences with very thin open spreading branches, 
and pedicellate flowers. The veins are often flat- 
tened and expanded at the vein axils and form pit 
domatia distally. The leaves drying grayish and 
the red fruit are characteristics of subgenus Psy- 
chotria. This species resembles P. marginata with 
more acute leaf bases and is closely related to P. 
graciliflora and P. orosiana. 



Psychotria limonensis K. Krause, Bot. Jahrb. 54, 
Beibl. 119: 43. 1916. Figure 64. 

Shrubs, subshrubs, or small trees, 0.6-2(-6) m tall, 
leafy stems 2-6 mm thick, glabrous or sparsely puber- 
ulent, drying reddish to pale gray or dark; stipules 5-12 
mm long, 3-6 mm broad, triangular to ovate, rounded 
to acute, entire or shortly bifid, glabrous, drying reddish 
brown, caducous. Leaves with petioles 1.2-6 cm long, 
0.7-3 mm thick, glabrous; leaf blades 1 1-22(-28) cm 
long, 5-12(-15) cm broad, ovate-elliptic to elliptic-ob- 
long or broadly elliptic, apex acute to short- or long- 
acuminate with tip 5-20 mm long, base obtuse to acute 
(occasionally slightly decurrent on petiole), drying thin- 
to stiffly chartaceous, gray to dark reddish brown, gla- 
brous above and below, 2 veins (7-)10-19/side and 
distally loop-connected to form a submarginal vein. In- 
florescences terminal (pseudoaxillary), solitary of 3, 3- 
9 cm long, 5-11 cm broad, broad panicles with opposite 
or whorled branches, peduncles 1-4 cm long, 1.2-2.5 
mm thick, glabrous and reddish brown, bracts 0.5-1.2 
mm long, triangular, flowers in congested cymes of 3-6, 
pedicels 0-1.5 mm long. Flowers glabrous externally, 
hypanthium ca. 0.5 mm long, obconic, calyx tube ca. 0.4 
m long, lobes 0.2-0.5 mm high, obtuse; corolla funnel- 
form to rotate, white, tube 2-2.6 mm long and 1-1.7 
mm diam., lobes (4-)5(-6), 1.2-1.5 mm long, triangular; 
stamens (4-)5(-6), anthers 0.8-1 mm long. Fruits 4-5 
mm long, 3-4 mm diam., ellipsoid, bright red; pyrenes 
with ca. 5 longitudinal ridges. 

Plants of evergreen lowland rain forest forma- 
tions on both the Caribbean and Pacific slopes, 
from near sea level to 200 m elevation (to 700 or 



256 



FIELDIANA: BOTANY 



rarely 1 700 m elsewhere). Probably flowering and 
fruiting throughout the year (fruiting primarily in 
June-January in Costa Rica). The species ranges 
from southern Mexico and Belize to Panama and 
Colombia. 

Psychotria limonensis is recognized by its large 
leaves with many secondary veins and arcuate 
submarginal vein, usual lack of pubescence, un- 
lobed stipules, small flowers with four to six co- 
rolla lobes, and small fruit. The red fruit and ten- 
dency for leaves to dry gray or pinkish are 
characteristics of subgenus Psychotria. The sec- 
ondary veins arise at almost 90 angles from the 
midvein. This species can be mistaken for spec- 
imens of P. panamensis var. compressicaulis with 
larger leaves and calyptrate stipules. It is also sim- 
ilar to P. mexiae with longer corollas and acute 
buds. 



Psychotria longipedunculoides C. M. Taylor, nom. 
nov. P. longipedunculata Dwyer, Ann. Missouri 
Bot. Card. 67: 389. 1980, non P. longepedun- 
culata (Gardn). Muell. Arg. in Mart. 

