ILLINOIS LIBRARY
,M URBANA-CHAMPAIGN
BIOLOGY
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rt.xv, no. 33
FLORA COSTARICENSIS
t\ iiliam Burger, Kditor
Family #202 Rubiaceae
vVilliam Burger Charlotte M. Tavlor
Kurcniber 30. 1993 'ublication 1454
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FTELDIANA
Botany
NEW SERIES, NO. 33
FLORA COSTARICENSIS
William Burger, Editor
Family #202 Rubiaceae
William Burger
Curator
Department of Botany
Field Museum of Natural History
Chicago, Illinois 60605-2496
Charlotte M. Taylor
Missouri Botanical Garden
St. Louis, Missouri 63166-0299
Accepted April 16, 1993
Published December 30, 1993 BIOLOGY LIBRARY
Publication 1454 101 "
PUBLISHED BY FIELD MUSEUM OF NATURAL HISTORY
© 1993 Field Museum of Natural History Library of Congress Catalog Card Number: 93-73814
ISSN 00 15-0746 PRINTED IN THE UNITED STATES OF AMERICA
Table of Contents
INTRODUCTION v
ACKNOWLEDGMENTS v
RUBIACEAE 1
KEYS TO THE RUBIACEAE OF COSTA RICA 2
Key 1 : Technical Keys to the Traditional
Tribes and Genera of Rubiaceae 2
Key 2: Artificial Key to Genera and
Illustrations 8
Illustrations of Rubiaceae 15
Description of Genera and Species (Aliber- tid) 82
Psychotria 220
LITERATURE CITED 324
LIST OF ACCEPTED SPECIES 325
INDEX . 328
List of Illustrations
1 . Twining shrubs (Manettia spp.) and subshrubs with small stiff leaves (species of Arcytophyllum, Declieux-
ia, and Diodid) 15
2. Twining herbs: species of Coccocyp- selum and Geophila 16
3. Herbs with small leaves and slender stems: species of Didymaea, Galium, Nertera, and Oldenlandia 17
4. Erect herbs with narrow lanceolate leaves and capitate or verticillate flowers: species of Crusea, Mitracar-
pus, Richardia, and Spermacoce 18
5. Erect herbs with narrow lanceolate leaves: Spermacoce spp 19
6. Erect herbs with narrow lanceolate leaves: Diodia spp. and two species
of Spermacoce 20
7. Herbs or subshrubs with larger leaves: species of Amphidasya, Hoff- mannia, Lasianthus, and Psychotria ... 2 1
8. Herbs or subshrubs with axillary flowers: unusual species of Hoffman-
nia 22
9. Herbs or subshrubs with axillary flowers: pubescent species of Hoff- mannia and H. congesta 23
10. Subshrubs with axillary flowers: spe- cies of Hoffmannia with larger leaves . . 24
1 1 . Subshrubs with axillary flowers: spe- cies of Hoffmannia with leaves ta- pering gradually to the base 25
12. Subshrubs with axillary flowers: spe- cies of Psychotria 26
13. Subshrubs with axillary flowers: spe- cies of Psychotria 27
14. Trees with very large or lobed leaves: three species of Pentagonia 28
15. Flowers with very long corolla tubes: species of Lindenia, Osa, and Poso- queria 29
16. Inflorescences with greatly expanded petal-like calyx lobes: species of Ca- lycophyllum, Mussaenda, Pogonopus,
and Warszewiczia 30
17. Inflorescences of involucrate heads: species of Psychotria (formerly Ce- phaelis spp.) 31
18. Inflorescences of involucrate or con- spicuously bracteate heads: species of Psychotria 32
19. Inflorescences of compact heads with flowers connivent at the base: species ofAppunia, Morinda, and Schroder a . . 33
20. Inflorescences long and narrow: spe- cies of Gonzalagunia 34
2 1 . Inflorescences long and narrow: spe- cies of Gonzalagunia and Rondeletia . . 35
22. Fruits usually terminal and solitary: species of Randia with small leaves ... 36
23. Fruits usually terminal and solitary: species of Randia with medium-sized leaves 37
24. Fruits usually terminal and solitary: species of Randia with larger leaves ... 38
25. Fruits usually terminal and solitary: species ofAlibertia, Duroia, Genipa,
and Hippotis 39
26. Fruits usually terminal and solitary: species of Borojoa and Genipa 40
27. Plants usually epiphytic: species of Cosmibuena and Hillia with smaller leaves 41
28. Plants usually epiphytic: species of Cosmibuena and Hillia with larger leaves 42
29. Trees with large open inflorescences: species of Ladenbergia and Conda- minea corymbosa 43
30. Inflorescences with clusters of long- tubular flowers: species ofAmaioua, Guettarda, Isertia, and Tocoyena 44
in
3 1 . Showy flowers: species of Coutarea, Crusea, Exostema, Ixora, and Pen- tas
32. Inflorescences with scorpioid or heli- coid branches: species of Guettarda ... 46
33. Flowers with narrow corolla tubes: species of Guettarda and a species of Chomelia 47
34. Flowers with narrow corolla tubes: species of Chomelia, Guettarda, and Hamelia 48
35. Inflorescences mostly axillary: spe- cies of Sabicea (vines) and Sommera (trees) 49
36. Inflorescences axillary or terminal: species of Chiococca and a species of Chione 50
37. Many small flowers in dense inflores- cences: species of Chimarrhis, Cin- chona, Machaonia, and Uncaria 51
38. Flowers in much-branched open in- florescences: species of Deppea, Rus-
tia, and Simira 52
39. Small flowers in dense or open pani- cles: species of Elaeagia 53
40. Small flowers and capsular fruits: species of A Iseis, Exostema, Ferdi- nandusa, and Macrocnemum 54
4 1 . Rondeletia spp 55
41 A. Rondeletia spp 56
42. Hamelia spp 57
43. Bertiera, Ixora, and Raritebe spp 58
44. Faramea: species with larger leaves ... 59
45. Faramea: species with smaller leaves. . 60
46. Coussarea and Rudgea spp 61
47. Coussarea spp. and two similar Psy- chotria spp 62
48. Coussarea: species with larger leaves . . 63
49. Palicourea: species with conspicuous bracts . . .64
50. Palicourea: species of lower eleva- tions and a species of Isertia 65
45 51. Palicourea: species with larger leaves
and yellow or orange flowers 66
52. Palicourea: species with larger leaves and blue, lavender, purple, or white flowers 67
53. Palicourea: species with smaller
leaves 68
54. Psychotria subg. Heteropsychotria: larger-leaved pubescent species and a species of Palicourea 69
55. Psychotria subg. Heteropsychotria: species with smaller leaves 70
56. Psychotria subg. Heteropsychotria: species with very small inflorescences ..71
57. Psychotria subg. Heteropsychotria: species with larger open inflorescenc- es 72
58. Psychotria subg. Heteropsychotria: species with conspicuous open inflorescences 73
59. Psychotria subg. Heteropsychotria: species of deciduous habitats and
some with smaller inflorescences 74
60. Psychotria subg. Psychotria: species with very small leaves and a com- plex of epiphytic species 75
6 1 . Psychotria subg. Psychotria: species
with smaller narrow leaves 76
62. Psychotria subg. Psychotria: high-ele- vation species and those with Ficus-
like stipules 77
63. Psychotria subg. Psychotria: densely pubescent species 78
64. Psychotria subg. Psychotria: large- leaved species 79
65. Psychotria subg. Psychotria: decidu- ous and unusual species 80
66. Psychotria subg. Psychotria: several unusual species 81
IV
Introduction
This is the eighth issue in the Flora Costaricensis series. The first dealt with the Piperaceae (Field- iana, Bot. 35, 1971). The second included families numbered 42 through 53, Chloranthaceae through Urticaceae (Fieldiana, Bot. 40, 1977). The third issue covered the Gramineae and was authored by Richard Phol (Fieldiana, Bot., new series, No. 4, 1980). The fourth issue included families num- bered 54 through 70, Podostemaceae through Car- yophyllaceae (Fieldiana, Bot., new series, No. 13, 1983). The fifth issue covered families 200 and 201, the Acanthaceae authored by L. H. Durkee, and the Plantaginaceae (Fieldiana, Bot., new se- ries, No. 18, 1986). The sixth issue included fam-
ilies 80 and 81, Lauraceae and Hernandiaceae (Fieldiana, Bot., new series, No. 23, 1990). The seventh issue included families numbered 97 through 103, Krameriaceae through Zygophylla- ceae (Fieldiana, Bot., new series, No. 28, 1991).
In the figures, leaves and leafy stems are drawn to the same scale throughout. Enlarged flowers and fruits are drawn to the same scale on an individual plate unless otherwise noted. The closed scales represent centimeters and the open scales repre- sent millimeters. The figures are somewhat dia- grammatic and represent the senior author's con- cept of a common or characteristic morphology.
Acknowledgments
We wish to thank the staff, and especially Pablo Sanchez, of the Museo Nacional de Costa Rica for their assistance over many years. A grant from the Museo Nacional allowed the senior author to work at the Herbario Nacional for several weeks in No- vember 1990. Charlotte Taylor received support for travel from the National Science Foundation (BSR 83-10702 and BSR 87-00068), the Fondos Institucionales Para Investigacion of the Univer- sity of Puerto Rico, and the Dee Scholarship Fund of Field Museum. Collecting programs by the Mis- souri Botanical Garden (MO), Institute Nacional de Biodiversidad and the Museo Nacional (CR), supported in part by grants from the National Sci- ence Foundation and the National Geographic So- ciety, have added significantly to our knowledge of Costa Rica's Rubiaceae. The recent collections of Jorge Gomez-Laurito, Michael Grayum, Wil- liam Haber, Barry Hammel, Gerardo Herrera, Quirico Jimenez, and Nelson Zamora have been
especially significant. Loans from the U.S. Na- tional Herbarium (us) and the Duke University Herbarium (DUKE) were important for our work on this family.
A number of our colleagues have been especially helpful in preparing this treatment. The annota- tions and advice of C. Dennis Adams, John Dwyer, Barry Hammel, and David Lorence were especial- ly important. In addition, John Dwyer and David Lorence have provided descriptions of new spe- cies, and Roy Gereau corrected all the Latin de- scriptions. The Flora Mesoamericana project un- der the leadership of Geritt Davidse (MO) has provided information and assistance on many oc- casions. We are also indebted to the Missouri Bo- tanical Garden for allowing Charlotte Taylor to contribute her time and effort to this treatment. Finally, we thank three anonymous reviewers who made many corrections and suggested useful im- provements for the text.
FLORA COSTARICENSIS
Family #202 Rubiaceae
RUBIACEAE
By William Burger and Charlotte M. Taylor
Herbs, shrubs, or small- to medium-sized trees (rarely vines or tall canopy trees), stems glabrous to pubescent with simple hairs, terete or angular; stipules of opposing leaves usually united across the stem (interpetiolar), sometimes united to the petioles and forming a broad sheath (Spermacoceae), rarely separate and paired at the leaf base, stipules of the same leaf sometimes also united above the petioles (intrapetiolar) and forming a short tube, often with hair-like or tooth-like colleters at the adaxial base or along the edge, persistent to caducous and leaving a scar across the stem (stipules transformed into small leaves in Galium and Sherardia). Leaves op- posite or sometimes whorled (very rarely alternate), al- ways simple, petiolate or occasionally sessile, glabrous or pubescent, nearly always entire and without lobes (pinnatind in Pentagonia spp., with small lobes in Simira spp., minutely serrate or crenate in some Spermacoceae), nearly always pinnately veined (palmately veined in some weedy Spermacoceae), domatia of pits or tufted hairs present in the vein axils on the lower surfaces of leaves in some genera. Inflorescences terminal or axillary, sol- itary to several at each node, very variable in form (open paniculate to cymose, racemose, spicate or capitate), branches of the inflorescence often opposite, bracts and bracteoles often present, the flowers often borne in distal cymes or dichasia, sometimes cincinnoid and 1 -sided (rarely solitary, fasciculate, or united), sessile or pedi- cellate. Flowers usually bisexual and radially symmet- rical (rarely unisexual and dioecious), most often 4- or 5-parted, epigynous, the hypanthium narrowly tubular to subglobosc. calyx tube usually present, calyx lobes usually present and equal or subequal (rarely with 1 lobe greatly expanded and colorful); corolla often salverform with a narrow tube (funnelform to rotate or tubular), corolla lobes valvate, imbricate or contorted in bud; sta- mens usually as many as the corolla lobes and alternating with them, nearly always borne on the corolla tube, fil- aments long to short, anthers usually narrowly oblong and basi fixed to dorsifixed, dehiscing longitudinally (with terminal pores in Rustia); ovary inferior (rarely half- inferior), usually with a ring-like disc on the upper sur- face, with 2 (1-8) locules, placentation apical, basal or from the median septum (parietal in some Gardineae), style solitary from the center of the apex of the ovary, stigmas usually 2 or solitary (clavate to capitate). Fruits capsular, baccate or drupaceous and often with 2 (4-5) pyrenes (a syncarp in Morinda and Schradera, a samara in Allenanthus), berries sometimes large with the seeds imbedded in a fleshy pulp, capsules opening along the locules (loculicidal) or along the septum (septicidal); seeds sometimes with wings or tufted hairs.
The Rubiaceae are one of the largest families of flowering plants, with an estimated 10,700 species (Mabberley, 1987). The family is best represented in the evergreen tropics and is often an important component of the lower strata of such forests.
In most cases the family is easily recognized. The simple opposite leaves are nearly always en- tire and pinnately veined. Only a few weedy spe- cies have subpalmate venation, and a few of our woody species have lobed leaves (Pentagonia spp. and Simira maxonii). The trichomes are never branched or stellate. Some species have domatia in the form of pits or tufted hairs in the vein axils on the undersides of leaves. Though variable in presence, domatia can be helpful in identifying species. Too small to be useful to ants, these leaf domatia probably offer shelter for predatory and fungivorous mites (Pemberton & Turner, 1989). The nodes are nearly always marked by interpetio- lar stipules or interpetiolar lines if the stipules have fallen. The stipules can be important in identifying species but may be apparent only on young shoots; they may be greatly enlarged when subtending in- florescences. Persisting stipules may be distorted or torn apart as the stem expands. Hair-like or tooth-like structures between the base of the stip- ule and the stem are called colleters. These are usually Anger-like with elongate axial cells and a palisade epidermis (Lersten, 1974). They are be- lieved to secrete mucilage, gums, or resins.
Inflorescences vary greatly in some genera. In some species the bracts subtending the first pair of opposite branches of the inflorescence may be replaced by smaller leaves. In this case an inflo- rescence that is solitary and terminal can be in- terpreted as being a group of three inflorescences: a terminal one and two axillary to the distal leaves (bract homologs). The flowers are often borne in distal cymes on opposite branches of the inflores- cences. Many species are distylous with long-styled (pin) or short-styled (thrum) flowers on different plants. The corolla is nearly always radially sym- metrical and with a conspicuous tube. Curvature of the tube or assymetry of the corolla lobes is rare. The inferior ovary is usually two-locular, and the number of ovules per locule has been used as
FIELDIANA: BOTANY, N.S., NO. 33, DECEMBER 30, 1993, PP. 1-333
a primary criterion for distinguishing the subfam- ilies.
While a very distinctive family, there are a few genera of other families that can be mistaken for Rubiaceae. Collections ofCassipourea (Rhizopho- raceae), Hedyosmum (Chloranthaceae), and Neea (Nyctaginaceae) are often found among specimens of Rubiaceae. There are also look-alikes in Acan- thaceae, Loganiaceae, and Onagraceae.
The Rubiaceae of Central America are relatively well understood, and their taxonomy is in good order. This is the result of intensive study by many workers, past and present. Among these, the work of Paul Carpenter Standley provided a solid foun- dation. His publications and many annotations have been particularly useful in preparing the pres- ent account. The treatments of the family for the Flora of Guatemala (Standley & Williams, 1975), Flora of Panama (Dwyer, 1 980), and Flora of Ven- ezuela (Steyermark, 1 974) have also been very use- ful. The recent studies by Dennis Adams, John Dwyer, Clement Hamilton, Joseph Kirkbride, Da- vid Lorence, and others have clarified many dif- ficult species groups and are cited in the text. Many collectors have contributed substantially to our knowledge of this family in Costa Rica (an index to exsiccatae is available on request).
tation, ovule orientation, and characteristics of seeds and fruits. This key follows those presented in the Flora of Guatemala (Standley & Williams, 1975) and the Flora of Panama (Dwyer, 1980). While often very difficult to implement, this key has wide application and places the genera into the traditional tribes. More modern keys can be found in Robbrecht (1988).
As Verdcourt (1976, p. 5) has stated, "[T]he family Rubiaceae contains so many genera and species, many of which resemble each other even when not closely related, that it is impossible to make a useable key which does not involve looking at small and difficult characters." To provide an alternative, we give an additional artificial key that is much simpler and attempts to make the illus- trations more readily accessible. Scanning the il- lustrations with the help of the second key will, hopefully, allow determination of many species without having to ascertain all the morphological details required by the technical key. Commentary under the genera and species gives characteristics that can be helpful in distinguishing the taxa; the detailed descriptions are useful in confirming a determination. Nevertheless, there is no substitute for careful comparisons with annotated herbarium collections to verify a determination.
Keys to the Rubiaceae of Costa Rica
We provide a technical key to the genera that requires ascertaining corolla aestivation, placen-
Key 1: Technical Keys to the Traditional Tribes and Genera of Rubiaceae (see Robbrecht, 1988, for a more modern system)
1 a. Ovules more than 1 in each locule of the ovary (subfamily Cinchonoideae) 2
1 b. Ovules solitary in the locules of the ovary (subfamily Rubioideae, except Naucleeae) 9
2a. Fruits fleshy and indehiscent (baccate or berry-like) 3
2b. Fruits dry and dehiscent (capsule-like) 5
3a. Corolla lobes valvate in bud 1 . Isertieae
3b. Corolla lobes imbricate or contorted in bud 4
4a. Corolla lobes imbricate in bud 2. Hamelieae
4b. Corolla lobes contorted in bud 3. Gardenieae
5a. Flowers in compact spherical heads 8. Naucleeae
5b. Flowers not in compact spherical heads 6
6a. Seeds with wings, tufted hairs or appendages, arranged vertically imbricate on the placenta
4. Cinchoneae
6b. Seeds angled but not winged, or if winged then arranged horizontally on the placenta ... 7
7a. Corolla lobes imbricate or contorted in bud 6. Rondeletieae
7b. Corolla lobes valvate in bud 8
8a. Seeds horizontal, usually many (> 25); stipules entire or bifid; trees and shrubs
FIELDIANA: BOTANY
5. Condamineeae
8b. Seeds vertical and imbricate, usually few; stipules usually setose; herbs and subshrubs ....
7. Oldenlandieae
9a. (from Ib) Seeds pendulous, the radicle superior; trees, shrubs, or woody lianas 10
9b. Seeds ascending, the radicle inferior; trees, shrubs, or herbs 12
lOa. Flowers in spherical/globose heads 8. Naucleeae
I Ob. Flowers never in globose heads 11
II a. Stamens usually borne at apex of corolla tube (the rarely encountered Vangueria of the
Vanguerieae will key out here; see text) 9. Guettardeae
1 Ib. Stamens borne at base of the corolla tube or from the disc 10. Chiococceae
12a. Corolla lobes contorted in bud; trees and shrubs 11. Ixoreae
1 2b. Corolla lobes valvate in bud; trees, shrubs, and herbs 13
1 3a. Ovules borne on base of the locule; mostly woody plants 14
1 3b. Ovules borne from the septum in the center of the ovary; herbs, shrubs, or trees 17
1 4a. Ovary with 7-8 locules; inflorescences globose; fruits multiple, of 4-50 united flowers ....
14. Morindeae
14b. Ovary with 1-5 locules; inflorescences various; fruits simple or with 2 united flowers if
multiple 15
15a. Ovary 1- or 2-locular and with a thin partial septum; fruits with 1 seed . . 12. Coussareeae 15b. Ovary 2-locular (5-locular) and with thick well-developed septum; fruits with 2(-5) seeds
16
1 6a. Stamens usually inserted near the apex of the corolla tube; flowers bisexual
13. Psychotrieae
1 6b. Stamens usually inserted near the base of the corolla tube; flowers often unisexual
15. Anthospermeae
1 7a. (from 1 3b) Stipules not leaf-like nor setose; trees and large shrubs; flowers united near the base;
fruits united or partly united into a syncarp 14. Morindeae
1 7b. Stipules either setose with awn-like appendages or leaf-like (and the small leaves apparently whorled and lacking stipules); herbs or small shrubs; flowers often congested but not united at the base;
fruits never united into a syncarp 18
18a. Stipules usually bearing 3-30 narrow setae or awns; leaves usually opposite . 16. Spermacoceae 1 8b. Stipules leaf-like; leaves and leaf-like stipules appearing as whorls of 4 or more leaves per node
. 17. Rubieae
1. ISERTIEAE (MUSSAENDEAE)
la. Leaves apparently alternate (a minute opposing leaflet often present, not known from Costa Rica)
Didymochlamys
1 b. Leaves opposite, both leaves of the node developed 2
2a. Leaves with the minor venation parallel (lineolate) 3
2b. Leaves with the minor venation not parallel 5
3a. Leaves large, to over 1 m long, with pinnatifid lobes in some species; rows of glands (colleters)
present on the interior of the calyx cup; plants often monopodial Pentagonia
3b. Leaves up to 35 cm long, never with pinnatifid lobes; calyx cup without glands on the interior;
plants often much-branched 4
4a. Calyx 5-lobed Sommera
4b. Calyx 2-lobed or spathe-like Hippotis
5a. Inflorescences axillary 6
5b. Inflorescences terminal 8
6a. Erect unbranched plants to 50 cm tall, with long closely clustered leaves; ovary 2-locular;
corolla more than 30 mm long Amphidasya
6b. Plants with leaves well spaced along the twining or creeping stems; ovary 2-5-locular; corolla less than 1 2 mm long 7
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
7a. Plants prostrate, herbaceous; ovary 2-locular; fruit bright blue Coccocypselum
7b. Plants erect or climbing, herbs or subshrubs; ovary 3-5-locular; fruits reddish becoming
purple or bluish black Sabicea
8a. Flowers and inflorescences drying black; inflorescence capitate; leaves and flowers semisucculent
Schradera
8b. Flowers and inflorescences not drying black; inflorescences subcapitate only in Amphidasya; leaves
and flowers not semisucculent 9
9a. Stipules fimbriate distally; herbaceous with erect unbranched stems to 0.8 m tall . . Amphidasya
9b. Stipules not fimbriate distally; woody plants with branched stems to 3 m tall 10
1 Oa. Inflorescences spike-like; shrubs Gonzalagunia
I Ob. Inflorescences cymose to paniculate; shrubs or trees 11
I 1 a. Anthers not transversely locellate (not divided by transverse walls); corollas less than 2 cm long;
ovary 2-locular Raritebe
1 Ib. Anthers transversely locellate; corollas 3 or more cm long; ovary (2-)5-6-locular Isertia
2. HAMELIEAE
la. Inflorescences always axillary; ovary with 2 or 3 locules; stamens with connective, rarely prolonged distally [corolla lobes imbricate or subvalvate] Hoffmannia
Ib. Inflorescences usually terminal; ovary with 4 or 5 locules; stamens with the connective often pro- longed distally 2
2a. Corolla lobes imbricate in bud; raphides present in leaves; inflorescences often with few cincinnoid branches Hamelia
2b. Corolla lobes contorted in bud; raphides absent in the leaves; inflorescences with many branches, not cincinnoid . . Bertiera
3. GARDENIEAE
la. Ovary with a single locule and intruding parietal placentas (but often difficult to see and the abutting
placentas appearing as a septum); pollen grains in tetrads 2
Ib. Ovary with usually 2 locules (the septum often thin or obliterated as the seeds develop); pollen
separate 3
2a. Plants native and wild, usually armed with spines on branches, frequently with short-shoots
Randia
2b. Plants grown in gardens for ornament, usually lacking short-shoots Gardenia
3a. Flowers bisexual 4
3b. Flowers unisexual 6
4a. Inflorescences with 1-few flowers, flowers not in a candelabra-like arrangement; leaves drying
black Genipa
4b. Inflorescences with several to many flowers; flowers usually in a candelabra-like arrangement;
leaves drying black or not 5
5a. Flower buds curved at the apex, corolla white and becoming yellowish in age; seeds with testa
cells more than twice as long as wide Posoqueria
5b. Flower buds straight at the apex, corolla bright yellow at anthesis; seeds with testa cells less
than twice as long as wide Tocoyena
6a. Fruits rounded or globose; terminal stipules not forming a conical cap, triangular and persisting
7
6b. Fruits oblong; terminal stipules forming a conical cap, caducous 8
7a. Fruits subtended by several whorls of persisting bracts (stipules), fruits more than 5 cm diam.,
pericarp thick, carnose, surfaces rough Borojoa
7b. Fruits not subtended by several persisting bracts; fruits to 3 cm diam., pericarp thin and hard, smooth . . . Alibertia
FIELDIANA: BOTANY
8a. Female flowers usually solitary; fruits hirsute Duroia
8b. Female flowers capitate or cymose; fruits not densely hirsute .... . Amaioua
4. CINCHONEAE
la. Vining or clambering with slender herbaceous stems (genus placed in the Hedyotideae in modern
systems) Manettia
Ib. Shrubs or trees, stems not slender and clambering 2
2a. Anthers dimorphic or trimorphic Ferdinandusa
2b. Anthers monomorphic (all alike within the flower) -3
3a. Calyx with 1 expanded (2-5 cm) white lobe in 1 or 2 flowers of the inflorescence
Calycophyllum
3b. Calyx lobes equal or subequal, inflorescences without expanded large white calyx lobes 4
4a. Plants epiphytic; leaves semisucculent, lateral veins often obscure when dried 5
4b. Plants terrestrial; leaves not semisucculent, lateral veins clearly evident 6
5a. Seeds winged at either end Cosmibuena
5b. Seeds with tufted hairs at one end Hillia
6a. Inflorescences spike-like; stamens attached at the base of the corolla tube [filaments hirsutulous]
Alseis
6b. Inflorescences not spike-like, various; stamens attached above the base of the corolla tube (except
in Coutarea and Exostema) 7
7a. Corolla lobes imbricate or contorted in bud 8
7b. Corolla lobes valvate in bud 9
8a. Stamens conspicuously exserted; corolla not inflated in bud; fruits not compressed or lenti- cellate Exostema
8b. Stamens not conspicuously exserted (may become exserted as corolla ages); corolla buds inflated in late stages (before anthesis); fruits strongly flattened, surfaces often lenticellate
Coutarea
9a. Flowers 4-parted; capsules rounded Bouvardia
9b. Flowers 5-parted; capsules oblong 10
lOa. Capsule splitting from below to the apex [a ring of hairs present within the mouth of the corolla]
Cinchona
lOb. Capsule splitting from above to the base 11
I la. Corolla lobes split at the apex; dehiscing capsules forming 4 coiled valves (not known from Costa
Rica) Joosia
I 1 b. Corolla lobes not split at the apex; capsules valves not becoming coiled 12
1 2a. Capsules opening into the locules Macrocnemum
1 2b. Capsules opening along the septum Ladenbergia
5. CONDAMINEEAE
la. Anthers opening by terminal pores; leaves with pellucid glandular dots Rustia
Ib. Anthers opening by longitudinal slits; leaves without pellucid glandular dots 2
2a. Inflorescences axillary 3
2b. Inflorescences terminal 4
3a. Flowers solitary or few, ca. 25 cm long Osa
3b. Flowers many, ca. 3 mm long Chimarrhis
4a. Calyx lobes equal or subequal, small; stipules large and bifid [leaves large and subsessile]
Condaminea
4b. Calyx with 1 lobe enlarged (2-6 cm) and colored in 1 or a few flowers of each inflorescence; stipules
small, not bifid Pogonopus
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
6. RONDELETIEAE
la. Corolla lobes contorted in bud 2
1 b. Corolla lobes imbricate in bud 5
2a. Corolla tube becoming very long (+ 10 cm), much longer than the corolla lobes; capsule valves
becoming coiled; shrubs of stream sides Lindenia
2b. Corolla never exceeding 5 cm, tube shorter than the corolla lobes; capsule valves not becoming
coiled; if woody not restricted to river edges and wet sites 3
3a. Small herbs of wet sites Sipanaea
3b. Trees and shrubs 4
4a. Corolla 4-parted, yellowish, glabrous within Deppea
4b. Corolla 5-parted, greenish white, villous within Elaeagia
5a. Calyx lobes unequal, often expanded into a broad lobe 6
5b. Calyx lobes equal or subequal, small and unexpanded 7
6a. Expanded calyx lobe bright red Warszewiczia
6b. Expanded calyx lobes whitish (in some spp.) Rondeletia
7a. Capsules ca. 1 cm diam., opening into the septum; seeds winged or flat and enlarged; wood turning reddish when cut and exposed (in ours) Simira
7b. Fruits to 5 mm diam., opening at the septum or locule; seeds lacking wings, not flattened; wood not turning reddish when cut and exposed 8
8a. Corolla tubes usually more than 8 mm long, stamens included; capsule valves usually not woody and bifid at apex; common plants in Central America Rondeletia
8b. Corolla tubes to 5 mm long, stamens usually exserted; capsule valves woody, bifid at the apex; rarely collected in southern Central America Bathysa
7. OLDENLANDIEAE
la. Plants subshrubs, leaves usually small and stiff (ericoid); seeds plano-convex or carinate; plants of
high montane formations 2800-3400 m elevation Arcytophyllum
Ib. Plants herbs or subshrubs, leaves not stiff and ericoid; seeds angular or winged; 0-2000 m elevation
2
2a. Garden ornamentals; flowers red, pink, or white Pentas
2b. Weedy plants of wet or moist sites; flowers white 3
3a. Flowers 4-parted, common introduced weeds Oldenlandia
3b. Flowers 5-parted, rare introduced weeds , Pentodon
8. NAUCLEEAE
1 a. Woody vines with recurved spines; inflorescences axillary, pedunculate, each with 2-5 globose heads
(some modern treatments place this genus in Cinchonieae) Uncaria
Ib. Trees planted for wood, without spines; inflorescences terminal, each with 1 globose head
. Neolamarckia
9. GUETTARDEAE
la. Woody lianas with vining branches [leaves with subparallel 3° veins; fruits fleshy; rare in Costa Rica] Malanea
Ib. Trees or shrubs, not lianas with vining branches (except in some species of Chomelid) 2
2a. Corolla lobes valvate in bud; some species with subparallel (lineolate) minor venation [spines/thorns sometimes present on stems and twigs] Chomelia
2b. Corolla lobes imbricate in bud; minor venation of the leaves not subparallel nor lineolate 3
FIELDIANA: BOTANY
3a. Fruits dry, separating into 2 indehiscent mericarps (cocci); flowers not secund on inflorescence
branches; spines sometimes present on stems and branches Machaonia
3b. Fruits fleshy; flowers often in secund arrangements; spines absent on stems and branches 4
4a. Fruits covered with a fine dense pubescence Guettarda
4b. Fruits glabrate (formerly Antirhea spp.) Chomelia
10. CHIOCOCCEAE
la. Flowers 4-parted; fruits dry, flat and broadly winged Allenanthus
Ib. Flowers 5-parted; fruits fleshy, not winged 2
2a. Fruits compressed laterally (oblong in cross-section); corolla lobes valvate in bud .... Chiococca
2b. Fruits rounded (terete in cross-section); corolla lobes imbricate in bud Chione
11. IXOREAE
la. Inflorescences axillary; floral bracts connate and calyx-like or involucrate beneath the flowers;
cultivated Coffea
1 b. Inflorescences terminal; floral bracts separate, not calyx-like; wild or cultivated for ornament ....
. Ixora
12. COUSSAREEAE
la. Ovules separate in a 1-locular ovary; seeds horizontal and fruits often broader than long, exocarp usually leathery; flowers blue or white Faramea
Ib. Ovules connate from a basal column, ovary 1- or 2-locular; seeds longitudinal, fruits longer than broad, exocarp spongy or carnose; flowers white Coussarea
13. PSYCHOTRIEAE
1 a. Plants with creeping prostrate stems and long internodes; leaves often cordate Geophila
Ib. Plants erect, herbaceous subshrubs to small trees; leaves various (rarely subcordate) 2
2a. Herbaceous subshrubs; fruits laterally compressed Declieuxia
2b. Shrubs, trees, or rarely subshrubs; fruits terete 3
3a. Stipules with a group of small digitate teeth at the apex; seeds with an incurved adaxial (ventral)
surface Rudgea
3b. Stipules without digitate teeth at the apex, simple to bifid; seeds with a flat but sulcate adaxial
surface 4
4a. Corollas usually yellow, orange, reddish, purple, or blue (rarely white), often swollen at the base,
corolla tube often long (+ 1 cm) and slender, with a ring of pubescence below the middle internally;
stipules usually persisting; most often found at higher elevations Palicourea
4b. Corollas usually white or green to yellowish, not gibbous near the base, corolla tubes generally short
(- 1 cm), with a ring of pubescence at or above the middle internally or glabrous within; stipules
persisting or deciduous; widespread with many species Psychotria
14. MORINDEAE
la. Base of hypanthium free, not united to adjacent flowers, not forming a syncarp in fruit; ovary and fruits with 7-8 locules . . . Lasianthus
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
1 b. Base of hypanthium united to adjacent flowers, forming a syncarp in fruit; ovary and fruits with 1 or 2 locules 2
2a. Flowers and fruits drying black; syncarps fleshy, more than 1 cm diam., calyx not elevated on fruits Morinda
2b. Flowers and fruits drying dark brown; syncarps dry, less than 1 cm diam., calyx tube prominent on fruits Appunia
15. ANTHOSPERMEAE
la. Wild plants forming prostrate mats at high elevations; leaves very small and rounded; fruits orange
Nertera
Ib. Cultivated ornamental small (< 1 m) erect shrubs with small narrow acute leaves; fruits brownish
. Serissa
16. SPERMACOCEAE
1 a. Fruits with circumscissile dehiscence Mitracarpus
Ib. Fruits indehiscent or opening by slits, pores or valves 2
2a. Fruits breaking with 3-6 separate indehiscent cocci (note that cocci are borne on the exterior of the
fruiting axis and may resemble seeds; compare Crusea) Richardia
2b. Fruits usually with 2 separate or united cocci, cocci dehiscent or indehiscent 3
3a. Cocci opening distally or longitudinally (sometimes 1 of the 2 cocci not opening in a fruit)
Spermacoce
3b. Cocci not opening or opening only at the base 4
4a. Cocci borne on and separating from a central persisting axis, indehiscent; calyx usually with rounded
lobes Crusea
4b. Cocci not borne on a central axis, indehiscent or dehiscent from the base; calyx usually with acute
lobes . . Diodia
17. RUBIEAE
1 a. Leaves opposite (interpetiolar stipules connate and small) Didymaea
Ib. Leaves whorled (stipules leaf-like) 2
2a. Native herbs; flowers on separate pedicels or solitary and involucrate Galium
2b. Rare introduced procumbent herbs; flowers 4-10 and subsessile in distal involucrate heads
. . Sherardia
Key 2: Artificial Key to Genera and Illustrations
Note that small plants, leafy twigs, and large leaves are all drawn to the same scale throughout the 67 figures. Closed scales represent centimeters; open scales are millimeters. Unless otherwise indicated, enlarged flowers or fruits are to the same scale on the same figure. The illustrations represent typical or common morphologies; they cannot show the range of variation.
la. Herbaceous plants or slender few-branched subshrubs, usually less than 1.5 m tall 2
Ib. Trees, shrubs, vines, or lianas 23
2a. Creeping plants with slender flexible stems, often rooting at the nodes [flowers usually axillary]
3
2b. Erect or prostrate plants, lacking slender consistently creeping stems, stems slightly woody if vining 8
FIELDIANA: BOTANY
3a. Leaf blades less than 1 cm broad 4
3b. Leaf blades usually more than 1 cm broad 7
4a. Leaves in whorls, often linear or narrow; flowers separate (Galium) or in terminal
heads (Sherardia, not illustrated) Fig. 3
4b. Leaves opposite, linear to ovate 5
5a. Leaves often linear; fruits dry capsules with few seeds; weeds below 1500 m
elevation (Oldenlandid) Fig. 3
5b. Leaves not linear; fruits fleshy, 1-2-seeded, rarely found below 1 500 m elevation
6
6a. Fruits orange to red, globose (Nertera) Fig. 3
6b. Fruits blue to black, usually bilobed (Didymaed) Fig. 3
7a. Fruits with many seeds, blue (Coccocypselum) Fig. 2
7b. Fruits with 1-2 seeds, red, blue-black, or black (Geophild) Fig. 2
8a. (from 2b) Leaves usually less than 4 cm broad and lanceolate, often subsessile 9
8b. Leaves usually more than 4 cm broad and petiolate 17
9a. Vining plants often climbing over other plants, with axillary flowers; fruits capsular
with many seeds (Manettid) Fig. 1
9b. Plants erect or if vining then close to the ground and not usually climbing over other
plants; flowers various 10
lOa. Leaves usually less than 15 mm long, stiff or coriaceous 11
lOb. Leaves more than 15 mm long, or thin and herbaceous when less than 15 mm long
12
1 la. Plants of high elevation often in exposed sites; leaves thick and blunt at the apex,
usually closely spaced (Arcytophyllurri) Fig. 1
lib. Plants of mid-elevations; leaves stiff and sharp at the apex (Diodia brasiliensis
and the cultivated Serissa, not illustrated) Fig. 1
1 2a. Ovules more than 2 in each locule; fruits with more than 2 seeds; rarely collected plants
usually found in wet or partly submerged sites (not illustrated) 13
1 2b. Ovules 1 in each locule; fruits with no more than 2(-3) seeds or mericarps; commonly
collected plants in many open weedy habitats, dry or wet 14
13a. Plants with sparse small hairs; corolla pink, tube 5-14 mm long Sipanea
13b. Plants glabrous, slightly succulent; corolla white, tube 1.5-4 mm long
Pentodon
14a. Stipules not clearly united to petioles, awns only 1-2 on each side of the node; stiff
erect plants from a woody base (Declienxia) Fig. 1
14b. Stipules united with the petioles forming a truncated sheath, the sheath usually with more than 3 thin erect awns on each side; fruits dry and usually 2-seeded (genera of
Spermacoceae; see the technical keys and the figures below) 15
15a. Leaves usually less than 2 cm long Figs. 1, 5-6
1 5b. Leaves usually more than 2 cm long 16
16a. Terminal capitula of flowers consistently present, often subtended by bract-like leaves
(Crusea, Mitracarpus, Richardia, Spermacoce) Figs. 4-5
16b. Terminal capitula rarely present (Diodia, Spermacoce) Figs. 1, 5-6
17a. (from 8b) Inflorescences terminal on short woody stems (compare also Psychotria spp. in
figs. 54-66) Fig. 7
1 7b. Inflorescences axillary on semisucculent or woody stems to 2 m tall 18
1 8a. Locules with 2 or more ovules; fruits with many seeds (Hoffmannia spp.) 19
1 8b. Locules with 1 ovule; fruits usually 2-seeded 22
19a. Leaves 3/node or petioles with vesicles Fig. 8
19b. Leaves 2/node, petioles without vesicles 20
20a. Species conspicuously pubescent Figs. 7, 9
20b. Species mostly glabrescent 21
2 la. Leaves larger and decurrent Figs. 9-10
21b. Leaves various, inflorescences smaller Fig. 11
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
22a. Ovary usually with 8 locules (Lasianthus) Fig. 9
22b. Ovary with 2 locules (Psychotria spp., but note that Psychotria aubletiana with sessile in-
volucrate axillary capitulae is not illustrated) Figs. 12-13
23a. (from Ib) Plants vines or lianas 24
23b. Plants shrubs, trees, or subshrubs 26
24a. Slender-stemmed vines (Manettia and Sabicea) Figs. 1, 35
24b. Woody climbers or lianas 25
25a. Inflorescences pedunculate globose capitula; stems with sharp recurved spines; leaf blades
without parallel or lineolate 3° venation (Uncarid) Fig. 37
25b. Inflorescences never globose capitula, with many small flowers in panicles (and leaves with parallel 3° venation in Malanea, not illustrated) or with larger (> 2 cm) flowers in few- flowered inflorescences in species ofHillia and Randia; spines sometimes present in Randia.