Herbaceous subshrubs, 0.5-1.2 m tall, usually un- 
branched, leafy stems 1.7-6 mm thick, semisucculent, 
glabrous or minutely papillate-puberulent; stipules 6-8 
mm long, 6-8 mm broad, broadly ovate, margin irregular 
or roughly 2-lobed, thin and translucent, deciduous or 
persisting. Leaves with petioles 14-33 mm long, 0.8-1.5 
mm thick (dried), glabrous; leaf blades 12-18 cm long, 
5-7(-9) cm broad, elliptic to elliptic-oblong or elliptic- 
obovate, apex acute or acuminate with tip 5-10 mm 
long, gradually narrowed to the acute base and slightly 
decurrent on petiole, drying membranaceous to thin- 
chartaceous, pale olive gray above, 2 veins 12-15/side, 
an arcuate submarginal vein present. Inflorescences sol- 
itary and axillary from distal nodes, 12-16 cm long, 1.5- 
3 cm broad, congested pyramidal panicles to subcapitate, 
flowering portion 1.2-3 cm long, peduncles 10-14 cm 
long, 0.9-1.4 mm thick, glabrous, bracts l-6(-12) mm 
long, linear-subulate, flowers sessile or subsessile. Flow- 
ers glabrous externally, hypanthium ca. 0.5 mm long, 
ellipsoid, calyx lobes 5, ca. 1.5 mm long; corolla to 2.5 
m long, funnelform, greenish white, lobes 5, stamens 5; 
anthers ca. 0.8 mm long. Fruits 4-6.5 m long, ca. 3 mm 
diam., oblong-obovoid; pyrenes with 4-5 rounded ridg- 
es. 



Recently collected on the Caribbean slope of the 
Talamanca mountains at ca. 700 m elevation near 
the border with Panama (Herrera 3255 CR, MO, 
usj). Flowering and fruiting in July. This species 
is only known from eastern Costa Rica, western 
Panama, and Choco, Colombia. 

Psychotria longipedunculoides is recognized by 
the single-stemmed semisucculent habit, glabrous 



parts, solitary axillary inflorescences with small 
flower clusters on a long peduncle, and membra- 
naceous ovate stipules. Similar in habit to P. mac- 
rophylla but differing by linear bracts, congested 
flowers, and stipules. 



Psychotria luxurians Rusby, Mem. Torrey Bot. 
Club 6: 50. 1896. P. berteriana subsp. luxurians 
(Rusby) Steyerm., Mem. New York Bot. Gard. 
22: 534. 1972. Figure 57. 

Small trees or shrubs, 2-8(-l 2) m tall, leafy stems 2.5- 
7 mm thick, glabrous or minutely (0.05 mm) papillate- 
puberulent, terete; stipules 3-6(-9) mm long, to 6 mm 
broad, broadly ovate rounded to obtuse, splitting in age 
or with a short ( 1 mm) sinus, usually glabrous, decidu- 
ous. Leaves with petioles 5-35 mm long, 1 .3-3 mm thick, 
glabrous or minutely papillate-puberulent; leaf blades 
1 2-26(-32) cm long, 5-1 1(-1 3) cm broad, elliptic-oblong 
to elliptic or oblong, apex acuminate with a narrowed 
tip 4-15 mm long, base obtuse to cuneate (acute) base, 
drying chartaceous, dark green or brown above (distinct- 
ly paler beneath), glabrous above and below or minutely 
papillate-puberulent beneath, 2 veins 8-14/side. Inflo- 
rescences terminal, solitary or 3, (7-) 12-26 cm long, 8- 
20 cm broad, pyramidal open panicles with mostly op- 
posite branching, peduncles 2-8 cm long, 1 .2-6 mm thick, 
minutely papillate-puberulent, bracts of primary branch- 
es l-8(-12) mm long and ca. 1 mm broad, triangular to 
elliptic, flowers congested in cymose clusters of 3-10, 
bracteoles 1.5-3 mm long, elliptic, pedicels 0-2 mm long. 
Flowers glabrous externally, hypanthium 0.7-1.3 mm 
long, obconic, calyx lobes 0.2-0.5(-0.8) mm long, broad- 
ly obtuse; corolla short-funnelform, white, greenish white 
or yellowish, tube 2-4 mm long and 1.5-2 mm diam., 
lobes 5, 1-1.7 mm long and 0.7-1 mm broad, obtuse or 
rounded. Fruits 3-5 mm long and 3-6 mm diam., oblong 
or globose, blue, blue-black, or purple; pyrenes 3 mm 
long, with 4-5 dorsal ridges. 

Plants in open secondary growth and light gaps 
in evergreen rain forest formations of the Carib- 
bean slopes and lowlands, 20-1000 m elevation. 
Flowering primarily in March-July in Costa Rica; 
fruiting in July-November. This species ranges 
from southern Nicaragua to Bolivia. 