26a. Epiphytic shrubs and small trees 27
26b. Terrestrial shrubs or trees 29
27a. Flowers small (< 15 mm), ovary with 4 locules; fruits baccate with 4 pyrenes (Psychotria
spp.) Fig. 60
27b. Flowers large (> 15 mm), ovary with 2-4 locules; fruits elongate capsules with many seeds
or united at the base into a partial syncarp 28
28a. Corolla tube less than 2 cm long; fruits united at the base, fleshy (Schraderd) Fig. 19
28b. Corolla tubes more than 2 cm long; fruits free at the base, elongate capsules (Cosmibuena
with winged seeds and Hillia with a tuft of hairs at 1 end of the seed) Figs. 27-28
29a. Larger leaves usually becoming 40-50 cm long, entire or lobed in a few species; fruits many-seeded
30
29b. Larger leaves not usually becoming 40-50 cm long, never lobed; fruits 1 -many-seeded 33
30a. Minor venation of the leaves subparallel (lineolate), some species with deeply lobed leaves;
fruits baccate or hard, seeds angular (Pentagonid) Fig. 14
30b. Minor venation reticulate, leaves entire or with small lobes; fruits capsular, seeds mostly flat
30
3 la. Stipules almost separate, 4/node; inflorescences with few 1° branches and no bracteoles
(Condamined) Fig. 29
31b. Stipules united, 1-2/node; inflorescences with many 1° lateral branches and small bracteoles
32
32a. Fruits small, ca. 4 mm long (Elaeagid) Fig. 39
32b. Fruits large, ca. 5 cm long (Simird) Fig. 38
33a. Flowers with corolla tubes more than 10 cm long, white (native species not found in gardens) . .
34
33b. Flowers with corolla tubes less than 10 cm long (or, if close to 10 cm, red and planted for ornament)
36
34a. Flowers funnelform distally, with a gradually expanded tube, corolla lobes broadly triangular
(Osa) Fig. 15
34b. Flowers salverform distally and with a narrow tube throughout, corolla lobes narrowly ovate
to oblong 35
35a. Leaves narrowly elliptic, to 14 cm long, low shrubs of streamsides (Lindenid) .... Fig. 15 35b. Leaves not narrowly elliptic, usually more than 14 cm long, shrubs and trees of forests
(Posoquerid) Fig. 15
36a. Inflorescences with some calyx lobes greatly expanded to form large leaf-like red or white "petals"
37
36b. Inflorescences without calyx lobes greatly expanded (slightly expanded in some spp. of Rondeletid)
40
37a. Inflorescences racemose, to 60 cm long, expanded calyx lobes brilliant red (Warszewiczid)
Fig. 16
37b. Inflorescences not racemose, to 1 5 cm long, expanded calyx lobes white to pinkish red or dull red . . 38
10 FIELDIANA: BOTANY
38a. Expanded calyx lobes densely pubescent, dull red (white in some forms); cultivated (Mus-
saendd) Fig. 16
38b. Expanded sepals glabrous to glabrescent; calyx lobes white or red; native and also planted
for ornament 39
39a. Corolla tube 3 mm long, white (Calycophylluni) Fig. 16
39b. Corolla tube 25 mm long, red (Pogonopus) Fig. 16
40a. Inflorescences dense heads of closely packed flowers 41
40b. Inflorescences lacking dense heads, if subcapitate the flowers becoming separate in fruit .... 43 4 la. Flowers united or connivent at the base, an involucre of bracts absent (Appunia, Morinda,
Schraderd) Fig. 19
41b. Flowers not united or connivent at the base, an involucre present or absent 42
42a. Inflorescences subtended by an involucre of colorful large bracts; native trees and shrubs
(Psychotria spp.) Figs. 7, 17-18, 56
42b. Inflorescences spherical, without an involucre; introduced trees (Neolamarckia, not illus- trated).
43a. Inflorescences long and narrow (racemiform to spiciform) 44
43b. Inflorescences not long and narrow 46
44a. Flowers arising separately and sessile, inflorescences spicate; rare in Costa Rica (Alseis sp.)
Fig. 40
44b. Flowers usually in small cymose groups, these often on short secondary peduncles, inflores- cences racemose 45
45a. Fruits fleshy (Gonzalagunid) Figs. 20.-21
45b. Fruits dry dehiscent capsules (Rondeletia) Fig. 21
46a. Flowers solitary or few at the ends of branches or short shoots, with short inconspicuous peduncles when few; fruits usually solitary at the tips of branches, usually large (-1-2 cm) and rounded; seeds many, often imbedded in pulp or horizontal (note: Faramea luteovirens and Rudgea monofructus, with few-seeded fruits and flowers resembling those in fig. 46, and Serissa, a small ornamental
shrub with short stiff leaves, are not illustrated) 47
46b. Flowers not solitary at the ends of branchlets, on well-developed peduncles when few; fruits rarely
solitary and terminal 48
47a. Ovaries unilocular with parietal placentation (but difficult to see, with placentas often fusing in the center), seeds variously oriented in white pulp turning black; spines present in some
species (Randia spp.) Figs. 22-24
47b. Ovaries usually 2-8-locular (but the septa often difficult to see), seeds mostly horizontal; spines absent in all species (other genera of Gardenieae; see the technical key) Figs. 25-26
48a. Corollas 3-10 cm long 49
48b. Corollas less than 3 cm long 58
49a. Fruits elongate and flattened or cigar-like capsules, seeds many and winged 50
49b. Fruits not elongated capsules, seeds not winged 52
50a. Flowers sericeous on the exterior (Ladenbergia and Cinchona) Figs. 29, 37
50b. Flowers glabrous on the exterior 51
51a. Seed with a tuft of hairs (Hillid) Figs. 27-28
5 Ib. Seed without hairs (Cosmibuend) Figs. 27-28
52a. Flowers usually axillary and solitary (Exostema caribaeum) Fig. 31
52b. Flowers neither axillary nor solitary 53
53a. Corollas inflated in bud; capsules flattened and opening on the broad side (Coutared)
Fig. 31
53b. Corollas not inflated in bud; capsules not flattened or with fleshy fruits 54
54a. Flowers red to purplish; garden ornamentals Fig. 31
54b. Flowers white or yellowish; native species 55
55a. Ovule 1 in each locule (Guettarda turrialband) Fig. 30
55b. Ovules many in each locule 56
56a. Corolla densely sericeous (Duroia and Amaioud) Figs. 25, 30
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 1 1
56b. Corolla glabrous on the exterior 57
57a. Corolla lobes convolute; ovary 2-locular; fruits ca. 9 cm diam. (Tocoyena) Fig. 30
57b. Corolla lobes valvate; ovary 2-6-locular; fruits ca. 1 cm diam. (Isertid) Fig. 30
58a. (from 48b) Inflorescences axillary; fruits from axillary peduncles; corollas mostly pubescent on the
exterior (glabrous in Chimarrhis and some species of Hoffmannid) 59
58b. Inflorescences terminal; fruits from terminal peduncles (or from pseudoaxillary peduncles when
lateral shoots continue growth); corollas glabrous or puberulent 69
59a. Ovule 1/locule; seeds 1 /chamber in a bony endocarp, fruits drupaceous; flowers often along 1 side of the inflorescence branches; minor leaf venation parallel in some Chomelia and
Guettarda spp 60
59b. Ovules 3-many/locule; fruits many-seeded berries and capsules; flowers mostly cymose;
minor leaf venation parallel in Sommera 65
60a. Fruits small woody capsules with many seeds 61
60b. Fruits fleshy, drupes or berries with 1-2 pyrenes or with a single stony endocarp . . 62 6 la. Capsules ca. 5 mm long, rounded; corollas 3-5 mm long (Chimarrhis) . . Fig. 37 61b. Capsules 9-20 mm long, elongate; corollas 7-14 mm long (Macrocnemum) . . .
Fig. 40
62a. Fruits usually with 2 pyrenes 63
62b. Fruits with a single hard endocarp with 2-5 locules [fruits not flattened or economically
useful] 64
63a. Fruits usually flattened laterally and with a lustrous white surface [corolla tubes
< 9 mm long; native plants] (Chiococcd) Fig. 36
63b. Fruits rounded, becoming red 64
64a. Corolla tubes > 10 mm long; widely cultivated (Coffea, not illustrated).
64b. Corolla tubes < 1 5 mm long; wild or rarely cultivated (species of Psychotrid)
Figs. 12-13
65a. Corolla lobes valvate or slightly imbricate in bud, often with a distal appendage (Cho- melia) Figs. 33-34
65b. Corolla lobes broadly imbricate with 1-2 exterior lobes, lacking distal appendages
(Guettarda) Figs. 32-34
66a. (from 59b) Plants generally with few lateral branches, weak subshrubs to 3 m tall (Hoffmannid)
Figs. 7-11
66b. Plants trees, shrubs, or clambering 67
67a. Fruits capsular (Chimarrhis) Fig. 37
67b. Fruits berry-like 68
68a. Stems clambering; locules 3-5 (Sabiced) Fig. 35
68b. Trees and shrubs; locules 2 (Sommera) Fig. 35
69a. (from 58b) Fruits dry and mostly capsules; ovaries with usually more than 1 ovule per locule . .
70
69b. Fruits fleshy or with arenchymatous tissue; ovaries with 1-many ovules per locule 76
70a. Fruits dry samaras with rounded circumferential wings, red and making a colorful display; rarely collected trees (Allenanthus, not illustrated).
70b. Fruits not flattened samaras with a winged margin 71
7 la. Flowers small (— 1 cm) and often closely congested in dense inflorescences or in small groups
on open branched inflorescences 72
71b. Flowers usually more than 1 cm long, not usually closely congested in the inflorescence . .
75
72a. Spines often present; capsules opening from the base (Machaonid) Fig. 37
72b. Spines absent; capsules opening from the top 73
73a. Anthers opening by terminal pores; leaves with pellucid dots (Rustid) Fig. 38
73b. Anthers opening by lateral slits; leaves lacking pellucid dots 74
74a. Corolla yellow; shrubs or small trees of higher elevations (Depped) Fig. 38
74b. Corolla white; medium to large trees of low and high elevations (Elaeagid) . . . Fig. 39
12 FIELDIANA: BOTANY
75a. Capsules usually small (2-5 mm) and rounded; corollas often pubescent (Rondeletia spp.)
Figs. 41-41A
75b. Capsules usually more than 9 mm long and elongated, rounded or flattened; corollas puber-
ulent (Cinchona and Exostemd) or glabrous (Ferdinandusa and Macrocnemum)
Figs. 37, 40
(from 69b) Ovaries with 3-many ovules per locule; fruits usually many-seeded 77
Ovaries with 1 ovule per locule; fruits with 1-2 seeds (usually 5 in Psychotria racemosd) ... 80
77a. Anthers with the thecae divided into small sections [ovaries 2-6-locular] (Isertid)
Figs. 30, 49
77b. Anthers with the thecae not divided into sections 78
78a. Ovaries and fruits 5-locular (Hamelid) Figs. 34, 42
78b. Ovaries and fruits 2-locular 79
79a. Corolla lobes valvate in bud; inflorescences with cymose branches (Raritebe) Fig. 43
79b. Corolla lobes contorted in bud; inflorescences with flowers along 1 side of branches (Bertierd)
Fig. 43
(from 76b) Fruits usually with only 1 pyrene (seed), rounded in cross-section; the ovary 2-locular in early stages, with a thin septum or with only 1 locule; flowers white to brilliant blue or lavender
(rarely yellowish) 81
Fruits usually with 2 pyrenes or seeds, the pyrene plano-convex in cross-section; the ovary 2-locular and with a well-developed septum in early stages; flowers white to yellow, orange, red, purple, or
bluish purple (rarely blue) 84
8 la. Flowers brilliant blue, blue-lavender, or white; fruits usually broader than long to globose;
stipules acute to long-awned at the apex 82
8 1 b. Flowers white to yellowish white; fruits usually longer than broad; stipules obtuse to acute,
not awned 83
82a. Larger-leaved species of Faramea Fig. 44
82b. Smaller-leaved species of Faramea Fig. 45
83a. Smaller-leaved species of Coussarea Figs. 46-47
83b. Larger-leaved species of Coussarea Figs. 48
Corolla lobes contorted in bud, corollas white to red; wild plants and cultivated ornamentals
(Ixord) Fig. 43
Corolla lobes valvate in bud, corollas white to red, yellow, or purple; plants not cultivated orna- mentals 85
Stipules usually rounded distally and with several short indurated tooth-like appendages at the
apex; inflorescences often few-flowered (Rudged) Fig. 46
Stipules not rounded distally and with thickened tooth-like structures at the apex; inflorescences
with few to many flowers 86
Flowers usually brightly colorful, yellow to orange, purple, or bluish purple (rarely white), corolla tubes often slightly enlarged on 1 side at the base, a ring of hairs present on the interior of the
swollen lower half of the corolla tube (Palicourea spp., see also fig. 54) 87
Flowers usually white or slightly yellowish, corolla tubes not expanded on 1 side at the base, a ring of hairs not present in the cylindrical lower half of the interior of the corolla tube (Psychotria
spp.) 88
87a. Palicourea spp.: inflorescences with conspicuous bracts and a species with spathaceous calyx
Fig. 49
87b. Palicourea spp.: lowland species Fig. 50
87c. Palicourea spp.: flowers mostly yellow-orange Fig. 51
87d. Palicourea spp.: flowers mostly purple-violet Fig. 52
87e. Palicourea spp.: smaller-leaved species Fig. 53
Fruits becoming blue, purple, or black (orange in P. racemosa with 5 pyrenes, red in P. haema- tocarpa with very small inflorescences); leaves drying greenish to brown (except when treated with isopropyl alcohol), domatia usually absent (except P. acuminata); stipules often persisting and not subtending a ring of reddish colleters (subgenus Heteropsychotria and other species) 89
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 13
88b. Fruits becoming red at maturity; leaves drying grayish, grayish pink, or reddish brown to black, domatia present in a few species; stipules usually caducous and enclosing a ring of reddish colleters
at their base (subgenus Psychotrid) 90
89a. Heteropsychotria: large-leaved and pubescent Fig. 54
89b. Heteropsychotria: smaller-leaved species Fig. 55
89c. Heteropsychotria: smaller inflorescences Fig. 56
89d. Heteropsychotria: large open inflorescences Fig. 57
89e. Heteropsychotria: conspicuous inflorescences Fig. 58
89f. Heteropsychotria: deciduous and other species Fig. 59
90a. Subg. Psychotria: species with very small leaves Fig. 60
90b. Subg. Psychotria: species with small leaves Fig. 61
90c. Subg. Psychotria: species with Ficus-\ike stipules Fig. 62
90d. Subg. Psychotria: species with dense pubescence Fig. 63
90e. Subg. Psychotria: species with larger leaves Fig. 64
90f. Subg. Psychotria: deciduous and unusual species Fig. 65
90g. Subg. Psychotria: miscellaneous unusual species Fig. 66
14 FIELDIANA: BOTANY
Manettia reclinata
Arcytophyllum muticum
FIG. 1 . Twining shrubs (Manettia spp.) and subshrubs with small stiffleaves (species of Arcytophyllum, Declieuxia, and Diodia).
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
15
Coccocypselum hirsutum
C. cordifolium
Coccocypselum lanceolatum
Geophila repens
G. cordifola
Geophila macropoda
FIG. 2. Twining herbs: species of Coccocypselum and Geophila.
16
FIELDIANA: BOTANY
Galium aschenbornii
Oldenlandia
corymbosa
FIG. 3. Herbs with small leaves and slender stems: species of Didymaea, Galium, Nertera, and Oldenlandia.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
17
Spermacoce verticillata
S. suaveolens
S. densiflora
Richardia scabra
Crusea parviflora
Mitracarpus hirtus
FIG. 4. Erect herbs with narrow lanceolate leaves and capitate or verticillate flowers: species of Crusea, Mitracarpus, Richardia, and Spermacoce.
18
FIELDIANA: BOTANY
Spermacoce exilis
Spermacoce prostrata
10 cm
Spermacoce ovalifolia
S. assurgens
latifolia
confusa
FIG. 5. Erect herbs with narrow lanceolate leaves: Spermacoce spp.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
19
D. serrulata
cJi
FIG. 6. Erect herbs with narrow lanceolate leaves: Diodia spp. and two species of Spermacoce.
20
FIELDIANA: BOTANY
Hoffmannia davidsoniae
Amphidasya ambigua
Psychotria guapilensis
Lasianthus panamensis ^ GJ3
FIG. 7. Herbs or subshrubs with larger leaves: species of Amphidasya, Hoffmannia, Lasianthus, and Psychotria.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
21
Hoffmannia
subauriculata
H. vesiculifera
FIG. 8. Herbs or subshrubs with axillary flowers: unusual species of Hoffmannia.
22
FIELDIANA: BOTANY
FIG. 9. Herbs or subshrubs with axillary flowers: pubescent species of Hqffmannia and H, congesta. BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE 23
Hoffmannia leucocarpa
11111111111 10cm
H. dotae
<J5
H. asclepiadea
FIG. 10. Subshrubs with axillary flowers: species of Hoffmannia with larger leaves.
24
FIELDIANA: BOTANY
H. hamelioides
H. longipetiolata
H. psychotriifolia
Hoffmannia pallidiflora
H. inamoena
H. laxa
FIG. 1 1 . Subshrubs with axillary flowers: species of Hoffmannia with leaves tapering gradually to the base.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
25
P. macrophylla
sycnotria aggregata Psychotria cartagoensis
Psychotria uliginosa 10 cm
FIG. 12. Subshrubs with axillary flowers: species of Psychotria.
26
FIELDIANA: BOTANY
FIG. 1 3. Subshrubs with axillary flowers: species of Psychotria.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
27
Pentagonia donnell-smithii
cm
FIG. 14. Trees with very large or lobed leaves: three species of Pentagonia.
28
FIELDIANA: BOTANY
Osa pulchra
Lindenia rivalis
Posoqueria grandiflora
FIG. 15. Flowers with very long corolla tubes: species of Lindenia, Osa, and Posoqueria.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
29
Calycophyllum candidissimum
Pogonopus speciosus
Warszewiczia coccmea
ussaenda erythrophylla
FIG. 1 6. Inflorescences with greatly expanded petal-like calyx lobes: species of Calycophyllum, Mussaenda, Pogono- pus, and Warszewiczia.
30
FIELDIANA: BOTANY
Psychotria poeppigiana
FIG. 1 7. Inflorescences of involucrate heads: species of Psychotria (formerly Cephaelis spp.).
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
31
Psychotria glomerulata
c-8
FIG. 18. Inflorescences of involucrate or conspicuously bracteate heads: species of Psychotria.
32
FIELDIANA: BOTA1
Appunia guatemalensis
Schradera costaricensis
FIG. 19. Inflorescences of compact heads with flowers connivent at the base: species of Appunia, Morinda, and Schradera.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
33
Gonzalagunia panamensis
ovatifolia
U9
FIG. 20. Inflorescences long and narrow: species of Gonzalagunia.
34
FIELDIANA: BOTANY
Rondeletia brenesii
Rondeletia buddleoides
••••••••••
Rondeletia urophylla
Gonzalagunia bracteosa
FIG. 2 1 . Inflorescences long and narrow: species of Gonzalagunia and Rondeletia.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
35
Randia brenesii
Randia loniceroides
Randia thurberi
Randia aculeata
R. armata
FIG. 22. Fruits usually terminal and solitary: species of Randia with small leaves.
36
FIELDIANA: BOTANY
Randia altiscandens
FIG. 23. Fruits usually terminal and solitary: species of Randia with medium-sized leaves.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
37
R. genipoides
FIG. 24. Fruits usually terminal and solitary: species of Randia with larger leaves.
38
FIELDIANA: BOTANY
FIG. 25. Fruits usually terminal and solitary: species ofAlibertia, Duroia, Genipa, and Hippotis.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
39
Genipa americana
FIG. 26. Fruits usually terminal and solitary: species of Borojoa and Genipa.
40
FIELDIANA: BOTANY
FIG. 27. Plants usually epiphytic: species of Cosmibuena and Hillia with smaller leaves.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
41
Hillia macrophylla
Cosmibuena macrocarpa
FIG. 28. Plants usually epiphytic: species of Cosmibuena and Hillia with larger leaves.
FIELDIANA: BOTANY
Ladenbergia brenesii
Condaminea corymbosa
L. sericophylla
FIG. 29. Trees with large open inflorescences: species of Ladenbergia and Condaminea corymbosa.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
43
Tocoyena pittieri
FIG. 30. Inflorescences with clusters of long-tubular flowers: species ofAmaioua, Guettarda, Isertia, and Tocoyena.
44
FIELDIANA: BOTANY
Exostemma caribaeum
FIG. 3 1 . Showy flowers: species of Coutarea, Crusea, Exostema, Ixora, and Pentas.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
45
Guettarda crispiflora
FIG. 32. Inflorescences with scorpioid or helicoid branches: species of Guettarda.
46
FIELDIANA: BOTANY
Guettarda foliacea
G. brenesii
FIG. 33. Flowers with narrow corolla tubes: species of Guettarda and a species of Chomelia.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
47
Chomelia microloba
Hamelia rovirosae
Chomelia recordn Chomelia spinosa
FIG. 34. Flowers with narrow corolla tubes: species of Chomelia, Guettarda, and Hamelia.
48
FIELDIANA: BOTANY
Sabicea panamensis
Sommera donnell-smithii
FIG. 35. Inflorescences mostly axillary: species of Sabicea (vines) and Sommera (trees).
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
49
FBCX 3*. b
species of Chucocca aad a species of Omome.
FIELDIANA;
Machaonia martinicensis
Chimarrhis latifolia
FIG. 37. Many small flowers in dense inflorescences: species of Chimarrhis, Cinchona, Machaonia, and Uncaria.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
51
Rustia costaricensis
A
Simira maxonii / \ Rustia occidentalis
FIG. 38. Flowers in much-branched open inflorescences: species of Deppea. Rustia, and Simira.
52
FIELDIANA: BOTANY
FIG. 39. Small flowers in dense or open panicles: species of Elaeagia.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
53
Alseis sp. aff. hondurensis
Exostemma mexicanum
Ferdinandusa panamensis —f^~ ^Macrocnemum glabrescens
FIG. 40. Small flowers and capsular fruits: species of Alseis, Exostema, Ferdinandusa, and Macrocnemum.
54
FIELDIANA: BOTANY
R. monteverdensis
FIG. 4 1 . Rondeletia spp.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
55
Rondeletia povedae
Rondeletia chaconii
R. hamelifolia
FIG. 41 A. Rondeletia spp.
56
FIELDIANA: BOTANY
axillaris
FIG. 42. Hamelia spp.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
57
Ixora nicaraguensis
Raritebe palicoureoides
FIG. 43. Bertiera, Ixora, and Raritebe spp.
58
FIELDIANA: BOTANY
FIG. 44. Faramea: species with larger leaves.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
59
parvibractea
6J3
FIG. 45. Faramea: species with smaller leaves.
60
FIELDIANA: BOTAN\
R. cornifolia
Rudgea reducticalyx
C. impetiolaris
R. trifurcata
FIG. 46. Coussarea and Rudgea spp.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
61
Psychotria umbelliformis
FIG. 47. Coussarea spp. and two similar Psychotria spp
62
FIELDIANA: BOTANY
Coussarea talamancana
FIG. 48. Coussarea: species with larger leaves.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
63
tilaranensis
FIG. 49. Palicourea: species with conspicuous bracts.
64
FIELDIANA: BOTANY
Palicourea guianensis
1 cm
P. crocea
FIG. 50. Palicourea: species of lower elevations and a species of Isertia.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
65
Ufl
P. lasiorrhachis
FIG. 5 1 . Palicourea: species with larger leaves and yellow or orange flowers.
66
FIELDIANA: BOTANY
Palicourea purpurea
FIG. 52. Palicourea: species with larger leaves and blue, lavender, purple, or white flowers.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
67
FIG. 53. Palicourea: species with smaller leaves.
68
FIELDIANA: BOTANY
Palicourea standleyana
i i i i i i i i i I i i i i i i i i i I
FIG. 54. Psychotria subg. Heteropsychotria: larger-leaved pubescent species, and a species of Palicourea.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
69
Psychotria goldmanii , ^
rt.
P. steyermarkii
70
FIG. 55. Psychotria subg. Heteropsychotria: species with smaller leaves.
FIELDIANA: BOTANY
Psychotria haematocarpa
deflexa
FIG. 56. Psychotria subg. Heteropsychotria: species with very small inflorescences.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
71
Psychotria tapantiensis
FIG. 57. Psychotria subg. Heteropsychotria: species with larger open inflorescences.
72
FIELDIANA: BOTANY
berteriana
FIG. 58. Psychotria subg. Heteropsychotria: species with conspicuous open inflorescences.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
73
2 mm
irticinaiis ^v^ — ---^ — -— ^»C*TI . .••••'_ . vi-i*ssss5Sfc.J/ T •x.'^-^^x" rgcsmosa
FIG. 59. Psychotria subg. Heteropsychotria: species of deciduous habitats and some with smaller inflorescences.
74
FIELDIANA: BOTANY
Psy. guadalupensis (sensu lato)
FIG. 60. Psychotria subg. Psychotria: species with very small leaves and a complex of epiphytic species.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
75
marginata
Psychotria monteverdensis
FIG. 6 1 . Psychotria subg. Psychotria: species with smaller narrow leaves.
76
FIELDIANA: BOTAN\
Psychotria mexiae
FIG. 62. Psychotria subg. Psychotria: high-elevation species and those with Ficus-like stipules.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
77
FIG. 63. Psychotria subg. Psychotria: densely pubescent species.
78
FIELDIANA: BOTANY
panamensis var. compressicaulis
FIG. 64. Psychotria subg. Psychotria: large-leaved species.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
79
Psychotria rosulatifolia
viridis
FIG. 65. Psychotria subg. Psychotria: deciduous and unusual species.
80
FIELDIANA: BOTAW
Psycnotna lamarinensis
FIG. 66. Psychotria subg. Psychotria: several unusual species.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
81
Alibertia A. Richard
Trees or shrubs, dioecious, lacking spines, glabrous or sparsely puberulent; stipules interpetiolar (and intrapeti- olar in a few spp.), acute, usually persisting. Leaves op- posite, petiolate; leaf blades entire, often with domatia. Inflorescences terminal, 6 flowers fasciculate or capitate, 9 flowers solitary or 2, flowers of both sexes sessile or subsessile, subtended by persisting stipule-like bracts. Flowers unisexual, (3-)4-5(-8)-parted, hypanthium hemispheric (in 9 flowers) to tubular (in <5 flowers), calyx tube truncate or dentate; corolla salverform, fleshy or subcoriaceous, corolla tube cylindrical, glabrous or vil- lous within, 3-8-lobed, lobes short to long, obtuse to acute, convolute in bud; stamens 3-8, filaments short or absent, anthers linear, dorsifixed, included within the
tube; ovary 2-8-locular, ovules 3-many on axile placen- tas within each locule, few-seriate to multi-seriate, often imbedded in a pulpy placenta. Fruits terminal and sol- itary, baccate, fleshy, sessile or subsessile, globose, usu- ally over 2 cm diam., 2-8-locular with thin septa, peri- carp firm and coriaceous, placentas becoming pulpy; seeds usually many, compressed to rounded.
A genus of about 35 species, ranging from Mex- ico, the West Indies, and Central America into South America. Alibertia is recognized by its dioe- cy, sessile or subsessile terminal flowers, and the large terminal subglobose fruit with a persisting terminal calyx tube. This genus is similar to Amaioua and Borojoa.
Key to the Species of Alibertia
la. Stipules 7-20 mm long; flowers 16-34 mm long; leaves elliptic-oblong; usually tapering gradually to the apex A. edulis
Ib. Stipules to 4 mm long; flowers 7-10 mm long; leaves usually somewhat obovate, abruptly narrowed to an acuminate or rounded apex A. garapatica
Alibertia edulis (L. C. Rich.) A. Rich, ex DC., Prodr. 4: 443. 1830. Genipa edulis L. C. Rich., Actes Soc. Hist. Nat. Paris 1: 107. 1792. Gar- denia edulis (L. C. Rich.) Poir. in Lam., Encyc. Meth. Bot. Suppl. 2: 708. 1812. Figure 25.
Shrubs or rarely small trees, l-4(-6) m tall, sometimes forming thickets, bark often peeling in longitudinal strips, leafy branchlets (1.5-)2-4.5 mm thick, glabrous or sparsely puberulent, reddish brown and becoming gray- ish; stipules 7-15(-20) mm long, 2-4(-5) mm broad at base, triangular to deltoid, acute or acuminate, longi- tudinally striate, acute or acuminate, brownish and gla- brous, subcoriaceous, persisting with older leaves. Leaves with petioles 2-5(-10) mm long, 0.5-1.8 mm thick, gla- brous; leaf blades 5-14(-20) cm long, 1 .5-5(-8) cm broad, narrowly elliptic-oblong to broadly ovate-oblong, apex acute to long-acuminate, base gradually narrowed and attenuate (in narrow leaves) to rounded and subtruncate (in broader leaves from Caribbean lowlands), drying stiff- ly chartaceous to subcoriaceous and often grayish green, lustrous above in life, glabrous on both surfaces (or pu- berulent beneath in material from Caribbean lowlands), 2° veins 6-1 2/side, small domatia of pits and tufted hairs often present in vein axils beneath. Inflorescences of subsessile <5 or 9 flowers, subtended by stipule-like bracts 7-15 mm long. Flowers 4- (rarely 5-) parted, 16-34 mm long, hypanthium 4-8 mm long, 3-5 mm diam., calyx tube truncated except for the short (0.3-1 .7 mm) narrow (0.3 mm) lobes; corolla white, tube 2-3 cm long, lobes to 2 cm long and 12 mm broad, triangular, minutely puberulent; stamens usually 4, subsessile, anthers ca. 1 3 mm long in <5 flowers; staminodes 5 mm long in 9 flowers. Fruits 2-3 cm diam.. obovoid to subglobose, yellowish brown, slightly truncated distally, persisting calyx tube
4-6 mm long and 4-6 mm diam.; seeds 3-8 mm long, oblong, slightly flattened longitudinally, striate.
Shrubs of both deciduous and evergreen forest formations, from near sea level to 500(-1000) m elevation. Probably flowering and fruiting throughout the year, with the main flowering sea- son April-July. The species ranges from Mexico to northern South America.
Alibertia edulis is recognized by its solitary ter- minal rounded fruit with persisting calyx tube, generally narrow oblong-elliptic leaves, stiff striate interpetiolar stipules, and generally shrubby habit. In Central America the species is most common in deciduous or partially deciduous woodland. Common names for this species in Central Amer- ica are lagartillo, trompillo, trompo, trompito, and "wild guava." The fruit is occasionally eaten by local people and sporadically cultivated. This spe- cies may intergrade with material currently placed under other names in South America; compare A. acuminata (Benth.) Sandwith and A. latifolia (Benth.) Schum. Specimens may resemble some species of Randia.
Material from the Caribbean lowlands placed under this name differs in having broader leaves that dry dark and have short straight hairs on their undersurfaces. This material, while quite different from that found on the Pacific slope, appears to intergrade with the more typical forms in Guate-
82
FIELDIANA: BOTANY
mala and Belize; it is not often collected in Costa Rica.
Alibertia garapatica K. Schum. in Mart., Fl. Bras. 6(6): 384. 1889.
Shrubs or small trees to 5 m tall, leafy branchlets slender (1-2 mm thick) with slightly thickened nodes, terete, brownish, minutely (0. 1 mm) puberulent and gla- brescent; stipules 2-4 mm long, ca. 2 mm broad at the base, persisting or deciduous. Leaves with petioles 2- 7(-10) mm long, 0.7-1 .2 mm thick, minutely puberulent; leaf blades 5- 1 2(- 1 5) cm long, 2-5(-7) cm broad, broad- ly elliptic-obovate to broadly oblong-obovate or elliptic- oblong, apex abruptly narrowed and acuminate, caudate- acuminate or rounded, tip 5-15(-20) mm long, base obtuse to cuneate, drying stiffly chartaceous and usually grayish in color, glabrous above and below, 2° veins 5- 7/side, domatia of tufted hairs in the vein axils beneath (in Colombian material). Inflorescences terminal and capitate, sessile or subsessile, with 4-8(-16) S flowers, the 9 flowers solitary or paired, flowers subtended by 2 triangular stipules (bracts). Flowers aromatic, 6-10 mm long, hypanthium 1-2 mm long, ca. 1.3 mm broad, ob- conic, distal margin entire, calyx cup and teeth minute (0.5 mm) or absent; corolla tube 3-7 mm long, greenish, lobes white, 1.5-3 mm long, ca. 1.5 mm broad near the base; stamens 4. Fruits solitary, sessile or subsessile, glo- bose or subglobose to obovoid, ca. 25 mm long and 30 mm diam., drying black.
Trees of evergreen and partly deciduous for- mations on both the Caribbean and Pacific slopes in central Panama, from near sea level to 500 m elevation. In Costa Rica it is known only from near Punta Mala on the Pacific coast (A. Jimenez 3912, flowering in March) and the Reserva Biol. Carara (Zuniga 232, fruiting in May), both in southern Puntarenas Province. The species ranges from Costa Rica to Colombia and occurs in south- ern Mexico.
Alibertia garapatica is recognized by its terminal subsessile flowers and solitary fruit, small inter- petiolar stipules, relatively broad and slightly ob- ovate leaves that dry chartaceous, and smaller flowers. A short tube may be present on some stipules. This species is poorly known; it may be mistaken for some species of Randia.
Allenanthus Standley
Small to medium size trees, branchlets glabrous or sparsely puberulent; stipules interpetiolar, persistent or deciduous. Leaves opposite, short-petiolate, leaf blades acuminate, entire, often with minute domatia. Inflores- cences panicles with opposite branching, broadly cor- ymbose in form, terminal or axillary to distal leaves,
bracteate, flowers pedicellate. Flowers bisexual, small (3- 6 mm), hypanthium obovoid to urceolate, truncated dis- tally, laterally compressed, calyx lobes 4, small; corolla whitish, tubular and with 4 spreading lobes, valvate to somewhat imbricate; stamens 4, borne in the throat of the corolla tube, filaments slender, anthers oblong; ovary 2-locular, with 1 ovule in each locule, style distally bifid. Fruits becoming dry and samara-like, flattened with broad lateral wings surrounding the 2 central narrow longitu- dinally parallel seed chambers, material of the wings slightly spongy, calyx lobes persisting; seeds laterally compressed, pendulous.
A small genus with two species, ranging from central Mexico to western Panama. Allenanthus hondurensis Standley is found in central and southern Mexico and in Honduras; our species also appears to have a disjunct distribution in Cos- ta Rica and Panama. The flattened fruit, resem- bling that of Ulmus or some Terminalia species, is unique among Central American Rubiaceae.
Allenanthus erythrocarpa Stand!.. Ann. Missouri Bot. Gard. 27: 344. 1940. Chimarrhis decurrehs Steyerm., Ceiba 3: 18. 1952.
Trees, 6-20 m tall, leafy branchlets 1.5-5 mm thick, internodes 4-8 cm long, usually glabrous, subterete; stip- ules 3-6 mm long, 2-3 mm broad at the base, apex acute, puberulent within, deciduous. Leaves with petioles 6- 15 mm long, 1-1.5 mm thick, sulcate with adaxial mar- gins with punctate (gland-like) projections along the edge; leaf blades 6-1 1 cm long, 3-5 cm broad, ovate-elliptic to ovate-oblong, apex tapering gradually and acuminate, tip to 1 .5 cm long, base obtuse and slightly decurrent on petiole, drying thin chartaceous and sometimes dark in color, 2° veins 5-7 /side, glabrous above or puberulent only along the major veins, with small (0.2 mm) as- cending hairs on the major veins beneath, usually with small tufted domatia in slight depressions in the vein axils beneath (with 2-lipped structures ca. 1 mm long at the vein axils in Zamora & Poveda 825). Inflorescences both terminal and sometimes also axillary to distal leaves and together forming a single conspicuous panicle (thyrse) to 15 cm long and 10 cm broad, becoming 20 cm long and 18 cm broad in fruit, primary peduncles 3-7 cm long, terete, shorter toward apex of the inflorescence, peduncles and rachis with opposite lateral branching, with 1 or 2 longitudinal lines of dense short (0.2-0.4 mm long) ascending hairs, bracts 1-2 mm long, pedicels 1- 2 mm long. Flowers with the hypanthium 1-2 mm long, somewhat flattened (compressed), calyx lobes 4, 0.5-1 mm long; corolla becoming 4 mm long, tube ca. 3 mm long, lobes 4, rounded; stamens 4, exserted on slender filaments ca. 1.5 mm long, anthers ca. 0.8 mm long. Fruits flat and samara-like, pink to red, 5-7(-8) mm long, 3-4(-6) mm broad, oblong-elliptic in outline, the base of the fruit decurrent on pedicel, body of the fruit ca. 0.7 mm thick, fruiting pedicels ca. 4 mm long; seeds forming an oblong area in the center of the fruit ca. 2 mm long and 1 mm broad.