Psychotria luxurians is recognized by its large 
essentially glabrous leaves usually drying greenish, 
the short and broad stipules that are entire or with 
two very short rounded lobes, large minutely pa- 
pillate-puberulent inflorescences with many small 
elliptic bracts, numerous small flowers, and small 
blue fruit. Ants may live within the stems of this 
species (Beach 1480 CR). Note that the persisting 
bracts subtending the flowers can be mistaken for 
calyx lobes. This species can be confused with P. 
angustiflora and P. berteriana, but those species 
do not have their flowers surrounded by as many 



BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 



257 



conspicuous bracteoles, and the flowers differ. 
Steyermark (1972) treated this as a variety of P. 
berteriana with larger bracts, but the stipule mor- 
phology is different in the two. Psychotria tapan- 
tiensis of higher elevations is closely related with 
larger bracteoles, corollas, and fruits. 



Psychotria macrophylla Ruiz & Pav., Fl. Peruv. 
2: 56. 1799. P. anomothyrsa K. Schum. & J. D. 
Smith, Bot. Gaz. 35: 3. 1903. P. macrophylla 
var. angustissima Standl., Publ. Field Mus. Nat. 
Hist., Bot. Ser. 22: 203. 1940. P. macrophylla 
ssp. anomothyrsa (K. Schum. & J. D. Smith) 
Steyerm., Mem. New York Bot. Gard. 23: 563. 
1972. Figure 12. 



Herbs or subshrubs, 0.5-1 .5(-2) m tall, leafy stems 3- 
1 1 mm thick, semisucculent, usually unbranched, gla- 
brous, becoming terete; stipules 2-5 mm long, triangular, 
usually with a narrow divergent succulent slightly bi- 
lobed apex, glabrous. Leaves with petioles 2-8 cm long, 
1.2-3 mm thick, glabrous or rarely puberulent in early 
stages; leaf blades 12-32 cm long, 4-14 cm broad, elliptic 
to elliptic-oblong, narrowly to broadly oblong, or lan- 
ceolate, apex acuminate with tip 8-18 mm long (rarely 
gradually narrowed and acute), base obtuse to acute and 
slightly decurrent on petiole, drying membranaceous to 
thin-chartaceous, dark greenish to dark brownish above, 
usually glabrous above and below, 2 veins 8-15/side, 
occasionally loop-connected near the margins. Inflores- 
cences axillary, usually I/node, 3-15 cm long (to 25 cm 
in fruit), 4-6 cm wide, usually open pyramidal panicles 
with opposite branching, peduncles 0.7-8(-12) cm long, 
glabrous or occasionally puberulent with hairs 0.1-0.4 
mm long, bracts 0.5-2(-10) mm long, triangular, flowers 
solitary or in small glomerules of 2-3 along the distal 
axes, sessile or subsessile. Flowers glabrous or puberulent 
externally, hypanthium ca. 0.5 long, obconic, calyx ca. 
0.7 mm long, lobes ca. 0.2 mm high, broadly triangular; 
corolla tubular-sal verform, white, tube 2-4 mm long and 
0.6-1.5 mm diam., lobes 4 or 5, 1-2 mm long; stamens 
4-5, anthers 1.2 mm long. Fruits 5-7 mm long and 3- 
4 mm diam., oblong, white and arenchymatous; pyrenes 
ca. 6 mm long, with 4-5 rounded ridges. 

Understory plants of wet sites in evergreen for- 
est formations, from 20 to 2200 m elevation in 
Costa Rica. The species appears to be more com- 
mon in highland forests than in those below 500 
m elevation; it is rare in sandy streamside soils at 
La Selva. Flowering and fruiting throughout the 
year (but flowering mostly in January-August). The 
species ranges from southern Mexico to Bolivia. 