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
83
Trees of evergreen or partly deciduous forest formations of the Pacific slope, at around 500-700 m elevation. The species is known from near Par- que Nacional Rincon de la Vieja in Guanacaste Province (Herrera & Rivera 843 CR, MO, Zamora & Poveda 825 CR, F). Flowers were collected in June (Panama); fruiting in August-September (Panama) and October (Costa Rica). The species is known only from Costa Rica and western Pan- ama.
Allenanthus erythrocarpa is recognized by its flattened reddish samara-like fruit with small dis- tal calyx lobes, conspicuous infructescences, and unusual lines of hairs along branches of the inflo- rescences. With their broad terminal inflores- cences and bright red or pink fruits, these trees are very conspicuous when fruiting (Zamora & Po- veda 825 CR, F).
Allenanthus hondurensis Standl. of northern Central America is a smaller tree found in decid- uous forest, with the leaves more puberulent be- neath and smaller (5x3 mm) yellowish fruit.
Alseis Schott
Trees or large shrubs, branchlets terete, glabrous or more often puberulent; stipules interpetiolar, triangular to subulate, caducous or persisting. Leaves opposite, of- ten clustered at the ends of branchlets, petiolate; leaf blades often narrowly obovate, drying thin-chartaceous, sometimes with domatia. Inflorescences terminal or ax- illary, solitary in each axil, usually spike-like or racemose and cylindrical in form, simple or with lateral branches (paniculate and racemiform), flowers lacking pedicels or the pedicels merging gradually into the base of the ovary. Flowers bisexual, small, white to yellow, protogynous; hypanthium obconical to subcylindrical, calyx lobes 5, deciduous; corolla tube cylindrical to campanulate or urceolate, villous within, corolla lobes 5, valvate (?rarely open) in bud; stamens 5, filaments attached near the base of the corolla tube, anthers exserted, oblong and sagittate, dorsifixed; ovary 2-locular, septum thin, placentas apical with numerous ovules in each locule, style long, distally bifid with recurved stigmas. Fruits capsular, cylindrical, 2-locular and bivalvate, dehiscing septicidally from apex to base; seeds numerous, linear-fusiform, the testa re- ticulate and prolonged at apex and base.
A genus of about 20 species found in Mexico and Central America and southward to Peru and Brazil. The often long (ca. 20 cm) obovate leaves clustered at the ends of twigs, long (15-30 cm) spicate or racemiform inflorescences with many flowers, and narrow capsular fruit splitting into two parts distinguish members of this genus. Su- perficially, these plants may resemble some species
of Gonzalagunia and Rondeletia. Alseis blackiana Hemsl., with leaves to 30 cm long, is found in central and eastern Panama. Alseis hondurensis Standl. occurs in northern Honduras, Guatemala, and Belize, while A. yucatanensis Standl. occurs in southern Mexico, Belize, and Guatemala.
Alseis sp. aff. A. hondurensis Standl., Trop. Woods 16:48. 1928. Figure 40.
Small trees, ca. 15m tall, trunk ca. 30 cm dbh with soft bark, leafy branchlets 1.5-7 mm thick, glabrous, pale brownish, smooth; stipules 3-8 mm long, 1-2 mm broad at the base, triangular-subulate, apex acute, caducous. Leaves clustered at the ends of twigs, petioles 10-55 mm long, 0.8-2 m thick, glabrous and drying dark; leaf blades (7-)9-19 cm long, (2-)3-8 cm broad, elliptic-obovate to obovate or ovate-elliptic, apex short-acuminate, tip 4— 7 mm long, base acute (obtuse) and occasionally slightly decurrent on petiole, glabrous above, glabrous below ex- cept for some thin hairs ca. 0.7 mm long near the vein axils (domatia?), 2° veins 6-10/side. Inflorescences ter- minal or axillary, solitary or 3, 1 1-18 cm long, ca. 2 cm diam., spicate with flowers sessile on the rachis or with opposite basal spicate lateral branches, peduncles to 4 cm long, ca. 1 mm thick, minutely grayish puberulent, bracts and pedicels not apparent. Flowers with minute (0. 1-0.2 mm) tomentulous grayish hairs, hypanthium ca. 1.5 mm long and 0.6 mm thick, cylindric to obconic, calyx lobes ca. 0.5 mm high and 0.6 mm broad at the base, triangular, brownish and mostly glabrous; corolla white, short-tubular campanulate, 2-3 m long, corolla lobes little differentiated; stamens exserted, anthers ca. 0.8 mm long, style branches recurved. Fruit apparently narrowly obovoid and splitting into 2 valves, each valve ca. 8 mm long and 2.5 mm broad, with a notch 1 mm deep at apex, yellowish and smooth-lustrous within.
This species is presently known from only two collections. Flowering material was collected in February 1989 east of Bahia de Drake on the Osa Peninsula (Q. Jimenez et al. 670 CR, F, MO). Old fruit were collected in July (Hammel et al. 17120 CR, F, MO) at the Reserva Forestal El Cangrejo (near the road from Puriscal to Quepos) at ca. 400 m elevation in San Jose Province.
Alseis sp. aff. A. hondurensis is distinguished by its sessile flowers on spicate inflorescences (rarely paniculate with one or two lateral spicate branches near the base), longer petioles drying dark, and narrowly obovoid capsule splitting into two sep- arate valves. The type of A. hondurensis differs in the shorter petioles, minute puberulence on the lower leaf surfaces, the leaf blades more often ob- lanceolate with a gradually tapering base, and much larger minutely puberulent inflorescences. In ad- dition, A. hondurensis is a species of the Caribbean lowlands, whereas our species is found on the Pa-
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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-
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sessile clusters of fruit often on three terminal branches, densely sericeous flowers with lustrous retrorse hairs on the corolla, and leaves often with minute domatia. The ring of colleters just above the encircling stipule scar are often hidden by the pubescence. Though often collected in central Pan- ama and in Nicaragua, we have seen only a few collections of this species from northern Costa Rica: Q. Jimenez 376 CR from near Liberia and Zamora & Chacon 1355 CR from Refugio Cano Negro.
pedicellate fruit in terminal umbel-like groups, flowers coming directly from the apex of the shoot on usually unbranched stalks (pedicels), and broadly elliptic leaves with long thin hairs on the upper surface in early stages. The unusual glan- dular teeth (colleters) above the stipule scar near the apex of the stem are also distinctive. At pres- ent, this species appears to be limited to a rather narrow altitudinal range on the Caribbean slope in Costa Rica.
Amaioua pedicellata Dwyer, Ann. Missouri Bot. Gard. 67: 30. 1980. Figure 30.
Trees 5-10(-15) m tall, trunks to 22 cm dbh, leafy branchlets 1.6-6 mm thick, with appressed-ascending sericeous hairs 0.5-1 mm long, glabrescent, with a ring of linear colleters ca. 1 mm long encircling the node just above the stipule scar but breaking off early; stipules 8- 16(-30) mm long, cap-like and caducous, with dense ascending lustrous sericeous hairs. Leaves with petioles, 8-18 mm long, 1-1.8 mm thick, appressed puberulent; leaf blades 6-1 3(-l 9) cm long, 3-6(-l 0) cm broad, broadly elliptic to broadly elliptic-oblong or elliptic-obovate, apex usually short-acuminate, tip 5-10 mm long, base obtuse (occasionally acute) and somewhat decurrent on petiole, drying stiffly chartaceous and dark brown above, upper surface of the young leaves with scattered slender whitish appressed hairs to 2 mm long but these quickly falling and the mature upper surfaces glabrous, lower surfaces with thin ascending hairs 0.3-0.5 mm long on the major and minor veins, 2° veins 7-1 I/side, occasionally with domatia in vein axils beneath. Inflorescences terminal fascicles of 6-12 pedicellate flowers, the flowers usually on unbranched pedicels (rarely on peduncles bearing 2- 3 pedicellate flowers), later forming a sessile or umbellate cluster of long-pedicellate fruit, pedicels 3-8 mm long, with dense lustrous ascending sericeous hairs. Flowers with hypanthium and calyx tube ca. 4 mm long and 3 mm diam.. calyx lobes 3-5, 0.5-1.5 mm long, subulate or linear; corolla rose with pale greenish tube 7-9 mm long, 1-3 mm diam., densely whitish sericeous, lobes 7- 9 mm long, 3 mm broad at base, narrowly triangular. Fruits subglobose to oblong, 1 2- 1 7 mm long, 10-14 mm diam., red to dark reddish purple (but drying black), sparsely and minutely puberulent near the distal end, annular calyx scar 3—4 mm diam., fruiting pedicels 20- 35 m long, 1-1.5 mm thick.
Trees of wet evergreen forest formations of the Caribbean slope in Costa Rica, and both the Ca- ribbean and Pacific slopes in Panama, from 600 to 900 m elevation. Flowering in June-July; fruit- ing in February, September, and December (in Panama). The species is known only from central and southern Costa Rica and Code and Veraguas provinces in Panama.
Amaioua pedicellata is recognized by its long-
Amphidasya Standley
Small shrubs or herbaceous subshrubs, woody at the base, stems unbranched; stipules connate/interpetiolar, large, lobed distally or deeply laciniate with filiform seg- ments, persisting. Leaves often closely clustered near the ends of stems, often long-petiolate; leaf blades large, margins entire, domatia absent. Inflorescences terminal or axillary, cymose to capitate, short, flowers usually closely crowded, pedicels short. Flowers bisexual, hy- panthium oblong to turbinate, calyx lobes 4-6, often unequal, persisting; corolla tubular-salverform, corolla lobes 4-6, valvate in bud; stamens 4-6, borne on the middle or upper part of the corolla tube, filaments short, anthers linear, dorsifixed; ovary 2-locular, with axile bi- lobed placentas, many ovules in each locule. Fruits fleshy, indehiscent, crowned by the persistent calyx lobes; seeds many, angular, testa reticulate.
Amphidasya is a genus of about seven species, ranging from Costa Rica through Panama to Co- lombia, Venezuela, and northern Brazil. Our rep- resentative is distinguished among Costa Rican Rubiaceae by the short herbaceous habit, large and long-petiolate leaves, densely clustered flowers with long calyx lobes, and long corolla tube.
Amphidasya ambigua (Standl.) Standl., Field Mus. Nat. Hist., Bot. Ser. 11:181.1931. Sabicea am- bigua Standl., Publ. Field Columb. Mus., Bot. Ser. 7: 49. 1930. Figure 7.
Herbs, 10-40(-90) cm tall, erect or decumbent, stems woody at the base, leafy stems 3-7 mm thick, terete, minutely puberulent, glabrescent, brownish; stipules 1 2- 20(-40) mm long, ca. 4 mm broad at the base, united basal sheath 3-5 mm long, with long linear acute lobes, minutely and inconspicuously puberulent. Leaves clus- tered at the distal part of the stem, opposite or subop- posite, often rosette-like, petioles 1.5-5 cm long, 1.9-2.8 mm thick, with few minute appressed hairs or glabres- cent; leaf blades 12-28 cm long, 5-10 cm broad, nar- rowly elliptic-obovate to narrowly oblong-obovate or oblanceolate, apex abruptly narrowed and short-acu- minate, base gradually narrowed and cuneate-attenuate,
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long-decurrent on petiole, drying stiffly chartaceous, es- sentially glabrous above, sparsely puberulent with mi- nute (0. 1-0.2 mm) ascending hairs on the veins beneath or glabrous, 2° veins 15-25/side. Inflorescences densely crowded in the axils of distal leaves, to 5 cm long, base of the inflorescence not usually visible, with 4-20 closely crowded sessile or subsessile flowers, floral bracts 1-5 mm long, acute. Flowers 5- or 6-parted, the hypanthium 5-8 mm long, glabrous or minutely and sparsely papil- late-puberulent in later stages, calyx lobes 8-18 mm long, ca. 1.5 mm broad at the base, often unequal, glabrous on the surfaces and with minute (0. 1 mm) hairs along the edge; corolla narrowly salverform, white, puberulent on the exterior, corolla tube 32-50 mm long, 1-2.5 mm broad (dried), corolla lobes 6-18 mm long, triangular, acute. Fruits 8-12 mm long, 4-6 mm diam., cylindrical- oblong, with persisting calyx lobes, surface of the dried fruit bullate from pressure of the seeds within; seeds 0.3- 0.4 mm diam., foveolate.
Plants of steep slopes in the shade of evergreen rain forests on both the Caribbean and Pacific low- lands, collected at elevations of 10-800 m (to 1 500 m in Panama). Flowering in August and Novem-
ber-December; fruiting in March-July, Septem- ber, and November. The species ranges from Cos- ta Rica to Colombia.
Amphidasya ambigua is recognized by its short stature, distally fimbriate stipules, crowded long- petiolate leaves, crowded flowers with relatively long calyx lobes, and relatively long corolla tube. These plants resemble species of Paradrymonia in the Gesneriaceae (but the latter have superior ova- ries). Costa Rican material was earlier thought .to be a separate species, distinguished in the follow- ing key. However, recent collections from Panama and Colombia have produced many intermediate variants and resulted in a broader interpretation of A. ambigua. Note, however, that the preceding description is based on Costa Rican material and does not represent all the variation found within the more widely defined taxon. The following key outlines the differences between the western and eastern collections.
la. Leaves often elliptic-oblong, petioles 2-7 cm long and densely appressed-puberulent, major sec- ondary veins 12-18 on each side; calyx lobes densely puberulent, corolla tube 2-3 cm long; central Panama to Colombia.
Ib. Leaves usually slightly obovate, petioles 1.5-5 cm long and glabrescent, major secondary veins 1 5- 25 on each side; broad surfaces of the calyx lobes glabrous, corolla tube ca. 4 cm long; Costa Rica and western Panama.
Appunia Hooker f.
Shrubs or small trees, glabrous or puberulent; stipules interpetiolar and united at the base with the petioles, subulate-acuminate, persisting. Leaves opposite, short petiolate, leaf blades often lanceolate and acuminate, entire, mostly drying thin-chartaceous, domatia absent. Inflorescences terminal or axillary, capitate with a few flowers congested at apex of a short to long peduncle, flowers free and subtended by bracteoles but without pedicels. Flowers bisexual, small, white; hypanthium hemispheric or oblong, calyx tube short-cylindrical, usu- ally truncate, lobes absent or minute (5); corolla fun- nelform to urceolate, corolla lobes usually 5, valvate in bud; stamens 5, filaments short, anthers dorsiflxed and included; ovary 4-locular, each locule with 1 ascending ovule, style slender, stigma capitate. Fruits fleshy and baccate, sessile and loosely aggregated on apex of the peduncle, each fruit with 4 (or fewer) nutlets, each nutlet with 2 unequal locules (a seed-bearing locule and an empty locule).
A genus of about 1 0 species in Central and South America. Most authors have placed this genus into synonymy under Morinda, but that genus has the
basally united flowers developing into a fleshy syn- carp and two stigmas.
Appunia guatemalensis J. D. Smith, Hot. Gaz. 48: 294. 1909. Morinda guatemalensis (J. D. Smith) Steyerm., Mem. New York Bot. Gard. 23: 385. 1972. Figure 19.
Low or slender-branched shrubs, (0.5-)l-3(-4) m tall, leafy stems 1.2-4 mm thick, terete, glabrous or rarely minutely (0.0 1 mm) puberulent; stipules 1.5-3 mm long, 2-4 mm broad at the base, with a narrowed simple or bifid tip ca. 0.5 mm long, glabrous. leaves opposite, petioles 2-5 mm long, 0.8-1.6 mm thick, glabrous or sparsely and minutely (0.05 mm) puberulent; leaf blades 7-1 6 cm long, 3-7 cm broad, elliptic-oblong to narrowly obovate, apex acuminate to acute, base acute, drying grayish green to dark olive green and often lustrous above, glabrous above and below, 2° veins 4-6/side. Inflores- cences solitary in leaf axils (2/node), borne on glabrous peduncles 3-27 mm long and 0.5-1 mm thick (dried), capitula with 3-12 sessile and congested flowers sub- tended by triangular bracts ca. 1.5 mm long. Flowers
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glabrous, hypanthium and calyx tube ca. 2 mm long, 1.7 mm diam. distally, calyx lobes not developed; corolla ca. 1 5 mm long, white or greenish, corolla lobes 5-6 mm long, 1.5 mm broad at the base, becoming recurved, greenish within. Fruits 6-8 mm long, 4-6 mm diam., subglobose, sessile, purple to brownish or black.
Plants of low elevation in open grassy sites or thickets, 0-300 m elevation. Flowering in Janu- ary-September in northern Central America. The species ranges from central Mexico along the Ca- ribbean coast to southern Nicaragua and has been only rarely collected in the Pacific lowlands of northern Costa Rica.
Appunia guatemalensis is distinguished by its small capitate inflorescences on slender peduncles in the axils of leaves, sessile flowers and fruits, and usual lack of pubescence. The inflorescences are at first borne on very short peduncles, but these elongate during anthesis and fruiting. This species resembles Morinda royoc (flowers fused at the base), Psychotria erecta (blue fruits), and Alibertia gar- apatica (terminal sessile inflorescences). Appunia seibertii Standley of Panama has cuneate-decur- rent leaf bases. It is possible that the few collections from near Liberia represent disjunct individuals and not well-established populations.
Arcytophyllum Willdenow ex Schultes
REFERENCE— P. Mena V., Revision of the genus
Arcytophyllum (Rubiaceae, Hedyotideae). Mem. New York Hot. Gard. 60: 1-26. 1990.
Shrubs or small subshrubs, stems woody, erect or prostrate, usually with short internodes and congested leaves, nodes thickened; stipules united and interpetio- lar, entire to bifid or setose distally, persisting. Leaves opposite, small, often closely crowded and imbricate, sessile or subsessile; leaf blades entire, thick-coriaceous, glabrous, venation often obscure, domatia absent. Inflo- rescences terminal (sometimes apparently axillary to dis- tal leaf-like bracts), with cymose or clustered flowers on short peduncles, or of solitary flowers, pedicels short. Flowers bisexual, glabrous externally; hypanthium hemi- spheric to obovoid, calyx lobes 4(-5), often with glands between the lobes; corolla campanulate to funnelform, corolla lobes 4, often papillate-puberulent within, val- vate in bud; stamens 4, free portion of the filament emerging from between the corolla lobes; anthers dor- sifixed, exserted or partly included; ovary 2-locular, pla- centas borne on the septum, ovules 4-12/locule, style slender, stigmas 2. Fruits capsular, turbinate to subglo- bose, usually dehiscing septicidally and basipetally, 2-locular; seeds few, oblong and plano-convex to con- cave-convex, punctate.
A genus of 1 5 species, ranging from Costa Rica through Panama into the Andes as far south as Bolivia. These plants are distinguished by their small stiff ericoid leaves, short internodes, and small stature in paramo or similar open high-al- titude vegetation types. Standley (1938, p. 1273) suggested that the genus might be congeneric with Houstonia.
Key to the Species of Arcytophyllum
la. Plants shrub-like, rooting only at the base, with many erect branching stems to 70 cm tall; leaves 4-8 mm long; flowers in cymose groups on short peduncles A. lavarum
Ib. Plants prostrate with main stems rooting at the nodes, short erect stems less than 20 cm tall; leaves 3-6 mm long; flowers solitary on short leafy stems A. muticum
Arcytophyllum lavarum K. Schum. ex Standl., Contr. U.S. Natl. Herb. 18: 127. 1916. Mallos- toma lavarum (K. Schum.) J. D. Smith, Enum. PI. Guatem. 5: 36. 1899, nom. nud. (based on A. lavarum K. Schum. in herb.). A. chirropoense Suesseng., Bot. Jahrb. Syst. 72: 285. 1942. Fig- ure 1.