Psychotria macrophylla is recognized by its usu- 
ally short stature with succulent unbranched main 
stems, the axillary inflorescences, small subsessile 
flowers on branched inflorescences, white fruit with 
four to five rounded ribs. This species is variable; 



leaves range from narrowly lanceolate to broadly 
elliptic in different plants; most plants are glabrous 
but a few are minutely puberulent. Some dried 
collections have crinkled leaf margins. The ar- 
rangement of flowers on the distal branches of the 
inflorescence varies greatly, from open-cymose to 
spicate. Steyermark (1974) interpreted some of this 
variation in Venezuela as hybridization between 
P. macrophylla and P. uliginosa, but this seems 
doubtful and it may be simply that individual 
plants of P. macrophylla can vary greatly in the 
expression of a variety of morphological features. 
Compare Psychotria aggregata, with densely ag- 
gregated flowers, slightly larger calyx lobes, and 
pyrenes with a single dorsal ridge. See the discus- 
sions under P. capacifolia, P. siggersiana, and P. 
aggregata. This group of species was recently stud- 
ied by Molly Nepokroeff (wis, 1992). 



Psychotria marginata Sw., Prodr. 43. 1788. P. ni- 
caraguensis Benth. in Oersted, Vidensk. Med- 
del. Dansk. Naturhist. Foren. Kjobenhavn 1852: 
34. 1853. Figure 61. 

Shrubs or treelets, (0.5-)l-3(-4) m tall, leafy stems 
1 .3-4 mm thick, terete, glabrous; stipules 5-14 mm long, 
to 5 mm broad, narrowly ovate and acute, glabrous, 
usually drying reddish brown, caducous. Leaves often 
clustered distally, petioles 4-22(-35) mm long, 1-1.8 
mm thick; leaf blades 6-17 cm long, 2-6 cm broad, 
narrowly obovate to elliptic obovate, oblanceolate or 
elliptic-oblong, apex acute to acuminate with tip 3-10 
mm long (rarely obtuse), base acute to cuneate and de- 
current on petiole, drying chartaceous or stiffly charta- 
ceous and grayish, glabrous above, glabrous beneath or 
minutely (0.05 mm) papillate-puberulent on the veins 
beneath, 2 veins 7-13/side, weakly loop-connected in 
the distal part of the leaf, margin thickened along the 
edge, usually ciliolate, midvein usually with small ex- 
panded flaps-like domatia near the vein axils beneath. 
Inflorescences terminal and solitary, 7-17 cm long, 5- 
1 2 cm broad, open pyramidal panicles with thin opposite 
branches diverging at 90 angles, peduncles 2-7 cm long, 
0.6-1.7 mm thick, glabrous, bracts 1-3 mm long, tri- 
angular, flowers 2-6 in distal cymes, pedicels l-3(-9) 
mm long. Flowers distylous, glabrous externally, hypan- 
thium ca. 0.6 mm long, obconic, calyx tube ca. 0.4 mm 
long, lobes 0.2 mm high, broadly triangular or obscure; 
corolla short-funnelform, white or yellowish, tube 2-3 
mm long, 0.7-1.3 mm diam., lobes 5, 1-1.5 mm long; 
anthers 0.8-1.1 mm long. Fruits 3.5-6 mm long, 3-6 
mm diam., subglobose, red or purple; pyrene ca. 3 mm 
long, with 4-5 ridges. 



Plants in secondary growth of evergreen forests 
and partly deciduous forest formations, from 10 
to 800 (-1 100) m elevation. Flowering and fruiting 
throughout the year (fruiting most often in Feb- 



258 



FIELDIANA: BOTANY 



ruary-May in Costa Rica). At La Selva, this spe- 
cies flowers in the wettest part of the year: July- 
November. The species ranges from Mexico and 
the West Indies to Peru and Bolivia. 

Psychotria marginata is recognized by its small 
stature, acute triangular stipules, usually obovate 
leaf blades often with pronounced domatia and 
ciliolate margins, small flowers, and small fruit on 
long thin pedicels in open inflorescences. The red 
fruit and leaves drying grayish (less often reddish) 
are characteristics of subgenus Psychotria. Com- 
pare this species to P. laselvensis, which lacks cil- 
iolate leaf margins and has a shortly truncated or 
rounded leaf base. 



Psychotria maxonii Standl., Proc. Biol. Soc. Wash. 
37: 53. 1924. Figure 60. 