Subshrubs with creeping and erect woody stems, 10- 40(-70) cm tall, sometimes forming mats, with many erect branches, nodes thickened with the bases of per- sisting stipules and leaf bases, internodes 2-7(-18) mm long, leafy branchlets 0.5-1.5 mm thick, glabrous, with 4 longitudinal ridges, becoming silvery gray to black;
stipules 1-2.5 mm long, distal margin entire to erose or spiny, thickened at the base. Leaves sessile or with pet- ioles ca. 1 mm long, glabrous throughout, articulate at the base; leaf blades 4-8 mm long, 2—4 mm broad, ovate- elliptic to ovate-oblong or oblong, apex obtuse or round- ed, base obtuse to subtruncate, drying thick and coria- ceous, darker and lustrous above, with a rim of lustrous tissue along the edge beneath, midvein impressed above, other veins not visible above or below. Inflorescences to 2 cm long, usually with peduncles to 1.5 cm long, branches of the inflorescence sometimes subtended by leaf-like bracts, glabrous, flowers usually in cymose groupings (fasciculate), pedicels 0.5-2 mm long. Flowers ca. 7 mm long, hypanthium 1-1.5 mm long, obconic (turbinate) to hemispheric, calyx lobes 4, 1-1.5 mm long, ovate-oblong to triangular and persistent, often with 1-
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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
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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-
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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.
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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
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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-
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like blade 2-4 cm long and 1.5-3.5 cm broad, the blade suborbicular or reniform to broadly obovate, rounded distally, obtuse to truncate or subcordate at the base, white or pale greenish white, palmately veined, the pet- iole-like base 1-2.5 cm long; corolla 5-7 mm long, white, campanulate to funnelform, corolla tube 2-3.5 mm long, ca. 1.5 mm broad at the base and 3 mm broad distally, often densely villose at apex of the throat with erect hairs ca. 1 mm long, lobes 4, 3-4.5 mm long, 2-2.5 mm broad, becoming reflexed; stamens 4, filaments 1.5-2.5(-3.5) mm long, anthers 1.2-1.5 mm long; style 3-5 mm long, stigmas 1.3-2 mm long. Fruits (6-)8-12 mm long, 3-4 mm diam., oblong-cylindrical, sessile or subsessile, gla- brous or sparsely puberulent, with 8 longitudinal ribs; seeds 3-5 mm long, fusiform with wings at 2 ends, body of the seed ellipsoid, 1-1.5 mm long.
Conspicuous trees of deciduous and partly de- ciduous forest formations in the Pacific lowlands, from near sea level to about 450 m elevation (to 700 m elsewhere). Flowering in November-Feb- ruary and May; fruiting in January-August. The species ranges from central Mexico, Belize, and Guatemala, along the Pacific slope of central and southern Central America to Colombia and Ven- ezuela; it also occurs in the West Indies.
Calycophyllwn candidissimum is recognized by the bright whitish petaloid structures developed from the calyx of some flowers. The trees bear many inflorescences over their crowns, and the bright petal-like sepal lobes make a striking visual display when in full flower. The large size of these trees in deciduous woodland also contributes to the effect. The species has been called madrono, salamo, and surra in Costa Rica. The wood is hard and highly elastic and fine textured and finishes smoothly; it has been used for tool handles, ar- chery bows, and many other purposes (Standley, 1938).
Cephaelis Swartz
A poorly defined genus of about 100 species in the American tropics and southern Asia. The ge-
nus was distinguished by the involucrate heads of flowers, two-locular ovary with solitary basal ovules, and drupaceous fruits with two nutlets. Most authors now agree that the species of Cepha- elis are polyphyletic and cannot be clearly segre- gated from Psychotria (Taylor et al., 1 99 1 , p. 1 39). See the treatment of Psychotria (key 3) for species formerly placed in Cephaelis, and Figures 1 7 and 18.
Chimarrhis Jacquin
Trees, often growing to large size and with buttressed trunks; stipules interpetiolar and intrapetiolar, leaving a scar encircling the stem above the node (and above the petiole attachment), caudate to acuminate, persistent or caducous. Leaves opposite, often clustered at the ends of twigs, short-petiolate; leaf blades large- to medium- sized, sometimes with domatia (absent in our spp.). In- florescences solitary or paired in leaf axils (rarely pseu- doterminal), paniculate and often corymb-like in form, flowers in open cymose groupings, bracts present. Flow- ers bisexual and radially symmetrical, monomorphic, protogynous in Costa Rica, hypanthium cupulate to tu- bular, truncated to dentate distally, calyx lobes 5 (4) and very small or none; corolla funnelform, white, corolla tube short and broad, villous within, corolla lobes 5 (4), valvate in bud; stamens 5 (4), borne on the throat of the corolla tube between the corolla lobes, filaments slender and villous at the base, anthers dorsifixed, often exserted; ovary 2-locular, placentation peltate on the septum, ovules many in each locule, style short, stigmas 2, obtuse. Fruits capsular, small and woody, oblong, dehiscing septici- dally from apex and 2-valved; seeds many, compressed or angulate, horizontal, testa reticulate.
A genus of about 14 species ranging from Costa Rica into South America and in the West Indies. Chimarrhis is recognized by the axillary inflores- cences, small flowers with poorly developed calyx lobes, corolla tube villous within, and small rounded bivalved capsules with many horizontal seeds. The buttressed trunks, great height of some individuals, and the stipular scar encircling the stem above the nodes are also distinctive.
Key to the Species of Chimarrhis
la. Fruit 4—5 mm long; leaf blades 1 1-24 cm long and 6-1 1 cm broad; evergreen forests of the Pacific
lowlands C. latifolia
Ib. Fruit 1.5-2.5 mm long; leaf blades 5-15 cm long and 3-7.5 cm broad; Caribbean lowlands
C. parviflora
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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.
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Chiococca P. Browne
Shrubs, woody climbers or small trees, the branches often pendant or clambering, branchlets terete, glabrous or puberulent; stipules interpetiolar and slightly intra- petiolar (to form a very short tube or cup), usually cus- pidate, persistent. Leaves opposite, petiolate, entire, membranaceous to coriaceous, pinnately veined, with- out domatia. Inflorescences axillary or less often ter- minal, racemose or paniculate, flowers opposite or along only 1 side of the rachis, pedicels present or absent. Flowers bisexual and radially symmetrical, usually 5-parted, hypanthium ovoid to turbinate, calyx lobes 4- 6, short and persisting, corolla campanulate to funnel- form, white to yellow, lavender or purple, corolla tube cylindrical to urceolate, often with longitudinal ribs in line with the sinuses between the lobes, glabrous within at the mouth, corolla lobes 4-5, valvate in bud, spreading or reflexed; stamens 4-5, inserted near the base of the tube, filaments pilose at the base, anthers linear, exserted or included; ovary 2-locular, with 1 ovule pendulous from apex of each locule, stigmas 1 or 2. Fruits drupa- ceous, fleshy to leathery, usually white at maturity, lat- erally compressed and rounded in outline (in Central America) or oblong-cylindrical when dried, with 2 py- renes; seeds pendulous and laterally compressed.
A genus of about 20 species, ranging from the southern United States through Mexico, Central America, and the West Indies to southern South America.
Chiococca is recognized by its often pendant clambering branches, the very short stipules slight- ly united above the petioles (and usually with an awn), and the unusual white fruit flattened on op- posite sides and rounded in outline (in Central American species). The ribbed and valvate corolla, the filaments free to the base of the corolla tube, the two-locular ovary with solitary pendulous ovules, and the white drupaceous fruits are also important distinguishing characters.
All our species are wide-ranging and quite vari- able; this may make them difficult to separate in the absence of flowers, since the fruit differ little among the species. In fact, the patterns of variation are so broad as to suggest that there may be hy- bridization between the species.
Key to the Species of Chiococca
la. Stamens usually well exserted at anthesis (with the filaments sometimes visible); corolla often campanulate-urceolate in Costa Rica; inflorescences usually with opposite branching, flowers sessile or with pedicels to 3 mm long; secondary veins obscure on the undersides of the leaves, petioles 10-30 mm long; (7700-) 1 600-2200 m elevation C. phaenostemon
Ib. Stamens included within the corolla tube or only the tips exserted; corolla usually funnelform; inflorescences with few alternate or opposite lateral branches, pedicels 1-6 mm long; secondary veins visible on the lower leaf surfaces, petioles 3-17 mm long; 0-1200(-1500) m elevation ... 2
2a. Hypanthium/ovary with thin erect hairs 0.1-0.3 mm long (rarely glabrous); leaves densely to sparsely puberulent beneath with thin straight hairs 0.2-0.3 mm long; inflorescences 2—4 cm long [petioles 1-4 mm long] C. semipilosa
2b. Hypanthium/ovary glabrous or with a minute (0.05 mm) papillate-puberulence; leaves glabrous or sparsely papillate-puberulent beneath; inflorescences (2-)4-10 cm long 3
3a. Leaves usually ovate-elliptic and drying grayish or greenish, rarely more than 4 cm broad, petioles 3-8 mm long; corolla 4-8 mm broad distally when open, calyx lobes narrow; fruit strongly com- pressed C. alba
3b. Leaves usually oblong-elliptic and drying dark brown, often more than 5 cm broad, petioles 5-17 mm long; corolla 7-10 mm broad distally when open, calyx lobes broadly rounded or obscure; fruit only slightly compressed laterally C. pachyphylla
Chiococca alba (L.) Hitchcock, Ann. Kept. Mis- souri Bot. Gard. 4: 94. 1893. Lonicra alba L., Sp. PI. 175. 1753. Figure 36.
Woody climbers, shrubs or less often small trees to 8 m tall and 10 cm trunk diam., distal branches often pendulous or clambering, distal twigs often opposite and held perpendicular to the main stems, leafy branchlets 0.7-4 mm thick, glabrous and terete, dark when dried;
stipules 1-5 mm long, the broad basal part 0.5-2 mm long and slightly (0.5 mm) united above the petioles (intrapetiolar), with an acuminate or caudate tip 0.5-3 mm long. Leaves distant along the stems, petioles 3-8 mm long, 0.5-1 mm broad, glabrous; leaf blades (2.5-)3- 9(-13)cm long, (l-)1.5-3.8(-6)cm broad, ovate-elliptic, narrowly ovate, oblong or lanceolate, apex long-acu- minate (sometimes bluntly acute to short-acuminate), base obtuse to rounded and slightly decurrent on petiole, leaves drying stiffly chartaceous to membranaceous, gla-
96
FIELDIANA: BOTANY
brous above and below or with a few thin hairs ca. 0.2 mm long beneath, 2° veins 3-5/side and weakly loop- connected distally. Inflorescences mostly axillary, (2-)4- 1 1 cm long, unbranched and racemiform or with few lateral branches and paniculate, peduncles (0.5-) l-3.5(-7) cm long, ca. 0.5 mm thick, usually glabrous, bracts 1- 1.5 mm long, narrow, the flowers usually borne along 1 side of the rachis, solitary and separate or in groups of 3, pedicels 1-8 mm long, slender, usually glabrous. Flow- ers with hypanthium 1-1.8 mm long, flattened laterally on opposing sides, ellipsoid in outline, glabrous or very minutely (0.05 mm) papillate-puberulent, calyx tube 0.5- 1 mm long, calyx lobes 0.2-0.6 mm long; corolla fun- nelform, white to yellowish or rose, usually glabrous ex- ternally, tube 3-8 mm long, 2-5 mm wide at apex, lobes 5 (4), 3-4 mm long, triangular; stamens 5, included or slightly exserted, anthers ca. 3 mm long; styles 5-8 mm long, exserted. Fruits 4-7 mm long, 4-7 mm broad, rounded-oblong (abruptly rounded at top and bottom) in outline and flattened laterally on the 2 opposite sur- faces, white at maturity, persisting calyx ca. 1 mm long and 1.5 mm diam.
Common clambering shrubby plants along open forest edges and disturbed areas, in both evergreen rain forest areas and in seasonally deciduous for- ests, from sea level to 1300(-1500) m elevation. Flowering in March-October (mostly June-Au- gust); fruiting in June-March. The species ranges from the southernmost United States (Texas and Florida), through Mexico, Central America, and the West Indies into tropical South America.
Chiococca alba is recognized by the clambering stems, the smaller often ovate to lanceolate leaves, the usually few-branched axillary inflorescences, funnelform usually yellowish white flowers, and white flattened fruit with round outline and per- sisting calyx. Most collections are glabrous, but a few have minute puberulence on the young stems, inflorescence, and hypanthium. The disc-like whitish seeds may have inspired two names used in Central America for the species: Idgrimas de Maria and Idgrimas de San Pedro.
Chiococca pachyphylla Wernham, J. Bot. 5 1 : 323. 1913. Figure 36.
Lianas and woody climbers (rarely shrubs?), l-5(-7) m tall, leafy branchlets 1-5 mm thick, glabrous, terete and drying dark or grayish; stipules 1-3 mm long, sub- acuminate to caudate at apex, slightly (0.2-0.5 mm) unit- ed above the petioles. Leaves well spaced along the stem, petioles 5-17 mm long, 0.6-1.2 mm broad, glabrous; leaf blades 6-12 cm long, 3-6 cm broad, oblong-elliptic to elliptic or ovate-elliptic, apex bluntly acute to short- acuminate, base acute to obtuse and slightly decurrent on petiole, drying stiffly chartaceous to subcoriaceous, glabrous above and below, 2° veins 3-5/side, the sec- ondaries usually darker than the lower surface and easily
seen. Inflorescences 4-8 cm long, mostly axillary and with few lateral branches, the distal axes racemose, bracts 1-2 mm long, linear, pedicels 1 .5-4.5 mm long, glabrous. Flowers glabrous externally, hypanthium 1.5-2 mm long, calyx tube 0.5-1 mm long, calyx lobes 0. 1-0.4 mm long, rounded or bluntly triangular, glabrous; corolla funnel- form, yellowish, corolla tube 5-7 mm long, gradually expanded to apex and 1-3 mm broad, lobes 2-3.5 mm long, 1.5-2 mm broad at the base, bluntly acute; stamens included. Fruits 6-8 mm long, 6-8 mm broad, broadly ellipsoid-circular to circular in outline, ca. 2 mm thick, green becoming white, glabrous, persisting calyx ca. 1 mm high and 1.5 mm diam.
Shrubs and climbers of evergreen and deciduous forest formations, from near sea level to 1 500 m elevation. Flowering primarily in May-Septem- ber; fruiting in August-December. The species ranges from northeastern Mexico to Costa Rica.
Chiococca pachyphylla is recognized by its more consistently vining habit, stiff usually oblong-el- liptic leaves, lack of pubescence, racemose inflo- rescence branches, and flattened white fruit. The secondary veins on the lower leaf surfaces are much easier to see than in C. phaenostemon. and the flowers and fruit appear to be a bit larger than those of C. alba. In addition, C. pachyphylla has a number of characteristics that appear to be in- termediate between C. alba and C. phaenostemon. Considerable variation in inflorescence and flower morphology adds to the difficulty.
Chiococca phaenostemonend Schlcctcnd., Linnaca 9: 594. 1834. Figure 36.
Shrubs or small trees (lianas), 3-14 m tall, often with separate trunks from the base, leafy stems 1.5-6 mm thick, glabrous, slightly quadrangular at first but becom- ing terete, older nodes conspicuously thicker than the internodes; stipules 2-4 mm long, the broad base 1-2 mm long, united around the stem for ca. 0.5 mm, with a narrow awn 0-2 mm long, the awn often breaking off to leave a shallow persisting cup at the older node. Leaves somewhat clustered at the ends of stems, petioles 7-30 mm long, 0.7-1 .5 mm broad, glabrous; leaf blades (4-)6- 13 cm long, (1. 2-) 1.5-4. 8 cm broad, elliptic to elliptic- oblong, narrowly oblong (rarely elliptic-obovate), apex gradually tapering and cuneate or acuminate, base ta- pering gradually and obtuse or acute, decurrent on pet- iole, leaves drying stiffly chartaceous to subcoriaceous, glabrous above and below, 2° veins 5-8/side and usually obscure on the lower surface, weakly loop-connected dis- tally. Inflorescences axillary or terminal, 3-12 cm long, paniculate with 3-4 primary branches (and 2-3 of these with secondary branches) peduncles to 4 cm long, ca. 1 mm thick and sparsely papillate-puberulent, bracts 1-2 mm long, flowers often in cymules, pedicels 0.3-3 mm long. Flowers with hypanthium 1.2-2 mm long, 0.7-1.3 mm wide, glabrous, calyx tube ca. 0.5 mm long, calyx lobes 0.5 mm long; corolla usually campanula to to ur-
BURGER & TAYLOR: FLORA COSTARICENSIS. RUBIACEAE
97
ceolate in Costa Rica (less often funnelform), white to yellowish, glabrous, tube 3-5(-6) mm long, 4-6 mm diam. at the mouth, lobes 5, 2-4 mm long, 1.5-3 mm broad at the base; anthers ca. 3 mm long, half to fully exserted. Fruits 5-6 mm long, 5-6 mm broad, broadly oblong or broadly obovate in outline, flattened on 2 sides (said to be thicker and rounded in northern Central America), persistent calyx 1-1.5 mm long and 1.5 mm diam., ped- icels to 3 mm long.
Trees of evergreen montane forest formations, from (?700-)1600 to 2100 elevation (to 2500 m in Guatemala). Flowering in July-September; fruiting in July-August and January-February. The species ranges from northeastern Mexico to the Chiriqui Highlands of Panama.
Chiococca phaenostemon is characterized by its highland habitat, larger and campanulate corollas (in Costa Rica), and the often exserted anthers. The glabrous often long-petiolate leaves with de- current base and the secondary veins usually ob- scure beneath also help to distinguish this species. While a very distinctive plant in the wild, some specimens of this species may be difficult to sep- arate from C. pachyphylla and C. alba. The com- mon name is chiraquilla.
Chiococca semipilosa Standl. & Steyerm., Publ. Field Mus. Nat. Hist., Dot. Ser. 22: 279. 1940. Figure 36.
Shrubs, l-3(-4) m tall, leafy branchlets 1-3 mm thick, minutely puberulent with thin erect whitish hairs 0.1- 0.2 mm long, soon glabrescent, terete; stipules 2-4 mm long, broad basal part 0.5-1 mm long, little (0.2-0.5 mm) united above the petiole, the narrow awn 1-3 mm long, minutely puberulent. Leaves with petioles 1-4 mm long, 0.5-1 mm broad, with lateral margins continuous with the lamina margins, minutely puberulent; leaf blades (l-)3-7.5(-12) cm long, (0.5-)l-2.5(-3) cm broad, nar- rowly ovate, ovate-lanceolate or ovate-elliptic, tapering gradually to the acute or acuminate tip, base acute to obtuse and decurrent on petiole, leaves drying stiffly chartaceous, glabrous to sparsely puberulent above with thin whitish hairs 0.2-0.3 mm long, sparsely to densely soft pubescent beneath with hairs 0.1-0.4 mm long, 2° veins 2-4/side, weakly loop-connected near the distal margin. Inflorescences axillary, 2-4 cm long, cymose or racemose with 3-9 flowers (rarely umbellate), peduncles 4-10 mm long, pedicels (0-)l-3(-5) mm long, puberu- lent. Flowers 5-parted, hypanthium 1-1.5 mm long, 0.7- 1 mm broad, little differentiated from the pedicel, dense- ly puberulent, calyx tube obscure, calyx lobes 0.5-1 mm long, acute and drying with little puberulence distally; corolla cream white to yellowish, broadly funnelform, usually minutely puberulent externally, tube 4-5 mm long, 1.5 mm diam. at the base to 3 mm near apex, lobes 1.5-2.8 mm long, 1.5-2 mm broad at the base; stamens included. Fruits 5-6 mm long, 4-5 mm broad, rounded in outline and flattened longitudinally, with thin erect
hairs ca. 0.2 mm long, persisting calyx 0.7-1.5 mm long, 1.8 mm broad, drying dark in contrast to the pale fruit.
Shrubs of evergreen and deciduous forest for- mations, from 200 to 1 600 m elevation. Flowering in June-July; fruiting in September-January. The species ranges from Belize and Guatemala to northwestern Costa Rica.
Chiococca semipilosa is distinguished by the short thin hairs on the hypanthium/ovary, the pu- bescence on the lower leaf surfaces, the short few- flowered inflorescences, and the prominent calyx lobes that often dry dark. There is the possibility that material placed here is no more than an un- usual form of C. alba. The figure is based on the Guatemalan holotype (Steyermark 31406 F).
Chione DeCandolle
Trees or shrubs, glabrous or glabrescent; stipules unit- ed (interpetiolar and intrapetiolar) and forming a cap over the shoot apex, leaving a scar across the stem and on the adaxial base of the petioles, small, caducous. Leaves petiolate; leaf blades often coriaceous, entire and pin- nately veined, domatia present or absent. Inflorescences terminal, solitary or 3 at a distal node, paniculate with opposite branching and cymose or corymbose in form, pedunculate, bracteolate, flowers pedicellate. Flowers bi- sexual and radially symmetrical, apparently monomor- phic, hypanthium turbinate, calyx lobes 5 or 6 or un- developed and the distal margin undulate; corolla funnelform, white or yellowish, corolla tube short, gla- brous within, corolla lobes 5(-6), broadly imbricate in bud with 2 exterior; stamens 5(-6), inserted above the base of the tube, filaments thick, anthers dorsifixed, ex- serted; ovary 2-locular, ovules solitary in each locule, pendulous from apex, style stout, stigmas 2, oblong, ex- serted. Fruits drupaceous, ovoid to ellipsoid, pyrene sol- itary and 2-locular; seeds elongate, the testa membra- nous.
A genus of about 1 5 species; fewer than 6 species are found in southern Mexico and Central Amer- ica; the others occur in the West Indies. The genus is distinguished by its glabrous parts, cap-like stip- ules (in some species), terminal inflorescences, short corolla tubes with broadly imbricate corolla lobes (in bud), and the fleshy fruits with two-locular pyrene (stone). The genus Oregandra is a syn- onym; Standley misinterpreted the ovules when he described that genus.
Chione sylvicola (Standl.) W. Burger, Selbyana 1 2: 138. 1991. Chomelia sylvicola Standl., J. Wash. Acad. Sci. 18: 182. 1928. Oregandra panamen- sis Standl., Publ. Field Columb. Mus., Bot. Ser.
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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.
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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.
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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
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oblongo-elliptica, 6-15 cm longa, 3-7 cm lata, venis lat- eralibus 5-7, venulis parallelis. Inflorescentiae axillares, ad 9 cm longae, pedunculo ca. 22 mm longo, ramis saepe dichotomis, floribus secundis. Flores puberuli, hypan- thio 1-2 mm longo; corolla alba vel cremea, tubo 6-9 mm longo, lobis 4. Fructus 13-15 mm longi.
TYPUS— E. Bella 414 (holotypus CR, isotypi F, MO), from Reserva Biologica Monteverde, Alajuela, Costa Rica.
Trees, 10-25 m tall, to ca. 35 cm dbh, leafy stems 1- 4 mm thick, sericeous or strigulose with ascending pale grayish hairs ca. 0.4 mm long; stipules 1 1-18 mm long, to 4 mm broad, lanceolate with an acute apex, sericeous at the base and along the midrib. Leaves with petioles (6-) 1 2-23 mm long. 0.8-2 mm thick, sericeous with pale grayish hairs; leaf blades 5-15 cm long, 3-7 cm broad, elliptic to elliptic-oblong, apex short-acuminate with tip 5-8 mm long, base obtuse to acute, drying stiffly char- taceous, dark brown above, much paler beneath, gla- brous or very sparsely pubescent above with thin whitish hairs to 1 mm long, appressed-pubescent beneath with thin hairs ca. 0.3 mm long and densely sericeous along the major veins, 2° veins 4-7/side, both the 3° and 4° veins at right angles to the secondaries, depressions (domatia?) sometimes present in the vein axils. Inflo- rescences solitary and axillary (2/node), ca. 5 cm long, to 9 cm in fruit, cymose with 2 lateral simple or bifid secund branches, peduncles 22-45 mm long and 1 mm thick, ascending sericeous, distal flowers along 1 side of the branch, bracts absent, flowers sessile. Flowers pu- bescent externally, hypanthium 1-2 mm long, 1 mm diam., calyx tube minute, 1.4-2 mm diam., calyx lobes 0.2-0.8 mm high; corolla tubular, yellow or yellowish white, fluted distally, tube 6-9 mm long, 0.8-1.5 mm diam., densely sericeous, lobes 4, 1-2 mm long, 1-2 mm broad; stamens 4, anthers ca. 3.5 mm long; style ca. 5 mm long, stigmas 1.2 mm long. Fruits 13-15 mm long, 8-9 mm thick, obovoid-oblong, with 6-8 longitudinal ribs, becoming dark brown, glabrous, persistent calyx ca. 1 mm long, pyrenes ca. 14 x 8 mm; seeds ca. 10 x l mm.
Plants of the wet evergreen forests of the Carib- bean slope of Costa Rica, at 600-900 m elevation. Flowers were collected in May; fruits were col- lected in October-November. The species is known from below Monteverde, Alajuela (84°43'W), and the southern Cordillera de Talamanca, Limon (82°59'W). Collections in addition to the type are Bello 17 2 & 872, Bella & Cruz 457, Herrera 3310, and Poveda 24.
Chomelia venulosa is recognized by its parallel minor venation, sericeous flowers on short inflo- rescences with two simple or bifid lateral branches, and restricted altitudinal range on the Caribbean slope. The two-celled deeply ridged pyrenes, the subimbricate to valvate corolla aestivation, and the parallel minor venation suggest that this spe- cies is best placed in Chomelia, as opposed to Guettarda. It may be allied to the " Anisomeris" group of Chomelia species with appendages on the
corollas. The overall appearance of the leaves is similar to that of Chomelia panamensis, but the minor venation is quite unusual. The 3° and 4° veins are little differentiated, parallel to each other, and usually perpendicular to the 2° veins.
Cinchona Linnaeus
Trees of medium height or occasionally shrubs, branchlets terete or tetragonal; stipules interpetiolar, tri- angular, often large, colleters present within at the base, caducous or rarely persisting. Leaves opposite, often large, petiolate; leaf blades with entire margins, coriaceous to chartaceous, domatia absent. Inflorescences terminal and solitary (apparently axillary when lateral branches are subtended by reduced leaves), open-paniculate, usually large and many-flowered, branching mostly opposite. Flowers bisexual, small in most species, aromatic, often puberulent externally, hypanthium turbinate, calyx tube with small calyx lobes (rarely with the calyx tube entire distally); corolla salverform to fimnelform, white to pink or purplish, corolla tube terete or slightly 5-angled, gla- brous or pilose in the throat; corolla lobes 5 (4, 6), spread- ing, valvate in bud; stamens 5, inserted in the corolla tube, filaments short or long, anthers linear dorsifixed, included or their apices exserted; ovary 2-locular, pla- centas attached to the septum and spongy, ovules many in each locule, peltately attached and imbricated, style narrow, stigmas short and obtuse, included or slightly exserted. Fruits woody capsules, 2-locular, subcylindri- cal to ovoid or oblong, dehiscing septicidally from bot- tom to top; seeds many, peltate, thin and flat, testa with a broad thin peripheral wing.
A genus of 20— 40 poorly defined species, ranging from Costa Rica southward to Bolivia, mostly along the Andes mountains. The capsules opening up- ward from the base helps separate Cinchona from closely related genera, such as Ladenbergia and Joosia. This genus has played an important role in the history of medicine as the source of the antimalarial drug quinine. The major commercial sources of quinine are cultivars of Cinchona cal- isaya Wedd. grown in Indonesia, which originated from the eastern slopes of the Andes. Species of the genus were introduced and have become nat- uralized in Guatemala (see the discussion in Standley & Williams, 1975, p. 38).
Cinchona pubescens Vahl, Skr. Naturhist. Selsk. Kjobenhavn 1: 19. 1790. Figure 37.
Small or medium-sized trees, (3-)5-20 m tall, trunks 8-30 cm dbh, bark pale brown and roughened, leafy branchlets 3-8 mm thick, distinctly 4-angled, minutely puberulent or glabrous; stipules 4-12(-20) mm long, 3- 8(-12) mm broad at the base, glabrous or minutely ap-
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pressed-puberulent, deciduous. Leaves with petioles (9_) 1 2-35(-60) mm long, 1 .3-2.5 mm thick, sparsely and minutely (0.2 mm) puberulent; leaf blades 10-32(-40) cm long, 7-17(-26) cm wide, broadly ovate to broadly elliptic-oblong or suborbicular, apex broadly obtuse to subacuminate, base abruptly cuneate to rounded and subtruncate, usually slightly decurrent on petiole, drying stiffly chartaceous or chartaceous, glabrous or very sparsely puberulent above, sparsely pubescent beneath with slender hairs ca. 0.3 mm long, 2° veins 7-12/side. Inflorescences terminal or axillary to distal leaves, 9-40 cm long, 8-24 cm broad, open paniculate with few op- posite widely spaced (3-1 1 cm) branches, distal branches minutely tomentulose, the flowers in congested distal clusters, pedicels 0.5-3 mm long. Flowers white, cream, or pale pink, densely tomentulose externally, hypanthi- um 2-3 mm long, 1.3-2 mm diam., calyx tube l-2(-3) mm long and 2-3 mm diam., calyx lobes 0.2-1 mm long, acute; corolla 15-16 mm long and funnelform, tube 10- 1 3 mm long, 1.4-2 mm diam.. glabrous within, lobes 5, lanceolate to oblong, 3-5 mm long and 1.5-2.5 mm wide, tomentulose externally and villose on the interior mar- gins; stamens 5, filaments ca. 2 mm long, anthers 2.5-3 mm long; style 6-13 mm long, glabrous. Fruits subcy- lindrical to narrowly oblong, 12-35(-50) mm long, 6-9 mm diam., glabrescent or minutely (0. 1 mm) puberulent, brown, lustrous yellowish brown within; seeds 5-1 2 mm long, 1.5-3 mm broad, flat and oblong-elliptic in outline, with a thin membranaceous winged margin, surfaces re- ticulate and the margin erase, dark center of the seed 1.8-3 mm long.
Trees of evergreen forest formation on both the Caribbean and Pacific slopes of Costa Rica, rang- ing from (500-)800 to 1700 m elevation. Flow- ering mostly in June-September, with solitary col- lections in November, February, and March; fruiting in February and November-December in Costa Rica. The species is apparently rarely en- countered in southern Central America. Our col- lections come mostly from the Caribbean slopes of the Cordillera de Tilaran and the Central High- lands in the Provinces of Alajuela and Heredia. In Panama the species is known only from the Chi- riqui Highlands. This species ranges southward to Venezuela, Peru, and Bolivia.
Cinchona pubescens is recognized by the larger often broadly rounded leaves, large terminal in-
florescences with small puberulent flowers in distal clusters, the narrow woody two-locular capsules, and the seed with a thin elongated membranous wing. The flowers are said to have the aroma of Gardenia or Cananga odorata (Annonaceae). The rarity of this species in southern Central America suggests that it is not native, and collections may represent relicts of native pre-Columbian intro- duction.
Coccocypselum P. Browne, nomen conservandum
Herbs, annual or perennial, prostrate and creeping to erect-ascending, usually pubescent with multicellular hairs; stipules interpetiolar, sometimes reduced to a very short (0.2 mm) rim, small and simple with a single su- bulate lobe (2/node) and with 0 or 2-8 smaller lateral awns, persisting. Leaves petiolate, entire, domatia ab- sent. Inflorescences terminal or pseudoaxillary, usually solitary, capitate with (l-)3-20 flowers, sessile or pe- dunculate, bracts and bracteoles small, flowers sessile. Flowers bisexual, monomorphic or distylous, small, ca- lyx lobes 4, narrow and persistent; corolla blue to purple or white, funnelform, corolla lobes 4, valvate in bud; stamens 4, filaments borne on the tube of the corolla, anthers dorsifixed below the middle or near the base; ovary 2-locular, the placentas borne on the center of the septum, ovules many and horizontal, style with 2 short branches. Fruits baccate and arenchymatous, often mealy and hollow, globose to obovbid, bright blue; seeds many and small, angled or flattened.
A Neotropical genus of 10-20 species, ranging from Mexico and the West Indies into South America. The genus is recognized by its herba- ceous habit, usually broad puberulent leaves, small capitula with few flowers, four-parted flowers, two- locular ovaries, and blue fruit with many small seeds. These plants are often confused with species of Geophila. Coccocypselum lanceolatum is our most distinctive species; the others may be difficult to distinguish.
Key to the Species of Coccocypselum
la. Leaf blades with 8-13 pairs of secondary veins, narrower than long, ovate-triangular to lanceolate and usually acute at the apex; with 8-10 flowers in each inflorescence [1000-2000 m elevation]
C. lanceolatum
1 b. Leaf blades with 5-6 pairs of secondary veins, usually about as broad as long, ovate to ovate- lanceolate, acute to rounded at the apex; with 2-6 flowers in each inflorescence 2a. Inflorescences sessile in the axils of leaves; plants of evergreen lowlands, 0-1000 m elevation
. C. herbaceum
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2b. Inflorescences pedunculate; plants of evergreen lowland and highlands, 0-2000 m elevation ... 3 3a. Leaf blades rounded and cordate or subcordate at the base, often wider than long [ 1 400-2000 m
elevation] C. cordifolium
3b. Leaf blades acute to obtuse or truncated at the base, usually as wide as long and ovate to broadly
elliptic 4
4a. Stems and leaves with hairs 0.5-1 mm long (or occasionally glabrous); calyx lobes 3-4 mm long;
stipules ca. 5 mm long; common in Central America C. hirsutum
4b. Stems and leaves with hairs 0. 1-0.3 mm long; calyx lobes 1.5-2.7 mm long; stipules 2-4 mm long;
not known from Costa Rica (key based on Steyermark, 1972) C. guianensis
Coccocypselum cordifolium Nees & Mart., Nova Acta Phys.-Med. Acad. Caes. Leop.-Carol. Nat. Cur. 12: 14. 1824. Geophila pleuropoda J. D. Smith, Bot. Gaz. 52: 50. 1911. Geocardia pleu- ropoda (J. D. Smith) Standl., Contr. U.S. Natl. Herb. 17: 445. 1914. Tontanea pleuropoda (J. D. Smith) Standl., N. Amer. Fl. 32: 148. 1921. C. pleuropodum (J. D. Smith) Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 281. 1929. C. roth- schuhii Loessner, Bot. Jahrb. Syst. 60: 370. 1926. Figure 2.
Herbs, prostrate or creeping, leafy stems 0.4-1.6 mm thick, hirsute or villous with thin straight or crooked hairs 0.5-1.5 mm long; stipules 1.5-3 mm long, united only at the base (ca. 0.2 mm) and with 2 narrowly linear awns on each side (4/node). Leaves with slender petioles 4-28(-55) mm long, villous or pilose with thin hairs; leaf blades 1 1-35 mm long, 1 2-42 mm broad, ovate to ovate- orbicular or ovate-reniform, apex rounded and bluntly obtuse (and usually minutely apiculate), base rounded at the cordate to subtruncate base, drying membrana- ceous or thin-chartaceous, sparsely to densely pubescent, the hairs 1-1.7 mm long on the upper surface and ca. 0.7 mm long beneath, 2° veins 3-5/side. Inflorescences axillary, usually only I/node, 1.4-5 cm long, the capit- ulum less than 1 cm long and with 3 (2, 4) flowers, peduncles 4-38 mm long, villous, bracts ca. 4 mm long, linear, flowers sessile. Flowers with hypanthium ca. 1 mm long, with thin whitish hairs, calyx lobes 1.5-2.5 mm long, linear-lanceolate, sparsely villous; corolla lav- ender, lilac, pale blue, or whitish, tube 4-6 mm long, 2- 3 mm diam. at apex, sparsely puberulent externally, lobes 3.3-5 mm long, 1.2-2 mm broad at the base, narrowly triangular to oblong. Fruits 5-6 mm long, 4-5 mm diam., ovoid, blue, densely villous and with the persisting calyx 2-3 mm long; seeds 0.2-0.5 mm diam.
Plants of evergreen lower montane rain forest formations, from 1000 to 1600 m elevation (to 2000 m in Guatemala). Rarely collected in Costa Rica's major highlands. Probably flowering and fruiting throughout the year. The species ranges from Mexico to Panama, and it is also found in eastern and southern Brazil.
Coccocypselum cordifolium is recognized by the usually subcordate or truncated leaf bases, the small
pedunculate heads with only two to four blue flow- ers, and the long hairs on the upper leaf surface. This species appears to be rare or overlooked in Costa Rica and Panama. This species is easy to confuse with Geophila cordifolia, which has one- or two-seeded red fruits.
Coccocypselum guianense (Aubl.) K. Schum. in Mart., Fl. Bras. 6(6): 315. 1889. Tontanea gui- anensis Aubl., Hist. pi. Guiane 1: 108, pi. 42. 1775.
According to Steyermark (1972), this species is distinguished from similar species by its short (1.5— 2.7 mm) calyx lobes, short (0.2-0.5 mm) dense indumentum on stems and inflorescences, smaller (3—4 mm) stipules, and slightly longer (5.5-9 mm) corolla tubes. The species is said to range from Florida and the West Indies to Venezuela and the Guianas (Steyermark, 1972; Hortus Third, 1976). These plants have been used in ornamental hor- ticulture as a ground cover and in hanging baskets. Because of these uses, it is likely that escaped pop- ulations have become established in some areas. We have not seen material that can be definitively ascribed to C. guianense from Costa Rica.
Coccocypselum herbaceum P. Browne, Civ. Nat. Hist. Jam. 144, pi. 6. 1756; also cited as Aublet, Hist. pi. Guiane 1: 68. 1775 (fide Adams, 1972); Lam., Encycl. 2: 56. 1786 (fide Standley & Wil- liams, 1975). C. repensSv/., Prodr. 31. 1788 (not C. repens H.B.K. 1819, not Condalia repens Ruiz & Pav. 1798). Tontanea herbacea (P. Browne) Standl., N. Amer. Fl. 32: 147. 1921. T. hispidula Standl., loc. cit. 147. 1921. C. hispidulum (Standl.) Standl., Publ. Field Columb. Mus., Bot. Ser. 4: 281. 1929.