Small epiphytic shrubs with stems 0.2-0. 5(-l) m long, 
often pendant, leafy stems 0.6-2 mm thick, succulent, 
glabrous and often becoming pale grayish; stipules with 
united truncated membranaceous tube 1-2 mm long, 
glabrous, persisting and sometimes becoming thickened 
and pale-colored at base. Leaves subsessile or with pet- 
ioles 1-3 mm long and 0.3-0.5 mm thick, glabrous; leaf 
blades 12-23(-28) mm long, 3-5(-6) mm broad, nar- 
rowly lanceolate to linear-lanceolate, apex tapering grad- 
ually and acute, base acute or obtuse and slightly de- 
current on petiole, drying stiffly chartaceous or 
subcoriaceous dark gray above, glabrous above and be- 
neath, 2 and 3 veins obscure in dried material, margins 
slightly revolute. Inflorescences terminal or axillary, 
I/node, 2-4 cm long, 2-4 cm wide, open few-flowered 
3-branched pyramidal panicles or cymes with slender 
opposite branches often red in color, bracts ca. 0.4 mm 
long, peduncles 3-12 mm long, 0.2-0.5 mm thick, gla- 
brous, drying black, pedicels filiform, 1-5 mm long. 
Flowers glabrous externally, hypanthium ca. 1 mm long, 
obovoid to oblong, calyx ca. 0.7 mm long (including 
lobes), lobes 4, ca. 0.5 mm long and triangular; corolla 
salverform, white and often tinged with red or pink, tube 
2.5-4(-6) mm long, 0.4-0.8 mm diam., lobes 4, 1-1.5 
mm long, obtuse; stamens 4, anthers ca. 1 mm long. 
Fruits ca. 4 mm long and 4 mm diam., globose, becoming 
orange or red and finally black; pyrenes smooth. 

Epiphytic plants in montane cloud forest for- 
mations, from 1500 to 2300 m elevation. Flow- 
ering in February-March, May, and July-August; 
fruiting in January-March, July, and October. The 
species ranges from the northern part of the Mese- 
ta Central (near San Ramon) along the Caribbean 
escarpment and continental divide to the high- 
lands of western Panama. 

Psychotria maxonii is recognized by the epi- 
phytic habit, small narrow succulent leaves, small 
thin few-flowered inflorescences, and small flowers 
with slender corolla tubes. The leaves dry grayish 



and the fruit become red, then black (as do species 
of subgenus Psychotria). Panamanian collections 
appear to have more ovate leaves than are found 
in Costa Rica. No other Central American species 
of Psychotria has such small and narrow leaves. 
This species is closely related to P. guadalupensis 
and shares the same habitats in Costa Rica with 
that species. It is possible that the two species are 
actually conspecific, with P. maxonii being no more 
than a very unusual form of P. guadalupensis; see 
the discussions under/*, guadalupensis and P. pith- 
ecobia. 



Psychotria mexiae Standl., Publ. Field Columb. 
Mus., Bot. Ser. 4: 296. 1929. Palicourea nigres- 
cens M. Martens & Galeotti, Bull. Acad. Roy. 
Sci. Bruxelles 1 1(50): 136. 1844, not Psychotria 
nigrescens De Wild, 1924. P. schippii Standl. & 
Steyerm., Publ. Field Mus. Bot. Hist., Bot. Ser. 
23: 24. 1943. Figure 62. 



Shrubs or small trees, 2-5(-10) m tall, leafy stems 2- 
4 mm thick, glabrous (in Costa Rica) or sparsely puber- 
ulent; stipules 1 2-20(-40) mm long, 1 .5-3(-5) mm broad 
at the base, narrowly lanceolate and sheathing the apex 
(as in Ficus), glabrous, caducous. Leaves with petioles 
3-20 mm long, 1.3-2 mm broad, glabrous; leaf blades 
8-16(-20) cm long, 2.5-7(-8.5) cm broad, elliptic to nar- 
rowly elliptic, apex acuminate with tip 10-17 mm long, 
base acute to obtuse and decurrent on petiole, drying 
thinly chartaceous to stiffly chartaceous, dark above, gla- 
brous above and below (rarely with hairs along midvein 
beneath), 2 veins 10-13/side, domatia with small hairs 
often present in the vein axils beneath. Inflorescences 
terminal, 1-3, 3-6 cm long, 2-5 cm wide, rounded to 
broadly pyramidal panicles with usually 4 branches at 
the first node, peduncles 8-28 mm long, 1 .5-2 mm thick, 
bracts and bracteoles not apparent, cymules of 3-8 flow- 
ers, pe