Herbs, procumbent or trailing, leafy stems 0.7-2 mm thick, sparsely to densely pilosulous with erect or ap-
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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
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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
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to acute, drying chartaceous to subcoriaceous, glabrous above and below, 2° veins 7-10/side, minute (0.2-0.3 mm) domatia often present at the vein axils beneath. Inflorescences axillary, subcapitate or appearing verti- cil laic, ca. 5 cm long (including the corollas), with 1-9 flowers per axil, flowers subsessile. Flowers 1 2-20 mm long, white, calyx lobes 5, minute, corolla white, tube 10-14 mm long, lobes 14-20 mm long; anthers exserted, 8-12 mm long. Fruits 10-16 mm long, 8-13 mm diam., oblong and abruptly rounded at apex and base, green becoming red (drying dark), glabrous, calyx scar 2-3 mm broad; pyrenes ca. 10 x 7 x 3 mm, planoconvex.
Cultivated or rarely persisting in evergreen and partly deciduous formations, 800-2000 m eleva- tion. The flowering season is primarily in Febru- ary, with fruiting in November-December. This species, probably native to western Ethiopia, is now cultivated throughout the tropics.
Coffea arabica is recognized by its glossy dark green leaves (in life), lack of pubescence, axillary clusters of aromatic white flowers, and distinctive two-seeded fruit. This species is only occasionally found outside of cultivation; birds and bats have been described as dispersal agents. In addition to the stimulant caffeine, coffee beans contain glu- cose, dextrin, proteins, and the flavor-enhancing volatile oil caffeol. Cultivation in cooler temper- atures at higher elevations (ca. 1000-2000 m) is an important factor in producing high-quality cof- fee (cafe), and this may account for the excellent reputation of Costa Rica's most valuable export crop.
Coffea liberica Bull ex Hiern., Trans. Linn. Soc., Ser. 2, 1: 171, t. 24. 1876. C. excelsa A. Chev., Rev. cult, colon. 12: 258. 1903.
Shrubs or small trees to 5(-l 5) m tall, leafy branchlets 1.8-8 mm thick, glabrous; stipules 3-6 mm long, 4-8 mm broad. Leaves with petioles (4-)8-24 mm long, 1 .5- 4 mm thick, glabrous; leaf blades (8-) 1 2-24(-30) cm long, (4-)5-12 cm broad, elliptic-oblong to elliptic-ob- ovate, apex bluntly obtuse to bluntly short-acuminate, base obtuse to acute, drying subcoriaceous, glabrous above and below, 2° veins 7-10/side, pit domatia 0.2-0.8 mm long at the vein axils beneath and often with a few short hairs. Inflorescences 2-5 cm broad, with ca. 5-10 flow- ers, bracts 3-8 mm long, narrowly elliptic to lanceolate. Flowers 6-8-parted, 25-35 mm long, corolla tube 10- 14 mm long and ca. 1.5 mm diam., lobes 8- 12 mm long, 2-3 mm broad; anthers to 6 mm long. Fruits 1 .2-2.5 cm long, oblong-rotund, yellowish red.
Cultivated or escaped trees of evergreen forest formations, from near sea level to 1200 m ele- vation. This species is native to coastal West Af- rica and is now found cultivated around the world.
Coffea liberica is recognized by its larger leaves, glabrous parts, white axillary flowers, and distinc- tive fruit. Unlike C. arabica. which is essentially a highland species, C. liberica grows well at lower elevations. While C. liberica is more disease-re- sistant and can do well at lower altitudes, the seeds (beans) produce coffee of inferior flavor, and the species is not an important crop in Central Amer- ica.
( ondaminea DeCandolle
Shrubs or small trees, usually with few branches and large leaves; stipules interpetiolar and intrapeliolar, unit- ed above the leaf base to form a short sheath and with a broad distal 2-parted blade (sometimes appearing as 4 free stipules at each node), persisting. Leaves opposite and large, sessile or short-petiolate; leaf blades entire, lacking domatia. Inflorescences terminal and solitary, open paniculate with cymose or corymbose distal branching, usually with 3 branches at apex of the pe- duncle (lateral branches equaling the continuing rachis), bracts small, bracteoles absent, pedicels short or none. Flowers bisexual, large, hypanthium turbinate to cam- panulate, calyx lobes 3-5 or none and the calyx tube entire, corolla funnelform to salverform, thick-fleshy, corolla tube barbate in the throat, corolla lobes 4-5, valvate in bud, glabrous; stamens 5, filaments inserted in the upper half of the tube, puberulent at the base, anthers narrow, sagittate at the base, exserted; ovary 2- locular, ovules many in each locule from axile placentas, style narrow, stigmas oblong, exserted. Fruits woody capsules, pyriform to turbinate, truncated at apex with a circular calyx scar, with loculicidal basipetal dehiscence forming 2 valves; seeds minute, attached horizontally, testa reticulate.
A genus of four or five species ranging from Costa Rica to Venezuela, Peru, and Bolivia. The large, often subsessile leaves, deeply two-parted stipules, lack of pubescence, very large terminal open-branched inflorescences, greenish flowers, and woody bivalved capsules with hundreds of minute seeds make this a distinctive genus.
Condaminea corymbosa (Ruiz & Pav.) DC., Prodr. 4: 402. 1 830. Macrocnemum corymbosum Ruiz & Pav., Fl. Peruv. 2: 48, pi. 189. 1799. Figure 29.
Shrubs or small trees, 2-5(-7) m tall, leafy branchlets 4-1 2 mm thick, essentially glabrous; stipules apparently free and 4/node. united above the leaf base to form a short (1—4 mm) sheath adnate to the stem (but difficult to see), the free distal parts equal and 2-6(-9) cm long, 6-9(-l 2) mm broad, lanceolate, reddish brown, glabrous or rarely minutely puberulent, with many parallel strong-
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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
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2a. Capsules 40-65 mm long; leaf blades subcoriaceous to coriaceous, acute to obtuse at apex, secondary veins straight to curved; 200-1000 m elevation c. grandiflora
2b. Capsules 62-1 15 mm long; leaf blades coriaceous, obtuse to broadly rounded at apex, secondary veins straight; 0-500 m elevation c. macrocarpa
Cosmibuena grandiflora (Ruiz & Pav.) Rusby, Bull. New York Hot. Gard. 4: 368. 1907. Cinchona grandiflora Ruiz & Pav., Fl. peruv. prodr. 2: 54, pi. 198. 1799. Cosmibuena skinneri (Oerst.) Hemsley, Biol. centr. amer. Bot. 2: 12. 1881. Buena skinneri 'Oerst., Vidensk. Meddel. Dansk. Naturhist. Foren. Kjobenhavn 1852: 48. 1853. Cosmibuena ovalis Standl., Contr. U.S. Natl. Herb. 18: 137. 1916. Figure 28.
Trees or shrubs to 1 2 m tall, terrestrial or epiphytic, leafy branchlets 3-6 mm thick, glabrous, quickly becom- ing pale gray; stipules 8-30 mm long, to 1 2 mm broad, obovate to oblanceolate, the intrapetiolar tube forming V-t-Va of the length (reduced below inflorescences), round- ed apically. glabrous. Leaves with petioles (5-) 1 0—40 mm long, 1.2-2 mm thick, drying dark; leaf blades 7-19 cm long, 4-12(-16) cm broad, broadly elliptic to broadly oblong or obovate, apex bluntly acute to obtuse or short- acuminate, base cuneate and slightly decurrent on pet- iole, drying stiffly chartaceous to subcoriaceous, dark brown or greenish brown above, glabrous above and below, 2° veins 3-7/side (in Costa Rica). Inflorescences with 3-5(-9) flowers, cymose to subumbellate, primary peduncle 5-30(-40) mm long, 2-3 mm diam., glabrous, pedicels (5-)10-20(-30) mm long and often merging im- perceptibly into the calyx. Flowers 7-10 cm long, hy- panthium 7-1 1 mm long, 4-6 mm diam., drying dark, calyx tube 3-9(-l 1) mm long, entire or with lobes 0-8 mm long; corolla white, tubular-sal verform, tube 4-9(-10) cm long, 2.5-5 mm diam., lobes 5-6, (10-)20-35(-40) mm long, (6-)10-14(-20) mm broad, obovate; stamens 5-6, anthers 10-18 mm long; upper part of style densely puberulent, stigmas 4-7 mm long, greenish. Fruits (3-)4- 6.5 cm long, 6-13 mm diam., oblong-cylindrical to ob- long-ellipsoid; seeds 5-8 mm long, 0.5-1 mm broad, linear fusiform, with thin narrow wing, body of the seed 1-2 mm long.
Trees of evergreen forest formations, 50-1000 m elevation. Flowering in April and July-Decem- ber; fruits were collected in March. This species ranges from southern Mexico to Peru.
Cosmibuena grandiflora is recognized by the general lack of pubescence, large fleshy flowers with long tubes, and unusual stipules. The thinner leaves drying dark above and with the secondary veins readily visible help to distinguish these plants from material placed under C. macrocarpa. This species is not often collected in southern Central America. The flowers are aromatic in the morning and late afternoon (Herrera 1072 CR).
Cosmibuena macrocarpa (Benth.) KJotzsch ex Walpers, Repert. bot. syst. 6: 69. 1846. Buena macrocarpa Benth., Bot. voy. Sulph. 104, t. 38. 1844. C. paludicola Standl., Contr. U.S. Natl. Herb. 18: 137. 1916. Figure 28.
Shrubs or small trees to 12 m tall, trunks to 25 cm dbh, epiphytic or terrestrial, leafy branchlets 3-7 mm thick, smooth and glabrous, becoming pale gray; stipules 12-24 mm long, 6-12 mm broad, forming a tube ca. Vj of the length but later splitting, obovate to rounded- oblong, apex rounded to obtuse, glabrous and pale green drying reddish brown. Leaves with petioles 8-20(-25) mm long, 1.8-2.8 mm thick, glabrous; leaf blades 6-18 cm long, 2.5-8(-l 1) cm broad, obovate to obovate-ob- long or subrotund-obovate, apex bluntly obtuse to rounded, base cuneate and slightly decurrent on petiole, drying coriaceous, often grayish, glabrous above and be- low (or puberulous along the midvein beneath in young leaves), 2° vein 4-5/side but difficult to see. Inflorescence terminal, subumbellate with 3-8 flowers, peduncles 6- 15(-30) mm long, 2-4 mm diam. (peduncle and inflo- rescence rachis to 3 cm long), pedicels 7-20 mm long. Flowers 10-14 cm long, glabrous, hypanthium 6-1 5 mm long but not clearly differentiated from the pedicel, 2.5- 4 mm diam., calyx tube 2-8 mm long, calyx teeth 0.5- 4 mm long, triangular to minute; corolla salverform and fleshy, white or pale greenish, tube 5-9 cm long, 2.5-4.5 mm diam., lobes 5, 18-30 mm long, 9-12 mm diam., oblong; stamens 5, anthers 12-15 mm long, 1.5 mm wide, included; style exserted, stigmas 6-9 mm long, 1 .8 mm broad. Fruits (4-)6-8(-l 2) cm long, 6-8 mm broad, linear-cylindrical to linear-oblongoid, brown, borne on pedicels 1-2 cm long, disc forming an elevated (2 x 2.5 mm) projection on the truncated apex of the capsule, surface glabrous and drying dark, often with elongate (3 mm) lenticels; seeds 5-9(-13) mm long and ca. 1 mm wide, thin, narrowly winged.
Trees of mangrove and evergreen lowland forest formations on the Caribbean slope, from near sea level to about 400 m elevation. In Costa Rica the species probably flowers April-November; fruit- ing in February-March and July-September. The species ranges from Costa Rica to Peru.
Cosmibuena macrocarpa is recognized by the large fleshy glabrous flowers with long tubes, co- riaceous obovate leaves with rounded apices and obscure venation, long woody capsules with small winged seeds, and usually epiphytic habit. The name Cosmibuena skinneri (Oersted) Hemsley has been misapplied to this species in the past (cf. Croat, 1978; Dwyer, 1980).
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Cosmibuena valerii (Standl.) C. M. Taylor, Ann. Missouri Bot. Gard. 79: 897. 1992. Hillia valerii Standl., J. Wash. Acad. Sci. 16: 164. 1928. H. ligulifolia Dwyer, Ann. Missouri Bot. Gard. 67: 218. 1980. H. chiriquiensis Dwyer, loc. cit. 216. 1980. Figure 27.
evations and with Hillia tetrandra with smaller flowers, more greenish leaves (when dried), and tufts of hairs at one end of the seed.
Coussarea Aublet
Shrubs or small trees, 2-15 m tall, usually epiphytic, leafy stems 2.5-7 mm thick, semisucculent, glabrous, drying reddish brown, older stems grayish; stipules 14- 42 mm long, 6-1 5 mm broad, enlarged beneath the flow- ers, intrapctiolar and splitting along the sides, oblong to obovate, bluntly obtuse to rounded distally, becoming reddish, caducous. Leaves often closely clustered distal- ly. petioles 3-12(-20) mm long, 1.5-3 mm thick, poorly denned because of the decurrent leaf margins, glabrous; leaf blades 3-8(-10) cm long, l-3(-4) cm broad, nar- rowly obovate to narrowly obovate-oblong or oblanceo- late, apex rounded to bluntly obtuse, base cuneate and decurrent on petiole, drying coriaceous and reddish brown to grayish green, glabrous above and below, 2° veins 3- 6/side, strongly ascending but obscure. Inflorescences terminal, the flowers solitary, sessile or on peduncles to 3 mm long (and difficult to distinguish from the ovary base), bracts short (2-1 1 mm) and triangular or long (20- 40 mm) and spatulate (enlarged stipules), caducous or persisting. Flowers to 12 cm long, glabrous externally, hypanthium 6-14 mm long, calyx lobes 4-18 mm long, 2-3 mm broad, triangular (when short) to narrowly Un- gulate (when long), obtuse or rounded distally; corolla salverform with a long tube, carnose, pale green to white, pink or reddish where exposed in bud, tube 4.7-9 cm long, 4-7 mm diam., lobes 5, 17-31 mm long, 10-18 mm broad, rounded distally; stamens 5, anthers 12-15 mm long; stigmas ca. 6 mm long. Fruits 4-10 cm long, 6-10(-14) mm thick, narrowly oblong or tubular, dark brown with scattered lenticels; seeds 5-6 mm long and ca. 0.5 mm broad, body of the seed 1.4-2 mm long, margins erose.
Plants of evergreen montane forest formations, from 700 to 2300 m elevation. Probably flowering and fruiting throughout the year. This species ranges from northwestern Costa Rica to central Panama.
Cosmibuena valerii is distinguished by its epi- phytic habit, smaller narrowly obovate stiffleaves, long fleshy solitary flowers often marked with pink, and montane habitats. The foliage often dries a reddish brown. There are unusual variations in the development of both the calyx lobes and the floral bracts in this species, but neither seem im- portant taxonomically. This species may be con- fused with Cosmibuena macrocarpa of lower el-
Shrubs or small trees, rarely dioecious, glabrous or less often puberulent, stems usually quadrangular in ear- ly stages but becoming terete; stipules interpetiolar (in- trapetiolar and sometimes forming a cap over the shoot apex in a few species), obtuse to acute (never with aristate or subulate appendages), deciduous or persistent. Leaves opposite (rarely 3/node), petiolate or subsessile; leaf blades entire, domatia present in a few species. Inflorescences terminal, solitary, usually open paniculate with opposite branching to elongate thyrsiform, racemiform, umbel- liform or glomerulate, usually glabrous, flowers often in distal cymes, bracts absent or minute (< 1 mm), flowers sessile or pedicellate. Flowers bisexual (rarely unisexual), hypanthium turbinate or obconic or ovoid, calyx tube short and usually distally truncated (calyx lobes not clearly developed or with 3-5 small lobes); corolla salverform to tubular, white, corolla tube with glabrous throat, co- rolla lobes 4(-5), valvate in bud, oblong to elongate or triangular, often carnose; stamens 4(-5), borne near the mouth or near the base of the tube, anthers subsessile, linear, included or exserted; ovary 2-locular or incom- pletely 1 -locular, ovules 2 and sometimes partly united, erect from a short basal column. Fruits fleshy, coriaceous or spongy drupes, usually longer than wide (ellipsoid to globose), usually with only 1 ovule developing, becoming blue-black or white; pyrene solitary (2), erect, without or with longitudinal ribs dorsally.
A Neotropical genus of about 100 species, with the largest number of species in South America. The genus is characterized by its usually single- seeded fruit, four-parted (less often five-parted) white flowers, often bright white inflorescences with minute bracts and bracteoles or lacking bracts en- tirely, and whitish infructescences. The inflores- cences are rarely more than 1 5 cm long. The stip- ules are triangular to truncated and rarely bilobed. Many of our species have spongy tissues in the corolla and fruit; a number flower at night. Species of this genus can be very difficult to distinguish from some species of Psychotria, Rudgea, and Fa- ramea. The pyrenes of Psychotria have hard walls with ridges, in contrast to the thin smooth walls of Coussarea. Closely similar species in these other genera are referred to under individual species be- low.
Key to the Species of Coussarea
la. Leaf blades conspicuously pubescent beneath, at least along the major veins 2
Ib. Leaf blades glabrous beneath, pubescent only in the vein axils beneath when domatia are present
. 4
110
FIELDIANA: BOTANY
2a. Calyx tube 7-12 mm long with lobes to 3 mm long; inflorescences with 3-15 distantly spaced
flowers, ovary and fruit pilose . c. enneantha
2b. Calyx tube 0.5-2 mm long with lobes to 1 mm long; inflorescences with 15-50 proximate
flowers, ovary and fruit glabrous .3
3a. Leaf blades 11-28 x 5-15 cm; corolla tube 10-13 mm long, distinctly pubescent; in the
Caribbean lowlands, 0-300 mm elevation c. hondensis
3b. Leaf blades 6-19 x 3-8 cm; corolla tube ca. 9 mm long, glabrous or minutely puberulent;
cloud forests, (600-) 1 200-2000 m elevation C. austin-smithii
4a. Leaves subsessile, domatia of tufted hairs or pits often present along the midvein 5
4b. Leaves with petioles usually more than 4 mm long, domatia absent along veins beneath (rarely
present in C. chiriquiensis 7
5a. Corolla usually minutely sericeous, calyx tube 2-3 mm long; domatia usually narrow de- pressions along the midvein above the vein axils [leaf blades 7-18 x 3-9 Cm; Caribbean
lowlands of northern Costa Rica] C impetiolaris
5b. Corolla glabrous or minutely papillate puberulent; calyx tube 1-2 mm long; domatia of shallow
puberulent depressions in the vein axils or absent 6
6a. Inflorescences 3-10 cm long, paniculate with distinct lateral branches; leaf blades 7-16 cm long, domatia of puberulent pits in the vein axils; Caribbean lowlands of southernmost Costa
Rica C. sp. A aff. curvigemmia
6b. Inflorescences ca. 3 cm long, subcapitate (paniculate but with short, closely spaced, lateral branches); leaf blades more than 1 5 cm long, domatia present or absent; lowland rain forest
C. sp. B aff. curvigemmia
7a. Flowering portion of the inflorescences elongate-racemiform, distinctly longer than broad ..... 8 7b. Flowering portion of the inflorescences open paniculate, corymbiform to umbelliform or pyramidal,
usually with length equaling breadth 10
8a. Stipules narrowly oblong to linear, to 2 cm long and 4 mm thick, acute at the apex and Ficus-